CHANGE IN NUCLEOPROTEIN CONTENT IN PLANTS UPON VEGETATIVE HYBRIDIZATION

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• Release Decision: RIFPUB
• Original Classification: K
• Document Page Count: 269
• Document Creation Date: December 27, 2016
• Document Release Date: September 17, 2013
• Sequence Number: 1
• Publication Date: December 1, 1952
• Content Type: REPORT

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Semenenko, G. I.
(i)
Izmenenie Soderzhaniia Nukleoproteidov
v Rasteniiakh pri vegetativnoi Gibridizatsii
[Change in nucleoprotein content in plants
upon vegetative hybridization].
Biokhimiia, 17(6):655-659. November-December,
1952. 385 B523
(In Russian)
CHANGE IN NUCLEOPROTEIN CONTENT IN PLA-NTS
UPON VEGETATIVE HYBRIDIZATION
Trans. 511
(Conclusions
only)
By:
R. Adelman
[Experiments were conducted with tomatoes of the "Marglob"
variety; eggplant of the "Delikates" variety; and the seed
offspring obtained from tomato/eggplant grafts].
CONCLUSION
In vegetative hybridization, substantial quantitative changes occur
in tho phosphorus contont of nucleoproteins in the young growing organs
and tissues of grafts and hybrid progeny.
? In grafting tomatoes onto eggplants and vice versa, tho chango in
the phosphorus content in nucleoproteins in tho leaves of tho scion is
inclined to favor tho stock; and in tho case of ropeated grafting it
incroases.
In tho seed progony of vegetative hybrids, concurrently with a
vigorous manifestation of hoterosis, an increased nuclooprotoin content
is observed in the loaves, blossoms, tips of shoots and in tho young -
seedlings of hybrid plants as compared with tho original [plant] forms.
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?
7
Glushchenko, I. E.
Tran. 512
(Conclusion)
By:
R. Adelman
Gibridizatsiia Rastenii putem Privivki
[Hybridization of Plants by Grafting]
Uspekhi Sovremennoi Biologii 30(1)115-48
July/August 1950. 442.8 Er3
(In Russian)
HYBRIDIZATION OF PLANTS BY GRAFTING
General Conclusions
1. Vegetative hybrids show that simple heredity characteristic in
vegetative reproduction can, as Timiriazev asserted, turn into complex
heredity characteristic in sexual reproduction.
2. Experiments in vegetative hybridization show an inconsistency
in the chromosome theory of heredity the principle of which is cell from
cell, nucleus from nucleus, chromosome from chromosome, gene from gone.
Vegetative hybrids convince one that somatic cells undergoing
physiological changes will, in the end, produce altered sexual cells.
There is no immortal embryonic course. There is a course of qualitative -
transformation of soma capable of forming sexual cells at a certain stage.
3. There is-a similarity, a parallelism, between sexual and vegeta-
tive hybridization. It is a fact that the second method, as well as the
first, will transmit any characteristic, any property from one component
to the other. Those properties become fixed in tho seed generations.
4. Side by side with parallelism there exist differences. A typical
feature of vegetative hybrids is the different type of fission of
characteristics. Not only tho plants vary in thoir basic characteristics,
but a shdrply pronounced differentiation occurs [proxodit] within the
organism.- In vegetative hybrids tho mixed typo of heredity is basically
intrinsic.
5. A characteristic of graft-hybrids is the different form of
manifestation of property dominance. Plants with rocossive charactorics
often produce offspring with dominant characteristics.
6. Crossbreeding carried out between vegetative hybrids and parcnts
with graft components and recessive characteristics domonStratc, graphically
that dominant characteristics may appear in the offspring. This indicates
that separate characteristics in hybrids arc in a latent state and, under
appropriate conditions, develop subsequently in the offspring.
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?(2) Trans. 512
7. On the change in oxtornal morphological characteristics, a
profound roorganization of order takes place in hybrids, the character of
coil structure changes in particular, plastids of various typos appear in
the cells.
8. Biochemical investigations show that the majority of quantitativo
and qualitative indicators ofthe second graft component ? the mentor --
appear in the seed progony.
9. Frequently tho absence of visible changes observed in the year
of grafting does not donote an ab6enco of specific qualitative changes in
the generative cells of the plant. Honco it follows that the progeny of
grafted plants must always bo investigated, even though no phenomenon of
variation in characteristics had been observod in the year of grafting.
10. It mould be most priLitive to think that as a rosult of grafting
one always has to look for grafts of the stock in the offspring of the
scion, or vica versa, to discover prop6rties, characteristics of the scion
in the offspring of fruit of the stock. 'Living [substance] constitutes a
process of .development, and every biological process does not know of
straight, diroct changes. They are realized only in a long chain of.
transformations.
Besides the phenomena of straight hybridism (presence of the
characteristics of both parent components in the offsprint), vegetative
hybrids include also the phenomena of now formations, i.e. the creation
Of now chaacteristics frequently not inherent in either one of the graft
components. A pro-requisite for this is extromo instability, inconstancy
of form, frequently observed in ifitraspecific and, most often, in
distant, [intergenoric] graftings.
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(1)
Trans. 513
(In full)
By;
R. Adelman
Nikitenko, G. F.
0 nekotorykh osobennostiakh vegetativnoi
gibridizatsii zlakov metodom transplan-
tatsii zarodysha
[Certain characteristics .of vegetative hybridization
of cereals by the embryo transplantation method].
Doklady Akademii Nauk SSSR, 76(2)273-276
January 119 1951 511 P444A
? CERTAIN CHARACTERISTICS OF VEGETATIVE HYBRIDIZATION
OF CEREALS BY THE EMBRYO TRANSPLANTATION METHOD
Submitted by Academician
N. A. Maksimov, November 11,
1950
Research of recent years has developed technical methods making it
practicable for a large group of herbaceous plants (principally the
Cucurbitaceae and Solanaceae) to bring about exertion of a larger measure of
influence by the stock upon the scion, which, as is knolirn,.is one of the
decisive conditions in obtaining vegetative hybrids (8). The essence of
these methods is the influence brought to bear on the scion at various
stages of its development, and the regulation of its assimilating (foliar)
surface (1,3,8,14).
However, the method for obtaining vegetative hybrids among cereals
cannot yet be considered as sufficiently developed. With regard to Gramineae,
the more widespread and generally accepted method at present is vegetative
hybridization by means of embryo transplantation, although other processes are
also known; marching [ablaktirovka] (3,9), grafting onto the node above
ground (6,12). [p. 273, end para 2]
Apart from the greater technical accessibility, the embryo transplanta-
tion method perthits influencing the scion (embryo) at the very beginning of
its development. There are several known alternatives of this method
(4,5,7,12,15,16), but all are characterized by a common shortcoming,-namely?
the altering action of the stock (endosperm) upon the scion (embryo). The
latter, being selective as to the conditions of its life, will "reluctantly"
make use of the alien plastic substances of the stock -- endosperm -- and
will assimilate them in-building up its body only until it forms rootlets
and a regular leaf, i.e. until it becomes self-sustaining. With the formation
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("4) 'runs. 010
of embryonic rootlets and the bringing forth a leaf above ground, the influence
exerted by the endosperm on the embryo, via plastic substances, stops. Fre-
quently this occurs before the supply of the endosperm is all used up, and
thus, the influence of the stock (endosperm) upon the scion (embryo) appears
to be limited not only with respect to duration of the action, but also as
to the amount of the assimilated plastic substances.
Attempts to overcome the indicated shortcomings by increasing the amount
of the altering plastic substances in-grafting -- by doubling the endosperm
stock) (15,16) -- were not successful. In our opinion, the reason for this
lies in the fact that such a treatment of the question solves only the problem
of increasing the amount of plastic substances of the stock, but does not
touch upon the other, the more important problem.? forcing the scion-to
assimilate food inappropriate for it, and in large quantities at that. On
grafting, the plants show essential individual distinctions, and their re-
action to the influence of alien food substances is very strong. Consequently,
a simple increase in the amount of plastic substances of the stock is not
sufficient to obtain the needed changes in grafting by trans- [begin p. 27411
planting the embryo -- it is important that the latter be compelled to
assimilate them. In underestimating this requirement, we are inclined to
explain the failures which have occurred in the work of many investigators
who are engaged in iegetative hybridization of Graminea and use the method
under consideration. (Let us point out, particularly, the experiments
conducted by Shtingl [or Stingl] with Triticum, Hordeum, Secale, Avena,
cited by Winkler (2), in an attempt to prove that the influence of varyifig
plastic substances is incapable of causing specific changes in grafting).
In the course of experimental work of vegetative hybridization of
barley by the embryo transplantation method, we succeeded in detecting
conditions allowing, to a considerable degree, to overcome the fundamental
fault of this method. The method of vegetative hybridization of cereals
which we suggested, frequently led to a very profound influence exerted by
the stock upon the scion: we refer to a case of complete suppression of the
scion in grafting and the formation of a plant with the characteristics of
only the stock (10).
The further development of the problem under consideration was continued
under conditions lof a specially conducted experiment with biologically
contrasting graft component. The varieties of winter barley selected as such
components were -- [p. 274, para 2] Kruglik 21 and KrawiDar 2494 (Hordeum
vulgare L var.pallidum) and two varieties or winter re Kazanskaia 5 4. 6
and Dotnuvo Aukshtein Secale cereale L. var. vulgare). The graft was grafts
accomplished by use of the method described earlier (10,11). A total of
were carried out Kruglik 21 and Krasnyi Dar 2494 which, on
2 Dotnuvo Aukshtein 2 Kazanskaia 5- 4. 6
September 1, 1949, were sown in hothouse pots [vazony] containing seed bed
earth (50% soil and 50% hubilus) at a dept4of 8-9 cm. Shoots appeared on
September 12-14, 1949, i,e. on the 12-14 tay following the sowing. Of the -
250 grafted grains only 86 germinated; the rest of the 164 produced no shoots.
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(3) Trans. 513
?
During the phase of 2-3 regular leaflets, the experimental plants were trans-
planted to open ground where they were well able to develop runners
iraskustitisia] by the time autumn vegetation ceased.
Even the first observations of the post-grafting development of the
expetimental plants revealed a profound influence of the stock in the graft-
ings. In 26 out of 86 cases, the shoots which emerged proved to Contain
anthocyanin, which is characteristic of the stock (of winter rye). As the
grafts were developing further, a progressive increase was observed in
stock characteristics % conservation of anthocyan at the base of the [p. 274,
end para 3] plants, a slightly raised and expanded form of the branch [kust]
(the scion of winter barley has a recumbent branch), noticeably less crescent
form to the ears of the leaf sheath.
In the spring of 1950, following the overwintering, there were only 12
plants left in the nursery, and these were of the 26 ithich, in'the fall of
1949 had revealed a strong trend in the direction of the stock. The rest of
the grafts, exactly as the controls (Intravarietal grafts and ungrafted plants
of winter barley), perished completely as a result of the very severe over-
wintering conditions of 1949-1950. The following spring, the development of
the surviving plants was distinguished by considerable originality.. In these
plants the formation of new leaves and runners were accompanied by the dying
off of the old ones, while the newly emerging tunners tended ever more toward
the stock in an externally morphological sense. It created the itpression
that the plants, so to say, were discarding their hybrid "jacket". In
separate plants the dying off of the vegetative mass proceeded more intensively
than the process of forming new ones, so that of the 12 overwintered plants
only 3 survived until the time of fruitage. The other 9 died off at different
stages % 5 soon after spring vegetation had begun (May 3-8)0 3 -- in the phase
of stem formation [vykhod v trubku] (May 25-29) and one plant at the peak of
spike [end p. 274] formation (June 24). At this time it is important to note
that, as per our observations, the dying off of 4uafts was not due to a
deficiency of food elements, of light, moisture etc., nor to injuries caused
by insect pests, and much less to mechanical injuries, but apparently, to
a lack of balance of strain requirements of the graft components.
After spike formation, the development of the plants which survived
proceeded normally. At this moment they resembled completely the plants
of winter rye; their flowering was rather'protracted (June 28 - July 13)
and it proceeded in the "rye" manner, i.e. it was open. For the purpose
of increasing the percentage of the, binding capacity of grain., these plants.
were artificially backcrossed [poreopyleny] among themselves. In addition,
one spike on each plant was isolated (with the aid of parchment isolators)
under which no grain whatever formed.
Three plants produced a total of 29'spikes yielding a harvest of 867
grains with an absolute weight of 30.8 g. Viewed from the external appear-
ance; the spikes obtained and the grains in them were clearly of the rye
type.
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(4) Trans. olo
Thus, in our experiment, [p. 275, begin para 3] plants with externally
morphological and biological characteristics typical of the endosperm-stock,
-- winter rye -- developed from embryos of winter barley transplanted onto
double endosperms of winter rye, if the grafted grains were-planted extra
deep (8-9cm) and grown under field overwintering conditions. We explain
this, undoubtedly, most interesting case of strong influence exerted by the
stock upon the scion as follows. Thanks to considerable (with respect to
duration and amount of plastic substances assimilated) influence of the
endosperm-stock, there occurred in the early stages of graft development a
sharp shift in the direction of winter rye. However, the degree to which
this shift was expressed in the grafts varied by virtue of the individual
differences in the components. The subsequent development of the grafts
took place under severe overwintering conditions and caused in some of them
a further increase in the characteristics of winter rye, in as much as the
prevalence of such conditions evidently contributed to the development, in
the grafts, of properties and characteristics of the more hardy component,
in the given case winter ryo. The other grafts which leaned in the direc-
tion of the scion (of winter barley) and, therefore, were less hardy, perished
the same as the controlS" (minter barley) because they could not survive the
low winter temperatufes. As a result of this, a complete predominance of
stock properties [p. 275, end para 3] set in in the grafts which had sur-
vived the winter at the time of full fruitage there were left only the
grafts of the extreme variant -- "toward tho-stock"; grafts "toward the
scion" and intermediate ones were eliminated.
The data obtained may serve as strong eviderre of the feasibility of
transmitting the properties of one strain to another without the partidipa-
tion of sex elements, solely by transmission of plastic substances (8). It
seems to as, however, that this does not exhaust the importance of the facts
disclosed. On the basis of these facts ono can assert that to ensure the
success of vegetative hybridization of cereals by the process of embryo
transplantation, it is not necessary to create conditions that "will maximally
accelerate the germination of the scion embryo" (15) by covering the grafts
with a thin layer (1 cm) of friable soil, but on the contrary, to pursue the ,
course of prolongirig the period in which the changing action of the "strange"
food of the endosperm-stock can be exerted. Only in this case one can expect
to obtain a high percentage of success in vegetative hybridization of cereals
by the method indicated -- not in the sense of generally obtaining a large
number of plants from-gfafts, but of obtaining a high percentage of real
vegetative hybrids, i.e. plants with the properties of the scion and the stock.
[p. 275, end para 4].
It may be assumed that by regulating the depth at which grafted grains
are planted [zadelka] (taking into account the size of the seed grafted) and,
by the same virtues, regulating the size and duration of the stock influence,
combined with expedient cultural practices fof grafts, it is possible to
obtain stock influence of the needed strength.
Mordovskaia Gosudarstvennaia
Selektsionnaia Stantsiia
Received
9/9/1950
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(5) Trans. 513
REFERENCES CITED
(1) ? A. A. Avakian, Iarovizatsiia, no, 1, (1941)
(2) H. Ulnkler, Untersuchungen ueber Propfbastarde, Jena, 1912
_
(5) I. E. Glushchenko, Vegetativanaia gibridizatsiia rastenii..
M., 1948
(4) L. A. Golovtsev, Agrobiologiia, no. 1, (1948).
(5) V. I.. Didus', Ze1ektsiia i Semenovodstvo, no 4 (1950)
(6) V. F. Illarionov, ibid, No. 11 (1948)
(7) F. M. Kuperman, Iarovizatsiia, nos. 5-6 (1939)
(8) T. D. Lysenko, Arobi_s__22_.ogii21_.94r2.
(9) G. Kh. Molotkovskii, DAN, 24, no, 4 (1950)
(10) G. F. Nikitenko, Selektsiia i Semenovodstvo, no. 1, (1949)
(11) G. F. Nikitenlo, ibid, no. 5, (1950)
(12) I. G. Plotnikov, Iarovizatsiia, no. 3 (1939)
(13) P. F. Sekum, Selektsiia I Semenovodstvo, no. 2 (1949)
(14) N. V. Turbin, Bot. Zhurn., 36, no. 6, (1949)
(15) V. I. Khmelev, DAN, 70, no, 5, (1950)
(16) V. I. Khmelev, Selektsiia i Semenovodstvo, no. 3, (1950)
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(1) Rererence
30919
Trans. 514
(Not made in
USDA)
Rautenshtein, I. I.
Cand. Biol. Sci.
[On Utilizing Antibiotics in the
Food Industry]
Vestnik Akademii Nauk SSSR,
? vol. 24, no. 5, May 1954, pp s 44-48.
511 Akl4V
(In Russian)
USSR WORK OF TiE APPLICATION OF ANTIBIOTICS
IN THE FOOD INDUSTRY
[Numbers in parentheses refer to
appended bibliography]
The idea of using the antagonistic effects between microbes for combating
pathogenic microorganisms had been originally suggested by I. I. Mechnikov.
However, practical application of the phenomenon of antagonism between mi-
crobes was first attempted in the 1940's, when antibiotics (tyrothricin, peni-
cillin, and gramicidin) were isolated in a pure state.
Modern medicine and veterinary medicine have found in antibiotics a
powerful means of combating infectious diseases. It is understandable that
the question was raised in regard to the use of antibiotics for the control
of harmful microorganisms in some industrial fields as well, particularly in
the food industry for the preservation of food products from spoilage (1).
Many invest*g tors, particularly abroad, attempted to apply for this
purpose antibioti s which are known and have received extensive application
in medicine, namely penicillin, streptomycin, chloromycetin, aureomycin,
terramycin, subtilin, gramicidin, etc. The investigations that have been
carried out dealt mainly with the problem as to whether the antibiotics
mentioned are suitable for the preservation of meat, fish, and milk from
spoilage. Notwithstanding certain contradictions which are encountered in
published articles, it may be considered ,sla an established fact that small
quantities of antibiotics, after being added to the food product or to the"
ice in which the food product is kept, prevent spoilage for a certain time.
However, none of the antibiotics used in medicine have found practical
application in the food industry., The principal reason for this is the
natural apprehension that as a result of prolonged consumption of antibiotics
microbial forms which are resistant to antibiotics willarise. Experience
shows that persons who had been treated with antibiotics develop resistant
forms of bacteria in their bodies with relative ease. It is obvious that
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(2) Trans. 514
after the development in the organism of microbes which are resistant to
a definite antibiotic, treatment with this antibiotic of the disease
produced by the microorganism in question will not succeed. In other
words, the application of an antibiotic in the food industry may lead to
the result that an effective therapeutic agent will be lost.
On the other hand, there still is no information worth mentioning on
the nature of the effect which small doses of antibiotics administered for
a long time exert on the human organism.
The rather extensive data pertaining to the action of antibiotics on
animals are very contradictory. It was established that addition to the
feed of minute quantities of penicillin, streptomycine, chloromycetin,
aureomycin, or other antibiotics in some cases, but not Always, has a
beneficial effect on animals, particularly young animals. It was establish-
ed that-the identical anitbioticb exert a different reaction on different
animals. The best results were achieved with auroomycini_terramycinc, and
bacitracin. The mechanism of this prolonged action of antibiotics has
not been investigated adequately as yet. But a number of authors assume,.
not without reason, that. improvement in the growth of young anithals, which
has been noticed after the administration of an antibiotic in the feed, is
produced principally by profound changes in the ccmposition of the
intestinal microflora and that these changes are brought about by the action
of the antibiotic. It has been established that some antibiotics suppress
the development of the microflora, which is harmful to the organism, for
instance the putrefactive and toxicogenic microflora; while the useful
microflora (lactic acid bacteria and some others), among which theio are.
species which synthesize vitamins needed by animals, is stimulated,
However, there are indications that the prolonged administration of
antibiotics for therapeutic purposes may lead to a vitamin insufficiency as
a result of the modification of the intestinal microflora. Other undesir-
able results may also arise. It has also been established that the
systematic administration of antibiotics to ruminants, in the nutrition of
which microbiological processes are of great importance, processes taking
place in the stomach (or rather the compartment of the stomach which is
called the rumen) eliminate the useful microflora and bring about severe
disturbances of the digestion. In young ruminants, particularly calves
during the earlier, period of their life, when the microflora of the rumen
does not yet play a very active role in their digestion, those disturbances
are not observed.
Thus, the data of some authors, who point out the useful effect of
the addition of antibiotics to the feed of. animals, differ from the date
published by other authors, Who have not observed any such effect or who
indicate that there is a harmful effect on the organism. It is not
surprising that in a number of countries the sale of antibiotics to be
used as an ingredient of the feed of animals is prohibited.
What has been said leads to the conclusion that our knowledge of tho
action of antibiotics when administered together with food is still
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inadequate. For that reason one must be very careful in approaching the
problem on the utilization of antibiotics in the food industry. The Most
thorough and many-sided investigations on this subject are essential.
Very original and, from the standpoint of this discussion, very promis-
ing investigations are under way on the application in the food industry of
the antibiotic propertied of higher plants. Work on this subject has been
done in the Soviet Union.
The juice"of"many higher plants, as has been shown by Soviet scientists,
particularly B. P. Tokin, contains the so-called phytoncides? which have
antibacterial properties (2). The phytoncides, just like antibiotics,
comprise substances of the most diverse chemical composition. At present
the chemical composition of many phytoncides has been investigated and
these phytoncides have been isolated in the pure state.
It is obvious that in order to find out whether phytoncide preparations
can be used as preservatives for food products one must investigate the effect
of prolonged administration of these substances together with food. However,
with reference to some food products, this problem is not of particular
importance. For instance, it has been established that in the production
of some canned preserves, one may effectively use the.phytoncidic properties
of the raw material from which these preserves are made.
Of particular interest in this respect are the investigations which
have boon conducted at tho All-Union Scientific Research Institute of tho
Food Proscrvo Industry. Thoso investigations established that phytoncidos
aro contained in many vogetablos and fruit as well as in some plants which
are used as species (3). Thus, phytonclics are contained in tomatoes,
carrots, boots, horseradish, parsley, onions, popper, mustard, and coriander.
Tho phytoncidic properties of plants depend on their spocios,and ago. Tho
antibacterial activity of some phytoncides is more pronounced after heating.
? Under tho effoct of tho phytoncides of vegetables, the quantity 6f
micro-organisms in canned prosorvos is reduced prior to sterilization.
This makos it possible to reduce the tomporaturo and extent of tho
sterilization of vogotable preserves by heating. The preserves obtained
by the now method, which involves loss rigorous conditions of steriliza-
tion, wore found to have high indexes of bacterial sterility.
In many plants, for instance tho onion, tho volatile fraction of the
phytoncidcs is of particular valuo. However, this fraction may be used
up in a very short poriod of time. This circumstance must be taken into
consideration in establishing the toohnological conditions for tho treat-
ment of tho raw material in quostion.
Tho investigations described are of grodt significancofram-tho,stand-
point of the fundamental principlos involved. They made postiblcan entirely
now approach , namoly utilization of the antibacterial properties of tho
raw material being preserved, and at tho same time domonstratod that on the
basis of this principle ono may achieve an improvomont in tho quality of
production, increases in the efficiency of technological processes, and a
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reduction of the prime cost.
It is obvious that further work along this line is of importance. The
attention of investigators must be concentrated on the selection and
introduction into the food preserving industry of those species and varieties
of vegetables, fruits, and other plants which are rich in phytoncides.
Furthermore, the technological processes for the treatment of such raw
material must be improved with the-view to achieving maximum utilization
of the phytoncides contained in it. It is also necessary to conduct work on -
the possibilities of using phytoncides in other branches of the food industry.
From this standpoint, the capacity of some phytoncides to suppress the
development of the putrefaction micfoflora which brings about spoilage of
fresh fish is of great interest (4).
Soviet scientists are developing a new and original line of research
dealing with the utilization of the phenomenon of interbacterial antagonism
for the improvement of the quality of food products and of their stability
during storage.
Organisms which exert an antibiotic effect are found within all groups
of microbes, including yeast, lactic acid bacteria, acetic acid bacteria,
and other species of microorganisms on the life prr;cesses of which many
branches of the food industry depend. However, the products obtained with
the aid of such microorganisms are often subjected to spoilage produced by
extraneous microflora. Thus, sweet butter in storage loses many of its
valuable properties as a result of the activity of putrefactive micfo-
organisms, lipolytic bacteria, mold fungi, and other. microorganisms.
Soviet scientists have established that some strains of yeast have the
capacity of suppressing microorganisms which damage butter (5). These
strains of yeast may be used for the purpose of increasing the stability of
butter to spoilage. A number of strains of lactic acid streptococci,
which exert an antibiotic effect on the causative factors of the spoilage
of butter, have been isolated from natural sources (6). If these strains are
included in the production stock culture, which is used in the manufacture
of sweet butter and which consists bf a mixture of strains of lactic acid
streptococci, one may suppress the development of putrefactive organisms and
other harmful micro-organisms in the butter.
These investigations deserve serious attention. It is not impossible
that one may discover among lactic acid bacteria culture which have not
only antibacterial activity but-also antifungus activity, and the property
of suppressing bacterial phages. The discovery of lactic acid bacteria
that have the last-mentioned property would be of great significance for
the cheese industry; where the problem of the control of bacteriophage is
of great importance. The phenomenon of antagonism between microorganisms
may also be successfully used in other branches of the food industry.
In January 1954 the Institute of Microbiology,-Academy of Sciences
USSR, conducted a conference at which a report by N. A. Krasiltnikov was
presented. This report dealt with the scientific basis of the application
of antibiotics in the food industry and the prospects of such application.
A number of other reports on research in this field was also presented.
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(5) Trans. 514
The conference noted the importance of continuing work on the utiliza-
tion of antibiotics and phytoncides, and the application of the phenemona
of antagonism between microbes in various branches of the food industry.
In view of the great significance which must be ascribed to the elucidation
of the action of antibiotics on the human organisms, when these antibiotics
are used as preservatives for food products, the conference recommended
that the appropriate institutes of the Ministry of Pulgic Health USSR,
particularly the Institute of Nutrition of the Academy of Medical Sciences
USSR, take steps to investigate this problem.
It was especially indicated that there is a necessity for expanding
work aimed at the discovery of new antibiotics derived from actinomycetes,
mold fungi, and bacteria, as well as of phytoncides and antibiotics of
animal origin, suitable for use in the food industry.
Introduction into the food industry of new antibiotics which are not
used in medicine presents, as has been stated, definite advantages. It is
obvious that the selection of a new antibiotic preparation for use in-the
food industry must be based on all the properties of this preparation. In
medicine and veterinary medicine, an antibiotic usually serves the purpose
of controlling a definite causative factor of some disease. In the food
industry, on the other hand, one must as a rule preserve the product from
the harmful action of a number of microorganisms which belong to the most
diverse groups. It is therefore necessary that the antibiotic have a
broad range of antimicrobial activity. Furthermore, an antibiotic Which
is suitable for use in the food industry must have an active effect on
definite) species of microorganisms not only in vitro, but also in the food
product. It must be devoid of toxic properties,-and harmless to the
organism on prolonged consumption in small doses. It must not confer an
unpleasant taste or odor, and must not produce' any other undesirable
changes in the properties of the food product.
Antibiotics with properties which are useful for the food industry
can be found among the most diverse microorganisms. Bacteria, fungi, and
actinomycetes are among them. Actinomycetes must-be regarded as the most
promising group of microorganisms in that respect.
As a confirmation of this may serve the results obtained in a search
for antagonists effective against a number of speciOs of microbes ftich
produce spoilage of meat and meat products (7) Bact. Coli and Bact, proteus'
wore used as test cultures; also Penicillium, Cladoe'porium, and AG.Ucor fungi.
A total of 806 strains of actinomycetes were tested. Of this total 244
straina, or 30%, exhibited the capacity for suppressing the development of
either one or several of the harmful cultures used in the tests, while
several had an effect on all the harmful cultures used. Antibacterial
properties were exhibited by only 27 cultures, and antifungus proprtios
by only 134 cultures, while 83 cultures suppressed-the development of both
bacteria and fungi present in various combinations.
One must assume that cultures which have antifungus prop)rties will
prove to boa promising source of antibiotics for use in those branches of
the food industry where mold fungi have to be combated.
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ko)
?
Antibiotics which have a law thermal stability and a relatively high
degree of inactivation during storage with a food product May prove to be
very useful. Such antibiotics would be very suitable for application in
the canning industry and other branches of the food industry. Antibiotics
of this type would lower the bacterial count of the crude material or inter-
mediate products, and at the sametime make possible the application of
milder sterilization process. The subsequent inactivation of the antibiotics
as a result of thermal treatment or storage would remove all possibilities
of an undesirable effect of the antibiotics on the human organism.
It is understandable that introduction of antibiotics for use as food
preservatives will nec6ssitate precise methods for their quantitative
determination on foods,
The problem of the application of antibiotics in the food industry is
a new one which has not been investigated to any great extent as yet, and
requires extensive study. For the successful solution to this problem the
joint efforts of microbiologists, biochemists, technologists, hygienists,
and representatives of other fields of knowledge are essential. In
conclusion, one must point out that the expansion of work on new antibiotics
will not only be of use to the food industry, but may result in discoveries
which will be very valuable to medicine and 'agriculture.
BIBLIOGRAPHY
(1) N. A. Krasillnikov, Application on Antibiotics in the Growing of Plants,
Vestnik Akademii Nauk SSSR, vol. 24, no. 10 1954;
pp: 50-57,
(2) B. P, Tokin, Bakteritsidy Rastitel.'nogo Proiskhozhdeniia-
Fitontsidy [Bactericides of Plant Origin -
Phytoncides], Medgiz, Moscow, 1942
(3) A. I. Rogacheva, Mikroviologicheskii Kontrolf Konservnogo Proizvodstva
[Microbiological Control in the Canning IndustrY],
Pishchepromizdat, Moscow, 1949.
(4) Iu. A. Ravich-Shcherbo,
The Use of Phytoncides for the Preservation of
Fresh Fish in Storage,
Rybnoye Khoziaistvo, no. 10, 1949.
(5) V. PL Bogdanov, Application of Yeast Cultures in Butter Making;
Mikrobiologiia, vol. 21, no. 3, 1952.
(6)
(7)
Report presented by T. -G. Romanovich at the Confer-
ence on the Use.of.Antibiotics in the Food Industry
held at the Institute of Microbiology, Academy of
Sciences USSR, on 19 January 1954
Report presented by V. R. HirzoyeVa at the above-
mentioned conference.
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R. Adelson
[Contents on41
4111LInko, G. I., Ed.
Zertavye 1. sernobobovye kulstur
na boor*.
[Cereals and legumes in dry farming
in regions of insufficient rainfall.]
loomderstvennoe Igdat,1' stvo Uzbek,.
skoi 422, Tashkent, 1952* 169 p.
69 242
(In Ruitaiiial
DITROVICTICK
TABLE OF CONTF31/5
p*S
Min= I. Agroteohnies or ,oersal
craps in dry farming in regions
of insuifteient
Natural +renditions for dry
farming in Uzbekistan p. 9
Condition. essential to the
of plants 11
The grasomfield System or --
agriculture 14
? IS
16
Crop rotation
Tillage
System of late tall tillage--
Ito, sewser crops) 21
System of seedbed prepays.. -
tics
Treatment or fallow 9f. 24
Fertilisers for plants 26
Local fertilisers "I...,
lineral fertilisers
3asteria1 fertilisers
Supplementary fertiliser
for crops ?
Fall crops of biologies)"
ly summer speeies of
cereals ?
Swum ems
Seeding methods
26
26
21
Depth of seeding ?
New agricultural mastic* 040
flurrew seeding or winter
crops
54
$S
Review questinns 55
CMPTERIi. Charaoteristies of the
toe eireereais on 'tend.
ere precipitation it seasonal
Charactoristiss of land* under
eortingent ixrigitio*
&Hiding on lands under contingent
irrigation
46
48
Charaeteristies of the arrangement
-
of crop rotations ? 49
Review questions $O
CHAPTER III. Care of 'rope.
Baste principles or wiring tor
wheat and barley crops ?? ? ?? 61
narrowing or roll crops 55
Additional seeding 66
Italchiag 66
Characteristics in oaring for
ether 'weal crops ??? ST
Review questions 69
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CHAPTER /V. Snow retention and re-
retention et' they water under
conditions of dry farmin
iftialsonorotininwrearion
? 80
Arrangement of ponds and
reservoirs 65
Moisturemoharged irrigation es
Review questions 84
CHAPTER Y. Seed lets on kolkhoses.
Designating the seed lot on a
kolkhos 85
Agroteobnioal tasks on seed
lots .... es
Chcreaterieties of harvesting 69
? Storage of grain 71
Review questions 72
Cann VI. Diseases and Pests of
cereals antf legman and their
control.
?
Diseases 75
Stinking smut Nattier herdeil ..78
Loose smut [Ustilago nude]
Yellow rust
Brown rust
Linear or stem rust
Lac oshyta,
Pests
Locusts [Aoridiidae]
Earnests/. integricepe Put
?Firestorms
Tenebrionid beetle*
Lethrus apterus [beetles]
Cobweb mites
Review questions
81
82
CHAPTER YIL" Wheat the most important
food ems
Importance of shoat to the
national eeonomy
86
Botanleal description of
wheat 88
Description of the speoies
of wheat 91
Characteristics of wheat
species regionalised and
prospeetive for Uzbekistan 92
Review questions OS
CHAPTER VIII. Barley and Oats
industrial forage ereps
Importanee of barley toll's
national economy eeo
99
74
77
77
vs
TS
79
79
81
Botanioal description of barley... 99
Description of the 'posies of
100
Importance of oats to the
national aSoU4Ny 105
Botanical deseriptian of oats 106
Description of the species
of oats 108
Agri/to/ohm's/a characteristics 106
Review Questions 107
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tTable of Con.
41PTER EL. Spesies oroye and the
regionalisation of wheat and
bailey spates
Importance of specie crops
Regionalised species ? ?
Zones for dry farming and region.
a/istng of species
For the Kashka.partya
district [oblast') ???? ao
For the Samarkand Region
toblastgi
For the Surkban.Daera
district [oblastel
For the Tashkent district
toblawbq
? Review questions
CHAPTER X. Grit; crops
Millet [Purlieu* milisesumi ....
?
Corn
tents)
CRAFTER II. /Aguas Crops
1211
Chica.peue ?
Importance of ebiskopeas to
108
the national amour ?????
128
109
Biological oharseteristies ? ? ? ?
130
Description of the species
109
of ohisk.peas
131
A7oteoh--leal sharaeteristiss ,??
131
110
Lentils
133
112
Botanical description ?????
13$
Biological characteristics
134
113
Agreteohnioal eharacteristios
136
114
Beans [Phasoilue mun4e1 ? *
1.35
115
Botanical description and
species
136
Agroteehnioal characteristics ? ?
137
116
Revive Questions
138
Botanical description and the
species 116
Agroteehnioal
ties
eharecteric.
Botanical description and
species
Agroteshnical character
tattoo
Sorghum tBsbugaral
Botanical description and
species
Agrotoehnioal eharacteristierr
118
120
121
122
1215
124
? 3.25
Review questions 126
CHAPTER XI/. _erLtiB1rekeoping accounts,
of yield an#-iiiii?Kiiage
Accomplishing harvest in record
time and without losses 139
Organising the work of
harvesting
Determining when harvesting
is to begin
140
142
Organising *cabin. harvesting 143
Harvesting with ordinary
sachines
Fulfilling grain deliveries
to the Government
Harvesting charseteristics or .
different crops
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146
147
Trans. 51 5
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*Met 14?
Corn and sorghum tdahngaral th, 147
Beans
4. 148
Lentils ....? 148
Chisk?peas 149
Controlirg *rep lessee 149
Laportaras of the *cabins
in the eontrol of lasses .4, 149
Accounting and guarding
toprikhodavanie) of the
crop
Accounting of grain when
harvesting is done with
combines
151
151
A40ounting of grain *Ion
threshing is done with
threshing machines 152
Review questions lig
Addenda 159
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Chirvinskii, I. A.
Novoe v Sellskokhoziaistvennoe Nauke
[News in Agricultural Science]
Dostizheniia Nauki i Peredovogo Opyta
v Seltskom Khoziaistve, 1954(1)72-75,
January 1954
20 D742
[In Russian]
NEWS IN AGRIBULTURAL SCIENCE
USE OF ANTIBIOTICS IN THE CONTROL OF
INFECTIOUS DISEASES
OF PLANTS
Trans 516
[In part]
pp 72-73
By:
R. Adelman
An important role in the solution of problems, set forth by the
September Plenum of the TSK KPSS [Central Committee of Communist Party
of the Soviet Union] before -the workers of agriculture, is assigned to
leading agrobiological science.
Workers of scientific institutions use every effort to exhaust all
available and heretofore unutilized internal resources and to discover
new possibilities that will rapidly increase the productivity of agricultural
crops.
It is known that plants frequently become infected by various diseases
which redude their yielding capacity and cause enormous losses to kolkhoz
production. Particularly great damages are inflicted by plant diseases
caused by bacteria, pathogenic fungi and viruses.
Existing methods for the control of plant diseases are not always
effective and at times produce no results whatever. Hence, an important
medium in increasing the productivity of agridultural crops is the timely
protection of plants against various diseases.
The scientists of the Institute of NicrobiOlogy, Academy of Sciences
of the Union of the SSR have, produced substances called antibiotics. These
substances are used as medical preparations not only in medicine, but also
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(2)' Trans. 516
in agriculture -- in veterinary medicine; and in plant culture for the
control of infectious diseases of plants.
Such a group of antibiotics includes penicillin, streptomycin,
sintomycin and others. Investigations conducted by Soviet scientists have
revealed that these substances can be produced by various little fungi and
bacteria inhabiting the soil.
The works of the famous Soviet scientists, N. A. Krasil'nikov, and
his staff reveal that antibiotics can be used for the prevention and treat-
ment of infectious diseases such as wilt of citrus plants, gummosis of
cotton, etc.
R. Mirzabekian, a Fellow of the Institute of Genetics, Academy of
Sciences USSR, conducted tests with antibiotics on apricot trees infected
by wilt-on kolkhozes in-the Armenian SSR: The tests produced positive
results. [begin p. 731. Injection of the antibiotic solution into the
tissue of an apricot plant, or Spraying of its crown precludes the further
development of tissue infection. Following the spi.aying, the plants'
recover and in future continue to develop normally. Sick plants not sub-
jected to treatment with this preparation perished completely.
Experiments conducted by C. Askarova (SOIUZNIKHI) [All-Union Scientific
Research Institute of Cotton] under field conditions, on a 12 ha area of
cotton crop treated with the antibiotic solution, showed a 40-60% decrease
in plant infection by gummosis. At the same time, the cotton yield in-
creased accordingly.
Experiments conducted on kolkhozes in the Uzbek S8R have shown that
plant infection by gummosis reached 44% when the 3otton seed had been
treated ilith water, yet the number of infected cotton plants did not
exceed 2.5% when the seed were treated with the antibiotic solution.
As regards their action, antibiotics are not inferior to granosana
IITUIF-1 [Scientific Research Institute of Fertilizers and Insedtifuges],
and in some cases they have obvious advantages over the latter. Thus,
for instance, antibiotics accelerate the germination of seed, increase
their sprouting capacity. Furthermore, they are not injurious to plant
tissues; yet at the same time they kill microbes of infectious diseases.
They remain active within the plant from 7 to 30 days
. N. A. Krasil'nikov, Corresponding Member of,the Academy of Sciences
USSR, for antibiotic treatment of agricultural plants recommends the .
injection of antibiotics into the trunk of a tree through an opening'
made with a drill, and spraying the crowns of trees and their leaves.
Treatment of seed of agficultural crops before seeding is -more effective
method. According to N. A. Krasil'nikov's data, destruction of pathogenic
microbes', bacteria or fungi requires 5 to 100 ml of antibiotic-solution,
depending on the characteristics of the microbe and the plants. Anti-
biotics can also be added to composts. Antibiotic substances added to
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composts eliminate the incidence of infectious diseases of plants.
Trans. 516
The report submitted by N. A. Krasilnikov on this question at the
meeting of the Technical Council of the Ministry of Agriculture, USSR,
aroused-the interest of specialists in the new discovery in the agricultural
science. The meeting of the Technical Council noted concrete measures for
the further study and adaptation in produation of the new method for the
control of diseases of agricultural crops.
NEW TYPE OF FERTILIZER?
(k Summary)
by
R. Adelman
At the end of 1953, the Technical Council of the Ministry of
Agriculture; USSR, heard the report of Candidate of the Agricultural
Sciences, N. N. Sokolov, on the use of "juice water" -- liquid waste of
potato starch factories -- in fertilizing agricultural cropsi The term
"juice water" is applied to cell juice of potatoes diluted with pure
water. The quantity of "juice water" which accumulates during the season
when the starmh factories are in operation on but one central belt of
the Soviet Union, is large enough to irrigate field crops and vegetables
covering an area of 14 thousand ha. The report includes a chemical
analysis of "juice water" properties and shows the results of experiments.
In view of the importance of this waste product in raising the
productivity of agricultural crops, The Technical Council, following a
discussion, adopted a resolution that a new type of fertilizer be added
to kolkhoz production at an early date.
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Nikitenko, G. F.
Vnutrisortovaia i Mezhsortovaia Vegetativnaia
Gibridizatsiia kak Metod Polucheniia porodno
Uluchshennykh_Elitnykh Semian Ovsa i IAchmenia
(Intravarietal and intervarietal Vegetative
Hybridization as a method for obtaining genetically
[porodno] Improved Choice Seed of Oats and Barley]
Selektsiia I Semenovodstvo, 17(5)335-39. May 1950
61.9 Se5
(In Russian)
INTRAVARIETAL AND INTERVARIETAL VEGETATIVE
HYBRIDIZATION AS, A METHOD FOR OBTAINING
GENETICALLY IMPROVED CHOICE SEED OF OATS
AND BARLEY
Trans, 617
(In full)
By
R. Adelman
High genetic [porodnye] properties of choice seed, i.e. their in-
creased vitality, adaptability to living conditions and, finally, their
capacity to produce a higher yield, are the basic characteristics dis-
tinguishing these seed from the seed of mass reproduction.
Nichurin's paramount agrobiological science has developed methods,
already widely utilized in practice, aimed at increasing the genetic
properties of the seed of perennial agricultural crops. These are;
primarily, the methods of intravarietal and intervarietal crossing.
At the foundation of these methods lies the most important biological
regularity contained in the fact that the crossing of organisms, even
though there are slight differences between them, is biologically
advantageous, for it results in the production of offspring with increased
vitality, with a heightened capacity to adapt itself to ever changing ?
conditions of external environment, while self-fertilization, particularly
a protracted-one, entails subsequent reduction of vitality, degeneration
of offspring.
This regularity, first, as is known, noticed by Darwin, has been
demonstrated in exhaustive ekperiments and has been developed further
creatively in the IFilorks of I. V. Nichurin, T. D. Lysenko and their
numerous followers.
From the viewpoint of Michurin's teaching, the process of fertiliza-
tion is a process of reciprocal assimilation-dissimilization, a process
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(2) Trans 517
of incorpordtion, to a certain degree, of various sexual cells into a
.single body.
, As Academician T. D. Lysenko teaches, the biological nature of the
fertilization process is constituted by the process of reciprocal metabolism
between sexual cells fused in one zygote. FuSion of distinct sexual cells
into one [begin p. 36) cell creates a. biological contrast in the single
living body imparting to it greatef vitality, greater adaptability to
conditions of external environment.
However, the crossing of organisms with sexual cells, already at
variance, is not the only means of increasing the vitality of the offspring
of plant organisms i such organisms can be produced also through direction,
by the will of man.
The process of forming in the organism sexual elements with different
properties can be directed through substitution of appropriate cultural
"conditions and through vegetative flybridization based on variation in the
metabolism of the graft components.
A vast amount of experimental work in vegetative hybridization of
plants shows grafically and convincingly that transmission of hereditary
properties of one strain to another can be accomplished without fusion
of the elements, solely by transmitting the plastic substances; it shows
also that the characteristics acquired by the organism through grafting
are inherited by the seed progeny. Vegetative hybridization is accompanied
by increased viability of the progeny.
Thus, in grafting, as well as in sexual crossings, organisms are
formed with stronger biological contrasts and, therefore, with more
vitality. Analogy in the conduct of vegetative and sexual hybrids can be
understood only in the light of the well known theoretical hypothesis of
Academician T. D. Lysenkoi, according to which the variation in plants
prompted by changed environmental aonditiens of vegetative and sexual
hybridization has one common basis.
Such a common basis is the organism's adaptation to changed conditions,
changed nutrition, which lead to the formation of reproducing cell,
internally-heterogeneous, and contrasting in theirpotentials and require-
ments, i.e. a change in-the type of metabolism definitely and specifically
Meant for the organisms.
The method for growing genetically improved choice seed, developed
. 1 T. D. ?Lysenko - Agrobiologiia, 1948, p. 376
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Trans. 517
for most agricultural crops, and the fundamental pidnaiples-of which are
are stated in the article written by Acadethician M. A. 01'shanskii-and
Candidates of the Agricultural SOiences, F. G. Kirichenko and E. T.
Varentsa (see Zhurnal "Selektsiia i Semenovodstvo" No. 7, 1949) is based
on the widespread use of intra-and inter-Varietal crossing in free
selective fertilization, in conjunction with expedient cultural-practices
for hybrid seed, and continuous selection for their improvement.
However, since the method for intra-and intervarietal crossing is not
completely developed, it is recommended that the practically unalterable
(former) scheme for growing choice seed be used for a series of the most
important crops, such as oats, barley and others.
Yet, it is precisely these crops which, most of all, are in need .of
methods that will increase the vitality and improve the strain in the
culture of choice seed.
It is known that, in protracted self-fertilization, the reason for
depression of fertilization and, as a result, the decrease of vitality of
the offspring is the comparative homogeneity of the sexual cells which
participate in fertilization. In natural populations or in pure lines
of self-pollinating plants which blossom more or less openly (as for
example wheat), there is always the possibility of cross-breeding, which
sharply decreases the harmful effect of inbreeding.
The specific characteristics of the blossoming of oats, and partic-
ularly of barley, nearly eliminate a similar possibility. It is known,
for instance, that in intravarietal 'crossing of oats and barley by the
methods used at the present time [begin p. 37] the number of grains
formed iS often quite negligible in relation to the number of castrated
blossoms. The low percentage of success is a result of the circumstance
that the pollen falls on the stigma of castrated blossoms in limited
amounts. As a result, instead of obtaining biologically advantageous
selective fertilization, there occurs limited pollination, fertilization with
a negligible amount Of pollen which is more an extremely forced than a
selective fertilization.
The circumstance indicated should be noted also because of the fact
.
that in barley, for instance, in cases in which it becomes necessary,
for practical purposes, to resort to free backerossing, the depression
caused by limited pollination, due to pollen deficiency, destroys complete-
ly the favorable effect of crossing; the picture obtained resembles in
many-respects the process of fertilization seen in isolated self-fertiliza-
tion. Hence the progeny of seed obtained as a result of such fertilization
(in limited selection of pollen) frequently is characterized by poorer
development and productivity.
As ACtidetician T. D. Lysenko points out," the degree of embryo
vitality ..... equals; in many respects, the degree of fertility. Tho
less the fertility, i.e percentage of fertilization of the ovicells,
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Trans. 517
- the loss also the degree of vitality of the seed obtained"2.
Leaving the development of a more perfect method for intravarietal
and intervarietal crossing an open question -- though its urgent need for
crops like barley, oats, etc. is obvious -- and proceeding from tho facts
stated above, we make bold to propose another method for obtaining
genetically improved choice seed of oats and barley. In our opinion, this
could be the Method of Intravarietal and Intervarietal Vegetative
Hybridization.
Up to now, specialized investigations have not' been conducted for
the purpose of inquiring into the effectiveness of intra-and intervarietal
vegetative hybridization as a method of obtaining genetically imp'roed
seed. It is, however, known, for example, that iii Academician
Prezent's3 experiments, intravarietal grafts of summer wheat Liutescens 62
proved more vital and, as 'a result, more productive than control (not
grafted) plants. To a certain extent, this is indicated by the entire
enormous factual experimental material on vegetative hybridization of
plants.
A large number of works on vegetative hybridization of almost all
field crops, including cereals, have ensured the developthent of a
sufficiently simple and reliable hybridization technique. Of the many
technical methods of vegetative hybridization of plants, in the case
under inveatigation, we primarily refer to the method of embryo trans-
plantation. The vast amount of experimental work accomplished by employ-
ing this method, and accumulated by the workers of the State Selection
Station, permits its presentation without any fear that the intricacy
and unadaptability of the technigue of this method will preclude its
application at the rate required.
Therefore, we suggest that, in growing genetically improved-choice
seed of oats and barley, the method recommended by Academicizn M. A.
01'shanskii and Candidates of Agricultural Sciences, F. G. Kirichenko
and E. T. Barenitsa, be extended by the addition of-a new section: a
nursery for intravarietal (intervarietal) graftings. Then, in place of
the accepted use of ordinary seed, it will be necessary to plant in?
selection-nursery seed obtained from intravarietal and intervarietal
graftings. Lbegin p, 38].
2
Academician T. D. Lysenko - Threo-year plan for the development
of socialized kolkhoz and sovkhoz-productive animal husbandry and
the tasks of agricultural science. OGIZ Sol'khozgiz. Moscow,
1949, p. 23
3
?Academician I;? I. Prozont -- The biological importance of double
fertilization. "Agrobiologiia", No. 5, 1948
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Culture of genetically improved choice seed
on intravarietal and intervarietal graftings can
a) Nursery for growing intravarietal
grafts (Fo);
Nursery for selection (F1);
c) [v] Seed nursery (F2);
d) [g] Preliminary propagation (F
e) [d] Super-choice (F4)
f) [e] Choice (F5).
Trans, 517
of oats and barley based
be visualized as follows:
(intervarietal)
In the first nursery intra-and intervarietal grafts are grown. In
conformity with the method recommended by the abo-ae authors, seed of the
original components (parent forms) for grafting, should be taken from the
more productive crops of the needed varieties. Transplantation of the
embryo can be carried out by any of the known methods, but it is beSt to
use the method of transplanting the dry embryo onto a dry endosperm.
This method is technically easily accessible and, more important, it
ensures a high degree of acclimatization of the grafts and affords the
oppoi-tunity to begin transplantation work in good time, long before seeding
time. In addition, transplantation by this method allows direction of the
reciprocal influence of the graft components through variation in the
amount of the assimilable plastic substances of the endosperm-stock
(grafting on a double endosperm, seeding at considerable depth). Experi-
mental work conducted by this method at the Mordov State Selection Station
shows that the following technique is more acceptable:
In two dry grains of one of the graft components the embryos are
cut with the blade of a safety razor (the indision must be made at an
angle of 40-45? to the surface of the grain). Then a second-incision
is made in one of the grains on the side opposite the embryo. The
incision on the surface are made with a safety razor in two perpendicular
directions, after which tthe grains] are steeped in water, best in warm
water (room temperature), The action of the water causes the incised
surface of the cut to sWell up rapidly (in 1-2 minutes) and to acquire
the consistency of glue. On endosperm thus prepared the transplantation
of the embryo is performed; at first the embryo is glued to the endosperm
on which two incisions had been mades and then the second endosperm is
grafted onto the first] from below.
With adequate practice and skill,
50-75 embryo transplantations per day.
of the required volume of work without
one worker can easily perform
This will ensure accomPlishment
expending much manpower. The
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Trans. 517
glued components keep very well and can be preserved in any type of
container until seeding time; Petri dishes or those of Koch are best for
this purpose.
The grafted grain can be sown in a hotbed, greenhouse or-directly in
the ground at the time customary for planting the given crops, In the
latter case, the sector must be well tilled and fertilized -- best with
manure (50-60 T/ha) -- prior to planting. For the purpose of increas-
ing the influence of the endosperm, the planting depth for grafted grains
should be increased up to 6-7 cm for oats and 7-8 Nn for barley.
Each grain must be provided with the Most favorable conditions for
growth and development: optimal area of nutrition (10 x 30 cm), frequent
supplementary feeding with a full mineral fertilizer (beneath the marker)
watering -- if necessary.
-Careful phonological observations must be conducted during vegeta-
tion: Inspection (field and laboratory), customary in plant selection
for selection nurseries: on the strength of development, productivity per
plant, resistance to phyto- and entomological pests, stability of straw,
etc.
111 Seed of the S'elected plants are seeded the following year in a
selection nursery.
The subsequent work is carried on in full compliance with the re-
quirements of the method recommended by AcadeMician M. A. 01'shanskii,
Candidate 6 of the agficultural sciences, F. G. KiriChenko and E. T.
Varenitsa. [begin p. 39]
Using the method described, it will take a maximum of 2000 grafts
to produce the necessary quantity of F1 of rejuvinated seed 50 thousand);
the culture of these grafts will require an area of 75-100 m .
With the amount of work involved, it will be easy to ensure the best
possible care and observation of growth and development of the grafts,
not only under conditions of State Selection Stations, but also in
regional seed kolkhozes [raisemkhoz] and on sectors of experimenting
farmers.
It is apparent that the success of cultivating intravarietal and
intervarietal vegetative hybrids depends, in many respects, on the
weather conditions of the year, the characteristics of the farm, the
ability of the experimenters etc, and, therefore, the technics of the
work adducted here must be considered as being schematic, as having a
comparative importance.
Mordov State Selection
Station
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Aleev, N. P.
Opyty po izmeneniiu Prirody Pshenitsy
Putem Vegetativnoi Gibridizatsii
[Experiments in changing the nature of
wheat through vegetative hybridization]
Agrobiologiia, 1948(4)348-56
July-August, 1948 20 Ag822
[In Russian]
EXPERIMENTS IN CHANGING .TBE IjATURE OF WHEAT
THROUGH VEGETATIVE HYBRIDIZATION
Trans 518
In full
Brg
R. Adelman
In this report-we describe some of our experiments-in vegetative
hybridization of wheat conducted in the years'1940-1942. We concentrated
on the transplantation of embryos of the seed of certain varieties.upon
the endosperm of other varieties and species of plants. Here are the
results-of the transplantation of embryos of wheat onto the endosperm
of rice.
The choice of the embryo seed for a scion was based, first of all,
on the fact that the embryo -- the younger-phase of plant life -- is
more easily influenced by external factors. It has been established that
the embryo aeed can develop not only on its own endosperm but also on a
strange one.
The seed of wheat of Erytrospermum 0841 and Melianopus 069 were
selected for the experimental work. The seed of'riOe of the Karatal'skii
8679 variety was selected for the stock (endosperm). After causing a
slight swelling in all experimental seed, their embryos were cut off to--
gether with a small part of the endosperm (to avoid injuring the embryo).
The wheat embryos were pasted with wheat dough to the endosperm of rice
on whi6h the small area obtained after removing the embr9-o expanded a
little. The embryos grew firthly onto the rice endosperm. The total
number of grafts made was 100.
The grain of wheat thus prepared were planted at the end of April in
paper cups filled with sand and a slight admixture of soil. In growing the
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Trans. 518
plants by the method described, the wheat embryos enjoyed the nutrients de-
rived from the substances of the rice endosperm,
Almost all of them germinated, but from thence their fates differed.
Most of the sprouts perished early; sothe of them succeeded in develop-
ing one or two regular leaves and then perished s and only a few specimens
developed to the point of the formation of generative organs. Some sprouts
may have perished as a result of insuffioient nutrition due to the loss of
contact between the embryo and endosperm.
When the wheat had reached the phase of two-three leaves, it was
planted in the field. Here it was watered in proportionto-soil dryness,
and was fertilized three times with full mineral fertilizer. Nonetheless,
the growth of the wheat was poor, the plants looked stunted and were easily
susceptible to disease.
Up to the phase of heeding there were left 16 specimens of wheat. A
few of them formed branches, but most of them produced only one stem. Many
stems formed spikes which fluctuated in length from 1 to 10 cm. [begin p.
49.J The spikelets of all spikes had scales of spikelete and buds, but the
stamen and the pistils were absent from most of the bud. Although in
fourteen out of sixteen plants the spikelet and bud scales were well
formed, they began early to turn brown and to fall off, so that nothing but
the core was left of the spike. Only one plant of the Erythrospermum
variety (fig. 1) and one plant of the Melianopus variety produced one spike
with kernels each. Fifteen kernels were found in the spike of the first
plant and in the spike of the second plant -- 8 kernels. The grains in
both spikes were, generally very, small and shrivelleds.sometimes-a little
larger, but there was not a single normal, fully developed grain._ The
grains in the grafted Erythrospermum were considerably whiter as compared
with the original form. The only Erythrospermum spike which produced
grains was distinguished by a strong pubescene of the spike and flower
scale, by porosity, by a large number of flowers in the spike -- up to
11, counting the poorly developed and sterile flowers. Another spike of
the same specimen disintegrated completely.
The fruit-bearing specimen of Melianopus (fig. 2) produced 3 spikes.
Fig. 1 - Left -- Original form of :Erythro-
spermum 0841.
Right-- Grafted plant (Photo of
1940).
Fig. 2 - Grafted plant of wheat, Melianapus
069 onto the endosperm of rice.
One grain-bearing spike had abnormally curved awns and a more porous struc-
ture than is normal. [Begin p. 50j The number of flowers in the spike came
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Trans. 518
to 9, but only 6 were fertile. Another spike, also abnormally porous, had
no awns, was sterile and soon disintegrated; a-third spike -- compact,
somewhat club-shaped, without awns and sterile.
Such were the hybrid plants of the wheat obtained in 1940.
To demonstrate the diversity of the plants grown from the seed obtained
by vegetative hybridization, I shall, describe the results of the crep from
the seed harvested in 1940.? As indicated above, only 2 spikes of two plants
produced seed l the plant of Melianocas wheat yielded 8 grains, and the plant
of Erythrospermum wheat -- 15 grains, ?
These grains were sown in a greenhouse on April 21, 1941, In the case
of Melianopus all eight grains germinated, in the case of Erythrospermum
only ten. On April 30 the plants were transplanted in the field. On the
Melianopus plants 2 specimens reached the stage of seed formation, of the
Erythrospermum plants ---7 specimens.
The rest of the plants dried up and perished at certain stages of their
development, or they produced sterile spikes, Harvesting of the plants was
carried out on July 31, which means that the vegetative period lasted exactly
102 days.
I shall describe bi.iefly some-of the vegetative hybrids'derived from
Erythrospermum and rice. Plant No. 1, first-goneration (fig. 3), Height
of the plant 88 cm, length of spike -- 15 cm. Two spikelets are perched
side by side at the bottom of the second, third and fourth rachises each;
in addition to these three twin spikelets there are also 19 single spikelets.
The total number of spikelets in the spike is 25 (including the three pairs
of twins); grains -- 106. Qualitatively all grains-are satisfactory,
they are white of pibment and barrel-shaped in form.
Plant No. 2 (fig. 4) reached the height of 82 cm including the spike.
The spike contained 22 rachises, There were two spikelets on each --.the
first, second, fourth, fifth, sixth, seventh and-eighth rachises; on the
remaining fifteen rachises there was one on each. The number of graine
in the twin spikelet ranged from one to five, in the upper six -- none,
[several words deleted] The 77 grains found in the spike described were
poorly developed, thin, and of a white pigment.
Plant no. 3 had especially large leaves; the blades of the upper
left -- 30 cm long and 1.8 cm wide (at the part where the width-is broadest).
The blade of the leaf is thick and has sharply protruding veins. In the
one and only spike, twin sterile spikelets were found underneath the
second and fourth rachisoS.
? The first and third single spikelet are likewise sterile, In the
rest of the spikelets 1 four grains each (1 spikelet), 3 grains each
(3 spikelets), 2 grains each (6 spikelets) and ono grain each (8 spikelets);
total number of grains in the spike -- 33. The grains were full, large
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Trans. 518
grains, but there were also small and poorly deireloped grains, Color and
the form were the same as in the preceding ones.
? Knew 'characteristic which appeared in this generation is the
distribution of twin spikelets on some of the rachises in a number of
'spikes,' This characteristic waS found in the progeny in the same incidence
through the year 1946,
In 1941 new grafts were made. The embryos of wheat were grafted onto
the endosperm of Karatal'sk ride Tsezium 0111, Erythrospermum 0341,-Melia-
nopus 169 and Gordeiforme 0189. Altogether there were 400 graftings, The
seeding was carried out on April 29.
? On May 14, some plants with two leaves each were. removed from the
soil for inspection. A cythe could be clearly observed in the Wheat embryo
while it was in the ground. The part of its own endosperm which adjoins.
the cyme, left there when the operation was performed was completely
absorbedg,[begin p, 51] it had .left on the cyme cavities and depressions
containing reserves of nutrients. Nothing was left of the rice endosperm
except the'membrane; the wheat embryo had used up all the substances of the
rice endosperm,
41/ Fig. 3 -- First generation, of hybrid
wheat Erythrospermum grafted
onto rice.
Fig. 4 -- First generation of hybrid wheat-
Erythrospermum grafted onto rice.
Only 16 plants reached the phase of fruiting, and, in addition, three
plants developed with sterile spikes. Partly sterile spikes wore found in
fertile specimens; for instance, a- spike of Erythrospermum 0341 had a total
Of 14 spikelets, yet only 5 of these yielded 6 grains,
In hybrid plants of wheat emerged properties which are absent in the
ordinary plants of the original varieties, These new properties are un-
equally distributed not only among separate specithens, but also within the
bounds of a single branch and even a single spike. I shall cite examples
of certain variations.
For instance, in one specimen of Gordeiforme the upper internode bear-
ing the spike is 8 cm long; it ends with a spike 4,5 in length; both, the
internode and the spike were confined in a strong sheath 18 cm long. The
spike, markedly bent, partly crumbled and sterile, emerged together with
the internode that bears it not through the opening of the sheath apex,
but through an opening in the side wall of the sheath. In another speci-
men of the Gordeiforme the spike came out only halfways and was sterile,
There are perfectly shaped spikes which look normal, but are sterile.
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Trans. 518
[Begin p. 52] Alongside of the plants that yielded 6, 11 and 20 grains,
there are spikes which produced 75 grains in 15 spikelets, 78 grains in 18
spikelets etc. At times, the kernels were very fine, thin and poorly
developed, at times -- large and well filled out.
The kernels in the wheat referred to above as Erythrospermum 0341, were
distributed in the spike as follows in the first, fourth, eighth and ninth
spikelets -- one grain each, in the seventh -- two grains, in the remaining
nine spikelets no grains whatever. Spikelets with a varying number of
flowers, up to eleven were found in some hybrids. The-number of grains in
the spikelets was just as varied -- from zero to seven.
Awns are subject to strong variation. In some Gordeiforme spikes the
awns are very long (up to 23 em), curved, and, at the top, strongly diverge
in different directions -- the space between the tips of the awns measures
22 cm, In other Gordeiforme plants the awns project straight upward and .
rise above the spike only 5 cm. Finally, there was a plant with practically
no awns on the spike; only in two spikelets the flowers had awns up to 4 cm
in length, but in the rest of the flowers the scales taper off either with
awns up to 1 mm, or into a sharp point. The awns in experimental plants
of Meljsanopus wheat are also extremely varied as to length and pigment (from
black color to white), they are of a different length and color even in one
and the same spike, and even the same awn is differently colored in different
parts. -
An interesting phenomenon is the branching out of the stem. Branching
was Observed in experimental plants of two varieties Melianopus and Gordei-
forme. For instance, in Malianopus 069 (fig. 5) a side shoot emerges from
the axil of the third leaf; the stem is 4 cm long; it has at its tip ?a
spiko 2.3 cm long, and including the aWns -- 13 Nn long. The spike bears
a spikelet with eleven-sterile flowers. On the same plant, a sedentary
spike, 2 cm long and 1.8 cm wide-at the upper and broadest part, emerged
from the axil.of the fourth leaf. The single spikelet had 7 flowers With
awns up to 7.5 cm; the spikelet yielded two good and one Sickly grain. The
top spike of this plant .(length 5-cm, and with awns -- 13,5 cm) is found
on the upper internode, 4 cm long. The awns are yellow-green at tha.base,
yet at the central and upper parts they aro slightly gray-brown to brown
and even entirely black (different awns are differently colored). Long
awns are found only in the lower flowers, the rest have either none or some
that do not exceed 7 mm. A spike that has 19 spikelets; one of them had -
seven flowers and yielded 5 grains. The rest of the spikelets were sterile.
Out of ten plants, Gordeiforme produced three plants with ramoso stems.
One of these plants- is constructed as follows; from tho axil of the third
loaf omerge8 a stem with two internodes: the first -- 3 cm long, and the
second -- 1.7 cm. The latter ends with a spike 3 cm long, with the awns
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10 cm. The spike produced 8 Small but full grains. From the leaf axil of
this axillary shoot another stem grew out -* 2 mm long -- and ended with a
spike 1.8 cm long and with the awns 5 cm long. This spike produced six
grains. The top spike yielded 11 shrivelled poor grains. From three Spikes
of one stem a total Of 25 kernels were harvested.
In another plant of Gordeiforme 0189 (fig. 6), the lower spikelet of
the top spike-lags 3 cm behind the others; the length of the rest of the
spike is 6 cm.
The longest awns are those at the, top, up to 4.0 cm; in the-middle part
they are considerdbly shorter, and the lower spikes have no awns. The spike
yielded 19 grains. Abegin.p. 53] From the leaf axil a stem grew out ending
with a 6 cm long spike, The Spike had 15 spikelets which contained 48 grains.
The largest numbei- of grain-in a spikelet was five; the length of the grains
fluctuated ?from 4,5 to 8 mm, The longest awns were in the middle part of -
the soike; the spikelets at the top and at the bottom had no,awns whatever,
Fig, -- Plant of the Melianopus 069 vario44
grafted onto the endosperm of rice,
Fig. 6 -- Plant of the Gordeiforme 0189 variety
-
grafted onto the endosperm of rice.
Among plants with non-ramose stems were found spikes which contained
more than 70 kernels. Many spike1et6 of the same plants contained an in-
creased number of flowers and grains.
All seed of the 1941 crop were sawn in the field on May 11, 1942. The.
sowing was carried out-with a spacing of 65 x 35 cm. The plants were
harvested on August 29.
[Begin p, 54] An analysis of the first- generation of wheat plants,
spikes and apikelets of Erythrospermum 0841 produced a great variety. Re-
gardless of the fact that Erythrospermum has long awns, all the hybrid plants
of 1942 were nearly awnless; awns were found only inthe five-six upper -
spikelets, yet not in all of them, and -their length did. not exceed 1,5 cm.
The number of spikelets in the spikes fluctuated sharply i from eight to
twenty-five. The same in the preceding year, twin spikelets were found on
some spikes, i.e. not ono, as is customary; but two and sometimes even
throe spikelets wore located on one rachis, The number of these twins
fluctuated between one and eleven (there Was one spike on which 11 twins and
4 single spikelets wore perched on fifteen rachises). More often than not
these twin spikelots were sterile. Tho spikelet and flower scales were
strongly depressed.
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Trans. 518
The number of flowers in the spikelet also varied 4 There were spikelets
with three flowers, yet spikelets with eight flowers were also encountered.
There Were sterile spikes with but two-three grains, yet there were also
spikes containing more than fifty grains (the largest number was 85 grains,
and in a spikelet -- 7 grains).
The progency of wheat grafted in 1941 was marked by the absence of
ramose forms. Productive bushiness was varied; the number of stems with
reproducing Spikes fluctuated in individual plants between 1 and six; yet
concomitantly, entirely sterile spikes were found in'the very same shrubs.
There were also plants which had sterile spikes only,
The number of kernels in different spikes is also wite varied;
variations *ere observed also in the dimensions and in other properties of
the kernels, Much variation was observed in the awns.
It. should be noted that culture of the seed of Erythrospermum 0841,
first obtained in 1940 by growing its embryo on the rice endosperm, was
continued annually through 1946. Many variations survived seven generations.
Figure - 7 shows the mnitdIraralLity of spikes obtained in the sixth generation
(1946).
First of all, one notes the conspicuous absence of awns which, in the
original form of Erythrospermum 0841, were twice as long as the spike. Tho
spikes of hybrids are considerably longer than the spikes of the original
form grown, for comrarative reasons, under identical external conditions.
Thus, the spikes of the original form are most frequently 8-9 om longs yet
the average length of the spike of a vegetative hybrid is 11-13 cm. In
structure the spikes are dissimilar: there are spiles with twin spikelets,
i.e two spikelets (occasionally even three spikelets) are perched on one
rachis. At times the spikelets are multiflorous and yield 4-6 grains, at
times they are poor in flowers and yield but 2-3 grainS; seen from the
face, the first are considerably wider than the latter.
The spikelet and flower scales are thickly furred, this characteristic
is loss pronounced in the flowers at the top of the spikelet. The kernels
are closely encased in the flower scales and do not spill.
Big differences are observed in-the kernels. In the original form they
are red, comparatively thin and long, Hybrid ketnelS. are white with a
slight yellowish hue, shortened and thicker (fig. 8). Equal amounts of the
largest kernels of both forms were used for dimensional comparison: the
length of kernels from the original form equalled 8.3 mm, and the width ---
3.3 mm; In the hybrid form the corresponding sizes wore l 7.3 mm and 3.8 mm.
The maximum length [begin p. 55] of the kernels of the original form --
9 mm, maximum width -- 3.5 mm; the corresponding sizes in the hybrid formg
8 mm and 4.5 mm. Therefore the weight of the selected. largest seeds of
both forms of wheat is fairly close 1000 grains of the original form weigh
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51 grams, of the hybrid -- 55 grams.
Fig. 7 -- Variation in the form of spikes of the
vegetative hybrid Erythrospormum 0841 x
rice in the sixth generation '(1946).
Trans. 518
If the form and structure of the spike do not as yet represent stable
characteristics (inultiflorous and multi-grain spikelots, the presence of twin
spikolots on the rachis), then the form of the endosperm, its exceptionally
white coloring and groat softness and moaliness characterize complete
stability and constancy transmitted by inheritance in the course of seven
generations.
It is obvious from this report that the variations which are obtained
in wheat plants, grown from embryos nourished by a rice endosperm, are
extremely varied and extend to the generative organs as well a8 to the
vegetative ones, while they become fixed in the seed offspring.
Among hybrid plants are found superior, economically valuable forms.
For instance, plants, the grain of which have the absolute weight of 47 g,
arouse con8iderable interest.- The specimens which yield 5-6 reproductive
spikes etc. are also valuable. It is ourtask to se6ure in the following
generations these superior traits of altered plants.
On the basis of observations of the hybrids obtained, it is possible
to draw certain preliminary conclusions as to whether those valuable
properties.which would be desirable not only to preserve, but also to
strengthen in the new forms, are tapering out, Or are progressing While
the generations are inthe process of development.
The overwhelming majbrity of agricultural characteristics, such as
spike formation, high quality of grain (fulness, dimensions), productivity
per plant among the better specimens, produce in the offspring higher
indicators in comparison with the parent forms. The amount of grain in the
spikelets increases,[begin p. 56] but the number of spikelets in the spike
decreases somewhat, hence here is a decrease'in the amount of grain per
spike; but this does not occur in all plants.
It was mentioned above that culture of the progeny of a.plant grafted
in 1940 continued through. the year. 1946, and that it firmly retained many
positive indicators.
Among the-hybrid plants obtained are found quite a few shortcomings
(sterility etc.), but there are also many valuable properties. Yet of the
highest importance is the fact that by practicing, vegetative hybridization
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Trans. 518
. it is possible to obtain offspring of the kind of plants in which sexual
hybridization is t1p3ssible (rice and wheat). The practice of directed
culture and selection of the best plantsfor vegetative hybrids will
produce economically highly valuable form t with properties that will sharply
differ from those in the initial. material.
Fig. 8 --
1.
.Top row. '?
Kernels of the
Erythrospermum
Bottom row --
Kernels of the-
Erythrospermum.
original form of
0841; ,
hybrid form of
CONCLUSIONS
Introduction into the tissues of an embryo, that is be-
ginning to develop, nutritive substances unadapted to its nature,
diverts the course of the developmental processes from the normal
and, as a result, new propetties are formed in the plant cells; .
these properties appear in the form of distinguishing morpholog-
ical and physiological plant characteristics.
2. Various new characteristics do not appear immediately in
the year the grafting is performed, but they continueto form
and develop in subsequent generations.
3. The instability of characteristics, the disappearance of
former ones, appearance of new ones, indicate a mutability, an -
inconstancy of the inherited nature of vegetative hybrid plants.
4. Cells and tissues, the generative as well as the somatic,
consist of multiple properties within the limits of a single
plant, a single spike, spikelet, bud.
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.(10) Trans. 518
5. -In our experiments in addition to the general disturbance
of the hereditary properties of wheat, the hybrid plants were
fed some of the properties of rice. We refer to the change in
the color of grain from red to white, the compactness of flower
scales, reduction of dimensions and color of awns, appearance
of pubescence on the scales of spikelets and flowers, and some
others,
Botanicheskii Sad AN Kazakhskoi SSR
Alma?Ata
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(1)
Oeipov, A. E.
Opyty po vegetativnoi gibridizatsii
zlakovykh
[Experiments in vegetative hybridiza
tion of gramineae].
Agrobiologiia, 1949(6)141-146.
November/December 1949 20 Ag822
(In Russian)
Trans 519
(In full)
Byl
- R. Adelman
EXPERIMENTS IN VEGETATIVE HYBRIDIZATION OF GRAN1NEAE
The first experiments in vegetative hybridization of gramineae Were
begun [at the Riazanskaia Selektsionnaia Stantsiia] in the year 1938. To
begin with, it was necessary to determine on a grafting technique,. Oat
plants of various ages were used for this purpose, beginning with the phase
of develoPment of the first surface node and ending with the phase of panicle
emergence. Grafts were performed on a stem, a, node, in a cut or cleft. -
Binding was accomplished with gauze, saturated bast and isolation ribbons,
Of 173 grafts Coalescence of the scion with the stock was secured in only
four instances,
In addition, in a large quantity of oat grain, the embryo of one grain
was transplanted onto the endosperm of another. In 4-5 day t some of these
grains showed sign 8 of growth, but they soon perished for reasons unknOwn
to us at that time.
The first year's work showed that grafting of Gramineae should be per-
formed in the earliest phase of their ontogeny.
In the year 1939 the experiments were continued with attention center-
ing primarily-on the transplantation of embryos. Experimental technics were
the following. Seed reserVed far grafting were shelled of their hulls fyroi'k
was conducted chiefly with hulled forms). Then the seed Was placed on moist
sand and covered with glass saucers. In approximately 20 hours (at sufficient-
ly high tetperatures -- 12 hours) the seed of oatsat.elled up and showed signs
of growing, and those of bromegrass -- in 30-hours. After this, the embryos
of stock and scion were cut off with a razor,. -It was established that the
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Trans. 519
cut on the stock had to be larger than that in the scion, so that no part of
the embryo might be left in the stock.
The embryos of the scion were carefully cut off and transferred onto the
endosperm of the stock. The edges of the cuts were smeared with collodium.
Just as soon as the collodium was dry, the seed were-again placed on
moist-sand under glass saucers. The sand under the saucers were always kept
moist. The grafted embi'yos of ioats began to.graw in 2-3 days, and those of
bromegrass in 4-5 days. Transplantation into pots [vozmyj with soil was
carried out 5-7 days following the grafting.
In a series of cases it was established that the embryos of the oat scion
cut off carelessly (with the endosperm) did not grow onto the stock, but they
continued to grow and, having been transplanted into containers with soil, they
produced weak, but nonetheless fertile plants.
A 194d'transplantation of an embryo of smooth brome [Bromus inermis] onto
the endosperm of oats produced 5 plants, including 4 from a combination of
bromegrass and oat t of the Moskovskii A-315 variety, and one from bromegrass
and naked oats (Av. nuda).
(Begin p. 1421 In the fall of 1939, the plants obtained were transplanted
to the field, and in the spring of 1940 they were well advanced in growth.
Their development proceeded satisfactorily, and, subsequently, control plants
of bromegrass lagged behind the experimental plants by 10-12 days.
All plants obtained from grafted embryos were stronger than control plants.
The variations produced,by the influence of the stock affected not only tne
general character of the plants, but also their separate parts and organs.
The panicles and straw of these plants looked more like oats (fig. 1). The
number of spikelets in their panicles was also larger.
The plants obtained from the graft of a bromegrats embryo onto the endo-
sperm of naked oats, had a softer leaf than the 'scion.
The axil of straw was naked. Furriness was negligible beneath the axil,
yet above it, it was sharply promounced. Spikelets of the panicle 'were
larger and they contained more flowers than are found in smooth bromegrass
(fig. 2).
(x) Translator's notes it seems that 1939 was meant instead of 1949.
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(3 )
Trans-. 519
The kernels, with-respect to pigment, resembled broomgrass, the pigment
of the hulls was light. The internal hull of the flower was nearly light
The external flower hull was larger than the kernel and looked as though it
enveloped it, and was easily removed (fig. 3). Among the grain of this plant
were found specimens, the kernels of which hulled easily from the flower
chaff: The stem of the grain is markedly shorter and thicker than in broome,-
grass.
The grain of plants obtained as a result of grafting the embryo of
smooth broomegrass onto the endosperm of oats-, Moskovskii A-315, were fuller
and looked as if they had been pushed through the flower hulls which did not
join either at the top nor at the sides.
The grain obtained was not uniform. The size of the grain, their form,
and the character of the flower hulls differed rather strongly within the,
limits of the same cOmbination and even on the same plant,
[fig. 1] -- Left:
Panicle of smooth broomegrass;
Right
Panicle of oats, Moskovskii A-315;
In center:
Panicle of plant obtained as a result
,pf grafting the embryo of smooth broome-
grass onto-the endosperm of oats Moskov-
skii A-315.
[Begin p. 143]
[fig. 2], -- Left:
Spikelet of smooth broomegrass;
Right:
Spikelet of Avena nuda;
In center:
Spikelet of plant obtained as a result
of grafting smooth braomegrass onto the
endosperm of Av. nuda,
_
On the basis of the factual material obtained, we have arrived at the
conclusion that,' regardless of the short space of time in which the grain
embryo of one plant was nourished by the ready plastic substances of the
endosperm of another, there Occurred :marked variations in the characteristics
of the scion exposed to the influence of the 'stock. We were, however, unable
?
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?
(4) Trans. 519
to tell then whether or not these variations will be transmitted to the follow-
ing seed generations, During the patriotic was these experiments were dis-
continued, and they were resumed only in the year. 1947 with the partially pre-
served material. The greatest interest centered on the transplantation of
the barley embryo.
In the year 1941, the embryo of,Pallidam 45 barley was grafted onto the
endosperm of the trifurcatum form of barley., As a result, a plant was produced,
the spikes of which included grain with hulls and naked Oain, and as a rule,
the naked grain was found in the lower part of the spikes. In a single .spike
one could encounter awns that were normal in Palliamm.. 45, shortened awns,
and.awns that looked as if they had been fractured beneath the axil.
In the spring of 1947, the seed obtained was seeded in the field
separately, naked grain, grain with changed awns, and hulled grain with
awns normal for Pallidum 45, All of them produced normal germinatio4n, but-
the plants obtained-from naked grain were completely destroyed by fleas
and the twedish fly.
The hulled'seed with changed awns produced plants with multiple rows
of awned spikes. Individual plants had soft awns, and one plant had.ramose
spikes.
- Hulled seed which had awn t normal for Pallidum .45 produced: one plant
with changed awns, the grains in the lower part of the spike bearing awns
5 cm long and ending in a prong [furca], the rest of the grains having long
awns with a fracture at the center, all grains were hulled; five plants with
rough awns; the grain was hulled, naked and semi-naked; four plants with
shortened awns and naked grain; one plant with long awns and naked grain;
[begin p. 144]. one plant with short awns, 9% of the grain hulled and 5%
naked.
[fig. 3]
Left
Grain of smooth broomegrass;
Rightg
? grain of Avena nuda with flower hulls;
In ,centers
Grain of plant obtained as a result
of grafting the embryo of smooth broome-
grass onto the endosperm of Av. nuda.
_
In the year 1948, the grain. of these plants were seeded separately in
the field. Thus we got.a third generation from the grafts. As a result of
an analytis, a large variety of form was established in the third generation
(fig. 4).
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Trans, 519
Variations formed in the first generation were tranSmitted to the next
seed generations. In the following generations a splitting up is observed in
the characteristics.
Thus, a plant, the spike of which had short awns and long pf-on0, produced
in the progegv5 morphologically different groups of plants (fig, 5).
Hulled grain with altered awns were selected from first generation hybrids,
and in 1947 theirembryos were grafted onto the endosperm of trifurcatum for a
second time. 30 graftings produced but one plant, which, however, cannot
Lfig, 47-7? Spikes of plant.F3 obtained as a result
of grafting the embryo of barley Pallidum 45
onto the endosperm of trifurcatum.
[begin p. 145]
[fig. 5] Progeny of rspike which had long prongs
and short awns.
be wholly attributed to the imperfection of transplantation technics.
As a result of repeated grafting, a plant was obtained which had sturdy
straw, a broad, leaf blade, many rows Of awned spiked and naked grain, The
awns were coarse, notched, 10 cm long, The spike contained 68 grains, the
spikelets were trifloral.
The seed of this plant were seeded in 1948. The plants that developed
from these seed had the same morphological characteristics noted in 1947,
Thus, the form of barley obtained by grafting the embryo of Pallidum 45
barley twice onto the endosperm of trifurcatum has the characteristics of the
scion -- coarse long awns and tall-straw, as well as those of the stock --
a broadleaf blade and naked grain.
To introduce oat varieties resistant to loose smut [Ustilago.avenae],
we grafted in 1939 the embryo of Avena brevis onto the endosperm of the
Moskovskii A-315 variety, and-kn 1941 -- embryo of the variety 'q'obeda" onto
the endosperm of Avena brevis.
In both cases$ no morphological variations were discovered in the year
of grafting. Nonetheless; the material obtained from the graftings was seeded
again the following years. As a result it was established that variations
in size and coloring of grain began to appear, in the first case -- in the
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(6) Trans, b19
3rd year, and in the second -- during the 2nd year (see table on page 146).
The experiment conducted shows that variations resultirigfrom grafting
cannot always be seen in the year the grafting is performed. Moreover, they
may not appear even in the second year of re-seeding This can lead the re-
searcher to the false conclusion that it is impossible to obtain vegetative
hybrids. Meanwhile, in the second', and sometimes even in the third year
following the grafting, the variations become evident.
In the given concrete case, variations in the embryo exposed to the
action of the stock endosperm occurred, positively, in the year of grafting.
But they, obviously, affected a small section of the embryo cell, and, there-
fore, were unable to appear in the first generation.
White and brown grainsselected from a variant of the 1947 yield of
Pobeda variety on Av, brevis were seeded separately in the year 1948. As a
result, the white grains prjdueed plants with white grain, and the brown ones
-- with brawn grain. [begin p. 146].
Characteristics of grain by year of consecutive seeding
Variant
Year of
grafting
, Crop
years
19417
1948
1940
1941
Embryo of Av. brevis
1939.
Variations:
Grain
Grain
Grain en
oats grafted onto
none
white
small,
Mass, gra:
endosperm of Moskov-
and
gray, a
separate
skii A-315 variety
brawn
few large
and white
grains
grains,
large,
white
Embryo of oat variety
1941.
' ---
Varia-
Grain
Grain
Pobeda grafted onto
tions:
largo.
large, en
endosperm of
none
whItoand
mass brow:
Av. brevis
brown
separate
grains
white
No less interesting was the case of partial variation in embryo cells under
the influence of the stock (endosperm), which we observed in the transplanta-
tion-of,the embryo of Pobeda variety Onto the endosperm of the variety Golden
Rain, siisrs also no variations were noticed in the year of grafting, and in
the following year the grafting was repeated. Fifteen plants were obtained
and it was established that the color of their grain varied. In addition, a
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(7) Trans,. 519
a sharp variation in the color of gf-ain was observed on a single plant, except
that it appeared In different stems. Thus, in 14 Plants cut of 15, the grain
of the center panicles was yellow, and only in one case it was a pale yellow,
yet on the side panicles it was Consequently the pigment of grain
varies in one and the same plant.
CONCLUSION
On the basis of the material acquired on vegetative hybridization, we
aro of the opinion that vegetative hybrids of cerealscan be obtained
perfectly by embryo transplantation. ? ?
Experiments show that the embryo of grain of one form, feeding on the
ready plastic substances on the endosperm of -another form, changes its
hereditary properties in favor of the latter.
However; the variations are not always observed in the year the grafting
is performed. Moreover, changed conditions of embryo nutrition may produce
a different effect on the same plant.
Changes in the scion in favor of the stock depend on the length of time
the embryo feeds on the plastic substances of the stock, and on the degree of
influence exerted by substances of the changed section of the embryo upon
the general process leading to the formation of reproductive cells.
In our experiments, repeated grafting produced, as a rule, sharp changes
the most profound action exerted by the stock upon the scion was rovbaled in
tho reproductive organs and transmitted to the following generations.
A. E. OSIPOV
Riazanskaia Selektsionnaia Stantsiia
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0
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(1)
ILLARIONCV, V. F.
Vegetativnaia Gibridizatsiia Rzhi
s Ozimoi Pshenitsei
[Vegetative Hybridization between Rye and
Winter Wheat].
Selektsiia i Semenovodeti-o, 15(11)31-41
November 1948 61.9 Se5
VEGETATIVE HYBRIDIZATION BETTEN
RYE AND 'TINTER 71BEAT
[Fundamentals of Topic]
Trans. 520
(In full)
By R. Adelman
The study of varieties and forms derived by the Iaroslav State Selection
Station from the universal collection of Vir tAll-Union Institute of Plant
Industry] and from other localities of the USSR has demonstrated that it is
an illusion to anticipate the finding of i-eady-made, highly winter-hardy forms
fit for production in the Iaroslav Region.
In three years of work at the station (from 1937 through 1940), more
than 700 specimens of the 1029 planted in the original selection nurseries
perished completely while still in the ground, and the rest proved inferior,
according to a series of economic and biological characteristics.
Being familiar with I. V. Michurinfs work on vegetative hybridization -
of fruit trees, we decided to apply the same method to selection of Gramineae.
Methods and Technics of Vegetative
Hybridization of Gramineae
The Iaroslav State Selection Station had to conduct experimental work
independently, in order to develop grafting technics of Gramineae (1940-1941).
The work of grafting began August 15, 1940 and lasted until May 15, 1941.
The grafting was carried out by two technicians, under the supervision
of the author of the present article, and involved various modifications in
the characteristics of the experimental objects [Gramineae species used].
e searched for the following experimental conditions t [begin p. 32]
1. Stages of development of the graft coffiponents at which
normal coalescence of the latter is ensured.
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Trans. 520
2, Best method of incision of ,plants to be grafted (form,
angle and direction).
3, Places of incision on graft components (surface level
of incision),
4. Methods of junction of coalescent plants,
5. Substances strengthening the'junction of components and
ensuring their better coalescence.
6. Optimal conditions of external environment for the
grafting and post-grafting period tO ensure better coalescence
(temperature, light, moist chamber).
Without describing in detail the progress of our experimental work,
we shall pause here only to consider the conditions we have-found to ensure
normal coalescence of the graft components a rye and wheat.
It was found that ono-centimeter shouts are best for grafting, 'Under
laboratory conditions (at a temperature of 16-18ton moist soil) this stage
of development ia reached in rye on the 4-5th day after seeding, and in wheat
on the 5-6th day. To give the graft plants an equal change, rye was seeded
24 hours later than wheat.
The advantage of the given developmental phase of stock and scion for
grafting, we perceive in the weakneas of differentiation of the tissues
and in theil postembryonic character, The moment shoots have reached the
growth of 1.5 cm, the percentage of actual coalescence drops rapidly. Yet
unstable coalescence of the components fails to ensure a completely normal
cycle of development of the vegetative hybrid.
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6.
(3)
I -.17-. A , k
A
?."::* / 1
......::: 1 / s \ . ' ., e.,"t
= I - .1% -
4
L'
1,21,..i.. 0 ,
.)--:-; .f'y)i-,-
-----.' '4,?' it,
- -4?,1\ i
'??\. i ' 1:1,;(:,: ' st
.s. i ::?, .I.
i\ tE, ? it f - 1.....
1
i
..f.,s ,
; 1.:F:: vI'' ), -::4. I
' 1 ? \ I : j
i ?-i? - 4.1 f ' I'
- t I
;?"0 1 ? t? '4' /
N. '...s. ( 1 : /-',/, ?
1 -
1II _ , .....--
,. _, ?!.:1 -
?
? --?:::?? -- -> ? h -
? k?
1 * i,t,... -?-""- 't ..:..s. 4..... . ,,..4 4, .., -3---:
-.A.
--. -
....., ... ,......? 4.. ' '
.,.:4i.5- ,A. .
r -,-,... r.
e.-.1.,.., r ..., -"
,- 4.--?,....../ 0',.4.-,..1-7..2; 1_,
:-
..,,,,,.._
Fig. 1 Internal structure of one-centimeter shoot
of Gramineae (rye and wheat): (1) - central
embryonic rddicle, (2) - zone of actual
coalescence, (3) - epiblast, (4) .Axillary
(supplementary) bud at point of growth, in
embryo, (5)-cyme, (6) - first leaf, (7) -
coleoptile: [The work ENDOSPERM appears
under (2)]. [Begin p.33]
Trans. 520
Rye stocks and wheat scions have the struature depicted on fig. 1 when
they are in the phase of one-centimeter shoots.
In natural shoots the "zone of division" between the root system and
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(4) Trans. 520
the stem can be easily defined with the naked eye. This zone is very import-
ant ifi the performance of wedge-shaped (klinovidnyi] incisions on stock and
scion. The base of the wedge-shaped incision of the scion, out off in a
manner that it retains the largest possible part of the first leaf including
the tillefing point, must have the lower point of the junction precisely in
this zone. No real coalescence of the components will occur, if the incisions
of stock and scion are either too high above this zone or too low beneath it,
but instead other types of coalescence will take place: a) "dynamia" and b)
coalescence with the formation by the scion of its awn root system. The latter
types do not ensure coalescence between the components whereby the scion
could utilize the root system of the stock of rye during the entire ontogeny,
since,-to begin with, the life span of such hybrids does not exceed 1-2
months. However, in the second type of coalescence, the wheat scion will
utilize only the endosperm of the rye stock and later, having developed its
own root system, it will become self-sufficient as regards nutrition. The
reason for the origin of such two types of component coalescence can be ex-
plained by the tongue-shaped incisions on stock and scion, performed either
too high or too low.
Fig. 2 - Method and angle of incision of
sprouts of stock and scion.
In the case of an elevated incision of tle sprout in the stock, the first
leaf with the tillering point remains in the stock and develops into a
distinctly rye plant-which a certain lapse of time throws off the scion,
feebly grown onto it. In the end result, it is immaterial just how the
cutting-off of the scion had been performed in relation to the "zone of actual
coalescence". Nonetheless, in such a case, the cutting-off of the scion
either too high; normally, or too low does influence the character of the post-
grafting development of the components. It determines still other types of
coalescence and component development to be discussed below.
If the sprout of the stock has been cut too low, its first leaf (with
the tillering point) has been taken out of it, and the low cut scion, that
replaces it has at the base of its tongue-wedged incision its own first leaf
with the tillering point and a particle of root tissue. This particle of
root tissue of the scion develops into a distinct root system of the scion,
regardless of the fact that the coleoptiles of both components coalesce
normally. This explains the above mentioned type of component coalescence
It has the drawback that, in its particular case, the influence exerted by
the stock on the scion is temporary, qnd takes the form of a rye endosperm
being utilized by a wheat scion. The reconstruction of the physiological
basis is poor and loses its effect within the next 1-2 years.
411In this typo of component coalescence, the morphological variability
pertains only to the grain and also disappears rapidly.
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Trans, 520
Fig. 3 - Method of fastening graft com-
ponents with paraffin.
Equal in importance to the zone of actual coalescence of the components,
and to the orientation as to haw their cutting off affects the zone, is the
angle of incision of the stock and scion shoots. lffe have established tho
optimal size of the incision angle at 50-55' (fig. 2).
If this size is reduced, then the base of tho wedge takes the shape
of the point and thus no longer houses the significant part of the first
leaf including the tillering point of scion and stock. If, however, they are
left in their natural position when the incision is made, then they regener-
ate into distinct plants displaying the characteristics of coalescence and
subsequent development of vegetative hybrids,
[Begin p. 34] An angle of incision more blunt (above 55') determines
the impossibility of fastening the components at their junction, since the
coleoptile of the stock is then left at less than 2 mm which is insufficient
for component cementing, so exceedingly important to coalescence. Yet, if
the gaft components were out higher at tho given angle (so their coleoptilos
would roach 3 rum), then the base of the first loaf with the tilloring point
would not be cut out. In such a case component coalescence would proceed
differently with regard to type, and conditions needed for normal grafting
would not be created.
Following the preparation and selection of sprouts of the stock and
scion, they were taken carefully from the box, and each pair was cut
separately. The cutting was performed with a sharp blade of a safety razor
in a place well lighted with either electric or natural lighting, The cut-
off scion sprout was tondiodiately transferred to the place on the stock
specially fitted for it. After this [the scion] must be pressed with one's
finger against tho stock, and the junction of the cuts must be inundated
with melted technical paraffin. From this moment onward it is best that
two people perform the grafting operation; one presses the scion to the
stock, and the other pours a drop of melted pafaffin on the junction of the
cuts with a dissecting [proparoval'noil noodle. The paraffin congeals
rapidly, ocvering the ehtir6 junction of the cuts and firmly fastens the
grafted components (fig. 3).
Following the grafting, the plants aroplanted in separate boxes filled
with soil sterilized with boiling water. The top layer of soil in tho box
is carefully smoothed down and rolled out, so that following the planting
of the grafted plant, its incision junction cov6red by paraffin would be
located on tho surfac0 immediately above ground9 This is necessary for
purposes of obsofvation of post-grafting development and determination of
coalescence typo.
It is precisely the character of coalescence of the grafted components
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(6) Trans. 620
that determines the specificity of subsequent developtent and the degree of
ultimate reciprocal influence between stock and scion.
The coalescence character of graft components is determined not only
by the successful incision of the graft components, the size of tho incision
angle, and the orientation of the wodgc in its relation to the "zone Of
actual Coalescence, but also by the seloction of the stock and scion. In
addition to conformity of age, it is essential that tho coleoptilc diametcrs
and tho internal structure of tho first loaf of the components coincide.
Non-concurrence of the diameters of stock and scion coleoptiles is, as a
rule, connected with the nonconformity of their first leaves and tillering
points in their general contours, as well as in the vascular conductive
system of the leaves.
Nonconformity of the given elements of the stock and scion cause either
complete non-coalescence of the latter, or entirely different types of
coalesconce2 1) partial or ch4morical, 2) false, 3) doublo. Of all tho
given types, it is only "double coalescenco" that ensures a radical ro-
organization of the nature of the scion, while tho two other typos of
coalosconco mentioned determine tho development of abnormal or chimeric
plants with a life span that does not exceed 1.12 months. Later such plants
oither perish or develop a singlo stock -- rye.
As a rosult, of-tho enumerated six types of coalescence of graft
components, [bogin-p. 353 only two typos of coalescence roproduco real
vegetative hybrids. However, oath of these two types of coalescence produce
difforent typos of hybrid plants. Therefore, we cite the characteristics
of all six types of coalescence of graft components of the ryo stock and
the whoat scion, according to the grafting technics and methods doveloped
by us.
Fig. 4 - Schomc of incisions of stock and scion that ensures normal
coalescence between ryo and wheat:
(1-3) - shoot of the scion with tho tillering
point at the baso of tho incision (3);
(4) - aroa of stock incision.
Fig. 5 - Morphological variation of a vegetative hybrid
as compared with the otiginal forms: Viatka
rye and Diurabl' wheat.
From loft to right -- Viatka, Diurabl', and
vogotativo hybrid.
1. Normal or Actual Coalescence Typo.
This typo coalcscenco is obtainod, if the grafting conditions enumerated
by us are obdervod. Anglo of incision of graft components of stock and
scion 50-55?. At the base of tho wedge (at its edge) of the scion must be
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located the first leaf including the tillering point of the scion (fig.4).
The wedge-shaped incision of the scion and stock must be strictly oriented
with respect to the facet of division of the roots from-the stem of the
shoot or "zone of actual coalescence" of the components. The diameters of
graft components must coincide not only in tho colool5tile? but-also within,
tho internal contours of their first loaves (soe fig. 1, 2, 3). The
vascular-conducting system of tho first loaves of both components must also
coincide.
Fulfillment of those conditions establishos complete contact between
tho corrosponding parts of stock and scion, as a result of Which comploto
coalescence of the componefits occurs within 4-5 days, with common metabolism
during the entire ontogeny.
(Bogin p. 36] Identification of tactual coalescenco" typo and conduct
of hybrid generations. Actual coaloscOnco can be dotorminod by tho following
indicators3
Tho incision junction inundated by paraffin rosts ovonly on top
of the soil (tho samo as when the plant was plantod after tho grafting).
On the 4-5th day the first (green) leaf of the scion emerges from tho
colooptilo of the scion and in future, continuos to develop normally. In -
comparison with tho non-grafted stock and scion, growth is strongly rotardod.
The horizontally arranged rootlets of the rye stock continue to grow longer
and to develop; this becomes apparent if they are broken off carofully from
the ground and one of tho rootlots is placed inside a covered stalk whose
diameter at tho opening is larger than tho diameter of the rootlet.
The typo of plant that develops is morphologically and physiologically
essentially different from tho original parent forms, and possesses special
rogularitiOs a'rising in the subsequent dovolopmont of its futuro genora-
tions (fig. 5).
Tho rye stock fails to dovolop a single fast-growing shoot (podgonn]
until tho ond of the hybrid ontogeny. Tho scion forms a bushy growth
which is of tho rye typo, according to the number of tho general as well as
the productive stems.
On reaching maturity tho grain has a clearly intormodiato form and
structure and at times surpasses the parent forms by its size. It has a
doop furrow on the ventral [briushnoi] side and a transverse corrugation on
tho dorsal side.- In the zero generation the straw resembles more tho straw
of the ryo stock. Yet in the following generation (in F1), when the hybrid
plant develops on its own roots during the ontiro ontogony, tho length of
the straw is shorter by 40-50 cm under normal field conditions of development.
Spiko and structuro of tho straw also undergo essential changes in favor of '
tho scion, but the color of tho ripe straw and its girth are). sharply distinct
from the scion as woll as from all other variotics of whoat.
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On the Whble, the grain in F1 undergoesa sharp change for the worse? -
in out experiments the absolute weight dropped from 42 gr. in FQ to 9-10 gr.
in F1 By its external appearance, the grain of this generation remained a
sharp intermediate between the initial parent forms.
The stock of rye transmits the gray-green color of the grain with a
transverse crimp-on its dorsal surface. . In chewing the grain one perceives
the taste of rye. Segregation of new fofms of plants starts with the first!
generation in small amounts (less that 0.01% of the entire generation). The
hybrid deviatives [vyshchepentsy] are of only one type each year; but
morphologically they are completely different in different years. Until
1945 (up to F4) cur vegetative hybrid of 1940/41 had a preponderance of
hybrid derivatives of the wheat and the intermediate type; in 1945, however,
in the fourth generation of the hybfid, there appeared a "tplinter-nybrid
in a considerable quantity (up to 0.01%) from the entire generation. -The
straw of the latter was 45-50 cm taller than that of the hybrid plant:
Morphologically, the spike also was ryoShaped and completely sterile,
The hybrid derivatives of the wheat type produced but two fertile forms
-- in 1942 and in 1946. The first proved of little value economically, but
the hybrid derivative of 1946 displayed a series of economically and
biologicallSr valuable characteristics (high winter hardiness and productivity
in 1946/47). This hybrid derivative propagates.
From 1942 up to the present, the morphology of the spike and of the
plant as a Whole has undergone no essential variation in the given vegeta-
tive hybrid, with the exception-of son o variation in tho length of the
hybrid's straw in certain years. The grain, however, continues to improve
with each successive generation from F1 (1942) until the present time.
Practicing selection since 1945 and cultivation of the hybrid on a
high agricultural plane was responsible for raising successfully the
absolute weight of mass reproduction grain (unsorted)-to an average 37 gr,
with their physical properties being greatly improved. In 1947, when the
basic form of the vegetative hybrid was fully stabilized and the hybrid
derivative was absent from the plants as a whole, there evolved 10 now
derivative forms among the grain. The latter difforod sharply from the
mass reproduction of the hybrid of the 1947 yield, as well as among them-
selves.
They were developed by individual selection from 30,000 typical spikes
of the hybrid which has been selected preliminarily ffom mass reproduction
and had boon threshed spike by spike, in a laboratory. Variation of ono
.form from the other, as well as from the grain of mass reproduction was
distinguished by the following characteristic? [Begin p. 37]
a) the color of the grain varies from rod to reddish gray, thon
to yellow, and finally to white;
b) the structure of the grain varies from puro glassiness to
intermediate [promezhutochnuiu] and finally to a starchy one?
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a
c) the absolute weight of grain varies from 37 gr for the entire
given reproduction of 1947, to 62 gr in some newly isolated forms;
d) the shape of thegrain in hybrid derivatives varied from
short to normal and to long.
The extent of variability in the hybrid derivatives among grain is,
on the whole, no less than the variation occurring in this characteristic
in the individual varieties of diverse ecological origin.
The common characteristic of the grain in those hybrid derivatives and
of the basic hybrid form is merely the polyspermy of grain found in tho
spike (from 52 to 90 grains), while the deep furrow noted oarlier -- a
negative characteristic which so persistently accompaniod tho grain of tho
hybrid in every generation was not found in all hybrid derivatives.
This variety of form in tho grain of the hybrid derivatives of tho
1947 vogotativo hybrid shows clearly how deeply tho horoditary nature of ,
wheat plants is altered by the method of vegetative hybridization. It is
possible that this specificity omorged in forms newly isolated in 1947,
will remain a hereditary [characteristics] in subsequent generations-, and
not only with respect to tho grain, but also in the plant as a whole.
The latter we are ablo to confirm by tho fact that tho seed of tho given
hybrids derivatives sawn for tho purpose of forced propagation under
laboratory conditions produced a variety of sprouts. Under analogous
sooding conditions and environment a-6 a whole, some forms producod sprouts
by 1.5 full days earlier than others. However, in point of character of
plant structure, rate of growth, and development, each form of the hybrids
dorivativos was notably different from mass reproduction and also from one
another.
We aro deeply convinced that the hereditary basis of Graminoae is so
profoundly altered undor the influenco of vegetative hybridization that
in the courso of generations them evolve now forms of plants oven in
instancos when the seeding had boon carriod out in the form of pure lines
(by family). Hence Vegetative hybridization is interesting not only
from the production aspect, but also from the point of view of variety
formation within tho confines of the family Graminoao.
Fig. 6 Process of development of a vegetative hybrid
of the "double coalescence type".
From the left ?rye, from tho right -- wheat.
On fig. 1, left, is seen a gap and regeneration
of th6 colooptilo of tho stock into a distinct
plant.
On tho last figure, right -- colooptiles have
been removed artificially so that tho goneral
base of the hybrid could be soon.
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2. Typo of Double Coaloscenco of Components
Trans. 520
In regard to method and technics, tho characteristic conditions for
tho given typo of coalescence are the following: similar to first varient,
the incision anglo in tho stock is made a little larger (by 10-15 degrees)
than the inciHon angle of the scion, when stock and scion components are
being cut off. The scion, put in the wedge-shaped pocket of tho stock, fails
to fill it out and leaves an empty gap (as large as tho differenco between
tho incision angles in the stock and the scion). The given gap is loft
only on one sido, While the other side of the components' lunction (as woll
as thoir-first leav6s) was arranged to match perfectly at the junction
(fig. 6). [Begin p. 38]
Fig. 7 - Loft -- hybrid of "double ,:ocalesconce";
(1) - wheat; (2) - rye; (3) grain of
rye, (4) - root system of rye.
Right -- hybrid of "single coaloscenco";
(1) - wheat; (2) --grain of rye; (3) -
root system of rye. '
Paraffin coating of the components is hero applied solely on that
sido of the component junction where there is no gap between the colooptilos
of stock and scion., Coalescence occurs on tho sido of component colooptiles
that had boon fitted together on tho opposite side of the gap. Hero coalesce "
also the parts of the first leaves with the tillering points of tho components.
Yot the other sido, tho one with tho gap, which had not coalesced and had not
been inundated with paraffin, produces germination in the colooptilo of tho
rye stock which regenerates into a distinct plant, doveloping side by sido
with the wheat scion within tho limits of one hybrid plant. Double coales-
cence is produced in this mannor: the wheat scion on the stock and root
system of rye grows together "in the zone of actual coalescence", and in
addition maintains a lateral contact with tho stock of rye which had re-
generated into d plant on tho sido with the gap, from one half of the stock
colooptilo (fig. 7).
In their further development, the rye and wheat halves of such a
"chimeribal plant" exert groat influence on one another, as a result of which
a diminishing-of the differences in the color of loaves and in structure of
stem is notod.
The grain is of an intermediate typo, while its color is partially de-
rived from rye, but differs essentially from that of rye. There is an
oven greater deviation of the color of -blies? grains, as well as of their
form and structuro, from the grain of the scion grown on its awn roots.
Frequently the given type of hybrid plants (double coalescence) is
formed also by other moans: a plant of rye stock develops not by regeneration
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from the stock coleoptile in the lower part of the gap, but from the sprout-
ing of the first loaf of the stock, loft in part in thO stock while a shoot
was being cut off at the time of grafting (soo fig. 1). A certain part of
the first loaf ortho stock always remains when incisions are made by our
method. [begin p, 39] However, in component coalosconco of tho first
typo (actual single coalescence), this part of rye, having coaloscod is
depressed by the scion and fails to regenerate into a rye plant. Yet, in
the event of "double coalosconce, tho part of the first leaf loft in the
stock-bogins to sprout on the sido with the gap and develops into a distinct
plant. The latter, however, maintains complete organic relationship and has
common metabolism with the scion through the coalescence at the other side
and at the lower part of the incision.
In the zero generation, the seed of vegotativo hybrids of the first and
of tho given typos of coalescence aro, in all characteristics, ossontially
different from thoir parent-forms (Sandomirka-Alibidum and tho Viatka rye),
as well as from one another. In addition, a higher degree of hybridity is
acquired by the scod of "double coalescence" hybrids which, during their
formation, had boon exposed not only to the influonco of plasma and root
nutrition of therye stock, but also to strong action of the leafy surface
of the stock. If parts of a hybrid enjoyed common metabolism then the
latter [leafy surface] performed the role of a michurin mentor, which was
graphically expressed also in the morphological specificity of the hybrid
seed. The latter showed a strong change in all their basic characteristics,
such as color, form, structure and size -- as compared with the original
parent forms (Sandomirka-Alibidum and the rye Viatka). Besides, it must
be kept in mind that the conditions of their external environment were
analogous during their whole ontogeny.
The regularities for obtaining such vegetative hybrids of double
coalescence (chimerical plants) have been investigated by the Station and
[this information is] made available to us upon request.
3. Dynamic Coalescence Typo
In caso the shoot of a rye stock is cut rather high, so that the base
of its first loaf and the tilloring point remain partially in it, and if the
scion has also retained thoso very parts, then a special typo of coalescence
will occur, if contact of the corresponding parts is onsurod at the junction
of tho graft components.
Coalescence of plants grafted by such a method is stable, but there
occurs a double growth tho nlower growth" normal in coroals and tho un-
usual top growth. In plants grafted by such a mothod, an elevation is
observod in the paraffin bond on tho soil, since tho first loaf of tho
stock, which was not cut off, grows and pushes upward the scion which had
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grown onto it. Yet tho latter, having its own baso in the form of a first
loaf and tilloring point, grown together on those same parts of the stock,
is growing independently at tho bottom, but, with-rospoct to tho general
habits of tho hybrid, an upper growth is obtained. As a result of such a
typo of coalescence, normal dovelopment of the graft components is not
obtained during the period of ontogeny. Tho plant as a whole loses normalcy,
begins to look poor and fragile -- and perishes at tho end of 1-2 months,
or develops into a plant of tho stock typo. Somatimes the death of such
plants is hastened by the appearance of fast-growing shoots of tho stock
which form branches on the 8-10th day of life following germination. All
onorgy of development is concentrated hero at tho'oxpense of tho stem on
which the scion that has coalesced with it, grows.
4. Typo of Coalescence in which tho Scion Forms Its
Own Root System
Conditions undar which the typo of coalescence forms have, in part,
boon outlinod above. It occurs in lowered incisions of the stock and the
scion, when tho scion, having retained a particle of root tissue. in addition
to the first loaf with tho tilloring point, is placed in the lowered incision
of the stock (ovorlapping the sono of actual coalescenco). This particle
of root tissue regenerates into a distinct rootlet sprouting through tho
nook of tho root of the ryo stock and developing into a complete root system.
Tho colooptilo of both graft compononts coalesce well in tho absence of
coalescence of their first leaves; this complicates the identification of
this typo of coalescence, as, in certain respects, the latter begins to bear
a rosomblanco to the first type of "actual coalescence" of tho components
described by us.
Hybrids of those two types of coalescence differ as to the following
characteristics:
a) Besides accuracy of incision as to depth, in
the first typo of "actual coalescence", complete
contact vas established not only with the 6oleep-
tilo, but also with the first leavos, indluding
tillering points of the graft components. On tho
strength of this, common motabolism between the
stock and scion is established immediately after the
grafting, and on the 4-5th day the first loaf of tho
scion emerges from its colOoptilo. The rootlet of
tho stock (rye) also grows. [begin p. 40]
b) In tho latter type, the component coleoptile alone
coalesced, their first loavos with tho tilloring
points did not coalosco and common metabolism was
not obtained. Bocauso of this, tho first loaf of
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Trans. 520
tho scion doos not emorgo from its colooptile
boforo 20-22 days, not bolero its awn rootlet
has formed. In this typo of coalescence, tho
wheat scion merely utilizes the ondosporm of
the rye stock, but later it turns to its awn
root for "wheat" nutrition,-since tho root sys-
tem of rye fails to develop. In essence, the
given typo of coalosconco is analogous with
transplantation of a wheat embryo onto tho endo-
sporm-of a rye grain, with the embryo cut out
of it. Analogous are also tho results obtained,
their effect is negligible, for the influence
exerted by tho stock on the scion is a temporary
manifestation in tho form of a single ondospora
of a rye stock being utilized by a wheat scion.
In vegetative hybrids of rye and wheat derived from "actual coalescence",
tho stock influence of tho scion is not, however, limited to the endosperm
of the stock alono, but is reinforced, in addition, by tho root nutrients
of the stock in tho coalOncenco of tho component cells at tho incision
junction.
5. Typo of "Partial or "Chimurical" Coalescence.
The typo of "partial" or "chimorical" coalescence takes place in an
elevated cut of the shoots of stock and scion and tho subsequent completo
contact of their coleptilos, and the partial contact of their first loaves
including tho tilloring points. This typo of coalescence produces plants
basically of the stock, but there is on the first loaf of the latter a
part of the functioning leaf of tho scion. Tho loaf of the scion coalesces
either firmly (when the vascular conductive systems of both leaves coincide),
or infirmly (whon this coincidence has not boon achicved). In the latter
case, tho loaf of the scion functions but a month and then dies without fall-
ing off of the stock. From components thus coalesced it is graphically
apparent that conjunction (joining), not only of the general contours of tho
first loaves of tho components, but also of the vascular conductive systems
of the first laavos is a requisite for tho normal development of a hybrid
after grafting. This aspect complicatos, to a large extent, grafting
operations on cereals in the given phase of their development, makos them a
subject of accurato, qualified work in which experience, slowly gained, will
be of groat significance.
6. "Falso" Coalescence Type.
False coalescence between scion and stock takes place when the incision
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of a shoot of the stock is elevated and the incisions of the scion vary
(any way at all). The appearance of this type of coalescence can be ex-
plained only by careless handling of the cut-off stock when held between
the fingers during grafting. Pressing of the cut-off shoot slightly between
the fingers causes displacement of the incision level of its first leaf
situated within the coleoptilo though poorly attached to the latter. Hence
contact between the scion and the stock can be established only in a part
of the colooptilos which normally do not coalesce. The absence of contact
between the first leaves of tho components causes growth in tho first leaf
of the stock, while the first loaf of thoscion dries up and curls up within
its own colooptilo without leaving it. Later, the first leaf of tho stock
grows into the colooptile of the scion, penetrates it, and begins to
dovolop into a distinct-plant of the stock which has only ono colooptile of
the scion grown onto it. Then, due to pressure from below, tho curlod up
first loaf of the scion is thrown from its colooptile by tho developing
first loaf of the stock which has grown through the coleoptile of the scion.
The end result of such typo of coalescence is the dovelopmont of a stock
plant.. This typo forms rarely and is hard to identify.
All of those six types of coalescence have boon well invostigated by -
the Station and their thoorotical and practical production has been adaptod.
Five of thorn woro invostigated in 1940-1941, and tho sixth typo -- "'doublo
coalosconco" -- was adaptod in 1946-1947.
The valuable pi'actical results 6btainod aro of scientific and of
production interest. We are unable to dwell hero on tho conditions of
external onvironmont of tho post-grafting period which perform an important
rob o at tho critical point in the existanco of young hybrid plants. In-
fluence exerted on coalescence, on viability after grafting and on tho
subsequent development of hybrids, by such factors as light, hoat, moist
chambers, has been studied by us experimentally.
We shall sot forth thoso problems, together with a literary survey
of achievement and of criticism of vegetative hybridization, in a spocializod
work enhanced with dotailod illustrations.
It appears from our work that the six typos of coalosconco vogotativo
hybrids of wheat and rye, described and investigated by us, aro conditionod
ontirely by the structural specificity of the stem and tho growth of coroal
crops.
Absence of the point of apical [vorkhushochnoi chasti] growth in the
stem and any fullness within it, as it is found in dicotyledonous plants,
is rosponsible for tho complexity and tho flood of groat accuracy in'graft-
ing operations, including tho coordination of its separate olemonts. Regard-
less of this fact, a certain share of tho success in the porformanco of
grafting oporations, oven by an oxperioncod worker, is always subject to
hypothesis. Tho very existanco of the six typos of coaloscenco, which we
have described, and of which only two typos-produco real vegetative hybrids,
bospoaks the justice of the abovo statement.
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CONCLUSIONS
Vegetative hybridization of cereal crops is a potential in deriving
now, highly valuable varieties and to produce now forms through "splingering".
The first rye-wheat vegetative hybrid produced by the station in 1940-
1941 has become the Station's best variety, surpassing in productivity all
varieities in competitive testing in the course of 4 years, and exceeding
the productivity-of control plants [standard] by 32.2% on the average in
3 years of tests. By decision of tho Stato Commission, it now has boon
turned over for tests to tho State Selection network of tho Iaroslav,
Kostroma, Ivanovo and Vladimir Regions.
Tho hybrid has a leaf with a ryo pigment,lligh winter hardiness, is
resistant to sclerosis, develops rapidly after sowing, sprouts early inthe
spring, and has other economically and biologically valuable properties.
In 1948 it was seedod on kolkhozos near the Statioh on an area of 5.5 ha;
in addition, it is being propagated at tho Station.
A technological analysis of hybrid grain of the 1947 yield, carried
out by the laboratory of .grain tochnology at the Institute of Grain Economy
of the Non-Black Soil Bolt, has verified its superior properties as to
milling and broad-baking; in its wet gluton content, tho hybrid exceeded
its scion (Diurabl' variety) by 8%, and Sandomirka Iaroslavskaia
variety in tho Iaroslav Region with rospoct to thoso-propertios -- by W..
In dry gluten it surpasses the Diurabl' variety by 2.1% and Sandomirka by
in addition to other auperior hybrid-grain properties with respect
to milling and broad-baking. Tho Station has produced other vogotativo -
hybrids also: (F0 and F1 between Viatka rye and Sandomirka Iaroslavskaia).
Tho conduct of these hybrids, in the area of reorganization of their biolog-
ical basis and the inheritance of parent properties, is a iepotition of the
behavior history of our vegetative hybrid Diurbl' x Viatka. For instance,
tho zero generation referred to produced tall-growing plants, oqual in
height to ry6 grown right thoro for control purposes, with a modified form
of the spike. Seed of this generation, having been sown for the your 1949,
produced sprouts with violet colooptilos, resembling the color of rye
sprouts, while tho sprouts of the scion Sandomirka, as well as the majority
of wheat varieties have no such pigment whatsoever.
Our eight-year work of vogotativo hybridization of Gramineao enables
us to establish tho same regularities, in tho formation and development of
vegetative hybrids between rye and wheat, that wore so brilliantly doffion:
strated by the groat transformer of tho nature, of fruit perennials, I. V.
Michurin.
Iaroslavskaia Gosselekstantsiia
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[Collection of
Abstracts].
GOLOVTSEV, L. A. (1948)
Vegetative hybridization of Gramineae.
Agrobiologiia, 1948(1):153-157 [USDA 20 Ag822]
Abstract by R. Adelman
Experiments in Gramineae embryo transplantation were begun at the
Ivanovo Selection Station in 1939, but were interrupted due to World War II.
In the spring of 1946 the experiments were renewed with the following
varients:
1) the embryo of Ukrainka winter wheat was transplanted onto
the endosperm of Liutescence 62 spring wheat;
2) embryo of hybrid Pionerka Koga onto the endosperm of Arnautka
Kochina hard spring wheat;
3) embryo of winter rye Tarashchanskaia IV onto the endosperm of
the spring wheat Tsozium III.
The grain of the scion was cut with a razor blade crosswise into two
equal parts. The smaller part was removod with pincers and the larger part
containing the embryo was soaked in cold water from 30-40 hours; later the
saturated semi-fluid endosperm was 6xtracted fully from the grain, leaving
only the embryo with the grain coat. Then an embryo was cut out of a dry
grain of tho stock, and the remaining ondosperm was put into the grain coat
of the scion, so that tho incision could bo arranged to fit the cyme of the
embryo. On the surface tho grain coat was tied with thread so as to join
it firmly with the insortod stock endosperm.
FolloiRing this operation the graftod grains wore soodod in boxes in a
greonhouso.
Author describes their subsequent development carefully, giving time
of year when grains wore seeded and when now characteristics appoarod.
Author stresses tho fact that a change in characteristics can be obtained
only if ombryo transplantation is performed in a vegotating plant. A
conservative characteristic such as profusion of awns will not chango if
the embryo transplantation is carried out in a maturo grain. Tho excess
of awns begins to develop at a certain stage of tho embryonic process and
can be altered only at that particuldr stago of dovelopmont by tho sub-
stitution of unaccustomed conditions.
KHMELEV, B. I. (1950)
Transplantation of the ombryos of Gramineao.
Doklady Akadomii Nauk SSSR, 70(5):909-912. February 11, 1950. (In
Russian) [USDA 511 P444A1
Vscsoiuznyi Nauchno-Isslodovatel'skii Institut Sakharnoi Svokly, Kiev.
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(2) [Collection of
Abstracts]
Abstract by R. Adelman
Concomitantly with oxporimonts in vegetative raprochetent of whoat and
barloy, tho Institute undortook to improve the method of embryo transplanta-
tion. Tho incentive of the experiments was spurred by two factors -- first,
because the method of embryo transplantation has already gained widespread
use in the country's Soloction Stations, and second, bocauso most investiga-
tors havo boon gaining but a low porcontago of success.
The author repeats that the fundamentals underlying the method are
I.ysenko's teachings to tho effect that oxtornal conditions assimilated
by an organism cause a change in its heredity. Author concludes with the
assertion that the grafts accomplished according to tho improved method
(moistoning of embryo and ondosporm) increase not only the percentage of
success, tho rato of growth, and the viability of tho hybrids, but also
tho degree of influence exerted by the stock upon tho scion.. Article
includes table showing rosults.
OMELEV, B. I. (1950)
Influence of sizo and amount of ondosporm-stocks upon the growth
of vegetative hybrids of Graminoae.
Solektsiia i Somonovodstvo, 17(3)21-25. March, 1950. [USDA 61.9 So5]
Abstract by R. Adolman
Procooding from the promise that an appropriate change in the nutrients
foci a young plant produces an adequate change in its hereditary properties,
tho researchers concerned dccidod to uSo tho embryo transplantation method
in their oxporimonts with spring wheat.
Tho method consisted of tho following: tho embryo was cut off of tho
scion kernel with the blade of a safoty razor and with due caro so as not
to injuro the cymo, yot to leave on it as little as possible of the endo-
sperm. Tho separated ombryo was pasted with flour paste onto tho ondo-
sperm stock in dry form on the place whore -its awn embryo had boon removed,
and then it was planted in sand or in soil.
Concurrently with the work on intraspccific and intragoncric grafting
of wheat for the purpose of immediate improvement of regionalized varieties,
the experiments worked also on vegetative raprochement of wheat and barley
for the purpose of their subsequent crossing aimed at obtaining a more
producrive variety of spring wlinat. Bore the problem was a little different
due to the different chemical composition of the barley.
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[Collection of
Abstracts].
In a second experiment, the embryos and endosperms were moistened before
they were grafted, and an effort was made to select large kernels. Article
includes a comparative table.
All-Union Scientific-Research Instituto
for Sugar Boots [Vsesoiuznyi Nauchno-
Issledovatollskii Institut() Sakharnoi
Svokly]
KHNELEV, B. I. (1952)
Vegetative raprochmnont by method of embryo transplantation.
Solektsiia i Somonovodstvo, 19(6):10-15. Nay 1952. (In Russian)
[Library of Congress SB 13 541
Abstract by R. Adolman
In an effort to develop valuable hybrid material, the Ivanovo Experi-
mental Soloction Station conducted investigations as to the feasibility of
improving cross-pollination action by the method of embryo transplantation.
Hard and soft summer whoat served as experimental objects.
They grafted the embryos of one variety of wheat onto the ondosporms
of another variety of wheat. Tho vegetative hybrids obtained wore cross-
bred tith tho variety whose ondosperm had boon used in the capacity of the
stock. The vegetative hybrid was always used as the pistilate plant,
while the common plants of the variety used in the capacity of the stock
served as staminato material. Control plants were crossbred simultaneously
they wero maternal plants regraftod on their own endosperm, or the progeny
of such plants (in the investigation of the first gonoration of vegetative
hybrids).
Doponding on existing conditions, the actual crossing was accomplishod
by one of the following methods: 1) conventional artificial pollination,
2) "8o1f-pol1ination" in large isolators, 3) limited pollination by the
wind.
Tho article has 8 tables showing comparative results.
CONCLUSIONS (in full)
1. Vegetative raprochement by the method of embryo trans-
plantation increases cross-pollination capacity of hard ?
and soft wheats considerable; this incrOaso is observed
in straight and in reciprocal graftings.
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[Collection of
Abstracts].
Z. An increase in the percentage of fertilization is Observed
under various methods of pollindtion and under sharply
different ecological conditions.
3. In one-time graftings the increase in tho percentage of
of fertilization i8 observed in the crossing of Fo and in
tho crossing of Fl. -In tho latter case tho fertilization
percentage is higher.
4. A two-fold grafting raises the percentage of fertilization
higher than a ono-time grafting.
KUPERMA.N., F. IL (1939)
? Grafting technique for Gramineae.
Iarovizatsiia, 1939(5-6)0.47-150. September/December, 1939. (In
Russian) [USDA 20 Ia7]
Abstract by R. Adelman
In early 1939$ when the method of embryo transplantation was still
young, researchers at the Kabardino-Balkarskaia State Selection Station (In
City of Nalichik) found that transplantation of sprouting embryos was in-
expedient; and hence they concentrated on transplantation of embryos in a
dry state. They cut off the embryo, together with the cyme of a dry grain
and transferred it with a flattened needle onto the endosperm of a grain
whose own embryo had been removed. They pasted the eMbryo to its now
endosperm with paste prepared from a puro endosperm of a grain of the same
variety on which tho transplantation was being performed. They sometimes
transplanted two embryos on one endosperm and sometimes one embryo on a
double endosperm, and found that transplantation on-a double endosperm
produced stronger plants than did control specimens. For hybrid components
they utilized "Liutesconce 062" and "Timophoevi" wheats, barley, and
"Sterling" maize.
In tho fdll of tho same year they conducted similar experiments with
winter wheats. They took components from "Zemka", "Stravropol'ka",
"Sarybugda" and "Timpheovi" wheat; for the study of influence exerted by
nutrients of different ondosporms, they used ondosporms from kernels of
barley, couch grass, rico and maize.
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[Colloctioll of
Abstracts].
MEDVEDEVA, G. B. and BAZAVLUK, V. Iu. (1953)
An invcstigation of directs effectsof embryo transplantation upon
form development in hybrids.
Trudy Instituta Gonetiki, no. 20, 1953, pp: 100-105.. [USDA 442.9 P441
Plant Brooding Abstracts, v. 24, no. 3, 1954
Hybrids of the hard whoats Candicans 75/09 and Niatica-Valencia 89
with Kahetinskaia Votvistaia [Branching-Kahctian] gave segregates in respect
of grain characters and branching habit. Some F2 plants bore largo and
vitreous grain of a quality equal or superior to that of tho hard wheat parent.
The percentage of plants with good quality grain increased when embryos of
the hybrids woro transplanted on to the endosporms of tho hard wheat paront.-
This practice, hawevor, roducod tho percontago of plants with branching ears.
Similar grafts of the hybrid endosperms on Kahotinskaia Votvistaia increased
tho number of plants with brO.nching habit, but decreased the number with
grain of tho hard wheat type.
NIKITENKO, G. F. (1949)
News in mothod of ve,I.etative hybridization of Gramineao.
Soloktsiia i Somenovodstvo, 16(11)22=29. November 1949. [USDA 61.9 505]
(Mordovskaia Gossolckstantsiia MASSR).
Abstract by R. Adelman
In an effort to obtain vegetative hybrids of barley that could servo as
initial material in developing a highly productive naked-grain varietSr of
barley, tho researchers solcctod the method of embryo transplantation.
For graft components they sclocted tho best varieties of awnloss and
pronged [furkatnye] bal.ley, including those with naked grain as well as
those with coated ones. In addition, they performed repeated grafts-of tho
second seed generation on original forms -- Pallidum arid Trifurcatum.
Thoy used the following technics. Embryo was cut off of graft components
with a safety razor. Incision was made most carefully and close to cymo.
In such cases tho surface of tho embryo incision is a mealy whito and tho
slightly darkor spot of its own embryo can barely be soon through it.
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[Colloction of
Lbstracts].
Tho ondosporms, rolioved of their embryos, wore notched on tho incision
surface with a safety razor in two reciprocally perpendicular dircction. The
embryos and ondosporms were moistened with water (of room temperature) and
tho transplantation commenced. Embryos wore prossod tightly against the
surface of tho notched and moistened ondosporm incision which, puffed up
from hydro-action, took on the consistency of paste and thus facilitated
tho pasting on of the embryo quickly and fully.
To ensure success in: a) compelling tho embryo to assimilate unaccus-
tomed nutrients from tho stock-endosperm; b) prolonging ondosperm action
upon embryo; c) increasing the amount of plastic substances assimilated by
the embryo from the endosperm-stock in the first stages of its development.
Since the above objectives called for a strengthened reciprocal influence
of the graft components, the researchers planted the grafted grain at a
greater depth (7-Born), and increased artifically the amount of plastic matter
of the stock.
The conclusions drawn are:
1) that vegetative hybrids can bo obtained from barley;
2) that new methods such as "deep planting" are discovered
in the process of experimental work;
3) that directed variations can be obtained in the year of
grafting;
4) that the work concluded has produced valuable forms of
barley hybrids (naked grain, good straw, etc);
5) that now forms bon-ring tho character of new formations
raise certain questions of barley taxonomy and phylogony;
6) that vegetative hybrids constitute good initial material
for selection work.
PISAREV, V. E: and VINOGRADOVA, N. M. (1946)
Intorgonoric hybridization in the family Gramineac.
Trudy Zonaltnogo Institutd Zcrnogo Khoziaistva Nochornozemnoi Polosy,
13:124-133. [Selykhozgiz, Moscow, 19461 [USDA 106 Z721
Abstract by R. Adolman
In 1941, the authors conducted experimonts by hybridizing cultural
whoat, barley and rye with species of Elymus.
The work, commissioned by Tsitsin*, was to be conducted for tho purposo
*Editor of book.
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[Collection of
Abstracts].
of finding methods that would simplify distant hybridization.
Preliminary experiments to hybridize species of Elymus with wheat and
barley proved that the success of such crossing was frequently thwarted by
the difficulties involved in developing an embryo that is already in a state
of germination.
Other failures encountered by the authors and other rosearchhors, includ-
ing Quincke and Thompson, prompted them [tho authors] to look for a mothod of-
raprochement by biochemical characteristics of genera designated for crossing.
The Michurin mothod of preliminary vogetative rapprochmont was not toch-
nically developed for coroal gi-afting, and their experiments with this
practice also ended in failure.
Hence, they docidod to try reciprocal transplantation of embryos of
one species or genera onto the ondosporm of another. Their motivo for those
experiments Ivor? tho rocont invostigations conducted by Schandor, Kholodnyi
and others, on tho role of phytohormonos placed in ondosporms of cereals,
and of theik influence on tho development of the embryo as well as on tho
grown plant.
They worked with soft wheat and rye which included ecotypos producing
a high percentage of hybrid grain, as well as some that do not cross at all.
The article includes a seemingly adequate account of the method employed
and a description of the results.
Cytologically only one hybrid plant (summer wheat VEP2-X E. aronarius)
had boon investigated at the writing of the' present article. Taking into
consideration that the maternal form of Tr. vulgar() had the chromosome
2N 42, and Elymus aronarius 2N 56, the hybrid was found to have 2N 49
(Poddubnaia -Arnoldi).
The authors claim that they were the first to obtain nortal, mature,
first goneration plants fro& a cross between soft wheat and E. aronarius
in the history of selection.
There is further comparative data on the yield of hybrids and control
plants.
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[Collection of
Abstracts]
PISSAREV, V. E. and N. M. VINOGRADOVA (1946)
Intergonoric hybridization in the grass family.
Trudy Zondlinogo Instituta Zornogo Khoziaistva Nochornozcmnoi Polosy
SSSR, vol. 13, 1946, pp: 124-133. [USDA 106 Z721
Biological Abstracts, v. 23, 1949 [G. L. Stobbins, Jr., Abstracter]
The following method was used to facilitato hybridization between
distinctly rolatod spp. of grassos of the tribo Hordoao. Using a safety
razor blade, tho embryo and scutollum, with asthin a layer of ondosporm
as possible, were cut away from dry seeds of one species, thon transferred
to seeds of another apocios, from which tho embryo had boon cut away, and
attached with tho aid of a paste mado from tho flour of soeds bolonging
to the species which sorvod as the "stock" of tho "graft". Plants grown
from such "gfaftod embryos appeared normal except for tho character of
their grains. Plants of spring wheat var. Lutoscons 062 grown from embryos
grafted onto ondosporm Of spring rye, had seeds of a dull gray color, less
vitreous, moro angular, and more deeply furrowod than seeds of control plants
of Lutcscons 062 raised under the same conditions. Chemical analysis showed
highor content of-N, protein, sugar, and ask, and a lower content of starch
than tho controls. Gluton isolated from them was dark brown, as in rye,
due to different content of tyrosin and a different activity of tyrosinaso,
and they contained the carbohydrate trifrucatosan C1030015, found normally
in rye but not in whoat. Those changes arcexplained by the prosonco of
hormones dorived from the ryo ondosporm. Embryos from grafted plants woro
again grafted to the ondospcfm of rye, and the -resulting plants wore
pollinated with the samo var., yielding 25% of seeds as'comparod to 4% in
tha controls. Plants of Lutoscens 062 graftod onto var. Aurora-of rye gave
19.8% of seeds in crossing, as compared to 2.8% in the controls. Spring
wheat 10H68 grown from ombryos graftod to tho ondosporm of Elymus aronarius
gave 5.2% of hybrid seeds with that species as cotpared to 1.9% for tho
controls. Spring whoat from Finland grafted to E. ara1onsi6 gavo 15% of
seed when crossed to that spooics, while controls gave none. Spring wheat
VIR22879 twice grafted onto E. gigantous-gairo 6.2% seed when crossed to
that spocios; once grafted Pra7nts gave 2.3%. Embryos of Elymus grafted
onto whoat ohdosporm gave seedlings in 4-5 days as compared to 14-15 days,for
the controls. Pollination of wheat with pollon from Elyrs grafted onto
wheat ondosporm gave 17% hybrid grains, tho controls 0.4.. Most of thoso
grains had abortive embryos andfailod to gorminato, but one strong hybkid
of Finland spring wheat x E arenarius and one of spring wheat VEP2 x E.
aronarius grow to maturity. In vegetative- characters they resembled
Elymus, but their spikes wore intormediato. They had 49 chromosomes, tho
expected number.
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[Collection of
Abstracts].
PISSAREV, V. E. and VINOGRADOVL, N. M. (1944)
Hybrids between wheat and Elymus.
Cothptes Rendus (Doklady) de l'Academie des Sciences de l'URSS? vol. 45,
no. 3, Oct.'30, 1944, pp: 129-132. [USDA 511 P444]
Plant Breeding Abstracts, v. 15, 1945
A method of grafting the embryo of one component of an intergeneric
cross on the endosperm of the other has been-applied in hybridization ex-
periments with wheat, barley, rye and Elymus. Wheat plants of the variety
Lutescens 62 produced by growth on the=a-parm of spring rye were
morphologically similar to the controls. The grain, however, differed ffom
that of the controls by its dull coloration and less vitreous appearance.
The chemical content of the grain of the grafted wheat approached that Of
the spring rye. An increased percentage of grain was obtained from crosses
between grafted Lutescens 62 and the rye; growth hormones in the rye
endosperm are believed to be the chief factor responsible for its increase.
In hybridization experimonts involving wheat and Elymus tho best results
were obtained in those cases whore both male and female plants of one
component had boon grafted on the endosperm of the othor. The hybrid seed
gave a very low percentage ofgermination on account of the abnormal
development of the embryos. Vigorous plants, described as hybrids, have
however been obtained. These possess loaves resembling those of Elymus
and spikes of mixed typo.
PLOTNIKOV, I. G. (1939)
Grafting technique for Gramineac (a preliminary report).
larovizatsiiu, 1939(3):63-65. Nay/Juno 1939. (In Russian) [USDA 20 1a7]
Editor's comments only:
Plotnikov's article is interesting as a description Of the first
attempts to graft Gramineac for vegetative hybridization. It must be noted,
however, that the technique used cannot as yet be considered as fully
developed. Plotnikov's methods have essential shortcomings. In the first
method (transplantation of embryo) the action of the stock may be insig-
nificant, since the scion feeds on its own root. In the second method
(transplantation of shoot) the shoot may produce its own root and also feed
on its awn root. The third method (grafting in the surface node) is more
promising, but here the work was accomplished with adult plants and con-
sequently, the influence of the stock can assert itself in but a small degree,
Translator's note: It is interesting that editor questioned author's
methods in 1939.
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[Collection of
Abstracts].
RIURIKOV, N. A. (1948)
Summer wheat "Vegetative hybrid No. 1".
Selektsiia i Semenovodstvo, 15(7):36-39. July 1948. (In Russian)
[USDA 61.9 Se5]
Abstract by R. Adelman
Preliminary work was begun in 1939. Seed were soaked in water and just
as soon as, the embryos began to sprout -- to peck through -- they were
transplanted together with the cyme onto the kernel of another wheat
variety. With the operation completed the grain were put in Petri dishes
under glass, but in a faw days the grafts perished.
After an interruption, the work vas renewed in 1944. In the new
experiment the grafted seed were planted-in an open box in very moist soil
and covered with soil of normal moisture.- When the third leaf appeared,
the seed were planted in soil in the open.
The article contains 4 tables indicating positive and negative
characteristics of Hybrid No. 1 and analyzing its yield.
In conclUsion, the author states that the conduct and all indicators
of Hybrid No. 1 confirm the. theoretical principles of Soviet agrObiological
science, and that cereal crops with nositiire economic characteristics can
be obtained by moans of form Modification..
SEKUM, P. F. (1949)
Vogetative hybrids among cereals.
Seloktsiia i Somonovodstvo, 16(2):22-27. February 1949. (In Russian)
[USDA 61.9 Se53
(Moskovskaia Gosseloktstantsiia)
Abstract by R. Adelman
Author indulges to some oxtont in pro Michurin-Lysenko and anti Mendel- -
Morgan propaganda. His experiments are conducted along the lino of Khmelev's.
The embryo of ono form of vihoat was grafted onto the endosperm of another
form. The embryo was pasted onto the alien endosperm with paste made of
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[Collection of
Abstracts].
flour from tho wheat varioty of the stock ondosporm. To prevent the
formation of mold over tho grafted area while the grafts wore germinating,
tho experimenters poured collodium over it. Article includes 5 comparative
tables showing morphological charactoristics of original forms and those
of hybrids, of growth intQnsity and other variations.
Author cites other oxporimontors Brusnotsov, V. F. Illarionov,
Pisarov and Vinogtadova, etc. whose experiments have boon successful.
VATULIA, E. E. and KUCHUMOV, P. V. (1954)
New forms of wheat obtained by the method of vegetative hybridization.
Agrobiologiia, 1954(3):68-72. July/August 1954 (In Russian)
[USDA. 20 Ag822]
Abstract by R. Adolman
Tran6plantation of wheat embryos onto ondosporms of rico in intorgenoric
gtafting. Authors declare that this method is not a now one and they cito
U. P. Aloov, "Experimonts in Changing the Nature of !loat by tho Method
of Vagotative Hybridization", Agrobiologiia, 1948(4):48-56. July/August
1948.
Authors found that success depended on coordination botreon tho
accessibility of nutrients of the rico ondosporm and tho ability of tho
wheat ombryo to assimilate them.
Authors oxporimented on rico grain with sprouts 1 to 10 mm long; they
softened tho kernels by soaking then from 4 to 6 hours, since this procoss
fascilitatod tho removal of the embryo. After cutting off the wheat
embryo with a razor blade, they placed it on the area of tho incision
mado on tho rico ondosporm whon its awn embryo was rethovcd. Then tho
grafted kernels wore planted in vogotativo containers. Of tho 50 grains
planted 38 grow satisfactorily.
Tho variations discovered in tho plants of tho first generation con-
vinced tho authors that roal vegetative hybrids can bo obtained by trans-
planting whoat embryos on rice ondosporms.- The best and most productive
plants were those of the fourth generation.
Tho high productivity of wheat/rice vegetative hybrids indicates that
they may be of selection value.
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[Collection of
astracts].
[The following four articles aro included for general intorost,
although they do not doal especially with cereal vegetative
hybridization].
BORKOVSKAJA, V. A. (1941)
Grafting hybrids and chimeras.
Iarovizatsiia, 1(34)178-83. January/Pebruary 1941. (In Russian)
[USDA 20 Ia71
Conclusions.; [Translated by R. Adelmanl
1. In grafting, the most rapid and sharpest changes take place in
the young cells which have formed in the area where the scion and the
stock grow together, since here their selectivity with regard to the
usual conditions of life can be realized least of all.
2. Old tissues of the scion and the stock which have formed on their
own roots change considerably less and more slowly. But the number of
accumulated changes in a young shoot, develop from such tissue, can be
la rger in "incision" [arez] grafting than in grafting performed by the
usual method, since the removal of all assimilators of the scion-makes a
shoot, which is beginning to regenerate, dependant on the stock.
3. If sharp changes are to be obtained rapidly in a plant exposed to
the influence of the stock, then the method indicated for developing shoots
in the area where the scion and the stock gro* together can be considered,
at the given stage, as the more effective one.
4. Changed plants obtained by graftitg-Solanum nigrum onto a "Shtofert"
tomato "by incision" [so srezom] are hybrids. Their hybrid nature has been
proved by the intermediate character of all characteristics combing the
generic peculiarities of the stock and the scion, and also by the sterility
in self=pollination inherent in the first generation of generically distant
hybrids.
NEDINETS, V. D. (1951)
Properties of maternal organism in progeny of hybrids.
? Selektsiia i Semenovodstvo, 18(7):3-12. July 1951. (In Russian)
[USDA 61.9 5e51
Abstract by R. Adelman
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[Collection of
Abstracts]
Article devoted to reciprocal hybrids of various crops, vegetables and
animals, includes also observations on reciprocal hybrids of cereals.
[p. 7-10]
In examining reciprocal influences that takes place between the emtryo
and the endosperm, the author rejects the idea of "plasmatic heredity".
He subscribes to the theory that the organism develops new properties and
characteristics in accordance with the varying demands which it exacts of
its environment during the different stages of its growth. Considerable
space is given to the influence of the maternal organism and it is asserted
that the properties of the maternal organisms are more pronounced in cases
of free fertilization than in cases of artificial fertilization [pollination].
In conclusion the author justifies the opinion that the reproductive
process -- process of fertilization and development of embryo -- is an
entity [or-union] of two forms of hybridization -- the sexual and the
vegetative.
SEMENENKO, G. I. (1952)
Change in nucleoprotein content in plants upon vegetative hybridization.
Biokhimiia 17(6):655,659. November-December, 1952. (In Russian)
T*USDA Trans 511 [USDA 385 B5231
Conclusions only, by R. Adelman
[Experiments were conducted with tomatoes of the "Nargleb" variety;
eggplant of the "Delikates" Variety; and the seed offspring obtained
from tomato/eggplant grafts].
In vegetative hybridization, substantial quantitative changes occur in
the phosphorus content of mucleoproteins in the young growing organs and
tissues of grafts and hybrid progeny.
In grafting tomatoes onto eggplant and vice versa, the change in the
phosphorus content in nucleoproteins in the leaves of the scion is in-
clined to favor the stock, and in the case of repeated grafting it increases.
In the seed progeny of vegetative hybrids, concurrently with a vigorous
manifestation of heterosis, an increased nucleoprotein content is observed
in the leaves, blossoms, tips of shoots and in the young seedlings of hybrid
plants as compared with the original [plant] forms.
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Trans. 521
[Collection of
Abstracts].
GLUSHCHENKO, I. E. (1950)
Hybridization of Plants by Grafting.
Uspekhi Sovtemennoi Bio1ogii-30(1)a15-48, July/August 1950.
[USDA Trans. 512] [USDA 442.8 Er3.1
(In Russian)
Conclusion, By: R. Adelman
1. Vegetative hybrids show that simple heredity characteristics in
vegetative reproduction can, as Timiriazev assetted, turn into complex
heredity characteristics in sexual reproduction.
2. Experiments in vegetative hybridization show an inconsistency in -
the chromosome theory of heredity the principle of which is cell from cell,
nucleus from nucleus, chromosome from chromosome, gene from gene.
Vegetative hybrids convince one that somatic cells undergoing physio-
logical changes will, in the-end, produce altered sexual cells. There is
no immortal embryonic course. There is a course of qualitative-transforma-
tion of soma capable of forming sexual cells at a certain stage.
3. There is 6 similarity, a parallelism, between sexual and vegeta-
tive hybridization. It is a fact that the second method, as well as the
first, will transmit any characteristics, any property from one component
to the other. These properties become fixed in the seed generations.
4. Side by side with parallelism there exist differences. A typical
feature of vegetative hybrids is the different type of fission of character-
istics. Not only the plants vary in their basic characteristics, but a -
sharply pronounced differentiation occurs [proxodit] within the organist.
In vegetative hybrids the mixed type of heredity is basically intrinsic.
5. A characteristic of graft hybrids is the different form of manifesta.
tion of property dominance. Plants with recessive characteristics often
produce offspring with dominant characteristics.
6. Crossbreeding carried out between vegetative hybrids and parents
with graft components and recessive characteristics demontrate graphically
that dominant characteristics may appear in the offspring. This indicates
that separate characteristics in hybrids are in a latent state and, under
appropriate conditions, develop subsequently in the offspring.
7. On the change in external morphological characteristics, a profound
reorganization of order takes place in hybrids, the character of cell sti-uct-
ure changes in particular, plastids of various types appear in the cells.
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Trans 521
[Collectioh of
Abstracts].
8. Biochemical investigations show that the majority of quantitative
and qualitative indicators-of the second graft component -- the mentor --
appear in the seed progeny.
9. Frequently the absence of visible changes observed in the year
of grafting does not denote an ab6ence of specific qualitative changes in
the generative cells of the plant. Hence it follows that the progeny of
grafted plants must always bo investigated, even though no phenomenon of
variation in characteristics had been observed in the year of grafting.
10. It would be most primitive to think that as a result of grafting
one always has to look for grafts of the stock in the offspring of the
scion, or vica versa, to discover properties, characteristics of the scion
in the offspring of fruit of the stock. Living [substance] constitute a
process of development, and every biological process does not know of
straight, direct changes. They are realized only in a long chain of
transformations.
Besides the phenomena of straight hybridism (presence of the character-
istics of both parent components in the offspi-ing), vegetative hybrids in-
clude also the phenomena of new formations, i.e. the creation of new
characteristics frequently not inherent in either one of the graft components.
A pre-requisite for this is extreme instability, inconstancy of form,
frequently observed in intraspecific and, most often, in distant [Inter-
generic] graftings.
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Golovtsov, L. A.
Grafting Cereal Plants.
Agrobiologiia, 1952(5)180-90.
September/October 1952. 20 Ag822
(In Russian)
GRAFTING CEREAL PLANTS
Ar
Trans. 522
(In part)
By:
R. Adolman
The author believes that the Michurin method of vegetative hybridiza-
tion would aid in the production of valuable varieties of grain crops. He
states, however, that this method is not as yet being used in cereal selec-
tion because the hollow stalk of these plants presents a problem different
from that of woody plants.
' To investigate this problem, the author and his colleagues were
guided by the hypothesis, that the plant changes under the influence of
?
external evironment.
They based their experiments on two principles: first, modification
df organic nutrients during embryonic and postembryonic development of plant
by embryo transplalitation, and second, coalescing embryo particles of various
species of cereals.
In their seven-year investigations they developed three methods of
vegetative hybridization:
1) transplantation of embryo on a mature grain;
2) transplantation of an embryo particle;
3) method of the double mentor.
In the last five years investigators experimented with 92 different
combinations and carried out over 50 thousand grafts, but they describe only
the more characteristic samples.
(1) Transplantation of embryo on a mature grain.
Frequently the embryo is cut off with a razor blade and is pasted
onto endosperm with some adhesive substance without providing for contact
between scion and stock or for conditions for feeding of embryo on the endo-
sperm. Such a graft deteriorates under influence of moisture and soon
perishes.
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Trans. 522
(In part)
By:
R. Adelman
The author's method provides for cutting a dry grain with a razor
blade on the tufted side and for soaking it in water at room temperature for
24-30 hours, until the endosperm is in a semiliquid state, then it i8 re-
moved from the grain coat leaving only the embryo and the empty coat. Then
they take a dry grain on whose endosperm the graft is to be performed, tut
off the embryo and put the dry endosperm under the coat of the scion
(embryo) with the incision adjoining the cyme of the grafted embryo. They
use no adhesive substances. The place of junction of the embryo and endo-
sperm is covered tightly with the grain coat (of the scion). This close
relationship between embryo and endosperm is not disturbed until the endo-
sperm is fully utilized. The binding link is the gain coat itself, and
not an adhesive substance; coalescence reaches 100%. Seeding can be
carried out in a vessel containing moist soil. At optimal temperature,
sprouts appear in 5-8 days.
There follows a description of the development of combinations of
different varieties, of the influence exerted by the endosperm on the
process of vernalization of winter wheat, and of the modification of '
varietal characteristics under the influence of grafting on endoaperms.
(2) Method of transplanting particle of embryo (Mentor).
Experimenters use dry grain of both graft components and on the grain
of one of these, they partially cut off, with a razor blade, the coat sur-
rounding the embryo. Then they cut out of the naked embryo laterally, a
piece about 0.3-0.5mm thick on the demarcation line between the initial
pedicle and rootlet. The depth of the incision is extended to the very
endosperm; pedicle and rootlet of the embryo are completely separated.
Following the above, a particle of the embryo of living tissue (the
mentor) is cut from the dry grain of the other graft component in the form
of a wedge-shaped blade and is placed in the incision made in the first
grain. The grain coat is placed over the embryo and its incision is
slightly flooded with paraffin. The embryo operated on represents a complex
formation in which the rootlet and pedicle belong to a plant of one variety,
and the meristem tissue connecting them -- to a plant of another variety.
The grafts are seeded in boxes with moist soil covered with a 3-4
centimeter layer of well rinsed and calcined sand. The grafted grains are
in the layer of sand close to the surface of the soil, so that the germinat-
ing rootlets enter the soil directly.
At optimal temperature and moisture, the embryo begins to grow; the
mentor particle begins to germinate at the same time and grows into the
embryo. Coalescence occurs in 5-6 days. Exact determination of the spot
in the embryo on which the mentor particle is to be transplanted is of
utmost importance. A slight deviation may result in failure of the trans-
plantation.
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(3) Method of double mentor.
Trans. 522
(14 part)
Brt
R. Adelman
Vegetative hybridization of cereals by the method of the Double Mentor
represents the combination of the two methods described -- transplantation
of embryo onto an endodperm and insertion of a particle of the stock embryo
into it [first embryo].
Here, on one hand, the endosperm serves as a mentor, and on the other
hand, the embryo particle serves as a mentor. The grafting technics do
not change, but the experimenters perform two operations on one plant:
first, they graft an embryo on the endosperm, and then they insert an
embryo particle of that form on the endosperm of Ilhich the grafting is
performed.
By the double mentor method, they accomplished grafts betIVeen winter
wheat and maize, with Kharfkovskaia 23 maize serving as mentor. They
soaked the wheat grain in water of room temperature, extracted the softened
endosperm from the grain after 30 hours of soaking, and inserted the dry
maize endosperm in its place. Since the grain of maize is much larger
than the grain of wheat, and is differently shaped, the maize endospefm
was first rounded off and then inserted under the grain coat of-wheat.
Having completed the first operation, they began the second one. Using
the method described earlier, they inserted the blade from the maize
embryo into the sprouting wheat embryo. By this method they made one
hundred grafts, using three control variants.
CONCLUSIONS
1. Vegetative hybridization can be realized by embryo trans-
plantation and by coalescing embryo particles of plants of different
varieties, species and even genera and families.
2. By grafting an embryo onto an endosperm and, concomitantly,
inserting into it [embryo] mentor particles, it is possible to produce in a
plant deep hereditary variation.
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(1) Trans' 523
(In full)
By:
R. Adelman
Kozarehko, VL
Some results from Vegetative
hybridization of cereals.
Zemledelie, 1954(2)0.14-115. (1954)
20 Z44
(In Russian)
SO hE RESULTS FROM VEGETATIVE HYBRIDIZATION
OF CEREALS
In the Trunov Region, situated in the arid steppe of Stavropol'
Territory, spring wheat is often subject to incomplete respiration [zapal]
and, therefore, produces a low grain yield.
Proceeding from the principles of the agrobiological science of
Michurin, I set myself the task of improving the properties of grain in
winter wheat and the developing of a drought-resistant variety of spring
wheat equal to winter wheat in grain yield, by means of vegetative hybridiza-
tion.
For this purpose, I selected in 19460 the following parent pairs:
"Melianopus 69" x "Boroshilovskaia"; "Voroshilovskaia" x "Melianopus 69";
"Kooperatorka" x "Melianopus 69"; "Melianopus 69" x "Kooperatorka";
"Melianopus 69" x "Coach grass-wheat hybrid", and a series of others.
We accomplished vegetative hybridization by joining the endosperms
of two grains with germinating rootlets and sprouting pedicals of equal size.
We soaked the seed in water, making a point Of first soaking those
seeds that had been sprouting for a longer-period. We cut the sprouted
grain with a razor blade across the embryo. Thus we obtained two halves of ,
a grain: on one-half the rootlet, and on the other pedicel; we left more of
the endosperm on the half which we selected as scion or stock. Then, we
joined the halves of grain with the rootlet (stock) and those with the
pedicel (scion) together, and planted them immediately in moist soil ?at
a 3-5 cm depth. -Watering was carried out before a shoot appeared above
the soil surface. Further cafe was the same as that given to grain crops,
seeded under field conditions. By this method of growing together,
coalescence achieved was 80-90%. .
In cases where spring wheat served as the stock and winter wheat as
the scion, all plants tillered well toward winter; overwintered completely,
plant 8 developed better and ripened a few days earlier than their parent
forms.
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Trans. 523
(In full)
Br:
R. Adelman
lhenever winter wheat was used as stock and spring wheat as scion, the
plants branched out strongly in the fall, but by the time spring came there
were left only separate specimen, and most of the plants that survived had
the characteristics of vernality.
It must be noted that the characteristics which appeared in the first
generation were retained in all ensuing generations. Of a series of hybrids
which I obtained, only four vegetative hybrids of winter wheat and three of
spring wheat presented considerable interest. These hybrids have a number
of advantages over the regionalized wheat varieties in the Trunov Region.
The root system of vegetative hybrids is developed considerably better
than that in the "Voroshilovskaia" and "Odesskaia 3" varieties seeded
alongside of the hybrids. Besides, the 1952 crop of the vegetative hybrids
of winter wheat (No. 1, 3, 4, 5) sown for four years without being treated
[with fungicides] were not infected by loose smut or hard smut, while-the
varieties of "Voroshilovskaia" and Odesskaia 3" were affected by them.
According to morphological characteristics and grain properties, the
vegetative hybrids-inherited the characteristics of the scion as well as
those of the stock. Thus, for instance, the glassiness of the grain of
111 vegetative hybrids of winter wheat is considerably higher than that of the
winter wheat varieties of "Voroshilovskaia" and ultooperatorka", but less
than that of the spring wheat variety "Melianopus 69".
Vegetative hybrids surpass the regionalized varieties of "Voroshilov-
skaia" and "Odesskaia 3" in the yield of grain also, calculating in centners
of 8-9 per ha.
0
Table 1.
Variety
Yield by years
(in c/ha)
Voroshilovskaia--
Adesskaia 3
Vegetative hybrid No;
No;
No;
11.
No.
1
3
4
5
44444
1951
1952
917
18:0
18;7
14;3
15,0
1
1
1
1
1
160
19;0
199
22;4
22;6
25.5
[Fig.] Root system of vegetative hybrid No. 3 (right)
and of winter wheat variety Voroshilovskaia (left).
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(3) Trans. 523
(In full)
Bks
R. Adelman
In the fall of-1952? the-four forms of vegetative hybrids obtained were
seeded on lots of 0.5 ha each. The regionalized variety "Odesskaia 3", was
seeded on a similar lot in control capacity.
Hybrid No. 1 yielded 14 centners, No. 3 -- 16 c., No. 4 -- 14.5 c.;
while the yield of the "Odesskaia 3" variety came to 12 centners per ha.
In the fall of 1953 the vegetative hybrids were seeded on a 15 ha lot for
the purpose of utilizing them as initial material in developing new
varieties, since they have the tendency to shatter in the sente of seg-
regation] strongly during the first three years after grafting.
To improve the hybrids obtained in 1949, we carried out selection
of plants annually. In 1953 we derived from each hybrid two more varieties
which we seeded on separate test lots. Thus; I obtained six varieties with
distinguishing morphological characteristics.
In the Stavropol' Territory,-spring wheat which produces a low yield
is being replaced by gray cereals.
In connection with this I tackled the task of developing a variety of
spring wheat by vegetative hybridization that would not only be as good
as gray cereals, but just as good as winter crops.
For this purpose I selected in 1949, the following parent pairs for
vegetative hybridization.
I selected the "Dzhugara" variety of sorghum for the stock, and for
the scion -- the spring wheat variety of "Palestinka" which yields up to
25-30 centners of grain per ha in the damp regions of the Stavropol'
Territory. The vegetative hybrid of spring wheat possessed the character-
istics of both parents. It had a large spike, a stem as hard as that in
sorghum, completely filled within with parenchyma and twice as thick as a
wheat stem. The root system was also considerably more vigorous than that
in ordinary wheat and, in addition,-atirial roots appeared on many plants
in the form of a supplementary tier. The grain resembled that of spring
wheat in shape and color, but was considerably larger. In the absence
of rain in the spring and early summer of 1950, the hybrid of spring wheat
produced normal grain, while spring wheat developed no spikes whatever that
year. We select annually the best plants from the young crop of the hybrid
of spring wheat.
tin 19520 this hybrid reached the height of 1.5 - 1.6 m.
The absolute weight of a grain of the 1951 and 1952 yield was 59 gr.,
the might of grain obtained from a single spike 2.5 gr. In 1952, spike
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Trans. 523
(In full)
13S1-:
R. Adelman
formation and maturing of grain was noted in the spring wheat hybrid 10-12
days earlier than in spring wheat of the "Melianopus 69" variety.
In the second year of seeding, the plants of the vegetative hybrid of
spring wheat yielded shatter-derivatives and produced fundamentally three
varieties distinct as to color of spikes number of awns and fullness of
grain. Heterosis of the'vegotative mass of plants appeared in a higher
degree from year to year. In 1953, this hybrid of spring wheat was seeded
on a lot of 120-square m, and it conducted itself the same as in the
preceding years.
Judging this hybrid by its conduct, it can be expected that it *ill
provc fully adequate for cultivation of spring wheat in arid regions.
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Trans. A-524
(In full)
By:
R. Adelman
Ustinov, A. A.
Novoe v Izuchenii Gallovoi Nematody -
Heterodera marioni (Cornu. 1879) Goodey
[New aspects in the Study of the 1ogt74clot
Nematode - Heterodera marioni (Cornu, ji7p,) 092ay]
Trudy Zoologicheskogo Instituta, Izdatel'stto Akademii
Nauk SSSR? vol. 9, no. 2, 1951, pp: 405-446. 410.9 L543
(Biologicheskii Institut Khartkovskogo Gosudarstvennogo
Universiteta)
51 - WORKS OF THE ZOOLOGICAL INSTITUTE OF THE ACADEMY OF
SCUNCES, USSR - IK
The root-knot nematode belongs to the most important parasites of agri-
cultural plants and represents a widely distributed farm. Its glebal distri-
bution is conditioned by purely ecological factors: the nematode now has in-
vaded every area where conditions are favorable for it. The thermophilic
capacity of the root-knot nematode indicates that its place of origin was in
a very warm zone, but more in a subtropic than tropical one, since it survives
winter colds easily and is fodnd in the ground in White Russia and near Moscow,
While tropical organisms are usually very susceptible to low temperatures. In
colder areas it is found mostly in greenhouses. The absence of anabiotic
capacity in the larva of the root-knot nematode indicates a probability that
its sourde was a moderately humid place, without sharp fluctuation in soil
moisture.
There is no available material that would aid-in establishing the root-
knot nematode's place of origin with some exactness'. The first mention of its
prevalence was made in 1855 when the nematode was fbuhd in a greenhouse in,
England, but almost simultaneously (1857), it was identified in Florida, where
it already had been recognized as a dangerous parasite of agricultural plants;
and in 1885 the nematode was found in Java on sugarcane -- an old world plant.
Its occurrence on Vrild flora also fails to indicate the indigenity of the
root-knot nematode. It possesses a very effective, although passive, method
for spreading and is very common in Uncultivated soil.
Reports of occurrence of the root-knot nematode in virgin soil have been
received from America and Africa, and the author [of present article] has found
it in Ukraine in the 'Donets river area in meadows and pine forests where the soil
has never been tilled. The list of-hosts of the root-knot nematode includes
plants from every tart of the world.
Morphology 'and Development'
A more detailed work on the morphology of the root-knot nematode is that of
Nagakura (1930); it contains a detailed description of the male and the mature
female, the other stages of development, however, are treated poorly. Brief de-
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2 Trans. A-5,24
scriptions of the nematode can be found in old monographs: Mueller (1884),
Stone and Smith (1898) and Bessey (1911). The illustrations of Stone and Smith
were used by I. N. Filip'ev in his abstract on nematodes (Filip'ev, 1934) and in
"Handbook on Zoology" (1937). Like thbse of other authors, these illustrations
are extremely simplified and schematic. The embryonic development of the root-
knot nematode had not been investigated; the postembryonic development had been
interpreted incorrectly until recently, and its clarification has begun only
since the appearance of the work of Christie,and Cobb (1941).
The root-knot nematode is a heterotropic [geterotopnyi] parasite. The -
larva which leaves the egg lives for some time in the soil, even though it does
not develop there -- this is the preparasitic to invasion larva ,capable of
penetrating into the root of a plant. Within the root the larva becomes an
internal tissue parasite of the plant and loses its motility. In males motility
is restored after they have accomplished metamorphosis, ?mhile individudls which
are transformed into females remain immotile to the end of their lives. The
function of the organism as awhile changes during ontogeny of the root-knot
nematode: the basic function of preparasitic lar ve is to find and to invade a
host, that of parasitic larva -- intensive feeding and growth, and the function
of adult males and females -- propagation. Naturally, the structure of the
different phases depends on their functional characteristics.
Fig. 1. - Map of global distribution of root-
knot-nematode compiled from available
data.
Embryonic and Post-Embryonic Develop-
ment of the Root-Knot Nematode.
Sexual Cells. TheeggS-of the root-knot nematode usually are kidney-shaped,
slightly concave on one side. The dimensions of eggs fluctuate even in those
taken from one ovary [ooteka], consequently laid by one female for a short period
of time. The average dimensions of eggs were 92 x 40 unit* (all dimensions
given are as per author's meaSurements). The eggs are supplied with two mem-
branes: external and internal. The first is isolated [vydeliaetsia] by the
sexual organs of the female and is a smooth homogerneus shell, about 1 unit*
thick; the internal or yolk membrane is isolated by the ovicell itself and rep-
resents a very thin film. The ovicell contains a large cyst nucleus, and in
its protoplasm are found many yolk orbits of a yellow-brown coloring. Eggs are
laid by the female in the ovary before they begin to granulate or in the very
first stages of granulation. Only on a visible depression of the females do the
eggs pass through embryonic development within the sex tubes, since the process
of hatching eggs is difficult in these ffemales. Spermatozoons represent
spherical cells 7-8 units*in diameter.
Fig. 2 - Eggs of the root-knot nematode in
different stages of embryonic development.
Invasion larva is visible in last phase,
lying in egg under cover of skin of the
larva of the first phase; from the right -
invasion larva upon leaving egg.
* -
Macron
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3 Trans. A-524
DEVELOPNENT At first, granulation of the eggs of the root-knot nematode
is uffiform; root-knot nematodes show no noticeable difference in the size of
the first two blastomeres (described in the case of other nematodes). The
cleavage of both blastomeres occurs simultaneously, so that a four cell phase
forms immediately. Further, some cells granulate more rapidly, others more
slowly,-,and in the_phase of 10 or.more blastomeres, it is possible to distinguish
the larger and darker cells of the endoderm from the lighter and smaller of the
ectoderm. Further, the difference between these embryonic layers increases still
more. As the embryo lengthens, there occurs clearly at the anterior part of its
body, a separation of the light section-composed of ectodermal cells -- StOmadeum,
from which the digestive tract deVelops. Further elongation leads to tho forma-
tion of a worm-shaped embryo (fig. 2), coiled several times within the egg. The
final formation of the larva constitutes the longest stage of embryonic develop-
ment. The invasion larva which has developed and assumed form, molts and then
crawl from. the egg through the opening which it has made at one of its narrow
ends. In contrast to description pertaining-to the best nematode, no trophic
stimulants are needed for the exit of larvaes Larvae effect their exit also in
perfectly pure water. Eggs incapable of surviving are found very rarely, usually
not more than 1-3% of their total number in the ovary.
POST'EMBRYONIC DEVELOPMENT', Much confusion has been observed in the
designation and interpretation of the, developmental stages of larvae of the root-
knot nematode, [Begin p. 4081 hence it is necessary to dwell on this question
in greater detail.. To begin with, there is no unanimous. opinion as to-what
stage of development invasion larva canatitutes when it leaves the egg. It is
usually referred to as'the first, but I. N. Filip'ev calls it the second, be-
cause he believes that the larva, which molts within the egg, constitutes the
first stage. Having penetrated the ro4 and turned to parasitism, the invasion
larvae change markedly, but they do not Melt and still constitute the second
stage, if the larva within the egg is considered as the first stage. This is
the only prolonged phase during which the larva subsists and feeds. Nonetheless,
the changes which larvae undergo in their transition to a parasitic mode of
life arc so great, that within the root it is looked upon as the ensuing phase
developed froth invasion larvae by means of molting, and sometimes, even as two
larval phases.
Having reached full development and taken on the shape typical for the
root-knot nematode, with a pintail [ship] at the end of its body, the larva
separates from the old cuticle, which constitutes its Second molting, if the
molting within the egg is also taken into consideration. The larva lying with-
in the old skin will constitute the third stage; it molts anew during. transition
to the fourth stage which, under the dover of two skins, molts for the last time
and is transformed into an adult form.' To illustrate the facts stated above,
we will present them in the form of a table showing how different authors have
designated the atages of development of the root-knot' nematode.
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4 Trans. A-524
Table 1.
Stages of development of the root-knot nematode as presented by different authors
Actual Stages of
development of
the root-knot noma-
todo _
.
Designation of
thee stages by
Filiptev (1934)
"Handbook on
Zoology" (1937)
-
Designation by
othpr native
authors (Ustinov,
1939, Kobakhidze,
1941, Brodskii
and
,
Designation by
Nagakura (1930)
and Goodey,(1933)
Zemlianskaia
1946,
1. Larva in egg First stage
prior to its
Considered as
embryonic stage
molting for ?
invasion
of development
2. Invasion (pre- Second stage
parasitic) larvR
First stage
First stage
3. Same larva, but Third stage
in state of
parasitism
(parasitic
larva)
Second stage
Second Stage
.
i
4. Same, at end Fourth stage
of its develop-
,
Third stage
Second stage,
the final
? ment: strongly.
expanded and
with a pin-tail
.
5. Two moltings
within skin of
larva with tail
(third and
fourth larval
stages).
not noted

book
Not noted
Third stage (onlY
one molting noted)
.
*See errata at end of
Begin p. 409
COMPARATIVE OUTLINE OF STRUCTURE OF ROOT-KNOT NEMATODE AT
DIFFERENT STAGES OF ITS DEVELOPMENT
BODY FORM. Tho structure of the root-knot nematode changes strongly in the
process of ontogenotic development. Preparasitic Ixrva escapes the modifying
influence of parasitism and retains the structural characteristics typical of
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5
0 the group. Its body is cylindrically elongated, the anterior end is blunt, the
posterior end narrows dawn gradually and, at the very end, forms a thin conic,
rounded tail. Tho transition to parasitism causes a rapid modification of body
form. Thanks to the strong expansion Of the intestines, the body of parasitic
larva grows even more in width (fig.3). Considerable thickening occurs only.
in that part of it where the intestines rest, while the tail neither thickens
nor grows bigger, but turns into a tenon [ship] at the posterior end of the
strongly expanded body. The growth of larvae in length during their tranSition
to paratism is negligible. The bodies of young' females are bottle-shaped. Tho
posterior part is an elongated oval, the neck is slightly sot off from tho body.
Females in the state of puberty are pear-shaped; the anterior part of their body
is drawn out onto the neck, the posterior -- oval or spheric, widely rounded.
The body of the male is elongated.
?
?
Fig. 3.
Second larval phase of the root-knot
nematode after transition to parasitism.
Initial phase of parasitic larva; body is
expanded; bulbus Ibul'bus] has increased,
motility gone
BODY DIMENSIONS. The sizes of pre-parasitio larvae fluctuate from 380
to 460 units* in length and from 13 to 17 units* in width. The tail is about
65-70 units* long, L:6?. it approximately constitutes 15% of the entire body
length (alpha == 23). Measurements of 43 parasitic larvae in the final stage,
taken from various plants; have shown that their body lengths fluctUate between
380 units* and 650 units*. The average length of the body including the pin-
tail; measured 465 units* of which 40-45 units* were taken up by the pin-tail;
the average body diameter -- 86 units*. The size of mature females is more
inconsistant. The length of propagating females is rarely*t-..less.than 500-600
units* or more than 1000 units*. The females, however, stretch sometimes very
much and reach a length of 1500 and even 1700 units*, but there are also dwarf
females measuring 450 units* in length. The width of females is rarely less
than 400 units* or more than 800 units*, but there are also strongly bulging
females which are 1000 units* wide, and in dwarf females whose body length is
315 units*. The body length of females depends on the elongation of the neck,
which sometimes, is drawn out extremely long. The length of males varied from
1 to 2 mm, but e*treme sizes were rare and prevalent measurements ran between
1300-1700 units*. Measureing 53 males from various populations, the avei.age
length proved 1515 units*, and the average body diameter -- 30-35 units*. At
the anterior end the body narrows down to 11-13 units* at the base of the cap,
yet the tip of the cap is only 7 units* in diameter. At the posterior end
the body contracts less; at the base of the spicules its width reaches 20-22
units*. Looking at the tail from above or from beneath, there is no further
contraction, but in profile the tail narrows down to 8 units*. [Begin p.. 410]
The tail iS short; the opening of the cloacas is situated 8-15 units* forward
of its tip.
From the example cited, it is apparent that a tremendeus increase in the
dimensions of the nematode body occurs during ontogeny (fig. 4). Taking the
average measurements of various stages of development and assuming, for approxi
mat6 calculations, that preparasitic larva is a cylinder 0.42 mm high and 0.016mm
*micron
** See errata at end of book
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in diameter, and using the formula for calculation Of cylindrical dimensions,
we will find that its dimensions equal 0.000084 =3. In parasitism this phase
oxpands rapidly, and the average size of the ultimate phase of parasitic larva
can, approximately, be accepted as a Cylinder 0.43 tm-in height and 0.086 mm
in diameter; its dimensions will be 0.002496 =3s-i.e., as compared to prepara-
sitic [larva], they will increase almost 30 times. Such an important increase in
dimensions without molting in the same larval phase, is a.raro phenomenon and,
therefore, until of late, the parasitic phase was considered as the no?t larva
of the nematode. After metamorphosis the female bulges out still more. Taking,
approximately, the average size of tho female for a sphere 0.7 mm in diametov,
and using the Coftesponding formula, we will find its dimensions to equal
0.179594 mm, i.o., compared to preparasitie hrva, thod imensions of the female
increase more than 2000 times (2133 times). Such an enormous increase in tho
animal mass during ontogeny is typical of those parasites which settle down in
the early stages of development, since it is to their advantage that each individ-
ual settling down have a minimal biomass, but that their quantity be maximal,
and, in this direction, the evolution of parasites is strengthened by natural
selection. The male, developed in larval skins and being smaller than the ul-
timA phase of parasitic larva, ceases growing. Assuming; approximately, that
tho male is a very elongated cylinder; 1.5 mm long and 0.032 mm in diameter, we
will find that its dimensions equal 0.001205 mm3, i.e. 140-150 times smaller
than the dimensions of tho female.
Fig. 4. - Ultimate phase of parasitic larva of
tho root-knot nematode before the be-
ginning of molting. Body very expand-
ed, bulbus increased oven more, tail
reduoed to pin at posterior end of
body.
COVERING. Externally the body of the root-knot nematode is covered with
cuticle composed of two distinct layers: the e?ternal ono, transparent, divided
by furrows into rings, and a thick internal one. Beneath the cuticle is located
the hypodermis, adjoined by longitudinal muscular cells most of which degener-
ate in the parasitic phase. The thiokness of tho cuticle increases with age:
in larva it reaches d thickness of 1.3 units*, in tho male tho cuticle gains
in thickness up to 2.5 units*, and in the tAil-rogion -- up to 4.5 units*, but
in females -- up to 8-10 units*, and in the bulbus region -- up to 13-16 units*.
In proparasitic and parasitic larva thickness of the cuticle was identical [Begin
p.411] and Nagakura's assertion that it grows thinner in parasitic larva is
erroneous. Tho cuticle of males with distinct rings is 2.5 unitswide; annula-
_
tion in young females is also apparent over their entire bodies, but it is .
particularly distinct in the anterior part, and poorly visible at the posterior
In adult females the annuldtion is noticeable only in the neck region; the width
of their rings reaches 2-2..5 units*. Annulation in preparasitic larva can be
discovered only under groat microscopic enlargement; the rings are very indis-
tinct and very narrow, about 1 unit* in width.
HEAD-CAP. Tho head end of tho nematode wears a so-called head-cap, named
also head of tho neamtode, which is divided from the body by a ring furrow. The
head-cap is well developed in tho motile phase -- of preparasitic larva and the
male, and it-grows much shorter, though still remains, in parasitic immotile
* micron
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phases. In invasion larva the height of the cap reaches 4-5-units*; in parasit-
ic larva it is reduced to 3 units*, and in the female to 2-2.5 units*.In the
male, ,the height of the head-cap reaches 5-7 units*: A relationship between the
dimensions of the head-cap and motility is definite. Sometimes the cap is don-
sidered as an auxiliary organ of movement, assuming that it acts as a wedge. It
is possible that the reduction of the cap in parasitic phases is connected with
the reduction of the receptacles [Retseptory] found on it, and with the general
reduction of the body.
Fig. 5. - Young female of the root-knot nematode
soon after finishing the last molting.
Body not yet much bulged out; transparent
cuticle permits examination of internal
organs.
NUTRITIVE ORGANS. (fig. 5). They begin with the spear or stilet of the
mouth, situated in the cavity of the mouth. The spear represents a thickened,
hardened distal [distaltnaia] part of the chitinized internal tube of the
esophagus and is pianctured in the center by a canal passing directly into the
canal of this tube. The esophagus which proceeds from the spear is composed
of three sections: the anterior, the narrowest; the center or bulbue, of a more
or less powerful muscular formation, and the posterior -- glandular. Behind the
esophagus rests the center intestine which passes into the sharply narrowed
posterior intestine opening up as an anal aperture. The anterior in-take sec-
tions of the feeding organs are weak in invasion larva, the intestine is narrow;
upon transition to parasitism, the strength and d imens ions of all nutritive
organs increase sharply, but in adult females they grow most powerful. In males,
the nutrition organs again grow weaker. [Begin p. 412] In all phases of de-
velopment, the root-knot nematode can feed on liquid food only, absorbing it
through the Stilet with the aid of bulbus contraction. The esophagus glands
eliminate secretion which passes through the stilet and conditions extra-intes-
tinal digestion, disolving the food before it entei's the intestine; this method
of digesting food is highly prevalent in nematodes.
THE STILET. The stilet is long and thin in prepardsitic larva; its length
varies from 12 to 14 units*, its width is about 1 unit*: Basal protuberances
located at the base of the spear form an expansion of 2.6-3 units*. The spear
has parallel little walls and is pointed only at the top. The stilet remains, in
parasitic larva as it was in preparasitic larva, since no molting occurs dabiate/
the old spear could be cast off together with the cuticle and a new one could
form. Nagakura's claim, rewritten by Filip'ev (1934), that the spear in larva
within the root grows mailer is clearly erroneous. In females the spear be-
comes considerably more powerful; its length reaches 16-17 units*; the posterior
port of the' spear is aylindrical? the anterior part conic; basal protuberances
stand out at the base of the spear in the form Of three bulges. The thickness
of the cylindrical part of the spear measures 2:5 units*, but at the base, in
the area of basal protuberances -- 5-5.5 units*. The measurements of spears in
females cited by different authors are very distinct; this prompted Filip'ev
to presume that different races of the root-knot nematode are prevalent in dif-
ferent countries. The figures of some author, however, are clearly orroneous
and their measurements are,refutsdr by further investigations of nematodes in
the same country. The length of the spear in female ?nematodes was in Sukhumi
*micron
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the same as in the USA, according to later and more accurate measurements accom-
plished there. The stilet in males is longef., from 18 to 23 units*. Form and
width of the spear is the same as in females. Already Strubell (1888) described
the muscle protractors and retractors moving the spear in the beet nematode.
According to his description, the first were affixed to the basal protuberances,
and the retractors to the centcf of the spear. The cuticle was the Punctum
fixum for both pairs of muscles. The description is repeated for the root-knot
nematode as well. According to our observations, there are only protractor
muscles, and the-pulling of the spear inside the body is its passive return to
its former place.
ESOPHAGUS. The esophagus section in preparasitic lai.va represents a narrow
tube, 6-7 units* in width and about 50 units* in length. The esophagus is hard
to notice in this phase, with the exception of the internal, sharply protruding
cuticle tube, about 1 unit* in diameter. During transition to parasitism, the
anterior section of the esophagus expands considerably and changes into a fairly
powerful cone-shaped formation, very easily noticed on living object. In the
final parasitic phase, the width of the esophagus in front of the bulbus reaches
15-20 units*, thus growing almost 3 times as thick as in preparasitic larva. In
females the esophagus grows still more powerful, but retains the same, generally
cone-shaped form prevalent in parasitic lara. The internal cuticle tube of the
esophagus expands very strongly, reaching 2.5-3 units* in diameter. In males
(fig. 6), the anterior section of the esophagus becomes cylindrical and is not
easily noticed, the same as observed in preparasitic larva. Only the internal
chitinized esophagus lumen, 2 units* in diameter, is clearly visible.
The 6ther section of the esophagua, or bulbus, represents an oval, solid
formation. Radical muscles are situated in the wall of the bulbus, and in its
center is located an oval cavity with thick cuticularized walls -- hen. [Begin
p. 413] The lumen shrinks due to the reduction of i-adical muscles, but expands ,
anew later, and thus causes the absorption movement: During ontogeny of the
root-knot nematode, the bulbus, too, undergoes form and dimensional changes. In
preparasitic larva, the bulbus is poorly developed and less clearly pronounced,
Its measurements are about 13-15 units* in-length and 10 units* in width. The
measurements Of the lumen are 5 x 4 units*.
Fig. 6. - Male of the root-knot
nematode.
airing transition to parasitism, the bulbus rapidly develops into a vigorous
and easily noticeable organ, reaching,-in the final phase of parasitic larve,
30-32 units* in length and 28-30 unitst in width, increasing several times in
size compared to its size in preparasitic larva. The bulbus wall grows strong,
and the lumen retains the same measurements, 5 x 4 units* -- which-it had in
preparasitic larva. The bulbus attains the greatest vigor in females. :Its
Measurements vary from 39-52 units* in length and 34-50 units* in width. Measure-
ments occurring more frequently are: 47 x 44 units*. The measurements of the
lumen fluctuate between 18 x 11 and 16 x 13 unit*. In males the bulbus is poor-
ly developed and again becomes poorly noticeable. Its measurements vary-from
23 x 18 to 28 x 16; the measurements of the lumen are about 8 x 6 units*.
*
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[Begin p. 414] The third section of the esophagus contains three mono-
nuclear eeophagal glands; a dorsal one and two subventral ones bordering on the
intestine. The dorsal gland opens into the esophagal lumen =- at the base of
the stilet, and the subVentral one -- into the bulbus cavity. In preparasitic
larvae these glands are narrow; in parasitic larvae they increase indimensions
at the expense of growth in width: The Glands in females expand even more,
while those in males shrink again.
INTESTINE. In preparasitic larva the middle intestine is narrow, cylindri-
cal, about 10 units* in diameter, turning into a very thin posterior intestine
near the posterior end of the body. The anal opening is situated at a 65-70
units* distance from the posterior end of the body. The middle intestine is
covered with drops of fat the size of which reaches 8 units* in diameter. Dur-
ing transition to parasitism the dimensions of the middle intestine increase
very strongly. The figures cited for increases in larval dimensions character-
ize this growth, since the expansion of larva is due mainly to the growth of
its intestine. The posterior intestine retains its previous form of an oblique-
ly set narrow tube passing from the center intestine to the body surface. As
a result of reduction of the tail, the anal opening is situated closer to the
posterior end of the body. In females, the middle intestine expands even more;
in young females it takes up the .larger part of the body, and in mature ones
the intestine is pushed aside by growing sexual tubes. The-drops of fat cover-
ing the intestine are large, about 20-50 units* in diameter. The middle intes-
tine-narrows down sharply and turns into the short and narrow posterior intes-
tine. The anus rests at the posterior end of the body, not entirely terminal,
but somewhat to the side; it can be seen clearly in young females, but becomes
less visible in mature ones. There is no thickening of the cuticle around the
anus, and due to' this characteristic it can be easily distinguished from the
sexual aperture. The middle intestine of males looks like a straight cylindri-
cal tube taking up the larger part. of the worbil:.body (at a 33 units* body diame-.
ter, the intestinal diameter equals 23 units*). The intestine ends in a wide
cloaca which receives also the sexual organs of the male. The cloaca opens out
to the surface as a transverse aperture near the-posterior of the body, at a
'distance of 8-15 units* from thetip of the tail.
ORGANS OF ELIMINATION. In all nematodes, the function of the elimination
organs is performed by the cutaneous glands, the so called cervical glands.
Root-knot nematodes, the same as other representatives Of the family Anguillul-
inidae, are endowed with the tylenchoid [tilenkhoidnyi] type of elimination
organs characterized by the fact that the excretory canal and the gland cell
are not paired, and rest in one lateral field of the hypodermis, in the root-
knot nematode on the left side. The elimination organs have not been investigat-
ed, hence their variation during ontogeny can be indicated only with respect to
the arrangement of the excretory pore. In preparasitic larva the excretory pore
is easily visible; it opens in a level with the glandular part of the esophagus
at 80-85 units* distance from the tip of the head. In parasitic larva, the ex-
cretory pore opens outwards, also on level with the glandular part of the esopha-
_
gus. In young females the pore is situated on the level of the bulbus which
obviously,, is connected with the strong increase in the dimensions of the latter.
In mature females the Excretory pore is barely visible, yet in males it can be
seen very well, and. it opens, the same as in larvae, on the level of the gland-
ular part of the esophagus, at a distance of 140-165 units* from the. head-cap.
*micron
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SEXUAL ORGANS. The sexual embryo forms in nematodes very early, but be-
comes clearly visible only in parasitic larva, In this phase the embryo repre-
sents an accumulation of spheric sexual cells which under the microscope looks
like a flakelet [plastinkal. [Begin p. 415] The sex of larvae is undetermin-
able and the formation of sexual tubes from the embryo can be detected only dur-
ing metamorphosis.
The sexual organs oftho female are composed of two sexual tubes (fig. 7),
which merge at the posterior end into one short tube -- the vagina: the latter
opens outwards as a sexual aperture -- the vulva. The sexual tubes are long;
the biggest one of-the tube Which I measured reached 6 mm, with the length of
the female being 0.85 mm, i.e. the tube was-6-7 times, and consequently, both
tubes were 12-14 times longer than the body. In comparison with,the relatively
short sexual tubes of motile phytohelminths from the genus Anguillulina and
Aphelenchoides, their length in the root-knot nematode must be recognized as
very great. The ovary composes about 6% of the entire length of the sexual
tube, and is its thinnest and brightest part. The globular ovogonia seen in
the ovary are about 18 units* in diameter. Without any define borderline, the
ovary turns into a wider oviduct [iaitsevod] which takes up about 55% of the
tubes length. It contains ovocytes; at the beginning their form is globular,
further on the ovocytes stretch across the tube and take on a wedge-like form,
later they gradually stretch lengthwise along the-tube and assume the oval
shape of an egg in the distal part of the oviduct. The oviduct flaws into an
irregularly globular chamber termed semenreceptacle [semepriemnik]; it is 140-
150 units* in diameter.
411 Fig. 7. Sexual tube of an egg-hatching
female of the root-knot nematode.
?
The precise function of this organ is the separation of the shell surround-
ing the ovicells coming from the oviduct. The semenreceptacle (?) is followed
by the uterus which takes up about-37% of the-length of the sexual tube; in it
are lying the fully developed eggs: [Begin p. 416] The uteri of both tubes
merge into a short unpaired vagina. The sexual aperture is closed and surround-
ed by oval spindles [valiki] of the cuticle, forming around the vulva a sexual
swelling [bugorok], which is plainly visible in a young female, but disappears
in a mature one. The glandular cells, the secretion of-which forms an oothoca
during egg hatching, also open into the sexual aperture.
In males, the sexual tube takes up about 607. of the total length of the
body; its length reaches 800 units* in a male whose dimensions measure 1300
units*. Tho proximal part of the tube is coffiposed of the male sexual gland
[semennik] with small cells of spormatogonia. Expanding gradually, the sexual
gland of the male turns into a semen-duct [semeprovod], which contains larger
wedge-shaped spermatocytes. The distal part of the tubo is formed by the
semen-emitting [semeizvergatel'nyi] canal, the length of which constitutes 45%
of the entire length of the sexual tube; it contains spermatozoons lying very
loosely and freely in the lower part of the canal. The semen-emitting canal
opens into the cloaca; it contains two spicules 29-36.unit6* in length, on the
average about 33 units*, equipped with special musculature. The width of the
spicule base is 4-5 units*; they narrow down at the tip. The bases of the
spicules are separated from edch other by a space of about 7 units*, but their
tips, but their tips converge. Tho cloaca also contains .a "rod-ferrule"
micron
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[rulek] (gubernaculum) representing a thickening of the cuticle wall of the
cloaca and serving as a slide and a support for the spicules; it is-usually
8 units* long, and its width, at the widest part, measures 2 units*. Tail wingg
(bursa) are absent, but the cuticle surrounding the tail grows 'somewhat thicker.
MUSCULATURE AND MOTILITY. Skin masculaturo is poorly developed. The in-
vasion larvae are slow-moving and sluggish animals;-movements are accomplished
by bending-their long body. They lose motility rapidly during transition to
parasitism. Parasitic larvae whose form has changed very little at the time
they are extracted from the root, lie completely motionless withoutbending
their body or trying to ci.awl, although it seems that the tail which has been
retained would allo* this. Only the anterior part of the body remains motile
in parasitic Larvae. Larvae removed from galls continually make probing move-
ments with their head-end by bending it slowly in various directions and b3)-
pricking rapidly with the stilot. Females, too, are completely deprived of
their motility. In females removed from galls the following movements were ob-
served: (a) slow bending of the head-end of tho body in different-directions;
(b) pricking with the stilot; (c) rhythmic pulsation of thelpulbus. The Males
are capable of crawling slowly, similar to invasion larvae. Structural charact-
eristics of the nervous system and sense organs in the root-knot nematode have
not been investigated.
Tho cavity of the body stretches from the base of the head-cap to tho
posterior end.
METAMORPHOSIS, DEVELOPMENT OF MALES AND FEMALES. The first melting of
parasitic larvae occurs in such a manner that tho cuticle which has become
separated is-not cat off, but covers, in the form of a hood, the new larvae
lying inside. At first the separation of the body from tho cuticle is observ-
ed at the posterior end: the pintail is emptied, tho posterior end of tho larvae
moves forward and coils; somewhat later, it can be-noticed that the body un-
loosens from the cuticle in the anterior part also. The spear is cast off to-
gether with the cuticloi its contour is plainly visible at tho anterior of the
skin of parasitic larva. The apertures which form between the larval body and
tho old cuticle arc at first very narrow, but later on the body usually Moves-
far away-from the skin of parasitic larvae at the anterior and posterior ends.
[Begin p. 417] The molting described is the first following the exit from an
egg and the second one, if the molting within the egg is taken into considera-
tion, and tho larva formed within the skin will constitute the second phase
upon its exit-from the egg, and the third phase, if tho larva within the egg
is considered. We shall call it the third larva.. The development of the third
larva is not entirely completed. Tho cuticular formations intrinsic in tho
origin of the digestive system dre not developed: there is no spear nor internal
cuticular tubo Of tho esophagus. This is a rapidly passing larval phase, in the
words of Christie and Cobb "more theoretical than actual". The third larva
molts quickly and is transformed into the fourth larval phase. The fourth
larva lies beneath the cover of two cast-off skins: the thick external skin of
parasitic larvae equipped with a pinlike tail and the very thin internal skin
of the third larva . The latter can be seen only from the anterior and poster-
ior ends, but usually not laterally, clinging closely to the external skin or
to the IL-rval body. The further development of males and females is strongly
distinct.
* micron
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IIIFig. 8. - Young female of the root-knot
nematode, developed inside of
three larval skins.
?
DEVELOPMENT OF TBE FEMALE. In tho dovolopmont of tho female, the fourth
larva is just as theoretical as the third. I have boon unable to find any.
difference in their structure: This is a, similarly, not fully developed
organism without cuticularized parts in the anterior section of the o8ophagus.
The bulbus is retained in both larvae and can be seen very distinctly. The
glandUlar section of the esophagus has grown shorter, hence the intestine which
isdirk, thanks to the drops of fat, clings to the bulbus. Sexual tubes develop
from the soxual embryo, but sexual outlets (vagina, secual aperture) have not -
yet formed and it is impossible to distinguiSh any organs bohind the intestine.
The anterior end of both larvao is very motile: tho larvae twists constantly in
diffefont directions. It is possible that theso.movomonts help to cast off the
skins. The molting of the fourth larva leads to tho fofmation of tho young fe-
male which lios under tho cover of throe membranes (fig. 8). The thin skin of
the fourth larva resting at the very interior can be discerned only neaf tho
head-end of the female, sometimes also at tho posterior end of its body. On its
exterior is found a similarly thin skin of the third larva, and still more ex-
ternally -- a thick cuticle of the parasitic larva of the second age. When the'
young fothales development draws to an end, all her inherent organs are visible.
[Begin p. 418] Tho anterior section of tho esophagus is sharply distinct from
the corresponding parts of parasitic larva due to the strong development of the
buticularized parts. SoXual outlets are clearly visible; the vulva opens out
on a sexual protuberance. On the background of the intestine are observed
brighter sexual tubes. Naturally, the developed female will be someWhat smaller
than the final phase of parasitic larva, in Ahose skin it developed. Its
further development leads-to a strong increase in its dimensions, in length
and particularly in width. 'The dimensions of her sexual tubes increase most
during her growth.
_
Fig. 9. - Larva of the root-knot nematode male
of the foufth phase in the process of
elongation. The larva develops within
two larval skins: the thick external
skin of the larva in its second age and
the thin internal of the third age.
Fig. 10. Young male developed within three
larval skins, before coming out.
DEVELOPMENT OF THE MALE. In the case of male development, the fourth larva
which rests under the cover of two skins constitutes the prolonged development- .
al stage during which modification of form and structure of the animal occurs.
At the beginning of this phase, the body of the fourth larva is just as thick
and short as that of the third, but at the end of this phase it turns intoa
long and thin male body. The elongation of the body proceeds rapidly. The
cast-off skin of parasitic larva is soon too short for the larva growing in
length, and its posterior end coils. Later, the loop forms and, finally, the
body grows so long that it can remain in the skin only by coiling in it several
times. Such "cocoons" from the skins of parasitic larva of the second age,
harboring within coiled larva of the foutth age; or males, is very characterist-
ic of the ontogenesis of Heterodera (fig. 10**). Development of the male from
** See errata at end of book
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the elongated fourth larva takes considerably longer than the time required for
the elongation of this larva. The duration depends on temperature.and is indi-
cated on p. 423. 'Finally, the last molting occurs and the skin of the fourth
larva is cast off. It can be seen on the narrow ends of the worm where the
skin appears on the body of the male. The young male now rests, as does the fe?
male, under the protection of three cutucularized skins: The external thick
cuticle of parasitic larva of the second phase, the intermediate thin and wide
cuticle of larva of thethird phase, and the internal narrow and thin cuticle
of the fourth phase. The development of the male is completed fully during
metamorphosis: he reaches his maximum growth, and the sei organs -- full de-
velopment. In males, still resting within the skins (fig. 10), spermatoids are
clearly visible in the semen-emitting canal, hence it can be assumed that males
are immediately capable of fertilization. The males which have departed from
the skins ("cocoons") cease growing and, obviously, consume nourishment, In any
case, the possibility of their consuming nourishment is just as limited as in
preparasitic larva.
MORPHO-PHYSIOLOGICAL ADAPTATION OF THE ROOT-KNOT NEMATODE TO PARASITISM. As a
result of the simple structure of the root-knot nematode, parasitism does not
produce in it such important morphological variations as in animals constructed
more intricately, nonetheless, these variations are greater than in. the majority
of other plant parasitic nematodes: The shape of the body and of the organs
-- sexual and digestive systems, i.e those more complicated in the nematode
,change markedly. Yet, no specially big modification occurred even in those
systems, and a second set of organs, acquired in the process of parasitism, did
not develop. Skin musculature suffei. a reduction and as a-result of this the
parasitic phases lose their motility. The post-embryonic development of the
root-knot nematode produced greater changes. Even though it retained all four
larval phases, inherent in nematodes, they were separated from each other by
molting, but only the second one which at first, as the preparasitic one, pro-
ceeds in the soil; and later as the parasitic one in the plant, developed fully
and consumed food.
Thus oligamerism [oligomerizatsiia] of the life-cycle (Beklemishev, 1945)
which, by the way, is one expressed in a complete shedding of larval phases, but
in their rudimentation; and is inherent in many parasites, was not observed in
the root-knot nematode. Hence the duration of the existence of non-reproducing
phases is shortened and the prolonged phase is only that of the mature female.
A comparison of the post-embryonic development of the root-knot nematode with
the development Of other species of the genus Heterodera shows that it is the
most specialized. This, together with the considerable morphological adaptation
to parasitism, indicates that the root-knot-nematode had turned to parasitism
within the plant a long time ago. [Begin p. 420]
ECOLOGY OF THE ROOT-KNOT NEMATODE. ECOLOGY OF THE PREPARASITIC LARVA.
CONDUCT IN SOIL. The rate obtained in attempts to determine the speed of
movement of preparasitic larvae of the root-knot nematode under microscope was
extremely low -- fractions QC millimeters per minute. Because of their slow
and sluggish movements, larvae of the root-knot nematode are sharply distinct
from soil-saprozoons of the genus Rhabditis and from predators of the genus
Mononchus. We were unable to induce the larvae to quick movements of escape
even by using shaft), plainly unpleasant irritants: injecting a heated needle
or a drop of acid. With the aid of glass capillaries filled with various fluids
and buriod in sand with nematode larvae, we eucceeded in establishing a positive
reaction of the larvae to weak organic acids. Stereotropism of larvae -- their
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inclination to COMB in contact with hard Objects is also easily observed. Their
reaction to light could not be detected. After even distribution of larvae in a
Petri dish filled with agar and half of it darkened with black paper, no coh-
centration of larvae in any part of the dish was observed, not even after they
had been exposed for a whole day. There was no attempt to escape even from
direct sunlight, although the sun killed larvae oven in the autumn When heat
was slight. Literary data concerning the speed of movement of invasion larvae
under field conditions aro available only in the works of tho author (Ustinov;
1934, 1939): in the course of a season, the larvae moved over a distance of
but 25 cm. Larvae, however, penetrate depth very considerably: In Turkmen,
nematode galls are-found on the roots of mulberry trees at a depth of 120 cm
(Blinovskii, 1945). The larvae of some other parasitic nematodes also display
poor motility. For instance, the larvae of the wheat nematode are capable of
infecting plants seeded no farther than 30 cm from the infected grain (Leukel,
1924). Of representatives of the genus Hotdrodera, the more motile larvae wore
those of tho beet nematode (Naunacke, 1077-
-
Retcnsion of Invasion capacity. Consumption of food by root-knot nematode
larvae begins only when they establish c6ntact with the root, up to that time
they live on the supply of embryonic fat. It is known that soil infested by
the nematode remains in a state of black fallow [chornogo para] having been in-
vaded in the course of many months, but the variations undergone by larvae dur-
ing starvation were not investigated. Nor are there any indications that soil
can serve as a supplementary source of nourishment for larvae; yet, by analogy
with other soil animals, this is entirely possible. Invasion larvae aro
capable of remaining alive for a period of two months oven in watch glasses
with a small athount of water: they lived at our place from mid April until
mid June, 1946. Cinsiddrable variations have been Observed in the external
form of starving larvae. The drops of fat covering the intestine in young
larvae are used up as they grow older, and bright striae, deprived of the fat,
appear on the intestine. At first the intestine shows transverse striae, later
the striae grow oven bigger, but the dark ones grow thinner and, finally, the
intestine turns entirely light, with but a small stock of fatty drops. Tho
amount of fat characterizes the 'age of larvae and their capacity for invasion
which decreases gradually. Experiments in introducing starving larvae into
plants have shown that upon loss of fats on half of the intestine, the invasion
capacity of larvae wanes, and in the case of a still greater loss (for instance.,
on 751, of tho intestine surface), they lose their invasion capacity completely.
[Begin p. 421]
Having penetrated the root after long starvation, the larvae, ,first of all,
ropldnish their normal supply of fats, and this strongly retards their develop-
ment. Having opened, in July, roots infested with such larvae eight days after
invasion, we found that tho larvae had not expanded and still had the for:m and
structure of-invasion larvae, but with a dark intestine covered all over with
drops of fat. The degree of larva fatness can be ascertained with conventional
symbols which make it possible to estimate the age and invasidn capacity of a
given larvae population, which, in turn, would permit a more thorough study of
their ecology. Does, however, the loss of fat occur in soil just as rapidly
as in water on a watch glass? Do larvae find supplementary sources of mourish-
ment in soil?
To clarify this, the suthor [of the present article] conducted several
experiments with a quantity of proparasitic larvae in humus enriched soil and in
thoroughly rinsed sea sand. Larvae in amounts as equal as possible were placed
in both media; this was accomplished first by obtaining a uniform suspension
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from freshly derived larvae in Wator and, then by moasuring this suspension
with a pipette in equal amounts. After a certain period of starvation, both
media containing the Larvae were washed off into flower pots containing soil,
cloared of holminths by drying it in the sun, and indicator plants. In tho
first experiment larvae were starving for a month -- from July 9 through August
9, 1946, at high temperature, no lower than 25?C. No larva capable of invasion
was loft in the sand, but tho humic soil caused slight infestation: 8 galls
appeared on a plant. In tho second experiment tho larvae wore starving for
25 days: from July 29 through August 23, 1946; at the maximal temperature of
that year, reaching 35-36?C in tho laboratory. Sand produced no infestation
this time either,. but tho humus enriched soil produced a very slight invasion:
only 4 galls woro found on a plant. Tho galls which dovoloped wore very small,
and tho larvae found upon thoir olioning were strongly rotardod in growth. In
the third experimont tho larvae starved for 22 days; from August 27 through
September 16, 1946, at a temperature lower than in tho preceding experiment.
Larvae capable of invasion survived in tho sand, but in a considerably smaller
quantity than in humic soil: 100 galls developed on plants in the latter, but
only 5 in the sand.
Apparently, all experiments produced similar results: in humus enriched
soil the larva? retain their invasion capacity longer than in sand. This means
that they find in humus sources of sustenance lacking in rinsed soil. An oxamin-
ation of starving larvae under the microscope-showed that the loss of fat pro-
ceded more rapidly in sand than in humic soil.
Life cycle and fertility of fomalos. Conditions for Development of Males.
When galls are opened in tho summer, no dead fomalos are found and this loads
to the conclusion that they live through the entire season. When plants wilt
in the fall, females survive their hosts to a considerable extent: in Sukhumi
living females are easily found in December and oven in January in the roots of
tobacco plants withered long ago. By the same token, females-continuo to live
long in branches of roots if they are kept in a moist chamber. (Bogin p. 422]
When root tissue docomposos to such an extent that it falls apart when rinsed in
water, the females continue to livo and the rhythmic protrusion of-their spear ?
and contraction of their bulbus can be observed under a microscope. There is no
doubt that the capacity to live long on the supplios of non-vegetating and oven
dead plants, developed in mature females, is most favorable for the dissemination
of tho species with seed material.
The potential life cyclo of the root-knot nematode apparently is very great,
and its propagation may continuo very long. Indications as to the life cycle of
females aro available only in the works of the author (Ustinov, 1934, 1939). In
observing the hatdhing of eggs in boxes-with collapsible walls, it was noted that
the female hatched 1814 eggs in 70 days. But these magnitudes can in no way be
considered as ultimate, since the energy of the female was interrupted by the
dying of tho root. Tho average lifo cycle of tho female is measured by the
summer months; a few Of them overwintor and, under Sukhumi conditions, resume
hatching eggs, but perish afterward.
Parthenogenesis is inhoront in the root-knot nematodo; reproduction without
0 males is regular and normal.
A larbo number of males-appears only if living Conditions are worsened,
primarily thoso of nutrition. In Sukhumi their number changes during the season
-
as follows: in the spring males can be found developing from ovorwintorod larva?.
Only in rare oases do they finish metamorphosis successfully, in most cases they
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perish during the last molting. In the summer males aro very rare, but in
August and September, when invasion increases greatly; and it is clear that tho
plant is being thwarted, the number of males enlarges. Small end galls are
frequently inhabited by males only, and the rate of their development hero is
several times highor than that of females in a similarly small gall. Tho appear-
ance of a large number of males under worsened living conditions has boon estab-
lished also in the case of the boot nematode (Molz, 1920).
Climatic factors. Influence of temperature and humidiIL Temperature.
The influence of temperature on the root-knot nematode is so obvious that it was
noted even by its first investigators. At first particular significance was
ascribed to severe winters and the freezing of soil considered-lethal for the
nematode (Neal, 1889); (Stone and Smith, 1898); (Bessey, 1911). Furthor obser-
vations, however, showed that high propagation of the root-knot nematode is
interrupted not by winter temperatures, but by summer temperatures, namely by
insufficiency of the sum total of temperatures and the short duration of the
period of effective temperatures. Low winter temperatures produce no lethal
action on larvae in the soil, and the root-knot nematode overwinters success-
fully under field conditions in the Don Basin [Donbas], in White Russia-and
near Moscow, and also in the countries of northern Europe and in Canada. Exper-
imental work investigating the influence of temperature on tho rate of develop-
ment of the root-knot nematode was conducted by Taylor (1933). She established
heat totals required for the development of the root-knot nematode and to
measure heat she used the conventional heat unit equal to temPorature by 1?C
above 10?C, effective for an hour; degree hours above 10?C. Ten degrees con-
stitute the threshold of nematode development, while the retarding influence of
high temperatures is observed above 28?C; but temperatui-es which substantially
depress adult females begin only with 36.5?C. [Begin p. 423]. The authorls
observdtions in Sukhumi resulted in numbers very near to thoso obtained by
.Taylor. In our experiments, the development of a nematode within the root
required a miniumu of 7478 heat units from the time it invades the plant until
it begins to hatch eggs and, according to Taylor, this requires from 7000 to
8500 units. Between the different periods of development, this sum total of
temperatures was distributed as follows (table 2)..
TABLE- 2,
Total temperatures consumed in development of different phases of the root-knot
nematode within the plant
1 Duration ofthverage Tempera-
Period of Development 1 developmentiture (in ?C)
1 in daysAbsolute
!
Total temperatures in
degree hours above 1Q?C
t %
1. Development of i
larvae prior to molt-
it* ' 8
_
25.9
3070
41
2. Period of molt-
ing (metamorphosis) 5
-
25.9
1912
26
3. Development of
female from immatur-
ity to egg hatching
8
..
23.0
2496
33
21 ?
7478
100
Tayler, too, indicates very close figures: the development of larvae into
a female which formed within the skin of parasitic larvae required an average
of 3000 heat units, or 41% of the average temperature total needed for the en-
tire postembryonic development; to complete molting -- up to the appearance of
the young female -- 1800 heat units, or 25%, and for the development of the
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female from a young one to initial egg hatching -- 2400 heat units or 34%. Our
observations show that the totals of temperatures consumed in the development
of the root-knot nematode in Sukhumi are the same as those in California. Ac-
cording to some observations, more heat is used up in the development of the
nematode in somewhat hotter Florida, here the threshold of its development is
not 100 but 12?, (Townsend, 1937). It would be most interesting to conduct
similar observations in the northern regions of root-knot nematode distribution.
In the development of males the elongation of the fourth larva proceeds
very rapidly: it was observed that the larva transformed overnight from a wide
and short (length 450 units*, width 82 units*) into a long and narrow form
(length 1150 units*, width 55 units*); the average right temperature was about
22?C. But the development of the male from the elongated fourth Ilrva is pro-
tracted, even though the exact time has not been established, since all larvae
removed from the galls perished. Observations of simultaneously invaded plants
show that the development of the male within the elongated fourth hrva lasts
longer than the time required for the appearance of young females, and approx-
imately the same length of time is needed for the development of distended fe-
males, Upon opening the galls, the skin of parasitic larvae in which male de-
velopment takes place frequently breaks, the elongated larvae of the fourth
phase are found in water and can be taken erroneously for fully developed-males,
which can lead to an inaccurate determination of the developmental period. .
[Begin p. 424, para 1]
A protracted developmental stage i? the embryonic one; according to Tayler,
its completion requires 5000 heat units. However, the time indicated by Tayler
herself falls below this magnitude. Thus, according to her observations, eggs
developed at 27?C in 9 days, this means that their development-used up 9x24x17==
3672 heat units. Our observations produced very close figures. On August 12,
1946 eggs were collected in the phase of tWo blastomeres, and on August 22nd
inva'Sion larvae crawled out into the water. Development had proceeded in ten .
days. Total temperatures for these days were 3784 Tayler units.
consumed
The embryonic development/33.5% (1/3) of the temperature total required
for the development of the entire generation. Granulation development of the
embryonic leaflets [listok] and formation of Ilrvae of the first phase proceed
very rapidly. In the case of the observations indicated above,:this process
was finished within fourdays following the initial granulation. The total
temperatures for those days came to 1420 Tayler heat units,-or 37% of the heat
required for completion of the entire embryonic development. Thus, more than
60% of heat is required for the transition of the first larva into the second
one, this, however, includes in the cycle of embryonic development, the process
belonging to the period of postembryonic development.
In our observations, no definite relationship was noted between temperature
and the number of eggs hatched by the female At fairly constant summer temper-
atures, the number of eggs hatched per day varies considerably, and with differ-
ent females, observed in boxes with collapsible walls, these variations differed.
The number of Tayler heat units needed for one hatched egg, came sometimes to "
only 4, more often to 5-6 units, yet in Some cases it rose as high as 20 units.
Heat formula. Knowing the total temperatures needed for the development
of nematodes, and the lowest threshold of development, it is easy to work out
a heat formula fot computation of an approximate duration of development at a
given temperature. It is .known that heat formulas produce but orienting numbers
* micron
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and represent the relation between temperature and speed of development in the
form of a stright line, while it actually; is expressed by a hyperbola which co-
incides with a straight one [line] only at average temperatures. Thei.e is,.hoW-
ever, no doubt as to the great practical convenience of such formulas. In their
application, a substantial error can be expected only in instances of lower temp-
eratures, when the temperature of plant roots, probably rises very rarely to
magnitudes depressing nematode development. By taking the average, instead of
the minimal, periods of development and-by converting Tayler heat units express-
ed in degree-hours into degree-days i.e. dividing the temperature total by 24,
we will obtain for the post-embryonic development of nematodes within the rdot,
from their invasion320
and
to the beginning of egg hatching, the equations S--
500
Pier
and for the complete developmental cycle, from egg to egg, S == wPS
T==io'
is the average duration of development in days, and t- the average emperature
at which development occurs. Our observations of the duration of nematode de-
velopment in flower pots standing in the shade, so that the temperature of the
soil in the cots was ve/-7 near the temperature of air (isothermic)? have shown
the followings [Begin p. 4251
1. In the early summer of 1946, development proceeded in 30 days; inva
sion was introduced in soil oontaining a plant on June 10th, the first oviposi-
tors were noted on July 20th. The average temperature of the period equalled
20.8?C. As per formula, development should have proceeded also about 30 days
(29.6).
2. Two observations were conducted in the middle of summer: a) Inocula-
tion on July 21; beginning of egg hatching, August 14; duration of development
24 days; average temperature 23.7?C. According to the formula development
should have lasted 23 days. b) Invasion was introduced on July 26; beginning
of egg hatching, August 19; duration of development 24 days; average tempera-
ture 23.6?C. As per formula, development should have lasted upwards of 23 days.
3. At the hottest time of 1946; the duration of development was 21 days
---froM August 15 through September 5. Average temperature of this period-==
24.9?C. According to formula, development should have lasted also 21 days.
We can see that the developmental periods computed during the summer ac-
cording to the formula are very close to those actually observed.. In our own
experiments, we were, however, unable to note the time when the larvae penetrat-
ed the root, but only the introduction of invasion material into the soil. In
the spring, at lower temperatures, development of the first generation of nema-
todes lasted from April 20th, when-the larvae wereintroduced, until May 30th
when the hatching Of eggs began, 40 days. The average temperature of this
period equalled 17.1?C. According to the formula development 8hould last 45
days. The formula, as is to be expected gives a longer period.
? Accurate data on the duration of root-knot nematode development in other
parts of the USSR is not available; let us nonetheless dwell on certain observa-
tions for comparison with our data.
According to Kobakhidze (1941), In Tbilisi, at' a temperature with a 24-hour-
daily average of 26.7?C, the development from invasion to young females (as per
author "females in phase of copulation") lasted 13-,16 days. In 4-5 days, they
"fully completed their growth and development". and."soonw began to hatch eggs.
As we can see, accurate periods of development cannot-be established from this
text, but. it is possible that it-lasted about 20 days. As per formula, duration
of development should be 19 days.
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According to field observations of Brodskii and Zemlianskaia (1946), in
Tashkent the spring generation of root-knot nematode developed from the 6th to
the 31st of May. For this period the total effective temperatures according to
10-day averages of soil temperatures at a depth of 15 cm, was 334 heat units, a
little more than the average total temperatures (320 units) taken for the formu-
la. Hence, the period of development observed is very close to the one obtained
with the formula: days observed were 25, and by the formula there should have
been 24.6 days. Yet, the duration of development of the following generation is
indicated by the authors as being 32 days -- from 13 June through July 15, al-
though temperature was higher during this period, and the total of effective
temperatures added up to 528 units, eiceeding the total required for the pre-
ceding generation by almost 200 units. So great a prolongation of development
could occur only in a case when soil temperatures has surpassed the optimum and
has begun to depress nematode development. This, however, could have taken
place only during certain days, since, during the observation period, the ten-day
average [srednedekadnye]-of soil temperature at a depth of 15 cm fluctuated be--
tween 24.1 and 27.8?C, i.e. it did not reach magnitudes depressive to nematodes.
Hence, it is more probable that the authors failed to conduct field observations
with sufficient accuracy.
Moisture. All nematodes are hygrophilous animals, but many of them are
capable 6f falling into anabiosis at an impending decrease of environmental
moisture. [Begin p. 426] However, we established by Frank (1885), larvae of
the root-knot nematode are not capable of it. In contrast to a series of ani-
mal helminths, the eggs of phytohelminths lack also resistance of drying; this
is understandable, since they usually remain within plants, seldom find them-
selves in external environment and do not serve for spreading invasion and,
therefore, their great resistance is irrelevant to the continuation of the
species.
Although experimental work on the duration of nematode survival in drying
soil has been conducted (Jones, 1932; Godfrey, 1933 and others), soil moisture
strange as it may seem, was not determined in these works and, therefore, the
maximum drying survived by larvae has not been established. According to our
observations, oothecae and larvae 6f-the root-knot nematodes perished when
placed in soil dried in the sun, i.e. in actually air-dry soil in the summer,
within the first 24 hours. But a large volume of soil retains a certain amount
of water, in addition to that adsorbed, and this water maintains the moisture
of soil air in a state of Saturation and contains eggs and larvae.
Fig. 11. - Decrea6e in soil invasion upon
drying. Broken line -- invasion
capacity; continuous line --
water-holding of soil.
To determine the lethal action of dried out soil, the author ['of present
article] conducted a few experiments in 1946: in the moist climate of Sukhumi
these experiments are possible under laboratory conditions only. All experi-
ments produced similar results: loss of soil moisture up to 4-5% of the total
moisture-holding Rills all phases of nematode development and renders soil
invasion aversive. Reduction of soil invasion in an experiment with the larg-
est quantity of soil (about 10 buckets) spread-in a large aquarium and dried
in the shade is shown on the diagram (fig. 11). The experiments have shown
that the root-knot nematode survives soil dryness such as cultivated plants
could not survive. As a measure for nematode control, drying-out under field
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conditions is feasible only in the most arid region of the Soviet Union.
??Yearlyjpevelopmental cycle of the root-knot nematode it Sukhumi. Throughout
the wintwr, kthe root-knot nematode is easily found in the rOots of many plants,
net only the woody plants, but also in herbaceous ones Vegetating during the
warm Sukhumi wintef. The root-knot nematode is found in all phases of develop-
ment (exCept male). Nematode development retarded by winter chills is resumed
in April. Mature females again begin to hatch eggs; yet, their sexual activity
in the spring does not last' long: in May, all females Obviously die. Parasitic
larvae overwintered in roots proceed to molt; they; however, also perishin the
large majority of cases before they finish molting. Fully viable are the over-
wintered invasion /1rvae and eggs. Many more larvae ovefwinter -in soil than in
plants; [begin p. 427] invasion of plants begins in April.- The size of nematode
population in the soil decreases rapidly during the winter. Thus, in the fall
in the same garden, 2000 cm3 of soil produced 400 galls on a plant, yet in -
spring we were entirely unable to infest a plant with the same amount of-soil.
In May, population statistics of the root-knot nematode are at a minimum. In
June they begin to grow due to reproduction of females of the spring generation.
Once begun, reproduction continues steadily; females propagate for a very long
period and they still continue hatching eggs after several generations have
developed from the first groups of eggs hatched.
Thus,
all stages
"number of
the number
older ones
there is established a steady flow of reproduction of nematodes in
?of development; there are no separate generations, and the term
generations" should be interpreted as their largest possible number,
which we would obtain if we considered only the offspring of the
as indivisuals of each generation appear.
The number of generations can be established if the total of temperatures
is calculated as being higher than the threshold of development and is divided
by the temperature total required fer the development of one generation. The
quotient of this division will give the number of possible generations. The '
accurate number of generations developing during the season can be determined
experimentally by riasing nematodes on plants in containers-and subsequently?
infesting fresh plants with the eggs of each new generation. We used both of
these methods; first we, naturally, used the calculation method, for it is very
quick and simple. Calculations have to be conducted on air temperature, since -
the temperature of soil varies considerably due to insolation and other factors.
Table 3.
Total temperatures above 10? in degree-days according to air-temperature aver-
a e of many years in Sukhumi
Months
III
IV
V
VI ' VII
VIII I
1
IX 1 X
XI 1
Annual
Total
?[
Tempera,.
ti
'
tures
10
'84
226
350
.415
424
318 201'
78
2092
1
Having infested plants at the very beginning of the season, over-wintered
invasion larvae cannot finish their development, hor can they begin to propagate
before the end of May, when total temperatures in 320 degree days needed for
post-embryonic development, can be reached. Invasion larvae can develop from
the eggs hatched in mid June (about 160 degree days), and post-embryonic develop-
ment can be completed and propagation of the second generation begun during the
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second decade [10th-20th] of July, when total temperatures reach the necessary
500 degree-days. Eggs hatched by the second generation will produce the third
generation which will reach maturity about themiddle of August and will initi
ate the beginning of the fourth generation. The fourth generation will begin to
propagate during the second half of September. Total temperatures are no longer
adequate for the development of the fifth generation. Judging from the monthly
average of air temperatures, in Sukhumi the seasonal capacity is upwards of 4
generations. [Begin p. 428] If soil temperatures on the southern slope were
used in the calculations, then the seasonal capacity would increase up to 5 gen-
erations. In 1946 the number of generations was studied experimentally by means
of.successive intertwisting. Tracing the seasonal pro'gress of the development
of generations under observations (fig. 12) and comparing it with the chart on
generations computed by total temperatures, we were astonished by their great
coincidence, which may have been due to the fact. that the temperatures of soils
is very near to the temperature of air and, in 1946 (excepting autumn), the
latter varied very little from the-average of several years. In the fall, in-
festation of plants is at a maximum, the same as the amount of invasion larvae
in the soil. From 5 gr. of soil taken from a tobacco field, we removed in early
November up to 54 larvae of the root-knot nematode by the funnel fvoronochnymJ
method; this comes to more than 10 thousand Itrvae per kg of soil. There is no
doubt that all possible generations of nematodes develop suncessfully on annual
plants only in rare cases, since vegetation of many plants ends before low temp-
eratures set in and the reproduction potentials of the root-knot nematode are
not fully realized.
Fig. 12. - Yearly progress of development of
generations of the root-knot nema-
tode calculated (broken line) by
the total of average temperatures
and established eXperimentally
(Continuous line).
Fig. 12. - [Directly under graft] reads as follows:
April May June July August September October
Probable Developmental cycle of the nematode in other parts of the USSR.
The close coincidence between the progress of calculated and actually observed
nematode generations in Sukhumi justifies the use of the method for calculation
of total temperatures also for other areas of the USSR. The method is less
promising for northern rgions, since development proceeds more rapidly at lower
temperatures than it oug t to; according to the formula, and this means that
total temperatures will 8wer. In the republics of Central Asia, retarded de-
velopment, possibly, is the result of-superheating. For calculation, we used
the world agroclimatic Hanbook (1937). It contains computed total temperatures
for periods above 100 and indications of the duration of these periods. To
obtain totals of effective temperatures, it is necessary to subtract the number
of degrees below 10 from the figures cited; this is obtained by multiplying the
number of days of the period by 10.
Let us examine the probable cycle of nematode development in Turkman.
Totals Of-effective temperatures reach: at Ashkhabad -- 2200, at Bairam-kli --
2300, i.e. a little more than at Sukhuilli; these localities, however, are dis-
tinguished by an early and warm spring. Total temperatures adequate for the
development of mature females from overwintered larvae will be reached as early
as mid May, and invasion of plants by newly developed larvae will .begin at the
end of May. earlier than in any other area (naturally, with the exception of the
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bordering southern part of Uzbekistan where it is even hotter). [Begin p. 429].
The second generation will begin to multiply during the second decade [from 10th
-20th] of July, the third -- during the second nalf of July] Here mass nema4.4
tode propagation and mass invasion of plants will begin earlief. than in Abkhazia,
hence it is natural to expect greater damage from the parasite. Seasonal Capac-
ity -- up to 5 generations;
In Baku, total of effective temperatures -- 2290, the same as in Bairam-,
Ali, but their distribution by months is different: in the spring there is less
heat, in the fall more. The first generation of the root-knot nematode develop-
ed from overwintered larvae can begin to multiply only in June, almost a month
later than in 'Turkmen. The second generation will begin to multiply about the
middle of July. Seasonal capacity -- 4-5 generations.
.In the Ukraine, the total of effective temperatures is practically every-
where below 1500. The root-knot nematode has been distributed considerably in
the area of the Donets river.
Fig. 13. - Probable seasonal progress of
development of generations of
the root-knot nematode in some
parts of the Soviet Union, cal-
culated by total temperatures.
Continuous line -- Bairam-Ali;
dotted line -- Voroshilovgrad;
broken line of dots and dashes
-- Minsk.
[Directly under graft] reads as
follows:
April May June July August September October
To characterize this region we shall use the data on Voroshilovgrad. Here,
the total of effective temperatures is 1300. Propagation of females developed
from overwintered larvae may begin in mid June, and invasion of plants by new
larvae - about July. The second generation of nematodes will begin to multiply
about the end of July, in warm soils even the third generation can develop.
To characterize conditions of development of the root-knot nematode in
White Russia, we shall use the data on Minsk. The total of effective tempera-
tures in Minsk is 580. This permits overwintered invasion larvae to reach
maturity and to multiply for some time, but a second generation of nematodes
cannot develop. As per observations conducted at the White Russian quarantine
laboratory in 1935, reproduction of nematodes (apparently those developed from
overwintered larvae) began during the first decade [1st to 10th] of July. Con-
sequently, the total of effective temperatures for this very hot month comes to ?
232. Invasion of plants by new larvae is possible only in the last days of July.
These larvae cannot complete their development, since average temperature of the
third decade of August is below 15?. In annual plants, larvae gone over to
parasitism are doomed. Nematodes overwinter in the soil in-the form of eggs
and invasion larvae. Under these climatic conditions, mass reproduction of root-
knot nematodes is not expeoted, hence-a heaiiy invasion of plants under field con-
ditions is hardly g-obable. [Begin p. 430].
All of these calculations are made on air temperatures and can be consid-
ered only as tentative. In this form, they are however, very-useful in estimat-
ing the possibility of mass reproduction and its consequences. ?
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ROLE OF PREDATORS AND PARASITES. Among the inhabitants of soil are found
many enemies of the root-knot nematode; they all destroy only preparasitic
larvae;'enemies attacking nematodes within the galls have not as yet been de-
scribed. Thus, the root=knot nematode is safe from their exacting action during
a large part of its life. Fifty-two enemies of the root-knot nematode have been
found in Hawaii (Linford and Oliveira, 1938);"most of these parasitize on other
nematodes, as well, and some attack each other. The list of these enemies in-
cludes 18 species of fungi, 1 of protozoa, 24 of predatory nematodes, 6 of mites
and 3 of tardigrada. The normal activity of organisms limits the nematode pop-
ulation. Apparently, the animals which cause the most important damage to nema=
tode ;larvae injurious to plants are predatory nematodes of the genus Mononchus.
We had a Mononchus female living in a match glass in a moist *Chamber for a
month, feeding exclusively on larvae of the root-knot nematode. A report on an
observation of one specimen of Monochus Papillatus which destroyed 83 larvae of
the root-knot nematode in one day is available in literature (Proceedings of
the root-knot nematode conference, 1937). In Abkhazia, tho numerical strength
of Monunchus has the same seasonal progress as the root-knot nematode; these
predators are more copious in the fall months. Of a 1Lrge number of fungi
parasiting on nematodes, the most important are the nematode catching fungi of
the genus Dactylella. Tricellular rings resting on bicellular podicels form on
the hyphae of these fungi when nematodes are present. If a nematode enters one
of these ring, it causes a rapid swelling and a contraction of the ring cell,
and as a result the worm is tightly compressed. Later, the fungus hyphae grow
through the body of the nematode and destroy it. Fungi of the genus Arthro-
botrys and C stopa e catch nematodes With the aid of viscous substances isolated
by the hyphao Drechsler, 1937, 1944). The reduction in root-knot nematode popu-
lation, noted long age, can be explained by the increased number of its enemies
in the soil due to tho influence of an abundance of decomposed green fertiliz-
ers. Decomposition of plants facilitates the propagation of saprozoic nematodes
by supplying them an abundance of food, which, however, leads to an increase in
the number of their enemies. ,The latter attack also the larvae of tho root-knot
nematode which, at first constitute a considerable part of the nematode popula-
tion in infested soil, but later, their number ceases to increase, because they
are incapable of developing on decayed matter.
RECIPROCAL RELATIONS BETWEEN THE ROOT-KNOT NEMATODE AND THE PLANT. Investi-
gation of the morphology and the development of the root-knot nematode shows
that its historically developed adaptation to parasitism is greater than that
in other plant parasites of the genus Ditylenchus and Aphelenchoides. It is to
be expected that tho interrelations between the root-knot nematode and the plant
will also be more complex. [Begin p. 4311.
Conduct of nematode and reaction of plant. Penetration of root. Invasion
larvae are assisted in finding plants by the acid section lotdeleniia] of the
roots causing positive motive reaction (chemotropism) of the larvae and a tac-
tile influence (stereotropisth) prompting them to maintain contact with the root
once it has been established. Larvae may begin to feed on a plant while still
outside the root by piercing the cell walls with their stilets. Introduction
of invasion larvae in water into the root cannot be observed, because action of
the stilet requires that the body have a buttress. This phenomenon can be
studied on agar slides by preparing 3% agar and pouring it on petri dishes. Tho
larvae penetrate into tho root between cells or' through the gap made by the
stilot in the cell wall. They can gain access only in the area of young cells,
hence infestation can occur only in the meristematic area of the root. Within
the root, the larvae can roam two-three days, consuming food more energetically
and mnvino. mnsflu TCZC. ^n1 10 r;".11-
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fixed in a certain position so that their heads are lying in the pleromn,
[pleroma] and the body -- in the parenchyma of the rind. Larvae frequently in-
fest also the subsurface parts ?of stems and tubers, but their infestation of the
surface parts ofa plant occurs rarely and must be considered as aberrant cases
of parasitism.
Effect of larvae on plant. Mechanical injuries inflicted by larvae while
penetrating the root, as well as during their migration within it, are usually
insignificant which, undoubtedly, is most favorable for the further subsistenCo
of the parasites. Larvae fixed in a root, soon lose motility and are no longer
capable of leaving the root in case it dies.- Furthermore, to form giant cells,
nematodes must feed on undamaged young cells. Plant helminths which do not
lose motility, for example, Pratylenchus pratensis (de Man), inflict on the -
plant considerably greater injuries during their migrations within its root.
The chemical action of larvae on the plant is much stronger; it is exerted
by secretion of the esophageal glands isolated during feeding and begins to
develop while the larvae still migrate within the root. Secretoi-y action appears
the strongest on the meristem and on the parenchyma of root rind. The former
becomes depressed: division of Meristem cells is thwarted, and growth' of the
root is interrupted or retarded. In a case of heavy infestation, growth of the
root ceases forever, as a result of which club-shaped galls form on the ends of
small roots. More often, growth has boon observed to recommence after a period
of retardation, this is connected with the resumption of activity by meristema-
tic tissue. Dimensions of parenchymal cells of the rind increase due to secre-
tory action of the root-knot nematode; their hypertrophy is responsible for a
thickening of the toot which sometimes is discovered already on the second day
following invasion.
Action of precipitated nematodes. After fixation, the nematode body be-
comes immotilo, with the exception of the anterior part whi4 retains the capac-
ity to move in tho dorsal, abdominal and lateral directions. Nutrition consists
of constantly recurring cycles including three actions: 3.) puncturing the cell
walls with the stilot; 2) injecting secretion of the esophatoal glands into the
cell and 3) sucking out a certain part of the cell contents. The parasites
secretion falls directly into the pleromo cells bordering the nematode head,
and retards their differentiation; [Begin p. 432] further, some of the undiffer-
entiated cells begin to increase in dimensions, and their nuclei divide without
the subsequent fission of cells; then the barriers between the neighboring cells
gradually disappear, their contents merge and form the bbginning of a giant cell.
The huge cells Which at thebeginning are not largo, continue'to grow duo to tho
inclusioh of the neighboring cells, but their fusion with one another was not
observed. As per our measurements, the largest giant cells reached transversely
260 x 221 units*; lengthwise tho giant cells are larger. Mb counted 20 nuclei
in them, but reports in literature indicate over 500 nuclei. Elements of tho
central cylinder which surround tho giant cells also undergo changes; some
tracheata and vessels are ruptured in the gall area, others passing by the giant
cells are merely turned aside. The secretion of the root-knot nematode exerts
a stimulating action on the cells of rind parenchyma, causing considerable in-
crease in its dimensions. Hence, the thickening of tho root, obsered at times
during larvae migration, increases strongly after nematode fixation. Tho essen-
tial mass of root-knot nematodes is composed of parenchyma cells. Yet, the numb-
er of parenchymal cells within tho gall, and of those outsido-it is the same.,
but cell dimensions within the gall increase by several times. Thus, in cucumb-
ers, the size of parenchymal cells in the gall equalled 160.units*, and in the
* micron
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*same root outside the gall -- 75 units*, i.e. by assuming that the form of cells
is cubic, their dimensions increased by 9 times (1603g 753); in tomatoes dimen-
sions increased-by 15 times, in meet potatoes -- by 6,. and in the Acalypha
weed -- 4 times. Action of nematode secretion causes activation also in peri-
cycle cells they proceed to divide increasingly, and this leads to the forma-
tion of lateral roots which frequently branch out from nematode galls in large
number.
Tho external appearance of galls. The external appearance of galls changes
considerably during the season. As spring approaches, small separate galls, 1.5
rim in-diametor, cuasod by one, more often by two parasites, develop on thin
roots. At the beginning of reproduction, the walls of such galls burst open
and oothocao form outside of them. Tho dimensions of' galls expand as invasion
increases, and at the end of the season, the predominant type constitutes a
strong swelling on large sections of the root. The size of galls depends on
the magnitude of the root, in larger ,::plants, such as squash or tomatoes, they
reach, 20 mm in diameter. A special group is composed of club-shaped end galls,
usually small in size, about 1.5 mm in diameter, formed as a result of a simul-
taneous mass invasion-of the root by so large a number of larvae that they in-
terrupted root growth. On perennials, galls sometimes roach enormous dimen-
sions. In Florida, galls on Thunborgia laurifolia were described as being 45
and 60 tom in diameter; they had developed on the roots and at the base of the
trunk (Steiner and Buhrer, 1934). On the other hand, galls wore found to be
poorly developed and on some plants completely absent; in the latter case rind
parenchyma does not expand, but a yellow spot forms around the female, since
0 the root:.knot nematode does not cause these plants hypertrophy, but cell
necrosis. The absence of galls is characteristic of sweet potato and Gorbera:
Importance Of gall formation. The importance of the described reactions
of the host organism to the introduction of the parasite rests in the isolation
of the parasite and the localization of toxins discharged by it. There occurs,
at the place of nematode fixation an increased influx of nutritive substances;
[Begin p. 433] granular impUritios'accumulate in the protoplasm of gigantic
cells which, according to Kostoff and Kendall (1930), develop as a result of
molecular condensation and consist of protein granules. The influx of abund-
ant nutritive substances serves the plant as a means of localizing foreign
matter, discharged by the parasite, but it is also very favorable for the
parasite which obtains d largo amount of food concentrated in a small area, and
around its head at that. The formation of galls, occurring as ,a result of
hypertrophy of rind parenchymal colls,"serves as a means for isolating the
parasite from other parts of the Plant. Galls,,however, are very favorable for
nematodes since they protect them from predatory soil organisms, insure them
against drying out and establish around them a very constant external environ-
ment.
Thus, the protective reaction of the plant which is costing it a great
deal and is causing the expenditure of a large amount of plastic and nutritive
material; naturally, to the detriment of other organs, is very useful for the
parasite. This conditioned by the inadequacy of the reaction itself to profit
from the protective function of the host organism, but chiefly by the adaptation
of the parasite and its ability to adapt itself to specific living conditions,
As a result, the protective reaction of the host becomes a necessary requisite
for the very existence of the parasite.
Dimension & females from large and small galls. It is to be expected, that
conditions for the development of nematodes will be more favorable in largo
galls than in small ones and, therefore, females will orow hiPPnr in lftrcen
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Trans. A-524
and will hatch more oggs. Moasuremonts confirm this hypothesis, (Table 4).
Table 4.
Measurements of females of the root-knot nematode from large and small galls.
Plant and appearance
of galls
Number of
moasuremonts
Measurement of females (in
units*)
Length 1
Width
Tomatoes:
Large galls ?
Small galls
Tobacco:
Large galls
Small galls
Blue Eggplants:
Largo galls
Small galls
*mu '
100
96
141
.
143
57
36
983 4 111
643 T 100
_
?
928 4 116
MO T 80
_
990 4 170
713 T", 81
_
704 i 86
475 T72
682 4 ll6
470 T 83
?
637 4,- 91
490 4" 70
Varying curves of female dimensions of-the-root-knot nomatode from large
and small galls on tobacco are shown on fig. 14..
Comparative action of different nematode species on plants. Hypertrophy of
adjacent-colls is a'very frequent reaction of the organism to introduction of the
parasite. [Bogin p. 4343 Some holminths (for instance, nematodes of the sub-
order S irurata) cause tumors in animals; expansion of plant cell (holmintho-
cecidia) is caused by all nematodos of the order Anguina and Heterodera, while
other nomatodos, such as Ditylonchus-and the leaf parasites of the order
Aphelenchoides cause tissue necrosis. The latter reaction is, undoubtedly, more
primititro and very injurious to the plant, as well as to the parasite, forcing
it to movo from dead tissue to a living one. This is achiovod either by direct
nematode migration, or observed in chrysanthemum nematodes -- Aphelenchoidos
ritzemabosi (Schwartz), by their trangition into an anabiotic stato and by
spreading of invasion with dry loaves. If the parasite loses motility and be-
comes incapable of anabiosis, then tissue necrosis will cause its destruction.
A sign of an-even lesser adaptation of the parasite to the host is tho strong
mechanical dostruction of plant tissues, and the nematodes causing such destruc-
tion, as for instanoo Pratylonchus pratensis (de Man), must bo considered esmore
primitive parasites. The interrelation between the root-knot nematode and the
plant must on the contrary be considered as very complicated..
Fig. 14 - Variation curves of female measurements
of the root-knot nematelde from large and
small galls on tobacco. Continuous line
-- body length of females in small galls;
punctatod -- same in largo galls.
The internal structure of Hoterodera galls is distinct from the structure
of other nematode galls. A nutritive zone feeding the parasites and surround-
ing the internal cavity harboring the parasites forms in the galls caused by
Anpuinn ns iirrii ne in 4-1,^0,-,
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0 the function of tho nutritive zone is fulfillod by giant colls Tho ability
to causo formation of cell syncytia [sintsitiev] is a specific Characteristic
of the genus Heterodora, inherent in all of its representatives. .
There are no data in literature concerning the nature of socretions isolat-
ed by the nomatodo, not even a hypothesis has boon voiced on this subject. How-
ever the similarity of the action of socrotions,of the root-knot nematodo to that
of growth substances on plant is immediately obvious t the latter stimulate root
growth when their concentrations are very weak; yet their stronger concentrations
retard growth and cause thickening of the root. According to Kholodnyi (1939),
the action of higher dosages of growth substances interrupts the growth of a.
root in length almost completely, yet at the same time there forms a thickening
in the section subjected to thoir action, and after 24-hours it develops into
a rather considerable swelling. Anatomic investigations show that the thiaken-
ing consists of rind parenchyma cells strongly expanded in width; [bogin p. 435]
the central cylinder also thickens, but to 5. considerably lesser degree. The
development of literal roots is accoloratod. Maksimov (1946) believes that the
basic function of growth substances rot in their supplying water and nutritive
substances to the area of their action. Of great importance to the offectivo-
ness of growth stimulants is the ago of tissuot their action on young coils is
considerably strongor.
The similarity between the action of root-knot nematode secretions and that
of growth substances prompts the assumption that their concurrent action would .
produce a stronger reaction of plants and the formation of bigger galls. Ex-
periments confirm this hypothesis: by planting cucumbers in soil strongly and
very evenly invadod by root-knot nomatodo6, and by'wetting a part of them with
a solution of a-naphthyl-acotic acid of 0.01 and 0.00%concentration, wtY Obtain-
ed considerably larger root expansions on the plant S treated, than those caused
by one nematodo; the latter were 1.6 times thinner than the oxpansion5caused by
concurrent action of a nematode and growth stimulants.
QUANTITATIVE CALCULATIONS AS TO ROOT-KNOT NEMATODES IN PLANTS. The method we
used for a quantitative calculation of root-knot nematodos in plants was the
following: a piece, as correctly cylindrically shaped as possible, was cut with
a razor blade from a gall, and then cut up with preparatory needles [preparovalt
numi iglami] in water into piocos so small that the nanatodos it harbored washed
out. Tho water with the suspended nematodes was distributed with a pipette on
a slide [prodmotnoe stoklo] in a narrow strip and the nematodes wore counted
under a microscope. " Invasion larvae, not yet in a state of parasitism, wore
not taken into account. The dimensions of the investigated piece of gall was
calculated as per the formula used to determine the dimensions of cylinders.
Calculation of developing galls showed that, in proportion to nomatode develop-
ment, the average size of a gall, designed for one nematode, increases consider-
ably. Thus, in the early developmental stage of a gall containing only parasit-
ic larvae, the average space per parasite was 0.14 mm3, but at a later stage, -
when-galls-aro already harboring young females, the average space increasos up
to (:)5 mm3. In old galls containing egg-hatching females, the average gall
space per parasite was even bigger, although not consistently, as soon on table
5.
Tho population of club-shaped end galls is more dense than throughout
the rooti in these galls, the average spaces assigned to one parasite does not
exceed 0.67 mm.
To obtain an averago number of the root-knot nomatodos within the entire
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root system, we measured the galls and calculated thoir dimensions on two plants.
In tho case of a bluo eggplant, we obtained gonoral gall dimensions amounting
to '94341.3 mm30 and for a tomato (on which small galls were not counted) --
188627.6 mm3. Assuming, on the basis of data-On table 4, that the average
space of a gall assigned to one parasite is 1.6 mm3, we shall find that the
average number of neratodos parasiting the galls Of tho blue eggplant was about
59 thousand, and for those parasiting the galls of the tomato .1- about 118,000.
The a verago spaCo of a gall assigned to ono female is 3.4 mm3. Thus, the roots
of tho blue eggplant wore inhabited, approximately, by more than 27,000 females,
and the roots of the tomato by more than 55,000 fOmales, i.e. in strongly in-
fested plants they are found in tens of thousands. Since tho fertility of each
female amounts to more than a thousand eggs, thaso females will produce tons of
millions of eggs and invasion larvae. [Begin p, 436, below table 5] Just what
mass of living matter is composed of nomatodo6 parasiting in roots? Assuming
that tho averago dimensions of a female are 0.179 mm3, we find that thoir number,
as calculatod in tho case of blue oggplant$ will constituto a bulk of-about
5,000 mm3, and the amount calculated for tomatoos -- about 10,000 mm3. The mass
of living matter of females is, approximatoly, 79 times larger than tho mass of
living matter of other phases, and the principlo devouror of plant substances
are the females. On tho strength of the prolonged process of egg-hatching, the
females have to absorb a great-doal of nutritive substances systomatically, and
this alone is enough to exhaust and to weakon a plant. The amount of secretion
they isolate is also much larger than that isolated by larvae, hence intoxica-
tion on the part of females-must also be considerably greater. This is con-
firmed by Observations over the growth of galls and the beginning of plant
depression.
Table5
Quantitative calculations of tho root-knot nomatodo in large old galls on blue
eggplant, in October 1946
Size of pioco of gall
considered
Number of different
stages of nematode
development
Average gall space
(in mm3) taken up by
one parasite
Length
(in mm)
Width
(in mm)
Dimensions
(in mm3)
5
16
1004.8 Parasitic larvae
- 167
For all stages
2
Female
168
for females
6
165
[Ehles
Total
500
_
11
379.9 Parasitic larvae
3
For all stages
15
Females
109
For femalos
3.4
_ ,
Males
96
Total
208
192.3
Parasitic Larva?
7
For all stages
19
Fomales
91
For females
2,1
Males
4
?
Total
102
10
10
785
Parasitic larvae
110
For all stages
1:7.
Females
171
For females
4.5
_
Males
1.60
?
_
Total
441
11
3
35.3
Females
29
For all stagos
0.95
Males
8
For females
1.2.
Total
37
-
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[Begin p. 4371
INOCULATION OF MICROBES. Apparently, all nematodes -- pldnt parasites --
are capable of inducing pathogenic microorganisms into a plant. Thi 6 applios
also to facultative plant parasites, such as Apholonchus avonae Bast. which in-
troduce fungus hyphao by stilet piercing (ChrITEITTETTTrndt, 1936). So much
the greater must be tho role of obligate parasites which break into the plant
and violate tho wholeness of its intogumonts. As per observations in different
areas in England, tho strong fluctuations in the potato nematode's capacity to
injure dopond on tho diseases connected with it (Triffit, 1931). Root rot of
tobacco is considered a compound disoase caused by penetration of the root by
a complex of pathogenic organisms consisting of nomatodos, more often Pratylen7
chus pratensis (de Man) and of fungi. In the USSR maceration of tau-saghyz roots
is eiTrainod by the combined action of nematodes and microbes, with tho primary
role being ascribed to nematodes (Kalinenko, 1933, Sveshnikova, and Skarbilo-
vich, 1937). The fact that infestation of plants by tho root-knot nomatodo is
accompanied by fungus diseases was noted already by Jobert (1878) with regard
to coffee trees in Brazil. Since then, the combination indicated has boon con-
firmed by a large number of Conducted observations; obviously heterodorosis is
always a combination diseaso. An especially big role is ascribed to tho root-
-knot nomatodo in iho accumulation of Fusarium vasinfoctum -- the causal agent
of cotton wilt. Tho nomatodo is an accessory to wilt in tomatoes in which the
disease is caused by another form of Fusarium. Apparently, the connection be-
tween the root-knot nematode and Fusarium is closer than with other pathogenic
fungi, which is explained by the fact that hoterodorosis as woll as fusariosos
[fuzariozy]-belong to the first immunogonotic group, acaording to Dunin's (1946)
system, i.e., they attack ontogonetically younger roots. In tho USSR, a combin-
ation disease of the root-knot nematode and fungi was observed in a section of
lavender on a sovkhoz near Sukhumi, but the required investigation of tho di-
sease has not been conducted.
PATHOGENESIS OF THE ROOT-KNOT NEMATODE. As a result of root-knot nomatodo
infestation, the plant is stricken by a protracted disoaso -- gall heterodero...
sis. As is the case with other diseases, tho courso of hotoroderosis may be
either slight or serious. Subsequent to the chronic course of the disoaso and
prolonged isolation of foreign matter, the disease spreads rapidly and affects
a large number of plants, i.e., it has the tendency to assume an epiphyte
character. Frequently the disease provos fatal to tho plant and young plants
may perish before producing their first yield; this was observed in tho southern
republics of the-USSR on the seedlings of fig, peach and mulberry, and also on
vegetable plants. The direct reason for the destruction is decomposition of the
root; large galls always decompose, and whenever destruction of tho root system
is considerable, the plant perishes; in the case of annuals, destruction is
usually observed at the end of summer. In slight infestation the disease passed
unnoticod,though it is only very rarely that it does not exert influence on
plant growth and yield (later on this is true also of a slight infestation of
annual plants). In the USSR the root-knot nematode causes damage in tho
southern republics of: Gruzia, Azerbaidzhan, Turkmen, a littlo loss in Uzbokis-.
tan. Work conducted there recorded much nematode damage (Blinovskii, 1945;
Brodskii and Zomlianskaia, 1946; Klochotov, 1947; Kobakhidzo, 1941; Solivonchik,
1938;. Ustinov, 1934 and 1939). [Begin p. 438] Estimates made of yield losses
caused by nematodOs to various crops were quito considerable. According to our
calculations, in Sukhumi tho weight of tobacco plants badly infOsted by the root-
knot nomatodo dropped in different calculations by 31-71%, and in average by
57%. In a zone whore the total of effective yearly tomperatutros reaches 2000
(in degree days), and above, the groat harm done by tho root-knot nematodo under
field conditions cannot be doubted. In areas of a temperate climate, much harm
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caused by the root-knot nematode in the field is recorded for Alma-Ata (LitVi-
nova, 1939b) where the yearly total of effective temperatures is about 1300...
In our opinion, the figures of losses cited by the author are extremely high
and need checking. It is possible that here-the nematode acted in conjunction
with microorganisms mor+athogenic to plants. In greenhouses and in conserva-
tories the root-knot nematode can cause great losses and even destroy completely
the yield of any latitude.
PLANT IMMUNITY TO THE ROOT-KNOT NEMATODE. By immunity is meant the ag-
gregate 6f plant properties inhibiting and depressing the development of a
parasite. Until recent times, plant resistance to the root-knot nematode was
being explained by the structural characteristics of roots preventing larvae
from penetrating into the plant. Frequently, reference is made (Filip'ev, 19340
and others) to be unpublished work of Arzberger who indicates that the roots of
unsusceptible varieties of cow-peas have more protective tissue than suscept-
ible tarieties; their cork layer is better developed and has fewer ruptured sec-
tions. This theory of mechanical barrier was pursued also by Korab and Butov-
skii (1939) in explaining the action of plants antagonistic to the beet nematode:
"By stimulating the passing of nematode larvae from the cyst into the soil, they
(the plants) attract them to themselves en masse, but keep the larvae from enter-
ing their roots, as a result of which they perish from exhaustion in the futile
attempts to penetrate the tissue of the roots they were attacking".
The humoral reactions of the plant to-the nematode within the root was
ascribed of little importance until lately. It is true that Kostoff and Kendal,
investigating nematode galls on tobacco, back in 1930, expressed the hypothesis,
that the plant fights nematodes by isolating antibodies namely precipitins,
lysins and agglutinins; however, the protection role of these anti-bodies was
not clear, since both, galls and nematodes develop on tobacco very well. Only
in 1939 did Barrons (1939) come forward with the assertion that larvae of the
root-knot nematode penetrate susceptible plants and sturdy plants in the same
ease, but do not develop in the latter. The author sees the reason for this
in the action exerted by sturdy plants on larvae, namely by their isolation of
antibodies neutralizing the ferments of the alimentary glands of nematodes.
Barrons' viewpoint was met with approval; the production of antibodies by plants
served to explain also the resistance of some potato varieties to the potato
nematode (Gemmell, 1943).
Immunity, however, need not necessarily depend on the isolation of anti-
bodies, but may be due to the structure of internal plant tissues; structural
characteristics, namely the roughness of cells of the meristematic area in the
stem node served to explain the non-susceptibility of some wheats to isosomes
[izozomyl* (Phillips and Dicke, 1935). [Begin p. 439] The plant may be saved
from infestation also by the quick tempo of its development: according to
observations conducted by Chailakhian (1947), this was the reason why hemp
grown on a short day escaped infection by broom-rape.
Our immunity investigations began with observations of cotton. Its in-
spection was given much attention, and cotton proved to be very resistant to
the root-knot nematode. In Sukhumi, on a sector severely infested at the time
of cotton planting, no galls were found on cotton roots at the end of the season.
However, in a systematic investigation of the roots, beginning with the day Of
.invasion, it-was found that infestation had occurred, but that later the larvae
had perished.
* lProbably isosome-galls]
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Observations of cotton were conducted in several series of experiments and
always with the same results: during the first week following invasion, galls
harboring larvae already in the phase of parasitism appeared on the roots --
thick, with a powerful bulbus and immotile. Then the gradual destruction of
larvae began; in two weeks a large part of larvae was dead,.and the gall & began
to disappear. Yet some larvae do complete development and begin to molt. No
larva succeeded in completing-development up to conversion into a female: they
all died during metamorphosis. Nonetheless, there were individual cases when
males appeared. During an investigation of nematode 6 by the tens in cotton
roots, three developed males were found living there. Their measurements were
dwarf-like: (1) body length 860 units*. width - 26 units*; (2) length - 850
units*, width - 28 units*, and (3) length - 1100 units*, width - 27 units*
A study of the galls showed that they developed on account of the increased
size of rind parenchymal cells which reached 90-100 units* in the galls, yet on
the same roots above and below the galls - 40 units*, i.e., assuming that theii-
form is cubical, the size of the cells had increased by approximately 13 times..
The central cylinder of the root also thickens, but to a lesser degree. In some
cells of the central cylinder considerable granularity is visible, which accord-
ing to Kostoff and Kendal, serves as an indicator of occurfing immunological re-
actions. We were, however, unable to find any giant cells. We believe starva-
tion to be responsible for the dying of larvae: larvae are incapable of causing
the formation of giant cells, source of their nutrition, and although some
larvae continue to subsist for ,a while and even reach maturity, they cannot ac-
cumulate the supply of food needed to complete metamorphosis, and inevitably
perish during molting. Males, which require les 6 food, develop in individual
cases, but do not attain their normal dimensions.
Observing parasite fate in other resistant plants, we found that it was
similar to that in cotton. In testing resistant South American tobacco --
Nicotiana glauca Schk., invasion of roots was achieved easily. Small galls
appeared on roots as a result of feeding the secretion of larae. Larvae began
to develop, but subsequently all died, and the galls resolved. Not even males
could be obtained on this plant, all larvae died before molting. A similar
picture was obtained in infestation of the sturdier varieties of sweet potatoes,
with the only difference that individual larvae did complete their development
in these sweet potatoes and were transformed into mature females. However, a
large majority of the larvae which had invaded the plants died.before their de-
velopment was completed. Citrus plants never suffer from heterodefosis, but -
infestation of young roots in strongly invaded soil occurs rapidly. [Begin p.
4401 Small roots invaded by a large amount of brvae cease growing and assume
a club-shaped appearance. Rind parenchymal cells swell up to 70-80 units*, while
their dimensions outside the gall do not exceed 20-30 units*, i.e. cell dimen-
sions increase by 20 times. Giant cells do not form; the larvae thicken-some-
what and assume the appearance of parasitic phases-, but they.soon perish. The
disease ends in citrus even quicker than in cotton. Nematodes infiltrate just as
easily the roots of crotalaria, but they never finish their development within
the plant and, thanks to its resistance to the root-knot nematode,-this plant
is being recommended increasingly for anti-nematode crop-rotations.
We can see that the easily perceptible difference between susceptible and
resistant plants rests in the circumstance that the plerome cells of the latter
do not react to nematode excretions by forming giant cells. Reaction of the
* micron
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rind parenchyma cells of resistant plants is, however, no-less than that of the
cells of plants most susceptible to the root-knot nematode. It has also been
observed that in resistant plants, larvae secretions depress fission of meristem
cells leading to temporary growth interruption in small roots.
Infection of maize is not always successful; regardless of strong soil in-
vasion, larvae frequently fail to infiltrate roots; yet in other cases, especial-
ly in flower pots, the invasion of maize is almost as successful as that of sus-
ceptible plants. At first, the penetrated larvae calase swelling of parenchyma
cells increasing their dimensions by approximately-5-6 times, afterward there
develop giant cells with many nuclei in the center. Thus originates-a normal
source of nutrition and larvae development can complete successfully. Observa-
tions show that in many caees'maize resistance is due to tne fact that larvae
do not penetrate the roots. In trying to introduce larvae into the roots of
some other cereals (such as Cynodon dactylon Pers.), We did not succeed even
once, and Barrons_' assertion that invasion larvae infiltrate resistant plants
as easily as they do susceptible ones was not confirmed by our observations.
%ether or not the absence of infestation is dub to the inability of larvae
to surmount the barrier of protective surface tissues or to other reasons (for
instance, absence of positive reaction in larvae to these plants) is not known.
The dimensions of females developed in resistant plants on which large
galls failed to form are smaller than in strongly susceptible plants. In an
investigation of-the same garden female dimensions were found to be:
on squash: length - 1060 4 150 units*; Width - 715 4 76 -units*;
on maize: length - 680 T 95 units*; Width - 418 T81 units*;
Since a large part of the female body centists of egg tubes, there is no
doubt that small females will hatch fewer eggs. The duration of development of
parasite larvae in sturdy plants, in which development usually is not fully
completed, is protracted already in the first stages. Yet, there was no differ-
ence in the duration of development of parasitic larva into a. mature female,
when plants of such varied susceptibility to the root-knot nematode as the
cucumber, the tomato, peas, clover, lupine, vetch, nightshade and Acalypha were
grown concurrently in the same containers.
In resuming our observations of plant immunity to the root-knot nematode,
we shall, first of all, endeavor to classify the phenomena observed. There is
no doubt that the protectie reactions of plants to parasites are simpler than
those in animals, (Begin p. 441] but, in relation to helminths, they can easily
be adapted to the classification scheme provided by Soviet helminthologists for
animals (Shul'ts and Shikhobalova, 1935).
To begin with, plant immunity will be natural or inherent. The phenomena
was observed in cotton, in tobacco -- Nicotiana glauca Schk., in citrus, sweet
potatoes and, probably, intrinsic in many resistant plants, fit perfectly the
concept of partial immunity characterized by the partial development of parasites
and their subsequent destruction.
In less sturdy pla nts, such as maize, vetch and others is observed un-
sterilized [nesteril'nyi] immunity characterized by (1) less infestation, (2)
smaller dimensions of developed helminths and (3) decrease in there fertility.
* micron
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411 Sometimes in these plants parasitic development is also protracted.
?
Immunity of age is also observed: woody plants suffer badly from heterode-
rosis mainly at an early age. In the heavily nematode infested Sukhumi Botani-
cal Garden, old'tea plants were not affected, but the seedlings of tea were
easily infested.
Naturally, the immune state of a plant is not unalterable, but it depends
on many factors. In case the plants weaken, or, on the other hand, the para-
sites are reinforced as a result of extremely strong invasion of the soil, galls
will form also on highly resistant plants. The phenomenon is analogous with
that observed in animal helminthiasis: weakening of the host reduces its re-
sistance to invasion of intestinal worms. However, plants particularly sus-
ceptible to the root-knot nematode, such as tomatoes, cucumbers and others, are
infested by it under all growth conditions, oven under the best, such as are
provided for vegetable plants in hothouses and in industrial gardens.
In plant immunity to the root-knot nematode, the phenomena of group
immunity have been observed: the same legume varieties are resistant to the
nematode and to fusarium, American grape is more resistant to the root-knot -
nematode than European grape; the same plant is resistant also to Phylloxera.
In a study conducted in the USA of the resistance of 17 domestic and 14 foreign
cotton varieties to Fusarium vasinfectum and to the root-knot nematode, resist-
ance to both parasites varied in the same direction (Niles, 1939). The parasit-
es indicated have the common characteristic of infesting the root during the
same phase, namely the early phase- of antogenetic development, hence the
characteristics of this phase will exert influence on all of these parasites:
thus, the rapid tempo Of development of the-root and the quick hardening of its
tissues will be unfavorable for each of them.
-Mochriikov, back in 1892, defined the infection as a struggle between two
organisms. The nematode means of attack is a chemical weapon, -- excretions
of its esophageal glands; possibly, the protective means of the plant are also
chemical -- antibodies neutralizing these excretions and preventing the de-
velopment of giant cells, source of nematode nutrition. The presence of anti-
bodies has not been ascertained, however, as per analogy with the protective re-
actions of animals to the introduction of holminths in them, it is entirely ,
probable. The hypothesis of antibodies is, however, hard to reconcile with the
fact that only the action of secretions -- formation of giant cells -- is
depressed, while the rest (depression of meristem cells and hypertrophy of the
cells of rind parenchyma) appear very obviously-. Therefore, it seems more
probable that resistance depends on the characteristics of pleromo cells, name--
1y on the law reactivity of those cells to secretions of the root-knot nematode.
Reduced reactivity may be duo to the fact [Begin p. 4421 that ploromo cells of
resistant plants pass rapidly through those early tages of their development in
which they are capable of turning into giant cells. The rapid development of
roots renders those plants resistant to all diseases infecting roots in the
early stages of their ontogenotic development.
HOST PLANTS OF THE ROOT-KNOT NEMATODE. Number and systematic position of
hosts. By virtue of the extremely wide range of hosts, the root-knot nematode
belongs to ourytrophic [evritrofnym] parasitot; the. number of plants on which
it is registered is vast and keeps increasin't. To illustrate the above, a
course of study of hosts is shown in table 6. ?
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TABLE6
Chronological record of hosts of the root-knot nematode
Author and title
Year of
Publication Number of hosts
indicated
Material used in compil-
ing list
Monograph on the
root-knot nematodo.
Frank.
Monograph on the
root-knot nematode
Neal.
Monograph on the
root-knot nematodo
Narcinowski.
Abstract on the
plant parasitic
nematodes
Bessoy.
Monograph on the
root-knot nematode
List of British
Imperial Bureau
Goodey.
Abstract on plant
parasitic nematode
List of the US Do,
partmont of Agri-
culture (compiled by
Buhror, cooper and
Steiner)
First addition to pro
ceding list (compiled
by Buhrer)
Second addition (com-
piled by Buhror)
1884
1885
1889
1909
1911
1931
1933
1933
1938
1940
36 On plant from Brazil, the
rest from Europe
50 Same
64 Florida, USA
235 Abstracts from litcraturo
480
569
621
855
4477
4 55
Same
tt
Abstracts from literature
and material of the Do-
partmont of Agriculture
Same
tr
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Thus, we can see that by the year 1941 the number of kno*n root-knot -nema-
tode hosts had increased to 1387 plant species and subspecies. It must be noted,
however, that not all designations are authentic; part of the plants registered,
probably,.possess partial immunity and should not be considered as host of the
root-knot nematode. Not once was root-knot nematode infestation registered on
spore plants [sporovye rasteniia], but among flowering plants infestation was
found in representatives of angiospormae as well as in those of gymnospermae;
in the latter, by the way, very rarely: there are descriptions of infestation
of the ginkgo tree, juniper and the palm (Cycas).
Of the 280 families into which angiospermae are usually broken down, infes-
tation by the root-knot nematode is kno*n among representatives of 125 families,
or 44% of their total number. [Begin p. 443] In some families, the number of
species on which the nematode is found constitutes 100%, yet in ethers it is but
fractions, of one percent of the species belonging to a family. The families
among whose representatives infestation is not known include the parasitic and
aquatic ones which cannot be infested due to their ecological characteristics,
and also little distributed groups which; probably, hardly ever are found on
cultural fields infested by the nematode.
In the USSR, in one year's work, notably 1935, conducted on a large
scale by the External and Internal Plant Quarantine Department of the Ministry
of Agriculture, USSR, 241 forage plants were registered as hosts of the root-
knot nematode (Ustinov, 1939).
At present, after utilizing w6rks on the root-knot nematode published
since that time -- accounts of quarantine laboratories for the years 1936 and
1937, materials of the Laboratory of Lower Worms of the Zoological Institute,
Academy of Sciences USSR, the author's observations in Abkhazia -- we have
increased the list of known hosts to 358 species. The host plants registered
in the USSR belong to 74 families, making up 59% of all those families whose
representatives include hosts of the root-knot nematode known in world litera-
ture. This great variety in the systematic position can be explained by the
fact that gardens and greenhouses-were investigated, the reason for the many
introduced plants among the hosts.
Families are
arranged in
1: Compositae
2: Legumes -
3. Nightshade
4; Mint family -
5: Mustard family -
6. Olive family -
the order of host abundance:
7: Rose family - 12 species
8; Goosefoot family-10 "
9: Cereals 10
10; Carrot family - 9
11: Willow family.- 9
12. Mallow family - 9
45 species
36 "
20
18
15
13
t I
tt
Naturally, these figures do not characterize the degree of infestation of the
individual family, since the number of species within the family varies greatly.
The root-knot nematode is less important to compositae than to some other
families, and the large number of hosts registered-in this family can be explain-
ed by the wide distribution of its representatives. This applies even more to
cereals on Which the nematode is a rare parasite. Yet to the gourd family, in
view of its smallness, this parasite is ve'ry important, oven though only 7 nema-
tode hosts have been registered among them.
Thanks to the cooperation of many institutions in the work on the root-knot
nematode, its hosts in the USSR have been investigated in a short time much
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better than those in a number of other countries, Which is quite obvious from
a comparison of the list of hosts Compiled by us with the lists of other count-
ries, and warmer countrios at that. Thus, only 32 plant species have been
designated for the Philippines (Fajardo and Palo, 1933); Southern Rhodesia - 108
(Jack, 1943); Hawaii - 122 (Parris, 1940); Japan 4..41 (Hagakura, 1930); Southern
China (Canton vicinity) - 26 species (Li and Ley, 1938)'. The number of known
hosts in northern countries is not at all largevonly 8 species of plants are
cited for Canada (Crowell and Lavallee, 1942).
*The largest number of hosts is-known in Turkmen -- 190 species, next in
Western Gruzia -- 136 plant species. In Azerbaidzhan, 73 host plants are
registered; in accounts of the Quarantino Laboratory, on 9 plants were indicat-
ed for Uzbekistan, but Brodskii-and Zemliatiskaia (1946) have already found the
root-knot nematode on 52 plants. (Begin p. 444] The numbor of host plants known
for the RSFSR is 74; for the Ukraine -- 46; Mite Russia -4- 5; Kazakh SSR --
14, and separate instances in some of the other republics of the Union.
SUSCEPTIBLE AND RESISTANT PLANTS. Having familiarized oneself with the long
list of hosts of the root-knot nematode, it would be wrong to conclude thatbio-
logically all of these hosts have equal significance for it; on the contrary, to
some plants the nematode is the most significant parasite, yet to others it is
an infrequent and unimportant one. The families more susceptible to tho root-
knot nematode include nightshade, goard, and loguthos, although the last of these
families has many resistant species andvarieties. Among other families, strong
infestation is known in some tspocies] of the carrot family, compasitao, morning
glory, and the goosofoot family. Thoso infested worse than others among woody
plants are the representatives of Moraceao, of the willow family, the peach of
the rose family, pomegranate' of the pomegranate family and the tropical Rubia-
ceae coffee. and quinine trees.
Those fesistant to the root-knot nematodo among annual cultivated plants
are cereals. There aro no indications in world literature as to the importance
of infestations to cereals or horbacoous plants, but the number of representa-
tives of thi-t family registered. as hosts of the root-knot nematodo is rather
considerable. Ineddition, some species and varieties of legumes are resistant:
they are of the greatest importanco in the struggle with the nematode. Resist-
ant are also a large pdrt of the European fruit plants, with the exception of
the peach and the plum. Plants among subtropical fruits which do not suffer
from the hoteroderosis aro citrus, avacado and foijoa.
BIOLOGICAL FORIE OF THE ROOT-KNOT NEMATODE. The developmental course of
parasitism usually leads from polyphagia to monophagia. Many eurytrophic noma-
todes -- plant parasites, as well as plant parasitic insects form nutritivoly
specialized faces which frequently constitute the first stage in the formation
of a spocios. Tho best nomatode Heterodora schachtii. Schmidt was until recent-
ly considered as one spocios with several races, bit now this species is being
broken down into 5 dittinct species. Another ourytrophic plant parasite -- the
stem nematodo Ditylenchus dipsaci (Kuehn) Filipjev is also a complex species
including 7 different forms, part of which, probably, will also prove to bo a
distinct species. In animals entering into so close an interrelation with plants
as those producing galls, specialization is particularly strong.
These deliberations prompted many investigators, beginning with Frank (1885),
to search for forms of the root-knOt nematode specialized in nutrition; thoy all,
however, Obtained negative results. Invasion from ono plant to a series of other
plants occurs very easily under field as well as under laboratory conditions,
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own if the latter plants were separated very far systematically. There can be
no doubt that a considerable part of the root-knot nematode population is ex-
tromdly ourytrophic and can dovolop on a large nUmbor of different plant vario*
ties. This, however, does not exclude the possibility of thoir cooxistance with
spocializod races developing on but a limited number of hosts. [Begin p. 445]
Plants especially susceptible aro infested by all populations of the root-
knot nematode, and no instance of an unsuccessful infestation of these plants '
by any viable invasion material, taken from any plant whatever, has boon noted.
. Sturdy plants, however, become infested by some populations and not byothers. Thus, according to Christie and Albin (1914), in the um, in testing
9 populations of the root-knot nematode on clover - Trifolium resupinatum L.,
the plant provod susceptible to 4 populations, produced indefinite reallts in
the case of one, and was resistant to the other four populations. These obser-
vations served the authors as a basis for their assertion that the root-knot
nematode includes specialized races. The second tests conducted by these authors
with 14 nomatode populations on 10 different plants also showed variod suscepti-
bility of sturdy plants to different populations of the root-knot nematode and3
in the authors! opinion, at least 5 different forms can be isolated from tho
populations tested.
To verify those observations, we grew resistant plants in flower pots in
soil dehelminthizod by drying in the sun, and then infested these plants with
different nematodo populations; by the word "populations" we mean invasion
larvae taken from one plant specimen, although from different oothecao. In the
course of the season, cotton was infested many times with populations taken
from cucumbers, tomatoes, tobacco, balm mint, acalypha and, in the capacity of
a particularly aggressive population, with oothecae developed on maize under
natural conditions, in so doing, invasion-material was taken from different
plant specimens grown in different places. Experiments with all populations
produced identical results: invasion kIrvao easily infiltrated roots, began
to develop there Causing sthall galls, but died subsequently, because they were
unable to form giant cells. Identical results were obtained also with clovor:
all populations infested the plant suacessfully, completed there their develop-
ment, and females began to hatch eggs. No variations were noted in the develop
mental tempo of the different populations.
Invasion of maize, on the contrary, produced quite different results. In
many cases, we were entirely unable to infest.it; in some cases infestation
occurred easily and sometimes oven to a considerable degree. Of the populations
tested, taken from balm-mint, acalypha and tomatoes, only those taken from
tomatoes infested maize. This recurred in two series of experiments conducted
consecutively with different maize varieties and with populations taken from
plants of different places.
Can a conclusion bo drawn from these experiments as to the actual exist-
ance of different "racos" of the root-knot nematode, with only a few of them
being able to infest maize? 77c believe that it cannot be done and that the
difference in invaSion results is due rather to a varying activity and viability
of invasion larvae. A seasonal variation has been observed in the resistance
of maize; in the spring its infestation is unsuccessful, but at the end of
summer it.proccods successfully. In 1946, maize grown on a sector near the
farm was not infested by the nematode (with very rare oxceptions), but at the
end of summer it was easily infested by soil taken from this sector and contain-
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ing larvae dovolopod chiefly on tomatoes and eggplant. Soil takon from another,
vary small gardon in Sukhumi at the ond of summer resulted in invasion almost
ovally of maizo and of cucumbers; 402 galls doieloped on tho cucumbers planted
on 20003cm of this soil, and on maize 323 galls. Yot, in the spring this soil
failed to infest maize growing in flower pots, and among the few maize plants
growing in the little gardon, only one.was slightly infostod by tho root-knot
nomatodo, [Begin p. 446]
It is natural to presume that tho seasonal variations in infestation are
due to the big differenco in the number of invasion larvae prevalent in the
soil: if, in the fall, their number was upward of 200 specimens on 1000 cm,
then in the spring there could be only single spocimons. In the spring,
invasion capacity of larvae which had boOn starving since fall must also bo
much lower than in those freshly emerged. All invasion material usod to infest
maizo in pots was fresh, nonetheless, tho maize was infosted only by tho popula-
tions taken from tomatoes. One must assume that larvae which develop on tomato-
es are more viable and more aggrossive than those on balm mint and acalypha,
for larvao developed from eggs of large females on susceptible plants are larg-
er than larvae doveloped from the eggs of mall females on more resistant
plants, As per our measuroments, the average size of invasion larvae from tho
oothocao of .large fomalos was 429 4 14 units*, and from the oothocao'of small
ones -- 414 4 14 units* (upon measTring 122 specimens of each group).
An interesting phenomenon is tho difference in the suscoptibility of
cotton to tho root-knot nematodo in the USSR whore cotton is resistant, and in
tho USA whore the nematodo is considered a serious parasite of this crop. Has,
there really developed a specialized nematode race, or does tho matter concern
the host and not-tho parasite? It is known that cotton reprosents a complex
of divurso forms. A similar diversity is observed also in plums: in the USSR
they do not become infostod by the root-knot nematode, but in tho USA they
frequently are affected by hetorodorosis. Such differences, in susceptibility
in tho same plant varieties are, however, very few; in tho largo majority of
cases, references to plants susceptible or resistant to the root-knot nematodo
are identical in all countries throughout tho worlth It is true also that
different populations did not succeed in causing certain typos of galls infost-
od plants; in a case of slight invasion galls dovolopod Only sparsely in a case
of strong invasion -- galls woro large and numerous, regardless of the popula-
tions wo used to infest a plant. Thus, up to now, we have been unable to estab-
lish tho existence of foras of root-knot nomatodo that possess a specificity
with respect to nutrition. There is no evidence that tho differences in inva-
sion capacity are fixed hereditary characteristics of some populations, and
there is no doubt that in a scries of cases those difforoncos are conditioned
by changes occurring in tho physiological condition -- age and viability of
larvae and in tho condition of plants.
The exclusive polyphagia of the root-knot nematode prompts the hypothesis
that its evolution did not progress in the direction of increasing specializa-
tion of tho parasite in relation to its. hosts, but in the 'direction of a decreas-
ing specialization, and that tho root-knot nomatodo acquired tho ability to in=
fost plants extremely diverse systematically. It is, however, hardly probable
that a species with so wide a realm of distribution and so large a number of
hosts should roprosont something homogenous; it is, however, Much more difficult
to establish races among i-Oot-knot nomatodos than among the other species of
plant parasitic nematodes..
* micron
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"Novoe v urhenii o gallovoi nematode"
[NE q. IN THE STUDY OF _THE GALL NEMATODE)
A. A. Ustinov
pp. 446-452.
CONTROL MEASURES AGAINST THE ROOT-KNOT NEMATODE
Trans. A-524
. There hardly exists another plant parasite against which so many control
measures were tested as against the root-knot nematode. In foreign countries
the bourgeois regime does not allow expansion of state measures and therefore
the invasion of the fields by the nematode steadily increases. For example .
in Florida and California the cultivated [begin p. 4471 fields are so severe-
ly infested by the root-knot nematode that only cultivation of plants re-
sistant to it or ripening early is possible.
lathe USSR.a series .of effective control measures are applied against
the root-knot nematode.
Physical control measures
Among phvsiCel control measures the thermal ones are developed, first
of all -- partial sterilization of the soil with steam . The technical re-
quirement is a sufficiently deep heating of the soil up to 60-65?C. This.
method is used in the U.S.A. for control of the root-knot nematode in green-
house soil, but in other countries it has not spread; in open ground on a
somewhat extensive scale it is not adapted anywhere. Various temperatures
were suggested for heating of the soil; the most widely used for this pur-
pose are sets of perforated pipes through which super-heated steam is led
under high pressure from a steam boiler., Attempts were made to uso boiling
water instead of Steam for the heating of the soil, but this is a less
effective measure and requires an enormous amount of hot water -- more than
800 liters per cubic meter of soil.
Hot water was suggested more than once for "dehelminthizatien" of
living plants; in the Soviet Union corresponding experiments Were conducted
by Ikhtinskaia and Arkhangel'skaia .(1939) for treatments of "levandau
[lavender by SeliVonchik (1938) and Ustinov (1938). Warm baths were -
arranged in a barrel into which a pail. was dropped. Into both' containers ?
was poured water of equal temperature and during the work very little hot
water had to be added into the inner container in order to maintain needed
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Trans. A-524
temperature. Before dehelminthization the plants were kept for 30 - 40
minutes in warm water at 42?C, than they were immersed in the bath and after
the bath immediately cooled in cold water. A 50? temperature was effective
when the exposure was considerable (at least 15 minutes), According to our
experiments the most suitable method is the ithmers ion of roots in a bath for
5 - 10 minutes at a 51 7 530 temperature; but not all the plants can stand
this treatment.
Heat dehelminthization of plants in the control of the gall nematode
cannot be standardized in the same measure as was possible in the control
of the stem nematode in hyacinth bulbs. The assortment of plants affected
by the root-knot nematode is very large and their tolerance of heat varies,
and many of them can endure it considerably less well than the hyacinth
bulbs. It is very difficult to adjust the temperature such that the nematodes
would be killed and the plants would not be harmed. Therefore this measure
did not become part of established practice either as means of quarantine
or treatment.
Chemical control measures
An enormous number of chemical substances wore tested against the root-
knot nematode. Attempts were made to introduce "OV" [according to Callaham:
abbrev. "otravliaiushchie veshchostva" toxic'agents_ i.e. fumigants] into
the soil for -treatment of infected plants, but they were not successful;
all the fumigants are poisonous for plants and can be applied only for the.
treatment of the soil before planting. Soil fumigation .presents consider-
able difficulties. The interaction which develops between the fumigants
and the soil changes the effectiveness [begin p. 446i of fumigation in
various soils (Chigarev, 1936). A conslOerable portion of the chemicals
are not available for fumigation, being adsorbed by soil particles, and
some of the fumigant evaporates; therefore the amount of poison used for
soil fumigation should be hundreds of times larger than for surface treat-
ment. Carrying out of fumigation is more successful in warm (but not hot)
weather (about 20?C); the soil has to be well cultivated and relatively
dry, because the greater part. of the fumigant does not penetrate moist
soil. The poison should be introduced into the soil in large amounts at
25 cm intervals; in small areas the work can be carried out with the help .
of an injector or undera shovel; for the treatment of large areas
apparatuses are constructed which are attached to plows. lhen powdered
substances are used, it is better to distribute them all over the plot'
[continuously] cultivating the earth so that the poison will be beneath
f_4.. layer- of earth. For successful fumigation, immediately after introduc-
ing the chemicals the soil has to be covered -- this is a necessary
condition for the effectiveness of fumigation. A certain substitute of
covering can be achieved by watering the soil from the surface, which also
inhibits, evaporation of fumigant. The surplus of chemical substances is
toxic for plants; therefore after removing the covet, and 10 15 days
are required for airing of the soil before planting. The majority of
fumigants have a favorable effect or the growth of plants increasing the
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410 amount of nitrogen substances in the soil,
As a rule no substance exterminates completely the nematodes in the
soil, but only reduces the population to an economically unnoticeable level.
A negative characteristic of fumigation is that it destroys also natural
enemies of nematodes. It is pointed out that a valuable characteristic of
fumigants is that the majority of them are not only poisonous for animals
but are also fungicidal. And this means that they kill not only the devour-
ing pre-parasitic larvae of predators but also the fungi parasitizing
nematodes and thus considerably reduce the number of their enemies. It is
understandable that this makes it easier for the surviving specimens to
rebuild the population. The results of experiments with the control of
root-knot nematode conducted in the USSR were already published (Ustinov,
1934, 1939). After the publication of these works, the treatment of soil
with "tsianplav" r? cyan-fusion ?] was tested. At a dose of 100 g. per
1 m3 ( 1 ton per 1 hectare) in a sandy soil the utsianplavn [11 produced
a noticeable nematocidal effect by reducing the contamination of plants in
cultivated plots on the average 1-0 times as compared with centrel plots;
nevertheless about 3 . of the plants in treated plots wore infested by
the nematode.
?
?
In order to compare the effectiveness of various control measures it
is necessary to give a numerical evaluation of the degree of infestation
of the field plot with root-knot nematode; and for that purpose the in-
festation index Of the given 'plot has to be calculated. Are determined the
index in the following manner: the plants in the plot wore pulled out and
'examined thoroughly; if the plot was small, -- all the :::lants were examined,
in case of a larger field section -- every fifth or tenth plant. The most
precise method for the evaluation of the infestation of individual plants
is the tabulation of all the galls on their roots. Bnt even this method
cannot produce entirely precise results because th nails. cam be of very
different sizes; in order somewhat to rodu.cc;. thse ,::ifferences it has been
accepted to count each 2 mm (lengthwise) of large gall for one gall. How-
ever when the thickness of the gall is considerable, in 2 mm of its length
there are many nematodes so that such a section equals not one but many
individual galls. It is necessary to limit-field work to an approximate
evaluation Of the degree of damage [begin p. 149] of the roots expressing
it in units. Adding the units and dividing', the sum by the number of
plants checked, the mean level of infection of one plant in the plot is
obtained. Multiplying this by the percent of affected plants and dividing
the product obtained by 100, the index of infestatien of the plot is
calculated. Comparison of indexes of various plots made it possible to
express the effectiveness of measure in percentage. The most effective of
the now known nematocides is chloropicrin. In comparing the results
which we obtained from the use of this fumigant with data from literature,
it can be seen that for tho heavy and moist soils f Abkhaziia larger
dosages of chloropicrin are .required than those recommended in the
There, 25 to 50 g. per 1 m2 dosages are usual while in our experiments the
25 g. dosage produced a weak effect; a 50 g. dosage (which. equals 500 kg.
per hectare) reduced 4 times the infestation in comparison with the control
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Trans A-524
and freed the plants for one year frxii-a- serious invasion. Only very high
dosages of 100 g. per 1 m2 pi.oduce a considerable effect reducing the in-
festation more than 20 times.
Ethylene-dichloride and methyl-bromide were suggedted as fumigants
which are cheaper than chloropicrin. Both Substances are toxic to humans
and the former is, besides, inflammable.and explosive. American publica-
tions recommend:for ethylene-dichloride a dosage of 15 cm3. and for methyl-
bromide -- 40-g. per 1 m2 of sell. There are no reports yet on- tests in
the USSR of these substances against the root-knot nematode. Apparently
the ester of dimethyl-dithio7carbamic acids synthesized in our Scientific
Institute for Fertilizers and Insect,--fungicides represent nematocides
with greater promise; they are produced as a dust .(with kaolin) and are
powder-like substances which do not require the use ef a gas mask; accord-
ing to preliminary 'experiments in 1948 they cause a considerable decrease
in soil infestation. DDT and hexachlorane appeared to be ineffective in
the control of the root,knot netatode.
Progress in chemistry promises discovery of new fumigants which will
make the cultivation of soil a less expensive measure; nevertheless it is
hard to see how soil fumigation can be so attainable that it will be
possible to use it on a large scale in the field. Se far the chemical
method is being used only in a covered ..enclosed ?] ground. Soil treatment
with chloropicrin was successfully applied in the control of the root-knot
nematode-in 1935 in vegetable hothouses of the sovkhoz "Chervoni Zori" in
Khar'kov.
Agrotechnical control measures
The difficulty of control of the root-knot nematode by the chemical
Method compelled already the'first-researchers? to emphasize control with
the help of resistant plants. Resistant plants. in a suitable crop rotation
are the most practicable and .often the only possible method of root-knot
nematode control in the. field in relatively large areas.' Other agro-
techniCal measures such as fertilizers, dates of planting, clean fallow,
as well as "lovchie"[?] plantings and flooding or drying out of the soil,
are of lessor importnce and in ntljority of cases are only auxiliary
measures.
Data on plant resi.sta;Ice root-knot nematode are scattered in
very many works published in most diverse publications. Tn Russian'
language most of the matc-ril is gathered [begin p. 450] in the collec-
tion published by the All-Union Institute .gf Plant Protection "Collection
of articles on nematodes of agricultural plants", edited by'E. S. Kir'ian-
ova (Sellkhozgiz, 1939).
The problem of the cause of resistance was brought up only rarely
(Klechetov, 1947). Foreign papers, mostly short notes frequentlY in form
of letters, are limited to simple statements of instances of resistance,
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43 Trans. A-524
without efforts at their explanation and theoretical basis. And also the
foreign authors quite incorrectly consider the plant resistance to root-knot
nematode as an unchanging static property of the given variety which does
not depend on the plant's growth conditions, stage of its development and
state. Therefore the data from different authors on resistance and sus-
ceptibility of plants frequently contradict each other.
The most important task is the further study of immunity and the
establishing which resistance factors -- anti-bodies or a.rapid root-
development -- are of greatest significance. Under unfavorable conditions
of development the resistance is lower and sometimes highly resistant
plants are affected by the nematode as, for example, the peanut, a con-
siderable infestation of which was observed in the Kazakhstan and Uzbekis-
tan; even more frequently corn loses its resistance. It is true that not
all the observations of infestation of resistant plants are reliable, be-
cause susceptibility is sometimes mistakenly registered according tb the
presence in the roots of only larval stages of the nematode and we know,
that larvae penetrate resistant plants and begin to develop there, causing)
thickening of roots.
Anti-nematode crop rotations are built chiefly of resistant species
and varieties of legumes and of cereals as well as grasS'es. Susceptible
plants should be planted in contaminated soil after a 2-5 year interval.
The influence of a change of crops on the root-knot nematode was followed
up at the Abkhaziia zonal tobacco experimental station. Records taken in
experiments preceded with tobacco showed that even .a one,year cultivation
of resistant plants greetly reduces the damage. to plants (Ustinov, 1934).
Study of soil infestation with invading larvae confirmed it on plant-
indicators planted in 900 cm3 of soil from the tobacco field 976 galls
developed and from the adjoining .corn field only 5 ealls. The Depart-
ment of Field Industry of the station developed for Abkhariia's tobacco
farms a six-field crop rotation 7ith the following order of crops:
1st
2nd
3rd
4th
5th
6th
year
It
TI
- Aheat and perennial grasses (red clover and rye grass);
- perennial grasses of second year of use;
- tobacco over a layer of perennial grasses, in the fall white
lupine over green fertilizer;
corn over green fertilizer, in the fall vetch-oats mixture
for hay;
tobacco over stubble, in the fall peas with rye for hay;
corn over stubble, in the fall sowing of wheat. with
perennial grasses.
The experimental crop rotation was conducted in a plot infested with
root-knot nematode and with its introduction ethe'reet-knot nematode ceased
to multiply in mass and did not cause severe. reduction of tobacco yield.
In another plot of the same experimental station in 1935 71 - 95% of
tobacco was affected with root-knot nematode and damage to the plants was
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so severe that a further planting of tobacco had to be stopped. The plot
was occupied partly with cabbage and sugar sorghum and remained partly unused.
By 1938 the soil infestation was so much reduced [begin p. 451] that of the
plant-indicators planted here only a few were slightly affected by the root-
knot nematode. The grass field crop rotation which are being introduced now
due to the resolution of the Party and the government, have to be effective
both prophylactically as well as a treatment against the root-knot nematode,
especially when cereal grasses and not legumes are introduced. The vegetables
are. the ones which suffer mainly from the root-knot nematode in the Soviet
Union; on vegetable farms it is also possible to control nematodes by way of
crop rotation. On large farms crop rotation with introduction of perennial
grasaes is possible; in stall workers' and home gardens the earth has to be
planted every year with vegetables, but it is necessary to alternate the
planting of more susceptible plants with resistant ones. For the latter can
be used onion, garlic, corn, cabbage and other members of the mustard family,
spinech, sorel, Jerusalem artichoke (Very resistant plant), in the South --
resistant varieties of sweet potatoes.
In arid and hot areas nematocidal effect is produced by clean fallow
under the condition of several additional repeat plowings of the soil during
the, hottost period; Brodskii and Zemlianskaia (1946) checked the effective-
ness of this measure in the vicinity of Tashkent. Mineral and manure
fertilizers reduce harvest losses caused by root-knot nematode, but they
are not a control measure, with the exception of calcium cyanamide, large
dosages of which (7 t. per 1 h.) decrease soil infestation. Green
fertilizers are more effective. than - the nineral ones, because they are
a biological method nf nematede control.
Bin.) eeicn1 (1,ntrni rsesures
Practice long ngo demenetr'ted the effectiveness of biological control
measures against the root-knet mematede. It is frequently recommended in
order to reduce the harmfulness of the root-knot nematode to introduce intc.
the soil around the trees fertilizers of straw, grass or other plant materials,
The nematocidal action of this measure was explained by an improvement of
conditions of tree growth or by the separation, during the decomposition-of
the introduced. substance, of poisonous gases which destroy the nematodes.
It has been only relatively recently found out that the effectiveness of
plant residue is conditi-ned by the fact that they create favorable condi-
tions for the development ;nf a. larne number of parasites of the root-knot
nematode. Korab and Butkovskii (1939)point cut the harmfulness for the
beet nematode of plowing under -17 sweet clover as a green fertilizer, ex-
plaining it by the fact that the rottenmass of sweet clover is harmful for
the nematode. But the significance of sweet clover is hardly specific and
prebably the matter here is also in the. development.of nematode's enemies
in abundant deteriorating nr.,anic matter. Cur obsarvatiens in the Abkheziin
tobacco experimental statin ',1m.monstrated the favorable effect of "sideratisent'
[enriching land by plantirn legumn crop] on decontamination of plots infest-
0 with root-knot nematode. Particularly marked was the decontamination in
the plot where the tebecce suffered alsn-from soil erosion an. Seo
damaged that it pr-duce -1most After- e cemeleted "slderation?
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111 (mainly with lupine) the tobacco yield was satisfactory
with nematode was very limited
?
and its infestation
Outlines of control measures against the
root-knot nematode
In greenhouses the control measures come down to removing and destroying
roots of diseased plants and to destroying larvae of root-knot nematodes in
the soil. This can be done by partial sterilization of the soil with steam,
its treatment [begin. p. 452] with fumigants and, in an extreme case, by re-
placing the contaminated earth with non-contaminated and by thorough clean-
ing and coating with lime of the walls, corners and bottom of the frames.
Of great importance are prophylactic measures against introduction of
root-knot nematodes into the hothouses. The law of the Five-Year Plan
specifies the development of hothouse farms in the vicinity of large cities
which makes the problem of their protection against the nematode very real;
foci of the latter were found in many points, of the Soviet Union and
probably by no means all of them were disclosed because the damage caused
by the nematode under field conditions further north is negligible. But
when brought into hothouses the nematode can become a most dangerous
parasite and its control will be very costly.
The leading measures under field conditions are agrotechnical. The
decisive factor for annual plants in the control of the root-knot nematode is
the introduction of crop rotations preferably in combination with other
measures: green fertilizers, introduction Of inexpensive fumigants (for
example calcium cyanamide) in arid areab-Olrying of the soil byway of
'additional re-plowings in the summer. In order to reduce the state of in-
vasion cf the soil, the roots of severely affected plants have to be removed
from the field after harvesting and destroyed; it is particularly important
and easily accomplished in vegetable gardens. ?
Methods for destruction of the nematode in growing perennial plants
are not yet known. Higher resistance of particularly susceptible species
and varieties can be achieved-by way of their vegetative hybridization .
with more resistant varieties. Improved feeding of diseased. 'plants by way
of introducing fertilizers decreases the harmful effect of the nematode;
planting between the trees of cover crops, resistant to the root-knot nema-
tode or keeping the soil as a ,clean fallow reduces the state of invasion of
the soil.
A radical decontaminating measure is the planting, in a seriously in-
fested soil, of perennial plants resistant to Heterodera infestation. Such
action was taken in a sovkhoz near Sukhumi: lavander which was in the process
of destruction from "heteroderoSisu was liquidated and the plot was occupied
with Aurantiaceae. After a few years we could not find here any nematodes
at all even though a small amount of them probably remained on weeds.
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BIBLIOGRAPHY .
The root-knet nematode 1.:elens to the number of the r:1::,st d:Lngerous
parasites of agricultural plants and thercf.xe natur.-lly attracted the at-
tention of researchers in all the camtrios of the w7)rld. A limited number
of writings on this subject are scattered in works of institutions of higher
learning and zoploi;ical institutes of the Union's .Lcademies of Science, in
publications on plant protodtim etc. 'ic'de net ot have a poriedical which
would review these writirvs. An andex of cur c ,untry's U tcratur. on plant-
eutinv nematodes for 1936 was published bY E. S. Kir'inova (1939); 1::,t.;:r
writings on root-knot novatode are indicate belw. F...rei;Fn liter-tureca
root-knot nematode is represented uminly by sh-rt notes and rep.,rts, et
frequently on problems of nemat.)do oentr, 1, and they are sct,ter,:: in n'st
diverse publications. The last ,r1.:rh on root-knot (.ssi-y)
was published in 1911 and is r cnlv of histeric-'1 sinifieanee. BiIlicf?-
raphy of older (up t. 1931) w-rks on nematc:ctc of the ronus Hetroera
was published by the En,dis':.. Inprial EI:ru of Agricultural Parasitology
(helminth.qou), 1931. Sinc.:: 1932 and up to the present time the Imperial
Bureau pul-lishes a 1.:ury-11 "Hinth-)logical 1.Lstracts9
which reviews the vrio literature r parasitic worms. From 1930-1933
the bureau puCiis;,,.' bibliographical handbooks on holmintholocy ("Pibliei.;-
raphy of Holminthelqy") which became in 1.934 part of the above-mentioned
journal. [Eor.in, p. 453]
ARKPANGELISKII, P. P? 1937 -
The works of th,, o.wroatino inspection !:).
laratory Of the Narkmzem UzSSR clirit-t 1933-1937.
(Results of invostL..io f. thc, Uzb,.;k SSR in.rerd
to the rcot-k31.0tc.
1 t,,(-hnika, 10-11; 2',57!-.22
?. 145 -
On of comparative parasit.l .o in rlica-
'tion too blood-suckinc arthrp,.-Aa.
K.Dditsin. narasitn14.iia, 12(1):3.-11
-
BLINCVSKII, K. V., 1945
'The CiTet,.2,:r?,,acrfL Cornu)
? - pest :.J7 Trkm6.nlia.
ThrRn..:;115 SSSR, 5-6, 107-.113
BRODSKII, A. L. Z71r,1A3KIA, A. I., 1946 -
P,ioloy. r!or,Jtoc.R.e'-
marieni 0A-nu an it .o ooerol.
Tr. Sect. zooloil.? Inst. botn.
TI7bei.717. SSR, 7-407? ?
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47 ? Trans. A-524
'.7..;raITSKII, I. ii., 1943 -
Ev,.;lution of develcumt:r.? net
the Asearidata Qrtl.cr.
Izv. AN SSSR, sr. nauk: 4, 4,o-OuQ
? ? ? ?
GILIA.ROV,. N. S., 1939 -
Influonce of soil con.J.itions on the fauna (A.' soil
pests. ?
Poch7Q,,,EAnie,, 9, 121-137.
NT'
GORLEI, D., 1933 -
The root-knot nemfitodc. in Turkm,Trii:.
Biull. pc kultt. sukhikh aubtrop., no. 4(36)
Stalinabad, p. 11-16.
GURVICH, G..4., 1935 -
Root nematode (Heterra marioni Cornu, 1879)
(Hetercra radirAccla Greoti.1872)o crops of
the eastern shore of th BlfacOz
siJern, Zooloh-,. Inst. .L.7. SOS'.'f, %L. 5,
p. 317-337, fiL.7' 1-15.
. 194L1
lr:
uoiz
przTi;, s., 194C -
Inmunoiun,?sis fLr..1 it praotic1
Tr. 1;....2.-1.zcq, 4-, 1;7.
EGOROV, P. I., -
The r.-A-knc., marl.ni C.:q-nu)
unfler t.IL co. of the Ukraine SSR.
Prutsi INA-sh. Univ., IV(57):99-121.
ZTMET7In, L. A. (oclit.) 1937 -
Class ro.-,Ind worms (m..matodos).
Textl)u,:)kin zoology, 1,7557-627.
IKS7T3E171,, T. cov: N., 193:9
w.rirk in
;he study f thc. rt-krtc)t
n,:Titoc.1,3 on "1-vanda" 1:?] in.
S. r: :i..
E. Y.:.?
Seltkhozz,-7177,...
YIR'IANCUL, E. S.,. 1939 -
PJ,7iow of acquired kn.pwledge on nematode disest,s
3f pl,:'.nt7, in ti-:c USSR.
rc:LA S-Ta rasteniL
-
Slikft.;z,-iz, 5-26,
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48 Trans. A-524
KIWILNOVA, E. S., 1949 -
The root-knot nematode and its control.
Wedin sessiia Sokts. Zashch: rast. VASKhNIL
i S-Kh, nauk, AN Az-S-SR7IR77.--
Plenum) Tezisy dokladov, 1. Baku, -p. 29-33.
KORAB, I. L, 1949
The root-knot nematode and control rr.al3res aainst
it.
Obfedin. Sessiia Sektsii Zashchity rast. VASKhNIL
i Ctd. Biolor. i S-Kh. Nauk AN AzSSSR.
KOBAKHIDZE, D. N., 1941 -
ObserVati,ms. of ruot nelittode under Tilisi
firms. '
Tr. Zool, Inst. AN Gruz. SSR.
-
KORAB, -I. I. and BUTOVSKII, A. P., 1939
Principal results of study of 17-,ot nematode
(Roterodera schachtii Schmidt) and. of methods
of'its control.
Sh.- rahst po nematodam S-Kh. rastenji pod red.
E. S. KiPianova, -
Sol'khoz4z, 75-120.
_ .
KLECRETOV, L. Er.,
2,ffoctin of rcTtr. of the rubber p10 ot kc,17saghyz?
with the rc,A-knot nomat:xl.
S-Kh.- inst. 9, 123-125.
LEVASHOV,,I.,% Y., 1932
n,..matode in Abkhzaiia and on the shcs
5ack Sea in North Caucasus:
VII Yes. S'ozda po zashchito
Nu.. 5, p. 21.
LEV. SHOV, 0,M., 1935 -
Ehterials for the study of ilout nem.l.tdos ?.1
:ALkhaziia and n tho,shr:res of the 2.1ac3J Sea in
North Cauca&us, sh. Zoled,. Inst.
SSSR, n7:. 5, iT. 301-315.
LTTVI7W, N. r.,
namLt.; in tj:k.. i:.zakh SSP.
po nomtodalf, S-Kh. po0
, . _ .
:L. S. J-,17-ianc7i,
Stl'k % 65-70.
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49 Trans. A-524
LITVIFOVA, N. F. 19391)-
Harmfu1ness of root-knot nematode. under Alma-ata
conditions.
Izv. Kazakh. filiala AN SSSR , 1, 101-108.
NAKSTYDV, Y. A., 1946 -
Groirrth substncos, nature of their action and
practical use.
Sovr. biclogii, 22(2):162-180.,
?fIROVOI Spravechnik
fori agro-climtic gaidoi. .1937.
Gidro motoorologichoskoe izJate1'st71, 1-149.
PAVLOVSKII, E. N., 1945 -
Ecoloi.,,icn1 tendenc$ in Farr:xitoly.
6(2)t65-92,
PLOVSKII, E. N., 1946 -
Handbr)ck on parasitol, ,urns.
Izd. AN SSSR, 1, 1-521.
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Trans .a-525
Abstract and .
Table of Contents
B:
R. Adelman
Verigina, K. V.
Agrokhimicheskii Analiz Pochv v
Laboratoriiakh MTS
[Agrochemical Analysis of Soils
in the Laboratories of MTS]
Izdatel!atvo AktIdemii Nauk SSSR;
Moskva, 1954 _:LtSpPo
ABS TRIICT
Introduction to above title states in part. "The non-black soil belt con-
stitutes 3C% of the territory of the USSR and about 50% of the European part of
the Soviet Union.
"Among the soils widely distributed in the non,vbiack soil belt are poda,01-
ized..soils which form under coniferous mossy forests, peat-podzolized soils
which develop und6r mixed grassy w000dlande, and gray woodland soils adapted to
broadleaf forests.
"In the non-black soil belt, particularly in its northern part, are spread
also podzolized gley soils and peat soils of lowland and upland bogs#,fertile '
meadow soils develop in the river valleys of this aono.
"Podzolized soils cover the northern part of the non-black soil belt. It
is characteristic for them to develop in the upperpart of their proftle a whitish
podzolic horizon deficient of organic matter and nutritive elements. The
podzolic horizon always has an acidreaction, lacks structure, and is composed
chiefly of a fine quartz powder. The magnitude of a podzolic horizon is varied
and fluctuates from a few centimeters ta 40-50 cm. Substances-which are leached
from a podzolic horizon store up lowol-)in the alluvial horizon. The alluvial
horizon usually has a brown coloring,-a solid texture and a heavier mechanical
composition than the podzolic horizon. It frequently possesses an acid reactiOn.
The alluvial horizon gradually turns into maternal rock."
The author further discusses in detail thedifferences between podzolic
soils and peat-podzolized soils, and technics and fertilizers required for their
cultivation. Author explains importance of soil properties, nitrogen sources ?
other than by introduction with fertilizers, role and availability of phospftorus,
potassium and other nAritive elements essential to plant life. Methods for
determination of natural -content of above elements end of soil moisture appear
to be described adeauatel. Methods and procedures for analysis and formulas f
mixing fOrtilizers are worked out, and required equipment is listed in the
appendix.
Translator believes that booklet-containe valuable material, finds text
rather "scientific" and far from easy.
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2
K. V, Verigina
Agrokhimicheskii Analiz Pochv
v Laboratoriiakh MTS
Agrochemical Analysis of Soils
in the Laboratories of MTS
(For the non-black soil belts)
Izdatel 'stvo Akademii Nauk SSSR
Moskva 1954
CONTENTS
Preface 3
Introduction
Briof characterization of soils of the non-black soil belt,?, 5
Method for getting soil samples a d their preparation'for
analysis ' 7
I. Ascertaining total nitrogen content in soil and its motile-fol-M
- ----10
Ascertaining total nitrogen content by Kielidal method, 12
Ascertaining total nitrogen content by Iodltbauer mebEhd--- -15
total nitrogen content by Golubev-mathod - "18
nitrates with disulfophenol acid 18
absorbed ammonium with aid of Nessler reagent 21
motile nitrogen by Tiurina-Kononnova method 22
Ascortaining
Ascortaining
Ascertaining
Ascortaining
II. Ascertaing motile forms of phosphorus in soil 25
-
Ascortaining-phosphorUS'aVailable to plants by tho KirSdnoV
method 27.
Ascertaining forms of phosphorus by the Chirikov method 29
.Ascertaining deficiendy-OraSSitilab10-phoSpherOug-adid - in -the
method of shoots 31
-
III.JAScertaining motile forms of potassiumain soil --
-- 34
Ascertaining motile potassium by the Poivo method
Ascertainin6 potassium available to plants by the Kirsatov-mothod::40
Ascertaining potassium by the sedimentary metric method 43'
IV. Ascertaining ratio: of nutritive elements availavle to plants by
analysis of their juice and external appearance 48
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Ascertaining ratio of nutritive element S.availblt?lOnt
by analysis of juices
Ascertaining deficiency of nutritive sUbstances in soil-by the
external appearance of the plants
V. Ascertaining soil acidity and calculation of lime dosages
48
--55
??
?
Determination of pH with the Aliamovskii device [comparator]
61
Determination of intefohange hydregeh and-actiVe aluminum by
Sokolov's method-
65
Determination motile aluminum by Peive's method...
sp
Determination of hydrolytic soil ccoidity by Kr.ppsh s Method
70
Analysis of lime fertilizers
71
VI. Determination of soil moisture
.74
Determination of soil moisture by drying it at 105?C
74
Determination of soil moisture by Ivanov's method
75
VII.' qualitative methods for identification of fertilizers
78
VIII. Preparation of titrated solutions
70
Appendix. List of-Chemidal equipment and reagent re-toured-Mr MTS
? laboratories 83=
References 86
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1.
?Kirfianova, E. S..
Lakovaia Nomatoda- Ditylanchus allii
(Beijerinck)
[The Onion Nematode- Ditylenchus allii
-(Neijerinck)]
Trudy Zoologicheskogo Instituta v.9
No2 Collection of works on nematodes of
agricultural plants
Edited by E. S. Kirtianova
lzdatel'stvo Akademj.i Nauk SSSR
MOskva 1951 Leningrad
p.513-553 410.9 L543
Trans-A6Y-526
R. Adelman
The Onion Nematode- DitylenchUs Allii (Beijerinck)
(Zoological Institute of the Acadaly of Sciences, USSR)
The onion nematode or the stem nematode of onions- Ditylenchus allii
(Beijerinck) is an Onion pest of grave importance in a number of the European.
regions of the USSR. Thus, considerable decreases in onion yield were noticed
in the District-of Arzamas, Gorki Region in 1938-1940 and in the Penza and Kursk
Regions in 1939. These decreases in yield were due to injuries caused by the
onion nematode.
The wgrk described here was conducted in 1939-1940 upon request of.the
Lenoblplan [Leningrad regional project] and the Gorki Regional Experimental
Station for .Field Husbandry. The war years delayed the publication of this
work, but it is not outdated even at the present time.
Investigations were conducted in Laboratory for Lower Worms of the Zoolo-
gical. Institute, Academy of Sciences, USSR,'in greenhouses, and on the ex-
perimental field of the Leningrad Regional Station for Plant Protection at
Petrodvorets. .Simular work was done at the village of Kichanzino on the ex-
perimental lot of the Arzamas base of the Gorki Regional Experimental Station
for Field Husbandry. In addition, in the Arzamas area, Material was collected
in the following villages: Abramovo, Vozovatovo, Zabelino, Merlin?, in
Krasnoe Selo, and in Pushkarka whore inspections were conducted on kolkhoz
fields adjacent lots belonging to individual farm workers [Kolkhozniki]. In
Leningrad and in Petrodvorets all observations and experiments-were carried out
directly by the authoi- with tho help of a laboratory worker, M. M. GrichoniUk,
and temporary workers. The main place of work at Kichanzlno was the experi-
mental "nematode% base near the. farm which consisted of two neighboring plots
belonging to farm workers, and was treated as ono plot. In previous years, the
farmers had grown their onions without rotation of crops. Their onion yield
dreppod markedly, and in 1938 the ontire onion crop was ruined on both lots due
to its extremely strong infestation by the onion nematode. Thenftupon request
of the Arzamas base, the kolkhoz assigned the above-mentioned farmers adjacent
lots at another-place, and made their original allotments available to the ex-
perimental base. TIrkus, the Arzamas base acquired a lot for its investigations,
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measuring about 0.1 ha which was badly infested by the onion nomatodo. In tho
years of 1939-1940 a series of experiments was initiatcd on this lot for the
purpose of /flaking a study of the onion nematode and of discovering measures for
its control. Our wishos concerning, primarily, the identification of forage
plants susceptible to the onion nomatodo and the development of agrotochnics-for
its contfol, wore taken into consideration when those experiments wore begun.
(Begin p. 513) As a rule, we chocked field observation by microscopic analyses
of which Moro than 10 thousand wore made in Kichanzino and about as any in
Leningrad.
Working Methods. To analyse infestation of whole plants as well as their parts,
we used two methods: 1) examination of the more doubtful tissue section with
binoculars and under a microscope id a drop of pure water on a slide or watch
glass, or sometimes in Petri dishes. The infested plant tissues wore then torn
into the smallest possible bits with the aid of dissecting noodles before or
during the investigation process: 2) nomatodos were extracted from the plants
with the aid of pure raw water (water oxtraction of nomatodos) as follows: the
whole plant or a part of it was first cleared thoroughly of all soil particics
stuck to it (oxpecially if the root system was to be analyzed), cut up into
pieces of 2-3 cm in length, and placed in a glass of water or in-a largo supply
jar Imatorialimaia banka] depending on the amount to be analyzed. Then enough
pure watof was poured into the container, calculated to immerse the bulk of
the plant. Atter 6-16 hours, depending on tomporature conditions in the la-
boratory, the pieces of the plant were removed with pincers from the container,
with the liquid loft to settle for 30-60 minutes. During this process, the
nematodos sank to the bottom. Then the top layer of the liquid was poured care-
fully, and the remaining liquid was pourod into a Potri dish calculated to fill
up one or two of its halves. The contents wore examined with binoculars and -
the nematodes detected were taken from the water with a fine dissecting noodle.
In so doing, the needle was placed under the nematode body at its very center,
perpendicularly to the longitudinal axis; the nematode w6s lifted to the sur-
face of the liquid and removed similarly on fine needles. The point of the
noodle was immediatoly dipped in a drop of pure water on a slide whore the '
nomatodo remained. The prepared compound was examined immediately under d micrc?
scopo, at which time the nomatodes, more often than not, were still alive. The
above described change in the funnel (voronochnyi] method, used generally to "
extract nomatodos from plants, simplifies the work and makes it more convenient.
Tho analybis of soil aimed at detecting the onion nomatodo was implemented
by washing a specific bulk of soil through a sieve made of milling gauze No.18.
About 1 kg of soil was taken from the layer under investigation and stirred
thoroughly; then 25 cm of soil was separated for analysis and placed in a glass
cylinder with an amount of water that exceeded the bulk of the soil by 5-10
times. After 3-4 hours, the well soaked soil blended thormOily with the water,
poured quidkly into the sieve, and washed several times with fresh quantities of
pure water. Then, the sediment was transferred to a Petri dish and examined, in
part-with binoculars. Ncmatodos =re caught with a needle as in the preceding
caso. It is essential to note that there is no need to wash the entire soil
specimen to be analyzed through a sieve, for then its precipitate would be so
largO that its examination would take a groat deal of time and complicate the
work. Tho procedure of washing, on the contrary, must be accomplished so as to
retain in the sieve the smallest possible amount of soil sediment. The end
result will oven benefit from this procedure. A tall glass cylinder, about
40 cm in height should be used for this purpose and the soil should be stirred
in water in this cylinder, for it provides quick rotary motion before draining.
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As tho liquid with the nomatodos is poured into the sieve, large particles of
.soil will settle on the botton and on the walls of the cylindor, and thereby
the sodimont in'tho sieve will be considerably docreasod after the rinsing.
This will facilitato cxatination of the soil sample and will in no way impair
accuracy of the analysis.
Composition of tho onion - Allium Cora L.-- On cultivated onions in the Arza-
mas area, we found 16 nomatodo species listed in table 1. Of those, the
following four nematodes should be added to the list of injurious spcciosi
Aphclonchvidos ho1oN4l4s (do Nan), Ditylcnchus allii (Boijorinck), Pra:...onch121
pratcnsis (de Man), Rotylencl*s Mplticinctus (Coob.). Apparently, in addition ?
to these, the list of onion posts should include the half-parasitic ncmatodo
(Begin P. 514) Aphoienchus aVenao Bastian, encountered frequently in plants in
very largo nUbbors Under field as woll as under storage conditions. Noteworthy
among the nomatodes cited in table 1, is also Euccphalobus olongatus do Nan,
which usually accompanied ApholonohuSaavonac Bastian, especially when onions
aro infected by fusarium or'gray mold. Both of those species arc attracted in
exceptionally largo numbers. We shall restrict ourselves to this brief data
concerning the specific composition of nematode fauna of the onion and shall
report the results of inVestigations aimed at clarifying the biology Of the
onion nomatodo and at developing measures for its control.
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Trans. li.:526
Table 1.
List of'nsmatodos found on onion plants in thc Arzamas area
No. in
consoc. io.rw, of nomatodos
ordor 1
Parts of plant whore found
Bulb
root 1loaf
1 i Acroboloidosbuotschlii (do Man) 4
2 1 Acroboloidos .sp 4
3 ' Apholonchoidos ho1aphillig-0.-e-Edzi):;:- ;
4 1 Apholonchoidos sp t
5 i Apholonchus avonao Bastian. +
6 i Ditylonchus allii (Boijorinck) +
7 I D. allocotus (Steiner) 4
8 ' Pratylonchus pratonsis. (do Man)" 4"
9 1 Rotylonchus multicinctus (Cobb) it
10 Eucophalobus Olongdtug-dOlaihn i-
11 I Cophalobus'sp ' I'
12 ! Diplogastor loritiodlidd-Cldub +
13 i Diplogastor sp +
14 I Dorylaimus sp +
III 15 !
! Parapholonchus psoudopariotinus (M1;--
colotzky) -r
16 Rhabditis 9p 4 t
History!
Thp onion nematode was originally doscribed by Boijorinck (1883) who had
obsorV'ed its infestation of tho onion in Holland under the name of Tylonchus
We find it impossible to ascribe tho honor of discovery of this species
to Kuchn, as some authors (Nowhall, 1932; Chitwood, 19401 havo done it on tho
authority of tho famous work by Ritzema-Bos (i88) ho Aattor actually report
that, in tho years of 1877-1879, Kuehn-had fou.44 trichina: in onions and had .
doscribod it as Tylonchus putrofacions. This report had boon publiShad. in tho
"Ballosbho Zoitung" in 1878 or 1879. .This issue, however, soon bocamO un-
availabilo, clad Ritzema-Bos lost the opportunity to study this article. Hence,
ho writOs in one of his own works that ho. must "also loavp unsettled the
quostioln as to whether or not Tylonch which Kuchri had found in a sick onion
plant afid hdd named Tylcnchus putrofacions n. gp. is idontical with:Tylcnchus
allii n. sp. described by Boijprinck, (begin P. ?515) and :in my invottigation
incorporated as Tylonchug dovastarix Kuehn. I, however, still think that it is
one and tho same species." Hence, there is no basis for crediting Kuchn with
the original description of tho onion nematode, particularly since there are
about 30 nomatodo:spocios which can be found in tho onioni some of wfiich accu-
mulatto in bulb tissues in largo numbers at all stages of .dovolOpmLont. Further-
more, Kuehn himself had given another name to thIc nomatode ho had found in tho
onion, so as to distinguish it from the stem nomatodo specios established by
him earlier Anguillula dipsaci KUohn:rAnguillula devastatrix
lina dipsaci (Kuehn) Goodey, 1932--Ditylonchus: dipsaci (Kuehn) Filipjov, 1934.
Our offlorts to obtain tho "Bblescho Zoitung".proved futile. Thus, tho identi-
ficaticin of the onion nematode with Tylonchus putrofacions described by Kuehn
of study of onion nomatodo - Ditylenchuz allii (Bbijorinck)
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Trans. 526
remains an open question, and justifies our boliof that Boijcrinck's descrip-
tion is the first, fully authentic doscription of the stem nomatodo of onions.
The species name of tho stem nomatodc Anguillulina dipsaci--Ditylonchus
dipsaci, which up to now embraces the onion nomatodc or stem nomatodo of tho
onion, described by us, was established by Kuchn (1858) who had found tho
nomatodo in decaying grains from tho heads of fullor's-toazel (Dipsacus
fullonum Mill.). In a few years he, however, was forced to acknowledgo that
this name failed to imply tho importance of tha post which caused also the well
known stem disease of grain crops, including rye and oats, described by Schwofz
as early as 1825, end an analogous serious discaso of clover and other plants.
Therefor? Kuehn decided to change the original name to a more general one, and
in 1868 he suggested naming the stem nomatodo Anguillula dovastatrix (ravager)
and-hc proved oxporimontally (1869) that it can pass from fullcr's-tcazol to
-rye. In subsequent years the stem disease was described to other crops.
Nitschke (1868) described the stem disease of rye in Germany and named its
causal agent Anguillula socalis Prillioux (1881) described a discasc'of hy?
acinths in Franco caused by tho nomatodc Tylcnchus hyacinthi Prillioux.
Kuehn (1881) described a stem disease of alfalfa
opinion, by a particular nomatodc spccios - Tylonchus
Boijorinck (1883) described tho above mentioned onion
causal agent of which was named Tylcnchuss allii. In
Ritzoma-Bos published a series of reports in which ho
disease of onions-, hyacinths, rye, clovor, alfalfa, buckwheat, and of mant other
plants in Germany. He arrived at the conclusion that the causal agent of tho
widely spread stem disease of many crops was one and the same species - Tylon-
chu dovastatrix (Kuehn), identical with the'Spocios causing similar discascs in
different Crops described by various authors. Thus, ho believed that Anguillula
dipsaci,"A. dovastatrix Kuehn, A: sccalis Nitschko, Tylonchus putrcfacions
Kuehn, T. hyacinthi Prillicux, T. havonstoinii Kuchn, and also Tylonchus
askenasyi cxtractod'from moss (Buotschli, 1883) arc synonymous with Tylonchus
davastatrix (Kuehn). It scorned that the oxporimcnts of Ritzoma-Bos fully con-
firmed his hypothesis concerning the identity of those nematodes. He infested
onions-with nomatodos taken from hyacinths, and hyacinths with ncmatodes from
onions. All of this justified tho assumption that tho stom nematode T. davasta-
tkix (Kuehn) is tho causal agent of discus? in 34 plant species.
Tho opinion of Ritzoma?Bos is generally accepted oven at the present time
but the spocics name suggosted by him did not take root and was displaced by
tho name Ditylonchus dipsaci. The generic name had-boon changed earlier in
connection with tho work of Bastian (1865) (begin P. 516) who had established th
gonus Tylonchus, and IiRon suggestion of I.N. Filip'cv (1934), the name Ditylcn-
chus was used instead. At tho present time, the name of tho stem nomatodo used
most extensively is Anguillulina dipsaci (Kuchn) Goodoy, 1932 and Ditylonchus
dipsaci (Kuehn) Filipjov, 1934-1936. Tho list of hosts of this nematode in-
cludes more than 300 plant species, among them many cultivated plants: onions,
oats, rye, maize, potatoes, boots, clover, alfalfa, 6.nd wild floral chick-wood -
Stollaria'modia Cyricl, bcdstraw - Gallium aparino L., snowdrop - Galanthus
nivalis L:, bindwocd - Convolvulus arvangis L., Stornborgia - Stornbcfgia lutoa
Ker.-Gawl., oracho - Chcnotiodium album L., meadow grass ? Poa-annua L., couCh
grass - kgropyrum ropons P.B., crowfoot - Ranunculus ropons L. and so forth.
Tho vastness of this group of hosts can be soon in the works of Schneider (1937)
who discovered tho nematode Tylonchus dipsaci var. tobacnsis in the galls of
in Germany causod,-in his
havonstoinii Kuchn.
disease in Holland, the
the years 1887-1892,
described tho nomatodo
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6 Trans. A.4 626
pond wood - Potamogcton malayanus and parrotfcathcr - Myriophyllum spicatum in
Southern Sumatra, and in the works of Zoller (1937) who found it in the same
plants in the vicinity of Vienna. According to literary data, the stem nematode
has a world-wide digtribution and its hosts are not only surface plants, but
also aquatic plants. On onions it lives in very many countries; besides Hol-
land and Germany (Beijerinck, 1883; Ritzcma-Bos, 1888), the nomatodc in found in
Franco (Chatin, 1884), in England (Gbodoy, 1933), in America (Laidlaw and Pricc,
1910, and in Southern India (Barber, 1905). In recent times, Thorn? (1945)
-segregated the potato stem nematode as a separate species - Ditylonchus destruc-
tor Thorne, 1945. In thc USSR, the onion nematode was first mentioned by Fokin
TU24) who, without seeing the nomatode, attributed to an onion disease in
Viatka (now Kirov). Onion injury by the stem nematode in the CSM attained full
authenticity in '1935 when we received diseased bulbs from the "mint" sovkhoz
in the Rostov Region and from the kolkhoz named for Kaganovich in the Penza Re-
gion. In 1936 we received bulbs infested by -worms from the storaLo rooms of the
Leningrad Fruit-Vegotablo Marketing EntorPriso [Lonplodoovoshchtorg] and from '
the Pella Station of the Leningrad Region. NO noted with interest the very larE
dimensions of the worms as compared with those from potato tubers. Investiga-
tion confirmed the fact that specimens of the stem nematode from potatoes of
various geographical points arc considerably smaller than those obtained from
onions. This difference in dimension was obsci-ved in comparing adult specimens
as well as in comparing larvae phascs and cgs. In addition, permanent dif-
ferences in the nomatode wore obsorved in the position of the vulva and in size.
alpha (relation of body length to its width). There existed obvious differen-
ces between the stOm nematodo of potatoes and that of onions; a report to this
effect has boon made (Kirtianova, 1939). At that time, almost nothing was
known about the biological differences between those nematodcs, save Goodey's
indications that in England, onions, wheat, barley, ryo, oats, fodder, boots,
turnips and peas grown on a lot infested by these posts, wore not affected by
the potato nematode race, although, as per his van romarks; this could have boon
due to insufficient infestation of the soil (Goodcy, 1935). No One had under-
taken a thorough study of the 'onion race, and it was, and by manSratill is,
considered to be a parasite preying on various cultivated plants. Tho few au-
thors -(for instance, Steiner and Scott, 1934), who endeavored to subdivide the
species Anguillulina dipsaci into several varieties-, also listed the onion
nomatado together with many other forms under one variety A. dipsaci var. com-
munis. (Begin P. 517)
Morphology. Tho onion nematode (table I, 1) is characterized by a well-propor-
tioned-thread-liko body. On the avorago, adult specimens, reach .1500 micron in
length. The head and tail ends of the body are visibly narrowed down, with the
tail ending in a point. The rialos are a little finer and considerably bettor
proportioned than the females. Even Ritzoma-Bos had noted that the worms from
onions were visibly larger than those from rye and hyacinths, and he cxplainod
this fact by the abundance of nutritive substances contained in the bulbs.
According to data by Ritzcma-Bos, in Germany, males of the onion race dre, on
the average, 1510 micron in length and females - 1540 micron in length. Ouf.
measurements of ncmatodes from onion bulbs produced close figures (Table 2).
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Dimensions
7
Table 2.
of onion nematodc Ditylonchus allii (Boijcrinck,
Author
t Material
received
: from
material from different places
526
1883) as per
,
Sox Body Body
? longth :width
(
' (in u) in u)
Alpha
Beta
Gamma
Ritzoma-Bosi Germany
1887
Kir'ianova
1936
Kirlianova
1939-1940
- _
! 1430-173p
.?1a1 (1540)
c , 1430-1570
(1510)
Poll Sta-
tion, Lonin-4
grad Region 1(1:2
Kichanzino
village,
Gorki '
Region
1392-1680
(1525)
139?-100
(1476)
37-47
(41)
41-50
(45)
28-37 44.9=56.2
(30.7) (49.8)
,20-25.7.54-78
i(23.7) (63)
! 1130-1924 30-43
(1440) (34)
.1190-1685 20-30
(1405) (25.5)
5.5-7.3
(7.1)
6.1-9.3
(7.3)
12-17.5
14-16.5
11.8-15.7
(14)
13.9-17
(15.4)
34.8-55.4 6.7=8.6 14-20
(42.5)- (7.6) (16.7)
44.8-74.2 6.2=8 14-18.6
' (56) (7.2) (16)
Remark: Average sizes citcd in parentheses.
Th C basic bulk of measurements was accomplished on specimens from tho August
generation. We found the largest female specimen (1924 u) in tho September col-
lection, and the smallest (1130 micron) - among worms gathered in July from shootE
Preliminary observations give the improssion that the Juno and July generations
havo somewhat smaller dimensions compared with the August and Soptomor ones;
this apparently can be explained by tho difference in. temperature. Regardless of
fluctuation in dinen6ions,?tho onion nematodo is relatively lar:cr than-thc stem
nematodo of potatoos. Besides, it is distinguished by a shapelier body. In tho
onion nematode the proportion of body length to its width (alpha) is on the
average over 40 in females and over 45 in males (Table 2); in tho stem nematode' o1
potatoes an avera,e alpha equals 33 in females and 42 in males. The dimensions, ci
larvae of the fourth-ago grown on onion arc twice the size of larvae of tho stoma
ncmatodo of potatoes. In the onion nematodo, the size of eggs, too, is larger.
Having noted the large dimension of the onion variety, as compared with the di-
mensions of varieties obtained from rye and hacinths, Ritzoma-Bos (1888) at-
tached no importance to thcso differences. (Begin P. 518) Ho bolicved also that
tho racos-from ryc,'hyacinths and onions die not vary essentially as to bursa
structure. He did, however, point out that in males grown on,onions the bursa
has somewhat smaller dimensions and that it is not as near to the tip of the tail
as in those grown on rye and hyacinths, and that the bursa varies markedly in
males obtained from different plants: it can oxtond to-tho very tip of tho tail,
completely encircle the tail, or reach only its center. We are unable to check
Ritzoma-Bos' .data concerning bursa structure in the stem nematodo of rye and
"'hyacinths, but we can report that, according to our observations, in males of the
. onion nematode, the bursa in most cases, does not roach the tip of the tail by
27-42 micron, at a tail length of 75-90 midron, and encircles tho tail wholly or
comes close to its end only rarely.
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8 Trans. AV:-526
The nematode body is covered by a thin, translucent, transversely streaked
and exceedingly tight cuticle. Lt the centr Of the head and is situated the
oral aperture encircled by 6 merged, and hence little noted, lips with 6 lip-
nipples. The oral cavity is equipped with a snail, strong spear expanding knob-
like at thc basc; its size is-10-12, more often 11 micron. Tho esophagus has
the appearance of narrow tube- with a small oblong nuscular bulbus, behind which
the nerve ring is situated. The posterior part of thc esophaLus'is somewhat
expanded and has tree esophageal glands, One. of which opens at the base of the
spear'and thc other two within the bulbus.- The length of the esophagus varies
mostly within the limits of 180-220 micron. -There is a vory clear demarcation
lino between thc esophagus and the intestine. In live worms, thc head end of H
of thc body stands out sharply with its translucent contents in the area of the
esophagus, extending to whore it borders on the intestine. The intestine is
characterized by contents of largo granules of grayish yellow hue and, therefore
stands out well among thc other organs. It consists of a, long and wide tube--
the central intestine and small posterior intestine,, ending with thc anal outlet
on thc ventral side near the posterior end of the body; in females, the latter
opens out apart from thc sexual one, but in males it is combined with the latter
and opens into a common cloaca.. .The sexual organs aro upaired and stretch along
thc body in the form of long, straight tubes beginning near thc anterior section
of the intestine. Thus, only their anterior sexual tube is well developed.
In females, the rudiment of the posterior sexual tube is represented in tho
form of-a-posterior uterus performing thc role of a semen roscptaclo [some-
priomnik]. The vulva is pushed' far to the back and is at distance equaling .81%
of thc total length from thc head and of the body. Spicules are paired,
slightly curved and expanded at thc tasc; before their narrbwing down, they.
form two identical node-like islets set toward the interior; Their length
fluctuates between 22 and 32, with an average of 27. microns. The ferrule
[Ruleki is represented by a short, t'implc, slightly curved rod, 11-12 micron in
length. Prcanal nipples are absent. Tho eliminating pore is located in the
af-ca of thc posterior segment of the esophagus behind the nerve ring. (Begin
P. 520) Thus, the onion nematode, possessing a series of organizational
features common in thc stem nematode of potatoes, is distinguished from it by
some permanent morphological characteristics: by body length of about 1500
micron by dimension alpha which in females equal 40-50, and in males 45-63,?
by the position of thc vulva which equals 81%, and by thc rlatively short
bursa which, in most cases, fails to extend to the tip of the tail by 22-23
microns. Those distinguishing narks together with thc nematode's narrow
specialization in a definite subsistence plant - the oriion, qualify it as a
separate species, Ditylcnchus allii (Beijerinck, 1883).
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9 Trans. L: 526
[Page 518, bottom]
Table I. Onion ncmatodt - Dity1onchu6 (Bcijcfinck) and injuries caused
by it. (Illus. by N. Y. Kondakov).
1. - Onion ncmatodc. From left to. right: thc head end, tail of fcmalc, pos-
terior end of female body and two eggs: II - initial.stagcs of infestation of
seed-onion bulbs by onion nematodc. Intensely white spots on first three
bulbs indicate places of nom-Aodc penetration and concentration in bulbs
medium age (midsummer); in last two bulbs nomatodcs penetratod at point of
growth of projecting now rootlets at a later stage of. the .plants' develop-
ment. Infested points of growth arc abnormally enlarged and have an intensely
white color; III - Oblong'splitting of garden onion bulb heavily infested by
onion nematodo; when bulb dried Up, worms moved to surface in the form of.
a cream-colored incrustation (indicated by arrows); IV - small piece of cream.7.
colored crust composed of females, males, and larvae of onion nomatode;y
sprouts of seed-onion (seeding of chornu8hka), (left) two healthy ones and
(right) those infested by onion nematode. Infested plant 6 are deformed
distended and, in most cases, have no seed capsule at tip.
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[Page 519 contains illustrations of onions ancl?of organisms.]
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11 Trans. 44-526
DEVELOPYFUT AND MODE OF LIFE
Development. The onion nematode hatches eggs in leaf tissue as well
as in the meaty scales of bulbs, obviously, not in thc root-system. Larvae
and adults arc, however, likely to be found in root tissues. Upon leaving
the egg, larvae differ from flaturo spociemont only by their dimensions and
the absence'of sexual organs. Aftcr a few moltings, they aro transformed -
into adults, Following copulation, the females hatch eggs, and generations
follow continually one after another, so that the worm can always be found
in the tissues of a-host in all stages of its development. Correlation be-
tween males and females equal 1:1. Duration of-the developmental cycle has
not yet been sufficiently invostigatod, but there is reason to boli:.vo
that it lasts one month, on the average, and is retarded or accelerated in
relation to conditions; the latter have not been sufficiently clarified, but
laboratory observations indicate that optimal tomporaturo for the develop-
ment of Onionnematodos lies within the limits of 10-14? C, and egg hatching
can occur even at 9-10? C.
If an infested plant dies, the-nematodes leave it, enter the soil and
search energetically for a now host. A necos8ary condition-for their re-
moval and migration is humidity close to 100%. If humidity is low, they re-
main in the tissues and dry up together with thorn. In such a case, their
body assumes the form of a spiral wound sever61 times; with the coning of
,beneficial moisture, the body straightens out. All phases can be subjected to
drying without endangoring-life. The duration of a stay in a bulb in a dried-
up state is very extensive. Goodey 0933) succeeded in reviving worms which
had spent two years in a dry bulb; in our experiments, all phases survived
drying within seed-oiion for 15 Months and resumed activity in 2-3 hours,
after being transferred to water. Nonetheless, an attempt to roviv6 nematodes
which had spent 4 years and 8. months in a dry bulb enddd in failure. Ana.
biosis in nematodes can occur only if the air surrounding the ditintegrat-
ing bulb or othor parts of the infested plant is well ventilated. If, how-
ever, the infested plant is subjected to fermentation and decaying prior to
drying, then the nematodes will perish, for they cannot endure the injurious
action of decomposed tissuo 15roducts.of the host plant. Such decomposition
occurs mostly during storage. If the dying-off of bulbs occurs in relatively
dry storage, then the nematodes crawl out in masses,and accumulate on the
surface of bulbs forming a pale cream-colored, porous coating or little
lump (table III) composed wholly of worms in different stages of develop-
ment. Tho bodies of the worms arc wound in spirals and-aro entwined with
each other so tightly that they form close little knots. The removal of
nomatodcs was observed also on the wall of a glass cylinder [Bogin P. 521]
?containing sick bulbs (table III, V), on paper in a herbarium pastoboard box,
.and on the surface of onion tops of herbarium specimens of seed onions; on
the surface the loaves had not been injured, but as the nomatodos wore
leaving the bulb they formed a nodular coating on the leaves. Finally,
many nodular formations compOsed of nematodes were notod on the surface
of an entire bulb of a mother-onion [lukamatki] from which the jacket scales
(choskui rubashki] had been removed preliminarily: evidently, nomatodcs
are capable of getting out not only through cracks, but aldo through un-
damaged tissues of coverings where they are not voryugh. The ability of
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?
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the onion nomatodo to remain in a state of anabiosis not only within tho dry
parts of a plant, but also without them, had remainod unnoticod. The re-
moval of nomatodos onto the surfaoc of a plant, similar to the one desdribed
by us, has boon not-d solely by S. Rostrup (1926) who had obsorved the exit
of last phase larvae through the crack in a narcissus bulb: In leaving, the
larvae also famed a white or cream-colored porou6 coating; Rostrup trans-
forrod those larvae onto an onion and infested it.
The duration of nomatodc viability in the form of the described coating,
composed entirely of worms, is not known; At can be assumed that it lasts at
least-15 months* Since that time period was the deadline in our experi-
ments. If infostod dried up bulbs are shaken up during sorting, or under
other circumstances, the nonatode coating can easily crumble and spill onto
healthy bulbs whiah arc infested by the parasite when favorable moisture
conditions set in. In the soil itself, the worm, supply is not big; ana-
lyscs conducted at various times'at depths from 0 to 60 cm have shown that .
in the summer time only-singlo'worm specimens aro found in the soil, ond only
in horizons above 20 cm. Infestation of fields by the onion nematodc nay
be due to planting slighly infested bulbs, and also to the introduction of
worms, in a state of anabiosis (in the form of the described coating), which
are dropped in the soil accidentally together with healthy bulbs. Another
sourco of infestation is thc post-harvest waste (dry scales, tops etc.) has
boon removed, the percentage of infested plants and bulbs is considerably
lower than that on test lots where it has not been removed (tablo 3). It
must be taken into account that infestation may be duo to nematodes which
inhabit the soil itself, for there they can dispense with food for an ?
extended time. In laboratory experiments, with humidity close to 10 0 and
at a temperature of 9-12? C, nomatodoa wore without food, yet in a state of
continuous motility.for over 3 months. It can bo assumed that in nature
this time-period is oven longer thanks to the cdse with which nomatodos
drop into a state .of anabiosis and revive again. To illustrate this.fact,
we shall cite the following experiment. On October 23, 1939, about a thou-
sand nomatodes which prior to that date. had been about a month in an aria-
biotic state in a bulb, were placed in a drop of pure water on a slide.
Two flurs later, the worms wore already moving around energetically and
some of the females were hatching eggs; soon the water eralOoratod, the
worms ceased moving, and stretched out; on October 25, i.e., two days later,
a small amount of water was poured on the slide and all worms revived within
1-2 hours,-with the larvae reviving before the adults. The experiment
was repeated on October 29, November 15 and December 9, and mass movement '
of larvae and adults took place; in all instances only on November 15 and
December 9 a part of them died'off; the experiment was conducted in a'room
whoro the temperature was 10-12? C and the humidity of 51%. [Begin P. ?522,
paral]
Distribution of the onion nematode is, undoUblodly, aided also by seed.
In the year 1888, Ritzcma-Bos established that infostation of dark-onion seed
can roach 3%. According to an oral report of N. M. Sveshnikova, infostatin
of onion seed in Kichanzino in 1938 was a little more thdn 1%; according to our
observations, in the same place in 1939, it was under 0.5%, and in 1940
the ncmatodo was found in but one seed out of 2011. Infestation of the ovaries
is somouhat'groator; of 727 sterile flowers, the nematode was found in two,
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(D
0
w
CD
Trans. 11. 526
12 A
Table 3
Influence of post-harvest waste on degree of seed onion infestation by onion nematode in flollowing
crop year (Kichan:zino , Arzame s District, 1940)
0_
73 c)
o
CD
U)
0,
H
,
C)
-n
o d
(D
(T)
(D
n.)
o.)
0
19
0
-0
co
0
0
n.)
n.)
May
19
May
Aug.
1
Aug.
1
1 State roil
post-har-
vest waste
Removed
in
September,
1939
Not re-
moved
Ho. of
test
lot
Total
Total
Total
collected
Among them
Healthy
Diseased
plants
61
58
119
66
57
123
bulbs
109,
108
217
plants
43_
21
62
118 3
105 6
223
9
bulbs
80
43
123
5
24
plants
37
57
63
51
114
Remarks, 1' or analysis, the . nematodes were alloted 5 linear meters diagonally in each t es
0
(D
0
(D
-a ?
% 'of infested
(D
'bulbs plants bulb m
29 34
65 64.6
94
113
81
194
o .
47.5
95)4.
89.4
92.6
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one in each. The membranes of seed pods wore infested.the most; they wore .
found to contain 15 nematodes out of 1369 analyses (table 4).
The slight infestation of seed in the year 1940 can be explained by tho
large number. of sunny days during thc formation and ripening period of seed.
Tho nematode avoids direct sunrays; therefore-surface parts are less invaded
than bulbs even in cases of heavy infestation. General infestation of seed
depends on meteorological c5nditions and its rate will be highest- in years
with rainy and cold summers.
Capacity for active motility. It is claimed that the stem nematode
has considerable capacity far active motility. Slogteren (1923) established
experimentally that the stem nematode of hyacinths can cfawl upward through
a 90 cm layer of sand and infest hyacinths planted in it. I have.conducted
repeatedly the following experiment. I placed a heavily infested bulb on
the bottom of a glass Cylinder and covered the cylinder with a fitted stopper
[pritertaia plastinka]. Inside the cylinder soon developed excess moisture
from the evaporated liguid of the bulb, and the nematodes reacted to it by
gradually crawling out. Soon they moved in masses onto the bottom and the
walls of the cylinder. %thin-a few hours, part of the nematodes were
reaching the top quite rapidly. Thus, in a cylinder 40 cm in height ald at
an air temperature of 10-12? C, the nematodes- climbed 17 cm in thdfirsttwo
hourn andin-a-fahmore hours they had 'reached the rim; [Begin p. 5231 their
density on the walls which was at its thickest at the base kept decreasing
as they were approaching the top; this-was duo to the fact that new batches
of worms kept crawling out of the bulb. It is interesting that even in over-
cast autumn days the nematodes crawled exclusively on the side exposed to
light, forming-on the glass a unique pattern of many thousands al' worm
(table III,-V). On the side of the cylinder not exposed to light, worms
were absent. If the cylinder was turned, the nematodes immediately crawled
over to the wall exposed to light, and during an experiment, they did
this many times over on overcast as well as on sunny days. Nonetheless,
nematodes react negatively to the action of direct sunrays and they crawl
rapidly to the side of the cylinder not exposed to them.
The nematode is capable of moving fairly rapidly in soil as well; this
was confirmed by our observations in one of the greenhouses at Potrodvorcts:,
infested bulbs wore placed in empty fluw-er -pots that were sot under a rack
on which experimental young crop of the dark Onion was laid .11.t; neither
the soil nor the onion seed had been infested. After 2-3 months it was
proved-that the nematodes had crawled into the rack and infested the onion
plants. Thus, they had crawled no less than 50 cm and, indidentally, they
did this during two days of h6avy, incessant rains ,when air moisture in the
green house was close to.10Cfg.
Conditions of temperature. The action of temperature on nematode ac-
tivity has been studied but little. On the basis of some of Our observa-
tions and experiments and also on information available in literature, it-can
be assumed that relatively low temperatures arc the most favorable for it.
Thus, the greatest motile activity of the worms was observed at 0-12? C;
these observations are, however, incomplete and approximate. At a tem-
perature of 9-12? C, the nematodes not Only climb upward on the cylinder
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walls, bdt they dispense with food for quite a long time without losing
motility. In contrast, the observations at Kichanzino showed that at 30?
the worms moved considerably slowert after Having the bulb, it took them
2 hours to climb no more than 2-5 cm on the cylinder walls, and they noor
reached tho top, although the height of the cylinder measured but 15 cm.
There is also the possibility that, in addition to high temperature, adverse
action was exerted on tho nematodes by extremely bright summer light. At
a 30? temperature,? the worms remained visable within tho cylinder no mote
than 2-5 days, perishing as a result of tissue fermentation in the bulb.
The fact that moderate temperatures are more favorable for the onion namatodc
has boon?confirmcd.by artificial infestation of bulbs under laborat6ry condi-
tions. The most typical picture 1' disease is obtained at 10-12? C. In tho
field, the process of decomposition of infested bulbs proceeds more typically
at 9-14? (manifestation of whiteness and porosity of tissues of internal
scales). In instances of high temperatures, the bulbs decompose extremely
quickly, but mainly as a result of rapidly-proceeding fermentation within them.
Law temperatures delay decomposition of bulbs, but disease shows gra-
dual progress even at 1-3? C, although it develops at an obviously slower
tempo. Neratodo motility at 1-3? is slaw, but it docs not stop complotely.
Nematodes from three bulbs in the cylinder at a temperature of 14?, crawled
out and out and concentrated on a wall in nodular which rosembold very much
the pale cream-colored Coating that forms in onion storage rooms when
infested bulbs grow dry. The nematodes remained in this state about a
month, but just as soon as tho temperature had-risen to 7=8?, they separated
and crawled all over the wall exposed to light. .(Begin P. 525) Thus, tho
nematode easily survives a drop in temperature to 1?, retaining some, if
over so limited, motile Capacity. Normal motility, however, requires tem-
peratures not under 7-8?. Whether or not temperatures below 0? are detri-
mental and, if so, to what 'extent, is difficult to say: no experiments were
conducted in this direction.
We did, however, have a heavily infested bulb that had been in pit
storage [burty] under thc snow in Leningrad all wintei--long; efforts to
revive nematbdes taken from this bulb ended in:fkilure. From the looks of
the bulb it could be concluded that it was frozen and that the nematodes
apparently, had perished with it; nearby, healthy bulbs had not been damaged.
In contrast to this observation,. agronomist M. V. Krylova of the Arzamas
baso (oral report) asserts that nematodes had .survived the very severe
winter of 1939-1940 in infested bulbs abova ground at the experimental base,
as well as underground a depth Of 10-50 cm. Unfortunately, in Krylova's
experiments tho degree' of nematode motility after overwintering was not
sufficiently verified. Thus, the negative influence of temperatures below 0?.
on nematode viability remains an -pen question, but it is clear that their
ability to survive a sharp rise or drop in temperature depends to a con- -
siderable extent on tho moisture in their immediate environment. While in a
state of anabiosis, the worms, naturally, arc more rcsigtant to sharp
fluctuations of temperature than during normal activity. . Concomitantly with
the above, the surprising adaptability of nematodes to drying propts?the
hypothesis that they aro no los resistant to the action of law temperatures,
and that this circumstance may be connected with a definite phase in their
cycle.
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14A
Trans. A. 5
Table 4,
Results of analysis of onion nematode - Ditylenchus allii.Beij. - infestation of onion seed gathered
from heavily infested plants of the 1940 yield
Consoc. order
1
Degree of infestation of
different parts of
mother-onion plant
?r-
Bulb
Shoot strela.
Total
ovaries
ana-
lyze-d,
1 'JPper part Lower part
Heavily in-
fested
throughout
Same: bulb
entirely
destroyed
No nematodes found
Same
3 .Same . Nematodes
nematodes comparativly
few
Entire bulb No ncma- Very many
heavily in- todes nematodes
fested, with ,found
'
1 -
erp-craxk
A. it
p ?
bm,
?nck
5 ' g6110,.wi.thout single
r dralc nematodbs,
pnly half of
1 bulh heavily Ho
'InfeSt-ed nematodes
No nematodes
MLIV
nematodes
Total
116
92
98
S47
7)44
g41
2738
Degree of seed infestation
0
CD
CD
Infestation dogrca
of seed pod mem->
brane
8 -
of thee
o
Total
mem-
0 o
branes
o
ana-
lyzed (T)
ow
4.) w
CD
_
n.)
- No anal sis 0.)
co
including
those with
nethatode
in seed
those with
nematode in
sterile flower
1
3
116
92
98
81-1-5
7)43
841
2735
92
-- 78
1
1
os
466
516
760
---------
2
2010 2
214
14
379 466
227 517
81 386
725 1369
f t
le
5
15
If
(-)
0
-0
co
0
0
cb
1354
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15 Trans. A: 526
The problem of temperature influence on the development and viability ,
of the onion nematode calls for experimental investigation; its solution has
a direct bearing on the development of control measures, particularly on the-
extablishment of an appropriate regime of temperature in onion storage rooms.
INFLUENCE OF ONION NEMATODE ON PLANT ORGANISM.
With reference to the older works clarifying more fully the influence
of the onion nematode on the organism of the host-plant, recognition is due
the work of Ritzema-Bos (1888) who offers a fairly good description of the
disease in very young sprouts of the seed-onion and in older plants of the
garden onion. In recent works, this question is accorded the greatest atten-
tion in the investigations of Chitwood and Newhall (1940) and in an article
b'y the same authors written in collaboration with Clement (Chitwood, Newhall
a. Clement, 1940).
We have succeeded in obtaining more complete data concerning the nema-
tode disease of onions under field conditions as well as under'storage con-
dition and, therefore, find it necessary to dwell on them in greater detail.
Our Observations and experiments were conducted under natural conditions
in the years 1939-1940 on young crop at the Arzamas base of the Gorki Re-
gional Experimental Station for Field Husbandry in the villae of Kichanzino.
In addition to these, experiments wore conducted in greenhouses and on the
experimental field at the Leningrad Station for Plant Protection in Petro-
drorets. Besides these, some of the experiments were carried out in the
laboratory for Lower Worms of the Zoological Institute, Academy of Sciences,
USSR, at Leningrad. These observations show that the disease manifests
itself variously, depending on the developmental phase of the plant and the
place where the nematodes penetrate its-tissues, and, at the same time,
has a series of general characteristics. [Begin P. 526]
Disease of seed-onion, Nematodes can penetrate onion seed (onion-
chernushka) at the earliest phase of their germination, as a resiAA of
which part of the sprouts do not oven appear on the surface, ds was de-
monstrated by Ritzema-Bos (1888) and Laidlow and Price (1910). The very
sparse plants that do appear are sharply distinguished by their dwarfish
growth and abnormally distended, frequently twisted shoots (table 1, V).
In healthy plants, the cotyledons always bear an empty sced capsule at the
end; sick cotyledons, however, throw it off considerably earlier, and,
therefore, it is often absent in the sprouting first leaves. Shoots of
infested plants have at various places irregular swellings, more often at
the base, but sometimes also somewhat higher; in the latter case the young
plants soon cease growing altogether. The swollen part of the shoot be:ins
to dry at both ends and gradually becomes diseonnected from the root, but
retains its green color and continues to live. A large amount of worms
develops within it hastening the process of its decomposition. In some
cases sick plants continue to develop, but fallbehind control sprouts and
perish within one-two months. Plants infested but slightly live a little
longer, but the Symptoms of disease are similar: retarded growth, thickening
and malformation of shoots, shortening and thickening of internodes (table II,
I). Those infested at a later date, for instance, during the formation of
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16 Trano. k-626
bulbs, vary but little from healthly plants at the beginning of the disease;
if, however, such a plant were pulled from the ground, then an intOnsely white
spot indicating the locale of nematodes would be found on the bulb. The
worms infiltrate a bulb mostly at the bottom, hence this spot is usually
near the bottom, bordering on it with one of its edges. However, in view of
the softness of bulb tissues in young plants, nematodes can penctrato them
anywhere and the white spot can be observed at different places (fig.1).
Tho dimensions of the spot umistakably, indicate the deLree of infestation
since they arc determined by the amount-of nematodes that have infiltrated
the bulbs or have developed within them. Sometimes infestation occurs via
the leaves, through the stomata of which the nematodos make their way to
the internal layers and then migrate within the bulb. Damaging the deeper
a8 well as the upper scales of a bulb, the werms populate it evenly, (Begin
P. 627] as a res:-lt of which the whole bulb assumes an intensely white
color. The nematodes dostroy dhiefly intercellular tissue substances, but
they often damage the colls;as well; the latter become separated from each
other and acquire the form of a porous, brilliantly sugar-white mass, as-
suming later on a light brown hue. More or less deep cracks form during
this period or earlier, particularly if infestation is uneven, and break
jacket scales and the open succulent little scales lengthwise from the
bottom to the neck. Very often, cone-shaped closed bulb scales, project
themselves cut in the forms of a deep cleft and, together with the buds
which they protect, they form a likeness of nodes on the bulb surface.
Sometimes these protruding scales split in turn longitudinally, acquiring d
resemblance to the tuft from small separate scales of leaflets, (table II).
Such a picture is usually observed in cases of concurrent infestation of
bulbs by nematodes and the onion fly. Though they continue to stand up-
right, the sick plants arc gradually swamped by countless nematodes which
eventually destroy their tissues, and in 1-3 months the plants perish.
Tho tissues buried in soil decompose and disintegrate into a powder; the
surface parts (leaves) dry up rapidly in the sun and wind. The nematode is
found in large numbers in dried up loaves. If infestation occurs at the very
end of the vegetative period, then the picture obtained is some7rhat different
and the cOursto of the disease is typical of that characteristic for the gar-
,
den onion.
Fig. 1.
Sprouts of dark onion injured by onion nematode. Two upper boxes
Kaba variety, lower - Arzamas variety. The two boxes ontho
loft were infested infested artificially with the onion nemtode
at seeding, the two on the right - uninfcated controls. All in-
fested plants perished at an early ago: first the'lcss fosistant
Arzamas variety, and then Kaba variety. (Illus. by T. A.
Gavrilova from photos).
?
The rate at which nematodes penetrate young onion sprouts is very high,
they can accomplished it in a few hours. Thus, by introducing nematodes
into the soil containing onion sprouts in experimental containers,-a con-
siderably number of worms can be found in the plants after 24 hours.
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17 Trans A:526
Infestation by the onion nematode affects the nutritive properties of
onions. Upon instruction of farm leaders [kolkhozniki], infested bulbs are,
used almost exclusively for cattle feed and in the preparation of "lukovniki"
(a local dish consisting of fihely-minced onions with a small admixture of
flour baked in a Russian oven). Infested onions are not fit far other types
of food; they impart to cooked food an unpleasant odor.
A comparison of the nutritive properties of plants infested by the onion
nematode and of healthy plants used in food in the form of green onions has
shown that the sugar and nitrogen cOntent in healthy plants is somewhat
higher than that in infested plants. The results of a chemical analysis of
the nutritive properties of infested and healthy plants of the seed (alien,
harvested in 1939 at the experimental base in the village of Kichanzino,
are cited below. Figures represent percentage of substance content:
Invert Saccha- Mois-
sugar Sacharose 'Nitrogen Cellulose rosity turo
Onion infested
by nematode
1.78
7.42
0.49
0.89
9.3
84.4
Onion nOt - - -
fasted
1127
10.39
0.52
0.73
11.66
82.41
These analyses were conducted by the Agrochetical Laboratory of the Gorki
Regional Experimental Station for Field Husbandry. [Begin P. 528]
Disease of garden onion. Nematodes have two ways of penetrating plants
of the garden onion as well as those of the seed onion: throt0a the bulb and
through the leaves. Depending on meteorological conditions, one way of nema-
tode penetration may hace its advantages over the other. In times of Short
but heavy rains followed by bright sunny weather, nematodes generally pone- '
tratc the bulb near the bottom at the points of growth of sprouting rootlets.
In this case, the first syMptom of infestation is a marked swelling of the
points of growth which enlarge to such an extent that they are 2-3 times the
size of the neighboring healthy ones, and assume an intensely white color.
The disease spreads gradually to the nearby points of growth of rootlet
ombryos,'which in turn become enlarged and acquire a brilliantly white
coloring. Between those and the bottom end there soon appears a small
crescent-shaped Cleft which increates in size as the disease spreads and is
likely to take on a circular shape. The jacket scales on the infested area
glide Upward and arc easily blown away when the bulb is iyulled from the
ground. The upper, succulent scales (1,2,3 :Rid above, depending on phase of
disease) also glide below the circular or crescent-shaped cleft, moving over
farther form thc ruptured area and exposing the internal succulent scales
(table II, II). The longitudinal splitting of bulbs form the bottom to the
nick, including jacket scales and meaty tcales as well, is widespread and
easily detected when crops are inspected. Longitudinal cracks often con-
tinue at the bottom and split that, alto (table II, II and III); in such a
case, thc circular cleft may be,absent. A break in the upper scale fa-
cilitates the projection of a considerable part of the lower scales.
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18 Trans. A. 526
During long, drizzly and often cold'rains, nematodes crawl from the
soil onto the surface, climb energetically onto the plant, and penetrate
the leaves through the stomata. In such cases, the initial phase of the
disease proceeds more latently, because the woftis have attacked the in-
ternal tissues of leaves and bulb scales. The disease can be discovered
only at a later date, when the bulbs are heavily infested.
Leaves of infested plants become very brittle and tough; they crack
and crumble from a touch; thOir coloring lose's its evenness due to deforma-
tion of lamina, and segments of white tissue. With ciperience it is fairly
easy to detect infested plants by their external look. Invasion of plant
parts by nematodes is irregular. . A favorite place for the concentration of
worms is the bulb, -next the leaves and stem shoot. They roots but slightly
and seed even less. For the purpose of illustrating the degree of worm
infestation of the leaves and root system of the garden onion, whose bulbs
were very heavily infested by nematodes, the results of anakrscs conducted
at Kichanzino in 1940 arc cited on table 5.
Table II. Injuries caused by onion nemdtode - Ditylenchus allii (Beijorinck
(Illus. by N. N. Kondakov)
I - From loft, healthy plants, and from right, infested plants of seed onion
(young crop of chernushka) at an early age; II - infested bulbs of garden
onion: left - initial phase of infestation; enlarged, intensely white points
of growth of rootlet embryos and of adjacent bulb tissues; are visible;
at center - formation of cleft between bottom and bulb scales, with their
subsequent gradual splitting longitudinally and sliding upward; from right -
same, at later phase of disease; III - cracking of bottom of garden onion
bulb heavily infested by onion nematode; IV - external appearance of seed
onion plants infested concurrently by onion nematode and onion fly. As a
result of longitudinal splitting of bulbs, part of covered cone-shaped, scales
have projected themselves outwards in tho form of leaves; V - form of tip of
tail'in pea population of onion nematode.
Table II
[Page 529 contains illustrations described above.]
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Table 5.
Degree of worm-infestation of different parts of ga.rden onion as por obser-
vations at Kichanzino in 1940.
Date of
analysis
Total plants
analyzed I
Those infested by nematodes
Bulbs
Number %
Leaves Roots
Aug. 31 56
Aug.
52
Number i% Numbori %
56 100 Analysis not
available
-37 171.1
52 , 100
13 123.2
17 i32.6
Table 5 shows that oven in extremely heavily infested plants, the per-
centage of worm-infestation of rootlets is several times loss than that in
bulbs and loaves. Furthermore, the number of worms in the rott system usually
docs not exceed 10-20 8pecimens, yet inside of bulbs and loaves 'the-! number
many tens of thousands.
Disease in bulbs under sterage conditions. Bulbs slightly infested
bY nematodes usually acquire a completely, healthy look after they have been
dried in the sun, and they can easily' be mistaken for healthy ones and stored
together with those actuallythcalthy. Infestation, however, becomes evident
in 1-3 months, depending on the number of nematodes within the plants at
the time they wore stored and on humidity and temperature in the storage
room. Jacket scales darken gradually where nematode concentration is the.
greatest, and they form a line of demarcation between themselves and healthy
.tissuc segments; because of this, the scales on sick areas appear to be wet
and have an irregular cacao coloring; sometimes- yellow-white or muddy=gray
spots.of irregular contours and different sizes appear on these areas..
The bulbs grOw over softer as they decompose, yet their jacket'scales be-
come brittle and arc easily blown away. On a transverse crack of a bulb,
the internal meaty scales in sick areas arc composed of very brittle, white,
or cream-colored disintegrating tissues (table III, IV) which under the
microscope arc-found to consist of a groat number of warms and an admixture
of plant cells.
When the disease has spread-over a large part of the bulb, small clefts
-
serving thernomatodes as an exit. In some cases appear on its outer surface.
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20 Trans. A. 526
Table III. Concentration of thc onion nematode- Ditylonchu't allii (Bcijo-
rinek) in plants and on various objocts. (Illus. by N. N. Kon-
dakov)
I. II, III - formation of a cream-colored coating consisting of nomatodos in
herbarium specimens of union plants; I - accumulation of nematodes on the
surface of leaves and near the bottom of the bulb, 2 - on surface of trans-
verse cut of bulb, 3 - on hcrbarium paper; ry -.transverse cut of a garden
onion bulb heavily infosted by the onion nematode; darkened sections of
scales with jagged, uneven margins - areas most heavily infested by worms;
V - worms in motion on wall of glass cylinder oxposed to light, .forming on
glass a unique pattern; at bottom of cylinder - bulb heavily infested by
stem nematodes which they have abandoned; VI - longitudinal tut through part
of stem, root neck and root of a tomato infested by onion nomatodc; maximum
concentration of worms is observed in pith which acquires a brown coloring
and becomes badly disintegrated; VII bulb of garden onion heavily infested
by onion nematodo under storage conditions. In areas of worst infestation,
jacket scales have an irregular coloring and appear to be wet.
Table III.
[Page 531 contains illustrations described above.] ?
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21 Trans. A. 526
A deep longitudinal split of the entire bulb [Begin P. 532] is observed.
It reveals the internal meaty scales, part of which arc likely to be covered
with a nematode coating, as the worms move to the surface. Conditions fa-
voring nematode removal to the surface have not yet been sufficiently in-
vestigated; therefcre, it is hard to explain why in some cases it has a mass
character and form t a fairly thiok coating consisting of nematodes; yet, in
others it is not Observed at all. Observations not sufficiently verified
create the impression that the intensiV'o removal of nematodes is due to
temperature and, possibly, to moisture. -Apparently, a very important role
is performed also by a good proliminary drying of the bulbs in the sun
before they arc put.. instorage In the latter case, as 'observed on a small
amount of material, the nematodes hardly -over come out on the surface of the
bulbs, especially when clefts arc abspnt.
If decomposition of bulbs Occurs when moisture is low, then theydry
up and in the end there is nothing loft of them but the covering. Outwardly
this, destruction may remain unnoticed and may be -discovered only by touch.'
Tho length of. time required for the complete destruction of bulbs equals,
on the average, the devclopmen al cycle of 2-3 nematode generations. At -
a 10-12? C temperature and air humidity of 50-60% it takes about 2-3 months.
It humidity is high and tiemperaturc above 15? C, invasion may be compli-
cated by a rapidly developing formantation.in-bulbs, the action of which is
as destructive to nematodes as it is to bulbs. In such cases'decompoistion
of bulbs proceeds even-more rapidly, often forming a wet rot accompanied by
a very unpleasant odor. ?
Destruction of bulbs infested by nematodes can be hastened also by a
secondary infection caused by representatives of fungus or bacterial flora; the
activity of the latter causes mass killing and destruction of worms, and the
end result presents so atypiCalia picture of bulb destruction that some-
times its initial cause is difficult to determine. Decomposition of bulbs
infested by nematodes is accelerated also by the presenee of plant parasitic
mites which greatly change the picture of the disease. The mite Rhizogly-
phus echinopus Fumouie et Robin-LI-lot only injures the bulb, but eats the nema-
todes in its tissues. In the year 1940, the author [of present article]
observed repeatedly how this mite destroyed almost completely a coating on
the 6urface of a bulb that consisted of many thousands of nematode speci-
mens. One could see under the microscope scraps of worm-bodies with mites
of different ages moving among them and among their eggs.
1
Mites identified by V. V. Redikortsev.
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FOOD SPECIALIZATION
The onion nematode belongs to the list of organisms closely adapted to
feeding on certain plant species. As Goodcy (1933) pointe& out, it'can trans.,
for from the onion to'a series ot other spooies of th6 genus Alliut, Thus,
it injurOs garlic - A. sativdm L., look - A: porrum L., chives - A.'schoeno-
prasum L., Shallot onion - A. ascalonicum L. and the wild onion - A. vincalo L.
As per Our observations, it injures in the USSR the following varieties
of the garden .onion (11. copa L.)i the Arzarbas, Bessonov, Kaba, Tsitaus,
Pogar, Strigunov and the Mstersk. (Bogin p. 533] According to our expori-
ments, it injures also tomato and pea plants.
Susceptible tomato varieties (Solanum lycoporsicum 1,.) To ,invostiga-
the possibilities of the onion nematode'b taartsfer fru m onions to totatoes,
we tested three varieties: Chudo-rynka,'P'orotta and Sparks-Erliana. The
first two were grown in a greenhouse and on the test lot at Potrodvorets, on
the oxporimental-farm of the Leningrad Reional Experimental Station for
Plant Protection: Plants wore infested artificially with nematodes while
still very young. Sparks-Erliana and in part Chudo-rynka wore tested in ex-
periments conducted on young crop at the Arzamas base of thc Gorki Regional
Experimental Station for Field Husbandry under conditions of soil naturally
and heavily infetod by the onion nema , Tho o:Torimontal results are
.cited on talo 5. The highest ratc-of infestation was observed in Sparks-
Erliana (38.6g) and in P'eretta (12.6-3%); the least, infested was Chudo-
Rynka.
We arc not inclined to consider this inequity in infestation as an indi-
cator of varying resistance in the varieties tested, bocausc the conditions
of natural infestation at Kichanzino can hardly be compared with the arti-
ficial conditions at Pctrodvorots. Furthermore, observations of the
P'erotta variety have shown that plants of-the same varietr. may be suscep-
tible to notatode infestation in different degree, depending on the ago of
the sprouts. The Poretta variety became infested in different phases of
development: part of the plants were infested at the formative stage of 3-4
leaves', the other part - at the stage of 12-14 leaves. ?Older sprouts were
infested soon after the seedlings were transferred from a box to a greenhouse
rack; younger sprouts were grown dirOctly on the rack and were not trans-
planted before nomatodo introduction. Infestation was accomplished on hdy 31,
1939. Two weeks lator, on Juno 16, part of the plants was transplanted in
the opqn, and part was loft on the same racks. Infestation was not repeated
in opon ground. At the end of the vegetative period, it was established that
tomatoes of the P'erotta variety inoculated in the phase of-12-14 leaflets were
more worm-infested than those inoculated in the phase of 3-4 leaflets. Of
the first group of plants grown in the greenhouse, 19 were analyzed and 11 of
these were found to be infested, and in open ground, 8 plants out of 22
wore infested. At the samo time, Of 92 plants grown in the groonhou8o and ino-
culated in the phase of 3-4 leaflets, nomatodos were found in only 3. It
is possible that the weakened condition of the plants caused.by transplan-
tation was more conclusive to infestation. ?
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23 Trans. A. 526
In infested plants, the nematodes were usually found in the axial part
of the center root,'in the fleck of the-root, and in the part of the stem
adjacent to it (fig. 4, VI). In infestod sogments, the vlarms woro ropre
sontcd by a colossal number ? of eggs, larvae, and adult specimens. Tho
tissue were porous, had acquired a brown color,andwcre destroyed in some :
placcs. Outwardly the infested plants differed but littlefrom healthy ones,
only in some cases, longitudinal cracks of the main'stom in the area whore
side shoots branch out were noted on'Sparks-Erliana. .Infested plants ap.
poared more-brittle and broke easily. In the upper part of the stem, nena-
todes,usually,werc absent and potiolos of loaves were not infested gall
fox'mations on stems wore not found. It must bo noted that, according to the
work of 7111liams (1936), the only source of information on tomato injury
caused by the stem nomatodo, the parasite is responsible for a pocIlliar,
spongy gdll formation on plant stem and Totiolos at base of latina. Perhaps
[Begin P. 5341 this case should be treated as parasitism of. some other
spacics of the stem nomatodeldistinct from the onion nonatodo by virtue of
the character of. its injuries.
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-2,61 Trans. A. 526
The pea (Pisum sativum L.) as a subsistance plant of the onion nematode.
In the course of the work at. Kichanzino, in 1939, our attention was called to.
the relativoly'heavy infestation of peas by the onion. nematode (Uladovskii'
M III varictS'r). Of 16 plants onalyzcd, nonatodes wore found in largo numbers
in 11 'plants. The plants on test lots looked bad because they had rust and
dark spots- on the loaves indiOating infection caused by the nematode and
fungus diseases concomitantly. A similar condition was noted on the 1940'
crop, when the nematode was found in 34 plants out of-73 analyzed (46.5%).
In those cases worms were found in vary large numbers.
. Transfer of onion nematode to other subsistance plants. To explain
the contingent of subsistence p10 ts of the onion nematode, [Begin P. 536] .
4,2.
analyses were made of more than 4 4thousand plants (besides the garden onion)
which belonged to over 40 varieties and species (tables 7 and 8). In re- '
lation to the nematode, these plants can be broken down in three groups.
The first group includes, onions, tomatoes and, probably, peas in the tissues
of which the nematode can multiply. In the second group should be included
crbps whidh attract the.nomatode, but in tho tissues, of which it cannot
propagate. The third and final group includes plants on Which tho nematode
cannot multiply and which, in general, do not attract worms.
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Trans. A. 526
T a b 1 c 6.
onion nematode infestation
Results of analysis of different tomato. varictios at Kichanzino and at
Potrodvorets in 1939-1940
Number ofplants analyzed
6 t? in greenhouscs
4-1 'CI
gti 0 (0 W
'Io 0
Pl>z
N C)
r-4 ? 0 0^ ^ C) r-I
4") C-4
c.3 4-4
cl 2/00rd
O t0 -t-1 0
d
Name of
variety
Place of of - Year
analysis
I in open ground
4
i
0 cd
r
1
1 m
1 di
I 0 4-1 0 .r-i -P
ic.1,13 o 00
rd 0
t Tio .2 PI
-1-1 rct rot o- -4t-i3 , - 4
-r-I d 0 %-9... 0
4-4
Sparks-Erliana
Chudo-Rynka...
P'eretta:
? a) infosted--
plants:::
b)
P'crotta:
a) infestod--
plants:::
b) controls...
Chudo-Rynka:
a) infosted--
plants:::
?
? b) controls...
.Chudo-Rynka:
a) infested
--
plants:::
b) controls?
Kichanzino
Petrodvorets
It
If
IT
It
1940
145
1939 4
1939
1939
1939
1939
1939
. 1939
35
50
1939 61
1939 43
2
97
94
50
? _
12.6
_
-
i.6
50 89 38.6
:_;3 25
10 19 34
21
=PE*
3 56
42
5.2
Total
- _
893 --
1
Nomatodc found-in one control plant growing near thc demarcation lino of
those infested.
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Trans. A. 526
T a b 1 e 7,
Results of analysis of to onion nomatode infestation of cultivated plants and
wood grown on naturally hoavily infosted'soil in the Villago of Kichanzino,
District.
Nos.
consoc.
order
Name of plants
.LTatal. 4 _lb-IAA,*
plants noma-I
!analyzed! todc!
Plants
with-
out
noma -
tode
3
4
6
8
9
10
11
12
13
14
15
16
17
18
? 19
Cereals (Gramineac)
'Oats, Zolotoi Dozhd' variety (hyena 1 126
sativa L)
Rye (uncertified seed) (Socalc ccrealc L.) 15
Winter wheat Undortifiod seed) (TritieUM-
vulgar() Host.)
9
119
Spring wheat (uneertified seed) (Triticum
vulgaro Host.)
Millet (uncortifiod seed) (Panicum
miliacoum L.) i
186
.Barnyard millet (Panicum crusgalli L.)
6
1
Mustard family (Cruciferea)
Cabbage, Ranniaia No. 1 variety (Brdssica-
105
1
eloracca var. capitata)
Turnip, OstorundoftSkii-variety (Brassica
rapa
132
16
Turriip (uncortified seed) (Brassica rapa
L.)
43
19
Radish, Pink witnivhito tip variety -
(Raphanus sativus var. radicola Pars.)...
110
Mustard (uncertified scod)((Sinapus.junded-L
26
7
Hors() radish (Nasturtium scmoracia
2
Shopherd's purse (Capsolla bursa pastoris
6
Potato family (Solanaceae).
Tomato, Sparks-Erliana variety (Solanum
145 56
lycoporsicum L.) ?
Tomato, Chudo-Rynka variety (Solanum lycoper-
sicum 1,4
4 1
Makherca (Nicotiana tabacum L.)
102
Potato, Epicure variety (Solanum tuberoSum L.
114 3
Potato, Sciancts 423 dna -3398 Varioty
(Solanum tuborosum L.)
5 1
Eggplant (uncortifiod scod) (Solarium molon-
gcna
121 3
126
15
9
1861
.5-
104
116
24
109
19
2
6
89
3
102
111
4
118
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Trans. A. 526
411
Tab 1o7 Continuo
Nos.'
conscc.
order
Name of plants
Total with
plants noma-
anaWeddtodo
without
nomatodo
Plants
?
20
21
Loguminous plants (Loguminosac)
Alfalfa, Gritm-ZaikeVidh-Vatioty (mcdidago
Ymodia Pers.)
WhAto clover (Trifolium sp.)
84
71
2
5
79
713
1
Grain:crops wore found to have a paritcular varicty of the onion nematode
2 characterized by small dimensions (less than ,1 ram).
On capago, alfalfa and carrots, wrms were found chiefly in early July,
immedl.ately after heavy rains which had a Stimulating effect od then and
encouraged thoir coming to surface of soil. ?
3
Stem nomatode species particularly adaptcd.to white clover and cucumbers
wore found on those crops; the nomatodc found on cucumbers was distinguished
by a Sharply pointod tail and small dimensions.
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Trans. A. 526
[Table 7.] (Continued [from
pagc 536])
Nos. t
consoci
orderi
22
23
24
25
26
27
28
29
Name of plants
P1
plants
analyzodoma
with
todo
Rod clovor (Trifolium'sp.)
Peas (Pisum sativum L.)
Votch (uncertified scod) (Viola sativa L.)
Lentil, Pot.roVSkaia-4/105 varioty-(lonS ?sou.;
lonta Mnch.)
Geosefoot Family (Chenopodiaccae)
Boot, Egyptian variety (BetaiiulgariS L.)-. 74
Fodder boot (Bota vulgaris L.) 13
Mountain spinach (ChonoPodium.tp;) ll?
2
16 7
89 45
20 4-
13 3
(Amaranth (Amaranthus sp.)
Carrot family (Umbellifcrao)
? 30 Carrot, Yantskaia variety (Dadcds
31 Parsloy, Sakharnaia varioty
32 Dill (knothum gravcolons-L.)
Thistle family (Compositac).
33 Lettuce (Lactuca sativa L.)
Gourd family (Cucurbitaccao)
34 Cucumber, Myromskii varioty (Cucumis sativus L.
35 pucumbcf-; ViatnikOlitkii-Varioty-(CdodMiS-Sati
[ vus L
?
36 ?
37
38
l'aterloaf family (Hydrophylaccao)
Tansy (Phacelia tanacctifolia Benth.)
Flax family (Linaccao)-
Flax (Linum sativum L.)
Mint family(Labiatac)
Unidentified wod
71
102
16
6'
11
- -
/MOM.
152*
1
29 5
115
ithout
n Ma
tode
9
44
16
10
68
2
. 11
2
56
101
16
100 1 100
299 39 260
76 2 74
1 1
Total Oa '2578 _1 252 2326
1
*For footnotes sec preceding pao.
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Trans. A. 526
Tablc 8
Results of analysis of onion nematode infestation Of cultivated plants inocu-
lated artificially at an early ago and the grown in a greenhouse and in open
ground at Pctrodvorcts
Nos. if
cons cc
order
Name of plants
2
Corcals (Gramincac)
Qats, Zolotoi dozhdt:vdi-iety-(AVend - sati-vft-L;)
' a).infosted-pldnts
b) controls
Winter wheat. (uncertified seed) ?(Triticum vul-
?gar0 Host.):
a) infestcd-plantS.
b) controls
NUstard family (Crucifcrac)
3 Cabbage; Ranniaia No. 1 variety (Brassica ?lora-
cea var..capitata L.): ----------: -- _
b) controls.
Radish, Rynochnyi Krasnyi Variety (Raphanus
sativus var. radicold Pers.):
a) infested plants
.b) controls
5 Radish, Lcdianaia sosulka Varicty (Raphanus
sativus var. radicola-PorS;)i---------------
a) infostcd-platitt--- --------- -------
b) controls
Potato family (Solanaceae)
6 Tomato,-Chudo-rynka variety (Solanum lycopor-
sicum L.): _
--a7-infestcd-plantS-- 118
b) controls 72
-1r---
: manta-
.
Total with 'without
plants nom- /lama-
analyzed todo todc
62
50
21
5
43
72
159
.0000
???.11.
WWII/
- _
62
501
211
5
43
72
145 14
108 108
104 86 18
30 30
7 Tomato,-15'cretta variety (Solanum lycoper-
icum L.):
--annfestcd-pldntt-
b) controls
148
125
5
1
113
71
25 123
125
So c sort of partidular?stem nematode spccies the r mcnsi-ns
WO IC not above lmm.was found on grain crops.
of whiah
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n Trans; A. 526
[Table 8. Continued]
Nos. in
consec
order
Name of plants
_Total Plantn
plants- Wlth without
ianalyzed nema- nema-
tode tode
8
9
10
11
12
13
Potato, Ranniaia Roza variety (Solanum tubero-
sum L.)
-IT infested plants.
Leguminous plants (Leguminosae)
Alfalfa (uncertified seed)-(Nledicage
a) infested plants 11
Clover(uncertified seed) (TrifOlitim
a) infested'plantt 28
b) controls 29 '
[Begin P. 538]
):
1
Gposefoot family (Chenopodiaceae)
Spinach (uncertified seed)
a.) infested-plantS
b) controls ?
Beet, Egyptian variety:
2'
2
a) infested plantS
97
b) controls
45
Boot, Bordo variety:
a) infested plants-----
82
b) controls
67
Carrot family (Umbelliferac)
11 Dill (Anothum graveolobe L;)i7-
a) infested-plants
b) controls
15
Thistle family (Compositae)
Daisy:
a) infested-plant8
b) controls
3
8
1
1
28
29
2
2
494
45
81
67
41
50
Gourd family (C6curbitaccae) ?
411 16 Cucumber, MUromskii-variety (CucuMiS.SativuS-L;)
0.infested plantS 30 30
b) controls 8 8
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Trans. A. 526
[Table 8. continued from
page 22]
Nos. in
conscc.
order
Name of plants
Total
plants
analyzed
with
nema-
todo
with-
out
nema-
tode
17
.Cuomber, Klinskii vaftety-(CucUrilit -
Plant
a) infosted-plantt
41
4
37
b) controls
23
23
? 18
Cucumber (uncertified-toOd)-(Cucumis sativus
a) infostod-plantS
b) controls
28
13
1
WOMB
27
13
Total
1744 286 -
1458
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Trans. A. 526
[Bottom p. 536] To the list of plants which attract posts should be
added the radish, the fodder turnip [Brassica campestris rapifora], the
[rcpa - Brassica rapa],mustard, tansy, and in part Votch and lentil. Their
attraction for the nomatodo is hard to explain, but there is'no doubt that the-
nomatodo is glad to invade them and is likely to contontratd thcro by the tons.
In 1939,.under field conditions at Kichanzino, the nematode was found in all
16 fodder turnip plants analyzed, in 19 out of 43 turnip plants analyzed, in
7 out of 26 mustard plants and in 9 out of 22 tansy plants. In lentil it
was found in all throe plants concOrnod, and in vetch, in 4 out of 10. In
most cases the worms wore numerous All plants of this group looked entirely
healthy, and an external inspection of the roots revealed no traces of in-
jury. In 1940, the crops rOforred to showed a considerably smaller per-
centage of worm infestation. In tansy, nematodos were found in 30 plants
out of 277 plants, in the radish, in 1 plant out of a 102, and in witch,
lentil and fodder turnip (10, 10, and 100 plants respectively), they wore
not found at all. Nonetheless, under conditions of artificially heavily
infested radishes, in a greenhouse at Pctrodvorots, worms were found in
145 out of 159 radish specimens of the Rynochnyi krasnyi variety, and in 86
out of 104 of the Lcdianaia sosulka variety (Table 7). [Begin p. 539] In -
this particular case the worms remained in tho plants until the very harvest.
Their absence in the same cropAn Kichanzino can be explained only by the
late date of analysis, which was performed in the phase of scud maturing,
when plant tissues had already lost their freshness and succulence and
probably therefore no longer had any attraction for nematodes. Furthermore,
in the field, the worms had selective possibilities as to subsistence plants,
thanks to the presence of woods; in the greenhouse, however, there were no
other plants besides the radish. The fact that certain phases intho develop-
ment of the radish have 2;ot an attraction fer the nomatode is so much the
more certain, because the worms were found very rarely, or not at all, on
othci experimental plants, regardless of equal artificial infestation of the
soil.
The third group of plants which do not attract the nomatodo and are
not injured by it, includes but a small number of crops: Wheat, rye, oats,
millet, cabbage, horse radish, makhorca, potatoes, eggplant, alfalfa, clover,
boots, carrots, parsley, dill, cucuthbors, flax and spinach, and the woods
goosofoot, amaranth, and 71-ild daisy. The possibility that worms stay tem-
porarily in-plants belonging to this group is indicated by the following'
observation. In Kichanzino, in 1939, thenematOde wasfoundin many plants
of alfalfa, makhorca, carrots and flax immodiatoly after the heavy rains of
July 8 and 9th. Under the influence of moisture, the worms crawled all over
the lot and, in passing, climbed also on crops to which they are not adapted
and which they later loft; thus, on the day following the rains, the noma-
todo was found in all 5 alfalfa plants analyzed; yet a month and a half later
it was not found in a single plant of 18 analyzed; nor was it found on a single
plant out of 61 alfalfa plants in 1940. The nematode Was found mono
makhorca plant on tho day following the rains in 1939, and in none out of
3 a month and a half later; in 1940, it was found in nono out of 102 mak-
horca plants. Of 12 plants of flax analyzed after the rains, the nomatode was
found in One, and at the end of the vegetative season, it was found in one
out of 64. A similar picture was Observed on carrots whore the percentage
of plants inhabited by worms in 1939 dropped considerably toward fall,
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g3 Trans. A. 526
compared tho July, and in the year 1940 it equalled zero. Consequunrly, in
separate cases, the nematode ean be found on crops which do not belong to the
list of its subsistence crops. Hence, it cannot be Inferred that a plant
'belongs to thc list of nematodc hosts, just because tho nomatodo had boon
prcsont in individual cases.
The nematode was not found on wheat, rye, oats or millet (tables 6 and 7
Nonetheless, a special variety of tho stem nematode was found in 4 rye
plants, 13 millet, in 2 plants of spring wheat and in 1 of winter wheat in
1939 at Kichanzino; it was found also in 3 plants of oats out of 62 of the
infested series, and in 1 out of 50 controls. In addition, this variety
was found in 3 plants of winter wheat oat of 21 taken form the infosted
series (Table 7). In the case of weeds, tho namatodo was found only in one
plant - barnyard millet, at Kichanzino in 1939. Thi happened to be a
larva 1017 microns in length and 20 microns in width. Most of the Crucifcrac
plants investigated as to the nematode arc included in tho second group
by virtue of their attraction for worms (fodder turnip, turnip, radish,
mustard). Yet, some of tho experimental plants - cabbage and horse radish -
obviously must be added to the list of those which do not attract the pest and
are not attacked by, it. The nomatode was not found in horse radish, and of
148-cabbage plants, it was present in only one and was limited to one speci-
men. It must also bc noted that, regardless of heavy artificial soil in-
festation at Potrodvorcts, [Begin p. 5401 the 43 cabbage plants grown there
wore free from posts.
Those of the potato family investigated wore: tho tomato, makhorca,
potato and eggplant. Of these only tho Chudo-Rynka-and Pcrotta tomato
varieties proved subject to infestation (see above). Potatoes, makhorca and
eggplants may, on rare occasions, 'be nomatode carriers; tney arc not, how-
ever, injured by thom. Thus, at Potrodvorets, a potato plant was grown
artificially in soil extremely heavily infested; it produced more than
ton large, healthy tubers, and not one of thorn, nor the plant itsolf, was
infested.
The following experiment wds conducted on artificially infested potato
and boot tubers and onion bulbs. Groups of 20 tubers and bulbs were infected
by nomatodos by moans of introduction of small particls of an infested bulb
through a deep cut; tho incision edges were brought together with bandages
and then the tubers and bulbs were wrapped separately in paper. Half of the
bulbs wore kept in a cellar at 10-12? C, and half were placed under a shelf
in a greenhouse where the temperature fluctuatcd from 30 to 20'and lower.
In 2 months -time all bulbs were infested, and most of all those kept at low
temperature. No nomatodos were found in boots: obviously they all perished;
They were absent also in the parts of potato tubers not damaged by incision.
A certain amount of larvae were found in but 3 tubers in tho place of
incision whore they-apparently had survived over since they had boon intro-
duced in the tubers.
Field observation at Kichanzino showed that even tho EpiCure variety;
one of tho most susceptible to the stem nomatodo, was but mildly infested.
Thus, in the year 1940, we analyzed 114 plants of this potato variety and
only 3 of thorn_ proved to be vectors of a small number of nematodes. All
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4
Trans. A. 526
plants looked perfectly healthy, and their yield of mote than 1000 tubers
was of a high quality; no nematodes were found in them. In eggplants, too,
the nematode-was found in very limited numbers and only in 3 plants out of
110 analyzed. On the basis of the above and also as a result of the very
good condition of the crop in all repeated tests, it can Ile assumed that
eggplants are not subject to injury by the onion nematode. Naturally, the
possibility that the nematode may adapt itself to a life on potatoes, egg-
plants, and other plants is not excluded,.especially if a nematode genera-
tion grown on tomatoes and other related crops were to be transferred to
these plants either in natural environment or artificially.
Plants of the legume family investigated were the Grimm-Zaikevich var-
iety of alfalfa, white and red clover, peas, vetch and lentil. Peas, above,
are host plants of the onion nematode, vetch.and lentil possess the capacity
to attract it. As far as clover and alfalfa are concerned, they are neither
injured by the pest, nor do they attract it. Of the 84 alfalfa plants
analyzed, the nematode was found in 5 and in all cases on the day following
the heavy rains of July 9, 1939. Further analyses, conducted at the end of
1939 and during 1940, have disclosed their complete absence in alfalfd.
None. of the 71 plants of white clover analyzed had the onion nematode. Red
clover proved to be a nematode vector in 7 cases out of 16,-with 5 cases
being registered tho day after the heavy July rains in 1939. The same year,
we conducted an experiment in artificial infestation of red clover under
DMDoratoty conditions at Leningrad. Seed were planted in sterilized soil
[Begin p. 541] in which, a few days later, a largo amount of worms was in-
troduced together with fragments of meaty scales from strongly infested bulbs.
Sprouts had a normal appearance, and a microscopic analysis of approxiniately
40 plants relirealed tho presence of single nematodes in only two plants.
A second experiment in artificial infestation of rod clover was conductod
at Petrodvorets whore a considerably largo amount of seed had boon sown.
The experiment was conducted twiccaand nematodes were introduced in tho soil
the same as in other cases, under tho roots-of young sprouts, with segments
of meaty scales from heavily infested bulbs. Tho plants wore developing
normally,-and an analysis of 28 specimens disclosed a complete absence of
nematodes. Laboratory experiments, substantiated by field observation at
Kichanzino, justify fully the statement that the onion nematode does not
injuto clover, and that the latter is not a subsistence plant of the nema-
tode. At tho same time, it must be noted that we discovered a special
variety of tho stem nematode loss than 1 mm in-length in 1 alfalfa and in
3 clover plants at Kichanzino in the year 1940.
No subsistence plants of tho nematode were found among the goosefoot
family (tables 7 and .8). In beets, tho nematode was found in several
in-
stances. In Kichanzino, in tho year 1939, the onion nematode was discovered
in 6 boot plants of the Egipotskaia variety out of 74 plants analyzed, and
11 plants of the fodder beet out of13 plants inspected. The greatest in-
cidence (12 out of 17) occurred immediately after the July rains, indicating
that the worms had crawled on these plants accidentally. In 1940; no nema-
todes wore found in a single boot plant out of 57 plants analyzed. Tho
beet crop produced a very good yield, and a microscopic inspection of several
hundred roots under field and storage conditions failed to disclose any
traces of infestation. The nonsuscoptibility of boots is confirmed also by
experiments conducted in Petrodvorots on tho Egipetskii [or Egipetskaia] and
Bordo [or Bordeaux] varieties. One of those experiments - a futile attempt at
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root infestation - has alroady been described; two others wore conducted
in a greenhouse. In one of them,nematodes were introduced artificially in
unsterilized soil on a rack whore boots wore being grown; a part of infested
and of control sprouts was-lator transferred to open ground whore it was
growing until harvest time. The second experiment consisted of growing
boots in sterilized soil in boxes at a depth of 20 cm. All togethor, more
than a 100 plants wore grown in boxes, and they grew well and produced an
excellent yield regardless of whether or not they had been infested by the
nomatode. During harvest, 407 boot plants of the Egipetskaid variety and
374 of thu Bordo variety were taken for analysis and storage. A.large part
of the plants grown in sterilized soil in boxes was subjected to micro-analy-
sis. In none on the 41 Egipotskaia variety boot spocimens"or in the 57 of the
Bordo variety, not counting controls, wore found nematodes. Of those infested
and grown in a greenhouse and in open ground, 56 boot specimens of the
Egipotskaia variety and 25 of the Bordo variety wore taken for analysis: in
the Egipotskaia variety,"vory, few nomatodos wore found in. 3 plants and in the
Bordo variety in 1 plant. The relatively small dimensions of the worms
(length of females not above 855 microns), together with the fact that in
3 instances they wore found in the open ground and in one instance in a
greenhouse in plants grown in unsterilized soil, justify the assumption that
hero, pogsibly, was involved a stem nematode species other than tho onion
nomatodo. Tho results of storing 490 beet roots (Egipotskaia 265 and Bordo "
225) arc convincing enough that the onion nomatodo does not injure this plant.
The roots were stored all wintar in boxes in a Potrodvorots basement and
were inspected in Juno of 1940. [Bcging-p. 542] From the group of infested
plants 30 roots wore analyzed of"bach of the two beet varieties; the nema-
tode was abson-tin all specimens.
Plants of the darrot family [Umbelliforao] investigated were dill,
parsley and carrots. Dill was seedod'in a Petrodvorots greenhouse and in-
fostod with the nematode artificially'. -Growth of the plants was normal and
no symptoms of disease Were observed. Of the 49 plants used for analysis, the
nematodo was found in limited numbers in only 8. In Kichanzino, dill was
growing all over the experimental lot as a result of solf?=seeding; of the
16 plants concerned, the onion nomatodc was found in none. In all probabil-
ity, dill need not be included at all in the list of host plants of this
nematode. The Sakharnaia variety of parsley was transplanted onto the ex-
perimental lot at Kichanzino in the year 1940; 102 plants were used for
analysis and in one of them was female nomatodo; no symptoms of disease
wore noted. Apparently, pai-sley likewise cannot be considered as a sub-
sistence plant of this post. The Nantskaia carrot variety was grown in
1939 and in 1940 on the experimental lot at Kichanzino. The plants wore
analyzed while very young; in 1940, the nomatode was found in none of the 26
plants concerned, but in 1939 worms were discovered in 15 plants out of 45.
Of these 15, the nematodo was found in 12 plants out of 14 analyzed the ddy
after the July rains, and only in 3 plants out of 31 analyzed in the fall.
Thus, carrots are not injured by the nematode, but its accidental invasion
of this plant can occur.
Those of the thistle family [Compositae] investigated were lottucc
and the wild daisy. Lettuce was grown only in 1939 at Kichanzino. During an
analysis of 29 plants, nomatodcs were found in very limited numbers in 5
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plants. Wild daisy grown in artificially heavily infested soil in a green-
house (table 6) proved to be free of nematodes.
Cdtbaribers were the only species of the gourd family investigated as to
possible infestation. T46 cucumber varieties Mgromskie and Viaznikovskii
were grown in Kichanzitw. Both proved resistant. In the NUronskic variety,
a distinot species of the stem nematode was found in 2 plants out of 103
analyzed. Its dimensions wore not over lmth and, besides, it was plainly
distinguished from the onion type by a very pointed tail. Three cucumber
varieties were grown at Petrodvorots - Muromskie,.KIinskio, and an unknown
variety from Mite Russia. The nematode was discovered in-4 plants of the
Klinskic variety and in 1 of the unknown variety (table 7). We grow sprouts
by the tens undor laboratory conditions and infested them with, the nematode
at seedinc:. time. The worms were introduced in the soil by sprinkling it
evenly Id.ith water containing nematodes in allphasos of development, extracted
from heavily infested Bulbs. Three weeks later, tho first batch of sprouts
was taken for analysis. The last analysis were made a month and a half later,
and of 26 plants analyzed, 2 worms of very small dimensions were found in but
one plant; it is possible that these belong to a-special variety tf--
stom nematode, distinct from the onion type; seeding had been carried out in
unsterilized soil. Analysis of soil surrounding the roots of experimental
plants revealed an almost complete absence of the onion nematode, and the 88
specimens found there represented 12 nematode species, but included only one
badly damaged female of the onion nematode.
Tho above account leaves' no doubt that the onion nematode is a restricted
specific pest that belong to a particular species 'of the stem nematode
(Ditylonchus allii Beijor.) [Begin p. 543] and is adapted to prasitism on the
onion and probably ether plants of the genus Allium; from time to time it
attacks tomatoes ,and peas. It is not injurious to a series of the more im-
portant agricultural crops, such as beets, potatoes, grain crops and many
others; the nematode disease of these has long since been ascribed to a
single variety of the stem nematode - Anguillulina dipsacivar. cothmunis
Steiner ct Scott, 1934 Ditylenchus dipsaci--(Kuchn) Filipjcv, 1934.
CONTROL MEASURES FOR ONION NEMATODE
One of the more essential measures is crop rotatioh with selected
resistant crops, in which the onion and other possible subsistence plants of
the nematode (garlic, tomatoes and-pcas) must not be sown in the same field
before the expiration of 1-2 years. This can be accomplished easily on
sovkhoz and kolkhoz (state an fatmsl, fields, since the number
of werm-resistant plants.is fairly large.
Of utmost importance is also seeding of uninfested sed material so as
not to add to the soil a new supply of the onion nematode. The old supply
of worms can be eliminated by efficient crop rotations and careful clearing of
fields of post-harvest-onion refuse. Selection of seed bulbs is accomplished
best during harvesting. The appearance of infested plants immediately upon
their extraction from the ground is so typical that they can be picked out
without any difficulty. The remaining few contaminated bulbs must be re-
moved by-additional sorting before that bulbs are stored and again just Before
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.Trans. A. 526
planting. Thus, thcgardon onion resorved for seed can be cleared fully. It
is not so simple with the seed onion. It must definitely be cleared during
harvesting, as its sorting later on is very difficult. Experience proves
that infestation of theseed onion after winter .storage is frequently 1-3%,
and deheIminthing of billbs can be accomplished completely only with the aid of
wet thermic treatments. Experiments have revealed that to achieve this
objective, it is most desireable to apply the following temperatures and
exposures:
45-46? C for 15-10 min.
50-52 " 10-5 "
55-57 U 5-3 ft
Tho temperatures and exposures indicated arc quite Sufficint for
dehelminthing, and they often kill the nematodes together with the bulb, if
it is badly infested; ono naturally, need not regret the loss of infested
bulbs, for they would produce no yield anyway. Net thermic troataent
before planting has no ill cffect on the growth of healthy plants. Laboratory
and greenhouse experiments at Petrodvorets have on the contrary, shown
that growth and productivity have benefited from thermic treatment. Results of
wet thermic treatment of the seed onion grown in 1939 in a Petrodvorets
greenhouse are cited in table 9. Each plant grew to full maturity in a
separate pot filled with soil sterilized for nematodes. Sterilization was
accomplished by heating the soil for about an hour at 60-80? C. The bulbs
collected at the end of the-experiment-lay in storage the winter of 1939-40
without producing and waste. [Begin p. 5441
b 1 e 9.
Influence of wet thermal treatment on growth of seed onions grown in sterilized
soil at Petrodvorets in 1939
Nos.in Tempera- ipxposure INumber IPlants Yield ;Bulb diameter of new
i
consoc ture (?C) in mi- of derived (in ! crop (in cm)
order ; flutes) bulbs 11, bulbs) From - to Average
1 45-46 10 3 21 5 )..--1.75 1:5
1
Contrel --
3
1 3 5 ;1.5-2
175
2 50-50.5 10 5 5 16 11 --2
1:75
Control 6 i 51 16 1.---2 7
3 52 - 1:75
5 5 5
23 10.5--1.75 1.5
Control .4) 4 4 9 11 7-2:75 2'
i
4 . 57 3 6 1 6 21 !1--2,75 1:5
I
Control 3 3 7 iO.52 115
5 67 1 5 5 12 11"--2 - 1:5
Control 5 31 6 !1.5--2.25 1 1.75
1
Total
Fusafium and
tion.
I 45 41
rose rot infection was
120
observed in all cases of plant destruc-
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Trans. A. 526
Similar results were obtained under field conditions in Kichanzino in
1939, where thermal treatment of seed onions was accomplished shortly before
planting at a 42-45? temperature and 2-hour exposure. The experiment was
conducted in April and repeated twice on test lots measuring 9.5 m in length e
,and 1 m in width. Three hundred and fifty seven bulbs were planted in the
test lot in 3 rows parallel to its long axis. Results *ere evaluated by means
of external inspection Of the plants in each middle row. The plants collected
were sorted into groups of healthy, diseased, doubtful as to infestation and
others.
Specimens infected by fusarium, gray mold, and injured by the onion fly
and other pests were included in the last group; immature plants, too, wore
included in this group. The results of counting and weighing of experimental
plants by groups are cited in table 10. Its analysis is convincing that wet
thermal treatment had failed to exert any great influence on the weight of the
yield gathered; the plants produced an average 316 gr drop in weight per test
lot, and reduced the number of bulbs by 30. This can be explained by the ex-
tremely long exposure of the seedling to wet thermal treatment which can be
greatly reduced without impairing dcholminthing results. A decreased ex-
posure should raise considerably the general productivity of the plants
treated. The experiments conducted, naturally, require checking on a large,
industrial scale, but there is no doubt as to the feasibility of using the
wet thermal method in treating seed onions against the onion nematode. .
Apparently, it is alse entirely possible to apply the wet thermal trcat-
ment in dehelminthing onion seed (of the-chornushka onip0) reserved for sowing.
From the unpublished experiments of T, F, Nikitinae- conducted under labo-
ratory conditions at the Gorki Regional Experimental Station for Field Hus-
bandry, it can be concluded that deholminthization of the "chernushka" onion
by the wet thermal method produces abundatit germination fo those seed. In
field experiments at Kichanzino, [Begin p. 545] this sort of-treatment of the
chornushka" onion also produced positive result (table 11). Tho experi-
ment *as repeated twice on test lots measuring 9.5 m in2leneeth and 1 m in
width. Altogether, experiments were conducted on 114 m . On each tot lot,
47.5 gr of onion were seeded in three rows, parallel to its long axe'. Seed-
inewas carried out in 1939; experimental results wore evaluated from August
21 through September 2nd. One row of each test lot was selected for calcula-
tion. Duo to dry weather in the first half of the summer, germination was very
protracted and the plants began to sprout noticeably only after the torrential
rains of July 6-8th which were accompanied by chilly weather., The ains caused
the mass movement of nematodes and their mass infestation of onions. At?
harvest time the general infestation of the seed onion was fairly heavy, but
considerably loss than that of the garden onion (table 10), thich must be ex-
plained by the relatively late infestation of the seed onion.
Due to late planting of the seed onion, the plants were not 'quite ripe
at harvest time, and, therefore, they were weighed with the lea-lies. On the
treated test lot, tho average total weight of plants was 157 gr. more than
that of control plants: yet the number of plants harvested was by 52 loss.
It is to be expected that by reducing the time period for preliminary tott-
ing, plant growth will prove considerably better than that of controls.
Unfortunately, the problem as to whether or not the' treatment kills the
nematode within the plant remains an open qestion. However, by analogy of
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Influence of thermal treatment on ,zrowth of seed onion and on its infestation by
per field observations at Kichanzino in 1939
Test
lot
Con-
trol...
Heating
in water
Re-petition
II
Number of Weight of bulbs
bulbs collected (in kg)
44
-9
52
105--
90
4
0
(/)
? ri
106
157
7
1740
-p
19 25 1.702 2.955
15 19 0,222 3.619
27 39 1.380 2.902
83?3-.3o4 9.476
0 -P
P 0
1.162
0,445
l.094
64 25 15 2.100 1.9g0 0.550
129 17 25 0.163 4.S45 0.563
Total
bulbs
collecte
Weight
(in kg)
194 5.819
200 I4.286
195 I 5.376
589 5.481
194 4.630
175 5.571
at 42-45 III
35
79
14
--
0.823
3.056,0.452
128
4.331
C for 2
hours...
Total
129
272
56
40
3.086
9.881 1.565
497
4.532
Average of one
Trans. A. 526
Ta b 1 c 10.
onion nomrtede as
0
CD
CD--
test lot in three varietiesm
Number, of bu bs weight of 13?:)s
Remarks., One average row of each .test lot was uscd for analysis in all th:
September 9th.
Total Weight
bulbs
col-
lected
196
166
.160
4.844
Heal
thy Sick Healthy Sick m
35
ree reqtitions.-
43
-n
CD
113 1.101 3.159 m
N) -
CD
O.- )
CD
QD
1.029 3.294'- 1
Ana? lysis was made On
A
?
0
(E)
0
CD0
(D
8
CD<
-n
(D
(T)
(D
n.)
-10.)
0
0
-0
co
0
0
n.)
-10
n.)
-10
(E)
?
Tr4.K. A. -5L6
Table 11.
Influence of dry and wet thermal treatment of (dark) onion seed on their growth and infestation by the
onion nematode under field conditions at Kichanzino in 1939
?
*
)
:
)
Test lot
---,
Date of
observation
Number
of plants
.1
Weight of same plants
(in kg)
Total plants
collected
Weight (in kg)
Average per test Lo
in three variants
ccii
Healthy
v)
g
i -rez?i
,
Number of
-plants
Their wei
(in kg)
I
c-3
rn
r-
cr)
P
,
0
r-i
0 ,0
?ri
0
i
:al
q
A . ,D
A
A
Control
Aug.
21
I
322
174
gl
19
0.530
0.389 1
0.126
596
1.045
Aug.
21
II
430
286
73
16
0.777
0.956
0.206
go5
1.879
770
1.784
Aug.
21
III
660
140
50
59
1.650
0.575
0.204
909
2.429
Dry heating f
r Aug.
21
I-
380
119
117
24
1.01g
0.548
0.324
642
1.g86
24 hours at
Aug.
21
II
253
194
57 19
0.575
0.757
0.17g
523
1.090
616
1.773
50 C -
Aug.
21
III
504
64
49
66
1.430
0.332
0.1g0?
63
1.942
.
Dry heating for
Aug.
21
I
630
104
70
24
1.283
0.694
0.345
g42
2.340
24 hours at
Aug.
21
II
222
185
64
14
0.454
0.705
0.160
4g5
1.319
659
1.822
6o C
Sept.
2
III
456
82
--51
61
1.254
0.340
0.209
650
1.80g
4
Preliminary
steeping in
water at 300
,for 4 hours,
then heating
Aug.
21
I
524
177
65
26
1.401
0.722
0.191
792
2314
in water at
Sept.
Sep t.
2
2
II
224
484
259
S5
16
0.482
0.775
0.234
564
1.491
718
1.941
50 C for 10
minutes
III
174
62
57
1.035
0.767
0.215
777
2.017
?
0
CD
=Pi
CD
t
8
CD<
CD
(T)
CD
n.)
(r)
0
0
-0
co
0
0
n.)
-10
n.)
0
0
0
(E)
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40
,z,k1 Trans. A. 526
action exerted by high temperatures on other nematode species, particularly
on that of wheat 7 Anguina tritici (Steinbuch), it can"bc assumed that tho
temperatures indicated in table 11 will kill the worms.
We find it imposSible to dwell on all experiments and observations con-
ducted for the purpose of investigating measures to combat. the onion nematode,
because most of them: for instance liming of soil, different varieties of
of fertilizers, testing of hicroclements, different seeding dates, ex-
periments in turning over layers, [oboret plasta] and others, have produced
no positive results, and the experiments. rInd observations which did.
produce results have been described above. Nonetheless; there is an cx-
perimcnt which cannot ,bc left unmentioned; it was conducted in Kichanzino
for the purpose of clarifying in what form worms are when they pass tfirough
animal'intostine and whether or not they might be distributed with
manure, Tho author had taken under observation two sheep which for 2 days
(August 5 and 6, 1940) had been fed heavily infested onions. In this
settlement, many fainers use infested onion for animal feed, particulary
for sheep and ,oats. 'Some of them preserve also dry onion loaves as
winter food for cattle. Infested onions intended for food were picked
from plants of the experimental lot. These wtrc mixed with cabbage leaves
and weeds, but the sheep preferred the onions. Excretory analysis have
shown that tsn .-large majority of worms had boon completely digested in
the intestines. Aatogother, 37 nematodes were found in axcrotions, al-
though the sheep had boon swallowing them by the million. Three of them
were observed to move the head end of the body very slowly and briefly;
an atteffipt to revive the worms by transferring them to pure water ened in
failure. .This experiment justifies the assumption that the worms cannOt
he distributed with manure. But it must be kept in mind that manure
may always contain a certain amount of undigested feed, and, to ensure'
full onion nethatode sterility of manure, care must be taken that feed is
well digested. Since nematodes cannot survive fermentation, well fer-
mented manure becomes entirely harmless as f!71.r as their distribution is
concerned.
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Trans. A. 526
COMPARISON OF ONION NETMODE POPULLTIONS GROIN ON DIFFERENT PLANTS
Assuming that food properties cannot fail to influence thc organism
of worms, we conducted a thorough morpho-anatomic study of onion nomatodo
populations grown of different plants. For characterization of each
population; we usually soloetod 50 specimens farm the phase undcr con-
sideration. Each specimen was measured and the dimonsions obtained wurc
rocorod accoriding to thc Cobb [Kobb] formula. In addition, measure-
ments wero taken of thc spine [igla], the distance between thc vulva and
thc postcrior ond of the body in females, the spicules and the 'rodforrule
[rulck], distance between bursa and tip of tail in males, and likowiso
sizc alpha (relation of body length to length of tail). Measurements
were to. ken chicfly of fixod ma-Us:i-ial (4% formalin); a few measurements
were taken also of live material.. It must be noted that thc nematodes
utilizcd for measurement were not collcctod at exactly thesame time.
Tho August 1940 onion samples wore not fixed; hence, to measure
onion nematodo populations,-it was necessary to use specimens collected
in August-September of 193(',. Populations from tomatoes and peas, how-
ever, were 'collected on 20-22nd August, 1940.
When populations of the onion nomatode grown on diffpront subsis-
tence plants arc comparod,At.can be seen that the onion population is
composcd of individuals considerably larger than those of the tomato?
population,and of just a little laror ones than those of the peas
population (table 12; fig.5). The peak of the variation curve in ncmatodos
extracted from onions lics-within tho'limits of 1400-1500 microns, drop-
ping sharply on both sides. [Begin p. 547] The curve for ferellos (60
specimens) and that for males (40 specimens) look almost alike. In the
onion population4 larvae of to fourth stage also have large dimensions,
possessing an average of 1100microns in length and 23 microns in width
taken of 50 measurements. Individual variation in thoir length rests
within the limits of 950 to 1300. microns in width, with minimum dimon-
sions measuring 60 x 30 and maximum 111 x 33 microns.
The population of the' onion nomatodo grown in tomato tissues roachod an
average 1190 microns in length and 30 microns in width in females, and
a corresponding 1150 x 24 microns in males. The dimensions of females are
alpha==40, bota==6.6; gamma=16, and in males correspondingly: alpha...49,
bota=6.7, gamma==15. In the tomato population, larvae of the fourth stage
have an average of 885 microns longth and 22 microns in width." Minimal
larvae dimensions reach 770 x 20 and maximal 1000 x 25 microns. The site
of eggs is almost the same as in eggs of the onion, population, having a
78 x 32 micron average (out ofo50 measurements).
The pea 'population is characterized by larger dimensions than that
of the tomato-population, and, in this respect, it does not differ from
the onion one. In 25 females extracted from peas, the average body length
equaled 1460,-and the width 32 microns; in females: a1pha==46, beta...7.3
and gamma==.16. The males are a little smaller and average 1349 microns
in length and 25 microns in width (24 measurements), in males, alpha==55,
beta:1.7 and gamma==16. ?
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42 Trans. A 526
Thus, as regards size alpha, and the total dimensions of male-and
female bodies, the pea population is undistinguishable from that of the
onion .one. Fourth stage larvae of the pea population vary from 880 to
1220 microns-in length (average 1109), and from 20 to 25 microns in width
(average 23). Consequently, the dimensions of the pea population do not
differ from those of the onion population. Since we lack dOtails as to the
localization of the pea population in pea tissues, [Begin p. 549] it is
impossible to determine the dimensions of their eggs, provided they do
hatch eggs in this plant. Dimension similarity in the onion and pea
populations is quite remarkable, if the worms actually multiply in pea
tissues.
Worms extracted from radishes, mustard, cabbage,- beets, lentils, vetch,
flax, tansy, tobacco, eggplant, potatoes and dill are, in point of
ditonsions, very close to specimens of the onion population. Although
found in many plants, their number was very small. -
Of the list of plants which attract the nomatode; the greatest atten-
tion was paid to the fodder turnip and the turnip. The females oxti'acted
from turnip averaged 1315 microns in length and 26 microns in width. Tho
males were somewhat smaller attaining (of 21 mea6urcmonts) an average of
1273 microns-in length and 23.6 microns in width. -In foma1e64.. alpha
416; beta 6.2, gamma 14.7, and in males, aripb0 50.3, beta 6.4, and gamma
13.9. Tho length of females varied from 1000 to 1700, and in males - from
900 to 1600 microns. Tho great variation in worm dimensions indicates
heterogeneity of nomatodos found in turnips. It is entirely probable
that this phenomenon is duo to the absence of conditions in turnip
tissues ossontiallto the propagation of the onion nomatodo, and indicates
that the worms had climbed on this plant from without and at different
times.
Dimension analysis of nematodes from fodder turnip (17 females and.
13 males) produced the same results. Average female dimensions equalled
1232 x 27 and those of males 1310 x 22 microns. Tho length of females
varies from 880 to 1517 and that of m6los trot-1100 to 1628 microns.
Average in females alPha.:45, beta=6.2 and gammar.-15; in males alpha=58.2,
beta-6 and gamma:1.14. Body dimensions of the nomatodc are as hotorogenobus
as in the preceding case. This, obviously, can be attributed to tho fact
that tho-worms had developed in plants other than the fodder turnip.
[Begin p. 550]
The materials cited reveal that a subsistence plant leaves its imprint
on the parasite developing within its tissues. The onion ncmatodo is
afforded a more favorable habitat in the tissues of onions and it produces.
a population that is characterized by-maximal dimensions, compared to
populations developed on other plants.
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Trans. A. 526
T a b 1 e 12.
Comparativc dimensions of onion nematode spocimens from populations
grown on onions, tomatoes and.pcas, at Kichanzino in 1939-1940
Dimonsions of nematodes (in microns
Lcngth of
body
Width of
body
Alpha
Beta
Gamma
.
. ,
.
Aug. 20
pcas
ci.)T
1110-1758
25-45
37-59
6.1=8.4
13-19.4
1940
(1460)
(32)
(45.9)
(7.3)
(15.9)
(31-81
1210-1517
20-30
40.3-74
6-8.1
14.2-17.5
(1349)
(25)
(55)
(7)
(16)
.
, . ...
Aug. 20
Toma-
935-1320
25-37
27.1-49.7
5.6=8.7
12.4-24.6
1940
tocs
T
(1190)
(30)
(39.7)
(6.5).
(15.9)
ee
950-4300
16-28
35.7-62.0
3.8=8.3
13.1-17.3
11, .
(1149)
(23.7)
(48.7)
(6.7)
(15.2)
' Aug; 15
Onions
00
1130-1924
30-43
34.8=55.4
6.7=8.6
14-20
Sp' 2
-r

(34) .
(42.5)-
(7.6)
(16.7)
1939
dle
1190-1685
20-30
44.8-74.2
6.2-8
14-18.6
(1405)
(25.5)
(56)
(7.2)
(16.0)
R.cmarks:?iavcraL;c sizos arc cited in parentheso
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ti Trans. A. 526
In contrast, nomatodes in tomatoes have a acsS favorable habitat
and this fact is markedly reflected in their sizes. Eggs arc exposed to
the least variation and arc alMost of the same size as the eggs of the
onion population.
900 1100 1200 1300 1400 1500 1600 1700 1800 1900
Fig. 2. Variation in body length in onion nematodc populations rcarcd on
different subsistence plants.
Line with highest peak - length of female body, with a loss high peak--
length of male body; solid line denotes onion population, broken lino -
tomato population, line of dots and dashes - the pea population; figures
beneath denote length of body in microns.
Influence of environment manifests itself first of all in the total
dimensions of the 1:--orms, but in individual cases, significant variation is
observed in scparato organs. Thus, in the pea population, the length of
the tail, which can change from a pointed one to a dull one, varies ,notice-
ably.
Onc, among several thousand nematode specimens with normally developed
pointed tails, one male was observed with an absolutely dull tail (table
II, 5). Finally, this influence can extend to the bulbus and, apparently,
to other organs. In nematodos extracted from turnips, it was obscfvcd
that the bulbus changed its oblong form to an almost spherical one.
In most nomatodcs collected from turnips in August 1939, bulbus dimensions
were 16 x'll microns. In thc..onion'population it is considerably more
elongated and averages 20 x 10 micron's.
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(1)
hilrav'ev, V. P.
Izmenchtiosty Virulentnosti Mokroi Golovni
v Zavisimosti ot Prokhozhdeniia Tsikla ee
Razvitiia cherez Raznye Sorta Pshenitsy
[Variability of the virulence of stinking smut
in relation to the passing of its developmental
cycle through different varieties of wheat]
Doklady Akademii Nauk SSSR, Juneil, 1954
Vol. 96, No. 4, P. 857-859 511 P44.4A
Trans. A. 527
Ffom the Russian by
R. Adelman
Variability of the virulence of stinking smut
in relation to the passing of its developmental cycle
through different varieties of wheat
In the fall of 1951, seed of winter wheat were inoculated with -
the spores of stinking smut (Tilletia triticWbefore they were seeded.
The spore stock was entirely homogenous; it was obtained from wheat spikes
also infected'as a result of spore inoculations in the preceding vege-
tative period.
We gathered separately spikes infected by smut from five varieties
(belotserkovskaia 200, Ligovskaia 3612, Khartkovskaia 917, Liutesconce 17
and Ukrainka) and extracted from them spores of stinking smut which
had completed their cycle of devel4mont in one vegetative period in
different varieties of winter wheat.
? To determine whether Or not the passing of smut through different
varieties had had an effect on its prOperties, and to ascertain the
character of this effect, we, in the fall of 1952 prior to seeding, .
inoculated spores into seed of the same varieties of wheat from which
the spore stock had been obtained, and, in addition, three other varieties,
namely: Lesestepka 75, ErythrosporMum 15, and Liutoscence 9. Thus,
experiments wore conducted on 8 vafietios of wheat characterized by
sharply varying resistance to smut. This was demonstrated decisively
by infection indicators. obtained as a result of calculations made at
the end of the vegetative period in 1953 (see table 1).
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T a b 1 c 1.
infection of winter wheat by'smut-that had passed through different .
varieties (percentage of infected spikes)
Varieties tested
Varieties from which !ores
Belotser, govi Khaf'L,
kovskaia 2001 skaia ' kovska
507M 917
47P--
Belotscrkovskaia 200
L'govskaia 3612
Kharikovskaia 917
Liutescence 17
Lesostopka 75
Erythrospermum 15
Ukrainka
Liutescence 9
14:9
40:6
45:8
525
75:0
61:3
77:0
71.5
were obtained
Liutcs- Ukrain- Aver-
conce ka age
17
12:8 150 15:9 12:7 14:3
42:7 41:2 36:3 40:5 40;3
547 577 44:8 58:4 52:3
573 63:3 49:8 54;3 554
69:6 65:1 72:5 58:0 68:0
70:3 699 739 68:3 68:7
71:2 74:1 552 67:1 69:0
60.1 71.2 81.1 64.1 69.6
Belotserkovskaia 200 proved the most resistant to stinking smut, which
coincides with its well-known industrial characteristics. The data cited
in table 1 demonstrate fully that the relative infection of the varieties in
this experiment, determined by percentage of spike infection, fluctuates
greatly when seed is inoculated with spores derived from different varie-
ties of wheat. Besides, the range of infection fluctuations within the
limits of each variety is less considerable in cases of rolativelSr
resistant varieties and more so in cases of susceptible varieties. Thus,
in the Belotserkovskaia 200 and L'govskaia 3612 varieties, the difference
between the maximal and minimal infection - U.ki-ainka and Liutescence 9 -
the difference is 21.8 and 21.03 respectively.
, Table 1 indicates that regardless of which smut was, inoculated,
Belotserkovskaia 200 and L'Ovskaia 3612 varieties hold first and second
places in resistance to smut. Third and subsequent places are held by
different varieties; thus, Lesostepka 75, Liutescence 9, Khar'kovskaia
917, and Ukrainka are found in fifth place. Mth respect to the last
variety, it can definitely be said that the appraisal, which it received
during an infection by smut that came form. the Liutcscence 17 variety,
did not correspond to the well-known charactcrization'of Ukrainka as
one of the most susceptible varieties of winter wheat. Thus, the
relative susceptibility of varieties, and the place occupied by a given
variety among a number of varieties tested, depend on-the spore
material on which the degree of resistance was tested.
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(3) Trans. A, 527
The data cited-above indicate that smut changes after passing through
different varieties. The'very important property that changes in par-
ticular is smut virulence: This 'circumstance attracted the attention of
the phytopathologist V. N. Shovehonko. It is a circumstance that must
be taken into consideration in the work of selection and development of
resistant varieties. To obtain comparative appraisals, it is.nocegsary
to use spore material of uniform composition for spore inoculation.
To Obtain an appraisal of variety resistance to smut for industry, it
is necessary that, for the preparation of spore material, smut infected
Spikes be collected in the region for which the varieties under appraisal
are, designated, and that varietal environment be taken into consideration.
By analyzing data on the degree of infection caused by smut obtained
from different varieties, and by ascertaining tho variability of its,
virulence; we can establish the character of the variability of smut
virulence. For this purpose, we will compare tho susccptibilty of
varieties with tho virulence of smut which has pa6sed its cycle of
development through a given variety (sec table 2).
The degree of susceptibility in table 2 was takcnfrom tho last
graph of table 1. For the virulence degree of smut form the given
variety we took themagnitude of infection caused by inoculation of its.
spores in all of the other 7 varieties. From an appraisal of this sort,
the virulence of smut that had passed through the Bolotserkovskaia 200
variety proiod maximal it caused a 60.5% infection in the rest of the
7 varieties. The minimum indicator of virulefice (50.7%) was found in
smut from Ukrainka, the most infected variety. ?
Thus, one can arrive at tho conclusion that tho greater the sus-
ceptibility of the variety, the less nocuous is the virulence of smut
which has pagsed though the given:variety. So, the correlation coeffic-
ient r 0.91 4 0.08, i.e. the reversible correlation, is sharply
pronounced and has been fully proved mathematically.
Thus, if tho cycle of smut development was consUmmatod in a resistant
Variety, then its virulence increases. Such a rule, proved experimentally
in tho case of stinking smut of wheat, undoubtedly is true also with
respect to causal agents of other plant diseases.
An adequate adaptation of the parasite to varieties i6 one of the
reasons for the apparent "loss of resistance by a variety".. "Apparent"
- because in tho given case, the phenomenon taking place is of a different
order t the variety can fully retain its resistance, but the reproduction
of the resistant variety will contribute toward an accumulation of a
more virulent parasite in the surrounding area.
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(1)
Trans. A. 528
Fftm the Russian by
R. Adelman
Krivykh, F. P.
Pochva s Parovogc Polia kak Bak:terialfnoe Udobrenie
[Soil from Fallowed Field as Bacterial Fertilizer'
Doklady Vsesoiuznoi Ordena Lenina Akadeffidi
Sellskokhoziaistvennykh Nauk imeni V. I. Lenina
Gosudarstvennoe Izdatelistvo Sellskokhoziaistvennoi
LiteraturSr
Moskva, 1954, Vol. 19, No. 2, P.16-20 20 Akl
SOILS FROM FALLGYED FIELD AS. DAC TERIAL FERTILIZER
(Submitted by the Plant Breeding Section of the All-Union
Lenin Order Academy of-Agricultural Sciences named for
V.I. Lenin)
The modern Michurin teaching concerning plant nutrition is based on
the close-inter-relations between higher green plants and, soil micro
organisms. Cultivated plants cannot develop normally when they lack
the appropriate microflora. At present, it is established that each
species of higher plants has its specific, coresponding groups and races of
microorganisis. It has long been known that a fallowed field is the best
predecessor of many crops. There is no doubt that here favorable con-
ditions for plants are created not only by means of water and food
reserves, but also by iariation in the qualitatiire and quantitative com-
position of microflora. A fallowed field contains considerObly more
microorganisms than virgin soil or a potato field (table 1).
Table 1.
Amount of bacteria ( in millions per gram of absolutely dry soil
On unfertilized fallow 1641
It potato field ' 1106
tt virgin soil 723
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(2) Trans. A. 528
The numbers cited show that the athOuntOf bacteria in virgin soil is less.
than half.that on a followed plot. It must also be noted that the
activity of aerobic bacteria proceeds more intensively on a followed plot.
. Not only the number of microorganisms decreases in virgin soil, waste
land, and in the layer under perennial grasses on structuroloss soil, but
their-activity, toe proceeds in a direction other than that in a followed
field. The introduction of soil from fallow effects a rapid chane in the
composition of microorgainsms and, thus; causes that inhibit the normal
development of plants can be eliminated.
For the,purposo of ifivostigating this problem, virgin soil was
plowed on LugUst 25, 1949; It this time, the biological processes in
soil were already ceasing. Seeding of winter rye was accomplished the
same day. The experimental scheme was: 1) control; 2) test Plot ferti-
lized with superphosphate; 3) test plot fertilized with soil from a
followed field (calculated at 4 centners per ha) and with superphosphate
(2 c per-ha). The fertilizers wore introduced into the rows together with
the seed.
On the fertilized test plots sprouts appeared on September 1-2nd,
yet on the control plot only on September 10th. Besides, most seeds on
the test Plot produced no sprouts whatever. On September 14, when germina-
tion was evaluated, it was found that the test plot fertilized with soil
from the followed field had produced 252 plants per square meter, the
test plot fertilized with superphosphate, 140, and the control plot
only 56. Tho sprouts on the control plot were characterized by poor
growth. The following year,,only the plants on the tett plot fertilized
with soil from the followed field developed normally; [Begin P. 1.7]
they flowered normally and produced seed. On the other test plots,
rye developed poorly, it managed to flower, but produced no seed.
Evaluation arc cited in table 2.
T.a b le 2.
Variant
Aeight of plants
.from 4 m2 in
air-dry state
(in gr)
? Height of
plants
(in cm)
Control ? 156
Test plot fertilized with superphosphatO7 324
Test plot fortilited With-Soil from 1368,
followed field
78
91
]116
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(5) Trans. L. 523
On the test plot fertilized with soil froth the followed plot, plants
developed much bettor than on the control plot. Fertilizing with super-
phosphate changed the situation but littic.
This experiment convicod us of the oxpodioncy of utilizing soil
from a followed field as bacterial fortilizOr. In subsquont experiments,
field germination of seed incroasod.considorably-and plants developed
better when soil from a followed field was added.'
Spocial experiments conducted to test the offoctiveness of the
application of from a followed field upon field germination of seed (soil
was introduced together with seed) have shown-(table 3) that in many
cases it can increase by 14 and oven by 20% 1.
T a b 1 o 3.
Crop and
varioty
Seedlings Differ-t
from 600 once
Variant seeds
Average
field gor-
mination
(in %)
Experithont-ofI951
Udarnitsa wheat
Control...04,00,4ft
476
78
With introduction of
soil from followed field.
_
504
28
84
Sibirskii flax
Control
383
63
With introduction of-66i1
from fallowod field
463
80
77
Tulunskow mil-
lot 39/9
Control
With introduction of toil
380
63
from fallowod field
473
93
76
Foxtail millet
Control
311
62
With introduction of-tail
from fallowod field
413
102
69
ExperimoritThf-1952-
Tulunskoo mill()
Control
275
46
39/9
With introduction of 8Oil
from followed field
310
35
52
IF. P. Krivykh. Problem of Wrmination of Sood. num. 'Scloktsiia i
Somonovodstvo" No.3, 1953.
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(4) Trans. 528
To increase the biological activity of Oil taken from a fallowed field,
bacteria-breeding nurseries were established. On small plots measuring
200-300 m2, 500-600 tons of manure were applied according to per-hectare
estimates; it was plowed under and the plots wore maintained in a stdte of
pure fallow, with plowing being repeated 3-4 times during the summer. 'In
the summer the manure decomposed into a chernozem-like mass. (Begin P. 18]
The following year the soil from the nursery was applied as a fertilizer
for grain and other crops. This soil contained not only a largo amount of
microorganisms, but was also richer in nitrogen, phosphorus and potassium.
Bacterial nurseries mut be established either near cattle yards or near
the fields to be fertilized.
In the first instance, the dosage of manure fertilizer can be increased
to 1500-200 tons per ha, utilizing the nurseries as manure storage (the
manure is applied at intervals).
If nurseries arc organized near the fields to be fertilized, than
manure can be applied at the rate. of 200 to 500 ton per-hectare. One
hectare of the bacterial nursOry can produce from 1 to 3 thousand tons of
soil to be usod as fertilizer. The amount of fertilizer derived farm one
hectare depends on the depth at which tho soil is taken -.the arable horizon
from one hectare weighs about 3000 tons. The manure, applied must be plowed
in and harrowed immediately, and later in summer, the bacteria-breeding '
nursery must be kept in a mellow and moist condition (with 4-5 treatments).
The nursery can be established either in the fall or spring. In well
prepared murscry soil, the thanure must decompose completely and turn into
a. mass resembling'chernozem.
We arc citing some experimental data of the Irkutsk Agricultural
Institute, the Balydacv Experimental Station, and partial data of .industrial
experiments.
Experiments of the Irkutsk 1,gricultural Institute. Soil from a
fano-wed field and from the bacterial nursery was applied for wheat, barley,
oats and potatoes. Data on productivity arc cited in table 4 (in ccntncrs
per ha).
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(5) Trans. k. 528
Table 4.
Variant
1952
1953
"Meat after Barley after
perennial potatoes
grasses
Oats
after
pota-
toos
Oats after
potatoes
Control
77ith,soil from fallairod-fiOla-
13:35
26.25
27.60
2 tons per ha
15.22
? 16.22
ON OM
29.81
,lith soil from nursery
2 tons per ha
15.83
18.19
31.45
Same, 20 tons per. ha
_
3709
'jith humus, 20 tons per ha?
15.68
18.51
33.33
In the experiments cited all fertilizers were appliOd in pre-seeding
tillage and were plowed under with goosofoot cultivators. Application of
soil from a followed field produced an increase in yield in all oases; the
strongest action was .3)mi-tad by soils from the bacterial nursery.
Table 5.
Variant*
Potato
yield
(in c/ha)
. 1952 ,
increase
(in c/ha)
1953
Potato
yield
(in c/ha)
Increase
(in c/ha)
Control
With humus, 10 tons pdr ha
77ith soil from nursery,
10 tons per ha.
Same 4 phosphorobactorin
Phosphorobactcrin
1770
204.0
238.0
????
27.0
61.0
NOON
20544
219.55
24576. ,
26910
223,02
41.11
4032
63:66
17.58
*Fertilizors were applied in holes when potatoes were planted
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In experiments with potatoes, the best results were also obtained
when bacterial nursory soil wasapplied, and when potato-tubors were given a
supplementary treatment with phosphorobactcrin (table 5).
Experiments of the Balydacv Exporimcntal Station. At the Balydaev
Expoiimontal Station, activated soil from a followed field was used in
1951. For this purpose, the soil was kept in a moist and loose state in a'
warm compartmont for several months, and then applied in pro-seeding dik-
ing of soil. Tho experiment-was conducted oh a-timothy bed: [for wheat]. .
Results aro cited in table 6.
T a b 1 0 6.
Variant
Moat
yield
(in c/ha)
Incroaso
(in c/ha)
Control
2417
7fith manure, 40 tons per ha
28:14
3:97
711th soil from followed field,"6 contnors per ha
30:04
587
'Aith soil from nursery, 6 c/ha
30.24
6.07
. The 1952 and 1953 experiments produced loss fa-crorablo results, but the
in crease inthe yield of wheat was, nonctholoss, 2.6 and 3.9 centners per
ha (table 7).
T a b 1 c 7.
Variant
1952
? 1953
Yield
kin c/ha)
Increase
(in c/ha)
Yield
(in c/ha)
Incroaso
(in c/ha
Control
'.7ith soil from fallaW6d-field-
1.5 contner per hi
Jith soil from fal1oWea-fiO1d-4
azotobactorin
'lith soil from nursery, 3 c/ha:
ft
6 c/ha:
20 c/ha.
27.9
30:5
30:7
31.8
m1.10
2.6
28
3.9
11,13
13.72
14.23
2.65
3.10
Here, too, the application of soil from-tho followed plot combined
with azotobactcrin produced the best results.
411. Vary good results wore obtained also when seeds were trated'ilth a
liquid solution of soil from a fano-wed plot.
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(7) Trans. A.'528
Treatment of seed was carried out as follows. For every 100 kg of
seed 1-2 kg of soil from a fallowed field was uscd; this soil was disolvod
in 30 liters of water and the solution obtained was-used to steep the seed
(the. same as it is done-in a case -ct vernalization). Aftei, 24 hours, the
seed was dried and .sown. The results arc cited in table 8.
Seed treated witn a solution-of soil from a followed field produced
a yield increase of 3.16 centners.
Analogous experiments conducted at the-Irkutsk Agricultural Insti-
tute confirmed the prospects of this method.
Experiments on kolkhozes [Collective farms], In 1951, agronomist
V. Azhunov of the 'Ent' Illichan kolkhoz, Baiandaev aimak*? used granu-
lated soil. from a fallowed plot to fertilize spring wheat seeded after'
winter rye in a field plowed in the autumn for spring crops. [Begin P. 20].
The soil was used as fertilizer in pre-seeding tillage at 8 centners
per ha. On the control plat, he obtained a yield of 16 centners per ha,
yet on the plot fertilized with soil from a fallowed field, 20 centners per
ha.
In 1952, tovarishch Anan'ev, initiated an analogous experiment with
spring wheat of the Liutescence 62 variety. On the ?Komintern" kolkhoz,
Kuitun Region soil was taken from farm fallow and applied under pre-seed-
ing cultivation. The results of these experiments are cited in table 9.
The data cited show that particularly good results were obtained
when PK [potassium phosphates (?)] was added to soil from a followed field...
Tab 1 e 8
Variant
Wheat yield
(in c/ha)
Control (dry seed)..... .
Seed treated with pure water
Seed treated with solOtion of soil
from fallowed field
[*Localism for "region".]
2805
26.27
31.21
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I' able 9.
Yield ? Increase
in c/ha) in c/ha)
Variant
Control
With soil from followed field 2.5 tons per ha
With humus, 10 tons per ha
With soil from fallow field, 2.5 tons/ha-PK
CONCLUSIONS
1450
1750
16:00
19.75
I. Soil from a fallawed field can be utilized
lizor. Such soil exerts a favorable influefice upon
required groups of microorganisms within it. Small
of such soil seeded together with seed considerably
grain and other crops.
2. The effectiveness of soil from a followed field increases markedly
if it is first exposed to warmth either in a warm chamber or in the sun for
several weeks. Hence, it must be taken from fallow located on southern
slopes which arc exposed to warmth; from plots rich in organic substances or
fertilized with manure.
1:50
5:25
as bacterial fcrti-
plants, thanks to the
doses (4-6 centners).
increase the yield of
3. Soil from a followed field cannot be compared with other bacterial
fertilizers, it must rather be used to supplement others. The combined use
of soil from a followed field and other bacterial preparations is more
effective than their application separately.
4. Followed soil produces the best effect When it is introduced in
the soil together with seed. When it is applied with a cultivator, then the
amount of soil per hectare must be increased up to 2 tons; itis best
plowed under with disk tools..
5. Soil from a bacteria-breeding nursery is most effective because,
in addition to a large amount of needed microorganisms, it contains many
nutritive substances easily accosible to plants. ThereforO, the estab-
lishment of such nurseries must be considered as expedient.
Irkutskii Sol'skokhoziaistvennyi Received at editorial
Institute Hoffice -
.January 19, 1954
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Matskov, F. F. and Podrazhanskaia, B. S.
Stimuliatory Posta kak Nikroudobrcniia.
[Growth Stimulants as Micro-Fertilizers].
Doklady Akadcmii Yauk SSSR, Apr. 21, 1954
Vol. 95, No. 6, P: 1329-1331 511 P444A
Trans. A. 529
Fllom the Russian by
R. Adelman
PLANT PHYSIOLOGYY.
GRUITH STIMULANTS AS NICROFERTILIZERS
Stibmitted by Academician A. L. Kursanov
February 26, 1954
In experiments conducted in 1948-1950, it was demonstrated that in-
troduction in soil of very small amounts of some organic growth stimulat-
ing substances, such as hetcroduxin, o.-naphthyl acctie acid (AHY), 2.4-
dichlorphenoxy acetate acid (2.4-DY), in doses from 0.1 to 0.001 mg per-kg
of soil, increases considerably thc yield of the dry mass of plants (1).
In addition, the experiments of 1950 revealed that introduction of growth
stimulants (together with microfertilizors) into soil furthers the growth
and the development-of the root system of plants to a higher degree than
their surface parts. 'AC cite the results of,some.of those experiments in
table 1.
Table 1.
Results of experiment with oats in Rotmistrov's boxes (plants harvested at
50 days of life)
Experi-
mental
variants
Height of Length 117cight of dry s icight of dry Total weight
surface of roots mass of sur- mass of roots of dry mass
organs in cm face parts of plants
in cm
gr !
NPK
NPK1-AHY*
NPKthetero-
aukin
NPK-2,4-DY
756
91.0
870
87.5
*0.1 mg per kg of soil.
505 10:90; 100:0
62.5 12.261 112.4
_ ?? ! -
57:0 12:80 1 1174
65.0 ;1,11.80 I 108.3
i
gr
gr
%
4:34
100:0
15:24
100;0
5.26
_
121.9
17.52
114.9
5:18
1216
17;98
1181
5.30
122.1
17,10
112.2
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Table 1 shows that mixing a microdose of ABY with NPK increased the
weight of the dry mass of surface organs of 50-day oat plants by 12.4%,
and the weight of their roots - by 21.9%; under the influence of hetero-
auxin; the weigfit of the dry Mass of surface organs and roots'incroased
by 17.4% and 21.6% respectively, and under the influence of 2.4-DY - by
8.3% and by 22.1%.
Thus, on the basis of our threo-year experiments, we can assort that
the effectiveness of mineral fertilizers increases, if small doses of '
stimulating chemical preparations are applied together with fertilizers.
We were unable to ascei-tain haw long stimulants introduced into soil
continue to exert action, i.e. remain active without undergoing further
transformation. Seeking an answer to this question, we initiated an ex-
periment with oats in 1951.
Tack in the fall of 1950, vegetative containers were filled with soil
(4 kg to a container) to which (pursuant to the experimental scheme) a full
.hutritive mixture (NPK) in Prianishnikov's norms was added simultaneously
with an admixture of microdoses of growth stimulants [Begin P. 1330]
(hoteroauxin ABY and 2.4-DY) - at 0.1 mg per kg of soil, Some of the con-
tainers were filled with pure soil. All containers were divided into two
groups, ofte'of which was kept in a little cold vegetation house until
spring (i.e. at temper-tures below zero), and the other in a warm green-
house where temperature was kept at 15-17?.
In the spring, the containers Were nrefillee, i.e. ?the soil was re=
moved from them, loosened, moistened and replaced in the same containers.
In so doing, mineral fertilizers (NPK) with an admixture of microdoses of
growth stimulants were added to the ,8ontainers which had been filled with
puro soil in the fall, as proscribed. From the decription of the experi-
mental scheme, it is obvious that our objective was to compare the action
of growth stimulants added to the soil in the autumn with the action of
those added in the spring. In addition, we wanted to obtain an answer to
the question as to haw growth stimulants will alter in soil left for 576
months under conditions of negative temperatures (when microbiological
processes do hot function), and under conditions of room temperature wheh
growth stimulants may be subjected to the action of soil microflera.
The results of this experiment are summed up in table 2
Analyzing the data in table 2, one can draw the following conclusionS;
the different growth stimulants used in the experiment change during a
prolonged stay in soil in various ways. Factors of temperature in pro-
mises where soil containers with added growth stimulants arc stored, exert
much influence upon them.
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(3) Trans. A. 529
Hetoroauxin introduced in to the soil in the fall, obviously, disap-
pears almost completely by spring of the following year, and, therefore, it
does not exert its inherent positive action upon the plants, In contrast,
2.4-DY, if added to soil in the fall, acts on the accumulation of organic
matter by lants even better (419,81.) (Begin P.13311 than if added in the
spring (48.7).
It is true that the yield of grain is higher, if 2.4-DY is added to the
soil in spring. As far as AT/ is concerned, its stay in soil during the
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Table 2.
Influence of season and conditions of .domicile of growth stimulants while in
soil upon thier activity (results qf vegetative experiment with oats in 1951).
ISeaspn Storage ? Height o4 Weight of Weight of
Ferti-
lizers
when place of plants 'dry mass dry mass
ferti- contaih- in cm of straw of grain
lizers er in gr gr
added winter
Total weight
of dry mass of
plants
gr
NPK
(Sep.30
1950)
NPK-i-ARYI Same
2.4-DY
NPK1L
hetero-
auxin
NPK .Spring
(Apr.30
1951)
NPK4AEY Same
NPKt
2.4-DY
NPK4-
Hetero-
auxin
NPK Fall
(Sep.30
. 1950)
NPK+ABY Same
NPK Spring
1(Apr.30
j 1951)
WPK4AHyi Same
Veget.
house
Same
ft
tt
tt
Green-
house
Same
If
101.5 8.25 100.1, 6.15-
112:0 10.14122.4-6.00
107.0 10.50 127.3 6.75
112:0 8;17 991.(f6-:57
111.0 10.85 100.0k8.05
118:0 13.15 121.2
108.0 11.05 101.8
100:4141404p*ID 0010'
I -
97.7116.10+0.00 118.8
109.8! 17.20.95 119.8
106.8! 14:74+0:74 102.4
100.0118.904-0.70 100.0
9.25 114.9
9.50 117.9
112:0 12:82 118:2 9:45
100:0 7.17 100.05.40
108:5
110.0
110.0
9?:70
11.35
'14.17
22.40+1.00 118.5
20,55+0.78 108.7
1174 22:27092
100.0/12.571-0.12
135:3 7:27 134.6
100.0 8.22 100.0
122.0 10.03 121.9
117..8
100.0
16:97+1:57 135..0
19.55t1.07 100.0.
24.20f0.45 123.7
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(5) Trans. A. 529
winter (under conditions of the little vegetation house) did not result in
its.complete disappearance, but it decreased considerably its positive
action upon plants seeded in the spring. This was particularly pronounced
in the case of grain yield.
Keeping soil to which ABY has been adUed under greenhouse conditions
does not feduce the positive action of growth stimulants upon oats seeded in
this soil.
Conclusions concerning the duration'of action of growth stimurants
added to soil in the fall are not final,since they are based on experi-
mental data of one year only.
Received
Juno 29, 1953
REFEREFCES CITED
'F. F. Matskoi, B. S. Podrazhanskaia, DE-IN [Doklady Akademii Nauk] 66 No.5
(1949), 72 No.2 (1950).
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Trans yo. A. 530
Tablsi of contents
From the Russian by
R. Adelman
TURBIN, N.V,w Ed.
Voprosy Biologii Oplodotvoreniia
[Problems of the biology of fertilization]
Izdateltstvo Leningradskogo Universiteta
Leningrad - 1954
394 P. 463.6 L54
CONTENTS
FOREWORD ? 3
Turbin, N.V. and Bogdanova, E.N.
Experimental data on the study of multiple fertilization in plants..:. 9
Turbin, N.V. and Bogdanova, E.N. Experimental study of the influence of
repeated polination upon fertilized ovicells in tomatoes 48
Aizenshtat, ILS. Influence of fertilization conditions upon hereditary
Transmission 73
Turbin, N.V. Biological role of alien pre-pollination 141
Turbin, N.V. and Narbut, S.I. Problem of the biological role of repeated
pollination in plants 163
Turbin, N.V., Gorobets? A.M. and Narbut, S.I. Concerning unutilized
reserve for increasing the productivity of tomatoes 176
Turbin, N.V. and Palilov, A.I. Castration of maternal spikes in free
inter-varietal repollination of wheat 188
Lebedev, M.M. Certain regularities in reproduction of domesticated
animals ,212
Kurbatov, A.D. Influence of the physiological condition of animals and
the properties of their sex cells upon the sexual composition of the
progeny 245_
Kilrbatov, A.D. Transformation of the nature of animals in inter-
specific transplantation of the zygot, gonad and in blood transfusion
of animals of a different species 277
lakarov, P.V. Cytological and cytochemical analysis of gametogenesis and
fertilization in the light of Nichurin's teaching 319
0 References 382
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(1) Transl. A. 531
From-the Russian
By R. Adelman
SUKHOV, K. S.
Virusnye bolezni Sel'skokhoziaistvennykh
Rastenii i mery bor'by s nimi
[Virus diseases of agricultural plants
and measures for their control]
Izvestiia Akademii Nauk SSSR
Seriia Biologicheskaia, Moscow, 1954
49-61 511 Sa25
VIRUS DISEASES OF AGRICULTURAL PLANTS
AND MEASURES FOR THEIR CONTROL
In the light of decision adopted by the September Plenum of the
Central Committee of the KPSS [Communist Party of Soviet Union] on
measures for the advancement of agriculture, improvement in agricul-
tural plant protection against diseases and pests assumes great
importance.
A significant place among the diseases of. agricultural plants is
held by virus diseases some of whioh are distinguished by wide dis-
tribution and great perniciousness. Most of-these diseases have
been well investigated by Soviet virologists. However, the practical
measures for their iradication,'Or restriction of thoir injurious
action, suggested at various tim6s in native literature, have not
always boon utilized in industry. In connection with the above, there
arises a need to study this problem at the present time, focusing
particular attention on the thoorectical basis-of control measures
recommended for the principal virils infections.
Virus diseases Of agricultural plants, the control of which is
of groat economic importance, arc the following:
1) For technical crops - control of tobacco mosaic, crown
chlorosis of makhorka [Azted tobacco], loaf curl of cotton;
2) for cereals - pupation of oats and mosaic of winter wheat;
3) for fruit crops - psorosis or peeling of bark of citrus plants;
4) for vegetable crops -" stolbur" of tomatoos, popper and egg-
plant; mosaic and "streak" of tomatoes, mosaic No.2 of cucumbors;
5) for pbtatbes- -"stolbur" wilt and wrinkled mosaic, or streak.
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1.1
S
?
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, (2) Trans. A. 531
Thc causal agents of those diseases arc different viruses
whose biological characteristics determine to"a considerable extent
the opiphytological character of each disease. Home, conditions
for the development of different virus diseases of agricultural
plants are dissimilar, and methods for their control must be changed
accordingly. Tho biological differences of the agricultural crops
themselves are no loss important, and their properties of ithmanity
or resistance to virus diseases often determine the outcome.
Knowledge .of the biology of the causal agent of disease and of
the agricultural plant which it attacks helps to find moans for
decreasing substdntially the loss caused by disease, and to in-
crease the yield.
The investigation of tobacco mosaic conducted by D. I. Ivanovskii
at the close of the past contury initiatod-rescarch in a new sphere of
biology the science of-filtrabic virusos. Having made the greatest
thcoretical discovery, D. I. Ivanovskii at the same time :took great
pains to develop means that would aid in the conti-ol of diseases
which greatly impaired the quality of raw tobacco. He accomplished a
great doal in this rcspoct2 an accurate diagnosis of the disease
was worked out, the -spread of infection through contact was as-
certained, [Begin p. 50], and it was demonstrated that the first
cases of infection could be Caused by retention of infectious remains
in the soil, which indicated the nocessity of introducing tobacco
into crop rotations. It was proved that infection was not trans-
mitted via tobacco seed.
It can bo said in substance that D. I. Ivanovskii established
the main elements- of the epiphytoiogy of tobacco mosaic and deter-
mined the principal moans for its control Which have lost none of
their improtance up to now. This tradition of combining theoretical
investigations with the practical tasks of agriculture is characteris-
tic also of modern Soviet virology.
The All-Union Institute of Tobacco and Makhorka (VITR) continued
to develop methods for the control of tobacco mosaic in the ninetoen-
thirtio6; hero, the most important rosults were obtained by I. P.
Khudina.
The biological charactoristics of the virus causing this disease
complicated its control. The virus of tobacco mosaic is distint
guished by an unusually high degree of infectiousness. dnd'by very
strong resistance to unfavorable action of environment. Light con-
tact ar a porson's, 'hands,. soiled with infectious sap, -with the loaves
or stems of healthy plants causing microscopic wounds, is onough to
enable the virus to penetrate into coils and to cause infection.
Having entered the injured cells, the virus multiplies, spreads
through the adjacent cells and the vascular system, and then pone-
tratos all organs of the plant. Hence it is natural that trauma
producing methods such as thinning of seedlings, planting seedlings
in the filed, breaking off suckers and leaves facilitate the rapid
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spread of tho virus ovOn if infection nidi were few at the beginning.
Original cases of infection might be duo to penetration of plant
rOots by tho virus from soil containing infectious plant remains.
As a result of growth movements, epidermal root cells aro likely to
be subject to slight injury caused by hard particles of tho soil;
this, too, creates opportunities for infection. Although tho virus
that has onoterod the root usually remains there without penetrating
the surface parts of tho plant, there, nonotholess, aro known cases
of a root infection having turned into a general one. Infection by
contaminated soil occurs mostly through tho contact of soil with
the lower loaves. Tho most frequent source of first infectionsis,
however, man himself who transmits the virus by hand. It suffices
to say that tho virus is preserved in smoking tobacco, .and that tho
hands of tho smoker may always be soiled with it. Utilization of
implements that had already boon in use, for instance, hothouse
frames, baskets and boxes for the transportation of seedlings to a
field, is apt to infect a certain number of plants, unless tho
implements had been disinfoOted. Infection contacted-from those
first sources is later spread by the hands of workors.
Tho characteristics cf opiphytology of tobacco mosaic cited
make it clear that tho main efforts in reducing this disease must
be concentrated on prophylaxis. Hence, tho principal task involves
- neutralization or destruction of infection nidi. This can be
achieved by tho introduction of tobacco in crop rotations and by
disinfection of equipment with a solution of formalin. If the
disease has spread but little, not ovor 2%, thon the infected plants
are removed from the plantation.
The most important positive results in Mosaic control are ob-
tained by development of resistant variotios.' Iri this respect,
recognition is duo the VITIM brooder, Prof. M. F. Tornovskii (1950).
As a result of crossing Turkish tobacco with Nicotiana glutinosa and
of subsoquont back crossing Of tho hybrids with tobacco, ho"ob-
tained varieties which have inherited virus immunity from N. Glutinosa.
Recently he o")tainod a series of resistant varieties, by intra-
varictal vegotative hybridization (1951). Resistant tobacco varieties
wore obtained by the same method at the Iristituto of Genetics,
Academy of Scionces, USSR (Agapova, 1953). By extending the area
under imthune tobacco varieties, mosaic infection will finally be
overcome. This is not the first virus disease [Begin p. 51] the
oradication'of which on a specific crop is a task to be realized in tho
near future.
Tobacco mosaic is one of the first virus diseases subjoctod to
research. Other virus diseases nave boon identified only recently and.
some have not yet boon diagnosed. Therefore it is not surprising .
that tho urgent task of dotormining tho nature of a now disease
named top chlorosis and spread in some rogions on makhorka plantations
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has arisen comparatively recently. A. A. Popova was the first to
describe this discasc and to demonstrate its infectiousness. To
dovclop measures for its 'control, it was necessary to invoFtigate-
the causal agent of the disease and its moans of spread in nature.
For this purpose was organized a complex investigation of the disease
in-which thc Institute of Genetics, Akadumy of Sciences, USSR
(K: S. Sukhov), the:A114,Union Institute of Tobacco and Makhorka
(S.E. Grushevoi-and T.M. Matveenko),-the Moscow Station of Plant
Protection (GM. Razviazkina Butsevict),-and the Lokhvitskaia
Experimental Base of VITIM (A. A. Popova and P. M. Diadiuchenko) were
participating.
With the aid of an electronic microscope and, filtration of
infected juice through bacteriological'filters, it was established
that the causal agent was a filtrable virus with particles of d
spherical form measuring about 45 milomicrons in diameter (fig. 1).
The disease transmitter, tobacco thrips, was established concomitantly.
This helped determine the nature of the disease and to characterize
the asic opiph9tOlogical elemontg in the first-year of research
(K.S. Sukhov, G.M. Razyiazkina, L.A. Butsevich).. Clear orientation
as to the develotment of protective measures was obtained in the courso
of the same work. One of these measures is the chemical method of
dusting makhorka plantings with DDT dust, and of introduCing BOnzino
hoxachlorido [GICBTSG] into tic soil, under makhorka (S.E. Grushcvoi,
T.M. Matvcenko, A.A. Popova).
At present, to chlorosis of makhorka is one or the well in-
vestigated diseases. Tho control method developed is used widely in
industry and produces effective results in raising the productvity of
this crop.
In the year 1953 the author of the present article and G. M.
Razviazkina established also that the tobacco disease known by
the name of "virus top deformity" (S.E. Grushevoi, 1951), was
identical with top chlorosis of makhorka. Thus, the information
obtained concerning the conditions of development *of the discaso
epiphytosis on makhorka can be utilized also in tobacco culture.
Chemical control of thccarrior of top chlorosis - tobacco
thrips - is fully equal to the task of eliminating the serious in-
fection of this crop. But this special measure must definitcly be
combinca with agrotechnival practices increasing plant resistance to
disease. In this respect, splendid results have been achieved by
leading teams of workers of makhork&grawing sovkhozos and kolk-
hozcs. 'For instance, the team of. M.IA. Fudorcnko, hero of socialist
work (1952), harvests up to 85 contners of makhorka per hcctarc by
using the entire complex of control-measures against top chrlorosis
and superior agricultural practices. This indicates that the first
resolution of the-control problem of top chlorosis of makhorka has
been satisfactory.
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In the year 1936, D.D. Vordorevskii described loaf curl, a
virus disease of cotton in Turkmcn. Later thi8 disoaso was discovered
also in Azcrbaidzhan whore it was studied by S.N. Moskovots (1951).
The discaso manifests itself in sharp doprossioft of plant
growth, in lodging of the amin stem arid in loaf curl. The damage
caused by thc disease is considerable. Cotton yield of diseased
plants drops sharply. There is a 5% reduction in the number of
pods, the technological qualities of the fibcr lcngth arid strength -
arc impaired, and so arc the propOrtios of sed absolute weight, f
germination =orgy, and sprouting. [Begin p. 52]
A 'small percentage of the distaso is transmitted by seed.
Its chief method ?spreading is, however, transmission by melon
and cotton aphids. Permanent virus sources in nature are suscep-
tible weeds, for instance, mallei and rape on whieh-cotton aphids
frequently settle in mass. (D.D. Vcrdcrovskii, S.N. Moskovcts).
Clarification of the basic factors of the spreading and develop-
ment of the disease made it possible to work out a complex of
measures as a-result of which leaf curl no longer causes any serious
lossA.n yield. The solution of this problem is one of the indicators
of the success ?of Soviet virology.
In overcoming the harmful'action of. discasc,. solectioti of re-
sistant cotton varieties pox-formed an important role. F.M. Muer
developed Azl, Az29, 2966-1, and other vdrictios distinguished by
a high degree of resistance to leaf curl. ? These varieties aro
particularly valuable for thoir'high resistance also to the wide-
spread bacterial cotton disease - Gummosis.
?.Agrotochnics are of groat importance in maintaining cotten
resistance. Ina regimo'of mineral nutrition of cotton in the regions ?
of diScasc distriblition, an important role belongs to phosphorus
fertilizers. Phosphorus As introduced in soil during fall plowing and
again in supplomentary feeding. To ensure a steady decrease of
virus resorvos in cotton growing rogion8, the destruction of carrier
aphids and woods is of great importance.
In our country cereals arc attdckod by two.virus diseases -
pupation and thcsaic of winter wheat. The distribution of those
diseases vary. Pupation occurs chiefly in Siberia, Mosaic in districts.
of the Voronozh'ahd Rostov RogiOn8... Both digoascs have boon thoroughly
investigated (K.8 Sukhov and A.M. Vovk; V.K. Zazhurilo and G.M.
Sitnikova)..
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Pupation virus attacks mostly oats, yet at times grave epiphy-
totics are noted on wheat'andbarley. Rye is practically immune
to the disease. In nature, the xbain reservoir of infection is'the
dark cicada - Delphax striatella. The virus remains long in the organ-
ist of this insect which spendswinter in the nymph phase.. After
developing wings in the third decade of May [Nay 21-31], the dark
cicada migrates Onto sprouts of cereals and selectively. concentrates-
on those,of oats. Spreading of infection occurs this time: Following
the hatching of eggs, the ovrwintered generation dies off. Part of
the second generation remains in the same cereal fields, yet part of
it migrates onto the young crop of late spring millet and oats seeded
after a mixed forage crop. The third generation does not attain
maturity dnd goes off to spend the winter in the nymph'phase of the
third ago. The degree of infection of young crops on which last
generation nymphs food, and their number determine the rate of?
opiphytotics of the following year.
Different oat varieties arc highly susceptible to pupation. In
cases of early infection, oat plants proceed to tiller incessantly
until fall without forming spikes. In cases of late-infection,
tassels develop, but spikelets as a rule are sterile. There occurs
an abnormal development of the ovary which protrudes greatly from the
? spike. Plants infected in the stage of sprouting arc a total loss, to
yield, they prodUce neither grain nor straw.
A biological study ofthe'pupation carrier helped to determine
the necessary control measures. A radical means of extermination of
the dark cicada is deep, late fall plowing or all fields that had,
been under grain crops in the year concerned. Experiments con-
ducted for many years have shown that 98-99% of larvae of the dark
cacada perish in deep plowing- A study of tho realm of pupation
distribution shows that the virus breeds only in those regiOns of
Siberia where late fall plowing is practiced insuffiCiently. In
regions adjacent to areas where fall plowing is wide-spread or even
continuous, pupation is practicdlly absent and there is no mass
reproduction of the dark cicada.
LL plate inserted between pages 52 and 53 has illustrations
degcribcd as follows:I
Fig. 1. Agregate of spherical particals of the virus of top
_
chlorosis of makhorka. x 9000
? Fig, 2. MoSai6 on loaves of lemon during psorosis [peeling of ?
barkb
As a control measure for pupation, deep fall plowing meets all
requirements that could be made in similar cases.. This measure is
extremely useful even from the viewpoint of grain crop agrotoc4nics;
it, calls for no special expenditure of labor or funds exceeding the'
established agrotechnical quotas, and is efficient at the same time.
The growth of mechanization of agriculture leading to steady ex-
pansion of fields. under fall Plowing is the real factor in over-
coming the menace of pupation. In connection with the above, the
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primary task in regions whore this disease becomes wide-spread is the
thorough practice of deep fall plowing..
In contrast to pupation virus, the virus of wintor wheat mosaic
dges not survive in the carrier which passes the winter in tho-
cgg phase, but in winter cereals and primarily in winter wheat..
The carrier -of the disease is the striped little ciedd Deltocophalus
striatus (V.K. Zazhurilo and G.M. Sitnikova, 1941; K.S. Sukhov,
1942). Development of cpiphytotics of the given year was determined
by the degree of infection on fall sprouts of winter Wheat. During
spring growth mosaic plants have a c'onspicuous chlorotic coloring.
Similar to pupation of oats, those plants, too, have been observed -
to tiller excessively. Spiko formation is late Or completely absent.
In diseased plants, the drop in yield fluctuates from 40 to .100g. '
As a-rule, spring wheat is attacked slightly. This is explained
by the fact that embryonic and postombrYonic,dovelopment of the
striped cicada ends comparatitrilk late, when spring wheat is already
in the phase of stem formation. In late infections, the disease'
loses its noxiousness to a considerable degree. Hence it is par-
ticularly significant that spring whoat be planted as'early in the
season as 'possible in regions where mosaic has spread.
As far as winter wheat crops are 'conccrncd, the inciden6o of
disease can be reduced by observing the following conditions.
1. Seeding on dates.fixod.by agricultural.rogillations. The
earlier seeding is done, the heavier the infection.
2. Normal rate of seeding. Thinned crops are always attacked more
severely.
3. Shallow plowing of stubble of spring crops as ?early as possible
immediately after harvest; also-carly fall plowing particularly on
fields adjacent to winter crops. Fall plowing thust be carried out
beferothe appearance of sprouts of wintor crop. Under these condi-
tions a considerable number of carriers 'perish.
If the measures indicated are applied cai-cfully, the development
of mosaic cpiphytotics can be cut off to a considorablc dcgrco.
In selection nurseries it is oxpodiont to treat the young crop
with DDT dust.
In the course of several years lemon and mandarin? infection
was observed on some citrus farms; manifestation of the disease was
notod'in the pooling of bark and in a heavy drop in the productivity of
trees. The cause of the disease was unknown. In investigations ,
conducted at the laboratory of virus diScases of plants'of the
MOSCOW station of plant protection (K.S. Sukhov and V.14 Kozlova)
it was identified as psorosis - a virus disease .4i-cad in some other
country. Tho disease is unique in that the grave symptoms appear
after d very long incubation period lasting from 6 to 10 years and
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Trans. h, 531
longo'r. Yet tho tissues of the affected trees become infected
early. The use of grafting matorial from trees concealing the virus
loads to tho gproading of infection. A throat arises that at the ago
of 10-12, i.e.,at the timo whoa the young plantation is in a period
of high productivity, [Begin p. 54] part of the troos may suddenly
-
develop disease and reduce Or discontinue fruition in a short time.
Inasmuch as the only method of spreading disease under prevailing
conditions is budding, the sdloction of healthy mother trees servos
as the most important measure to prevent infection.
? It has, however, boon pointed out already that diagnosis by
symptoms on the bark cannot produce results, since pooling develops
late. We wore confronted by the task of working out a method of
early diagnosis of psorosis under conditions prevalent hero.
It is known from foreign literature that psnrosis infection in
citrus trees is conducive to a rather early emergence Of mosaic on
newly developed leaves. This 'symptom is rcliablo, but it is ephomoral
in the son ge that mosaic is manifost for a very short time and then
.disappears.
As a'rosult of investigations conducted by V.I. Kozlova, an
applicable method was dovoloped for mosaic diagnosis under our .
circumstances.
. A speoial feature of mosaic in instances of psorosis is its
transionco. .M6saic is more pronounced during the spring,poricd of
loaf formation. It is not noticed on the youngest lodflots, but is
more pronounced on the next formation of young loaves. Mosaic-dis-
appears in a few .days, in proportion to the maturing of loaves.
At the end of spring growth, it can no longer be found. Manifestation
of mosaic is not observed on all tho increment of new vegetation;
it is found more frequently onlSr on a small portion of the increment,
somotimoS only on single plants.
-In the presence of psorosis, the mosaic pattern is very
characteristic and cannot be confused with genoral chlorosis of-a
non-infectious nature, Which at times is found on citrus plants.
Mosaic appears in the form of bright little bands and is likely to-bo
accompanied by a lighter coloring of the veins of lower formations.
Tho bright bands are scattered over the leaf, or they form a regular
pattern in the shape of a zigiag-margin or the main leaf vein reminiscent
of the contour of an oak loaf. In this case, the tissues within
the margin have a normal color, whilo a mosaic coloring is observed
on the outside .(fig. 2). Frequently circular or semi-circular spots
of the same coloring as the bdnds or zigzag-shaped linos are found
though-out the mosaic pattern. Tho mosaic pattern is vory-delkbAte
and barcly noticablo in rofloctod light if the loaf structure is.
sturdy. The mosaicis quito visible if the leaf is examined in
penetrating, diffused light. Its examination is difficult in diroct
sunlight. ' ?
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(9) Trans. A. 531
Careful inspection of trees from which grafting material is
taken presents an opportunity to reject sick specimens and thus to
prevent the spreading of infection. Diagnosis, natufally, takes into
consideration also symptoms which appear on the bark. Since citrus
bark sometimes bears injuries in the form of so-called "burns"
caused by sharp changes in temperature between day and night in winter
or in early spring, a distinction must be made also between these
injuries and psorosis peeling.
In cases of psorosis, the bark begins to change with the ap-
pearance of small bulges or pustules which somotithes arc scattered
over, the trunk and sometimes combined into groups. In the area of
pustules-the bark begins to split and its exterior layer begins to
separate. As thediscase progresses, the splitting is likely to
stretch over the entire trunk in a continuous hand and to transfer
onto separate branches of the first and lowest formation. The in-
fection increases gradually. Bark prbceeds to come off in the
form of scales or shred. The bark beneath the peeling layers acqires
a brown or yellow color. In time, infection penetrates ever deeper
and finally reaches theweed which darkens and sometimes takes on a
dark-brown coloring.
[Begin p. 551 "Scalds" present a different picture of symptoms:
brown strips form on the bark, then split and come off the wood.
In the course of a summer the lesion is margined by excrescence
resulting from the activity of cambium, and frequently the wound
heals. Open wounds margined by excrescence are likely to form in
cases of multiple "scalds". Such a structure or the "scalds" dis-
tinguishes them easily, from psorosis infections.
Considering the gravity of psorosis as a fector in reducing the
productivity of citrus crops, it is necessary to inspect most carefully
and continually the farms situated in regions where the disease has
spread. Uprooting of diseased but still fruitbearing trees is in-
expedient and unnecessary since there arc no natural means of spreading
the disease under conditions prevalent hero. Careful marking of sick
trees becomes more important, for it fully eliminates the possibility
of utilizing infected buds for grafting. Essential measures in
organizing such inspection is the inclusion of psorosis in the group of
items under Observation of the quarantine service, and adequate
knew16dge of the diagnosis of the disease by citrus growing agrono-
mists.
Virus diseases cause great losses of vegetable crops. In the
still recent past the opiphytology of "stolbur" of tomatoes and other
Solanaccac appeared to be an enigma. Tho distribution. of this
disease is in the -southern 'regions of the country whore it occasions
serious infection. Ecavy outbreaks are not inffoquont and then most
plants on tomato plantings prove to be infected. Tho extent of
damage caused by the disease is determined by.a,sharp increase of
sterility in flowers which turn green, as well as by the' ligni-
fication of the pulp of fruit which renders it inedible.
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.Tho rapid growth in "stolbur" opiphytotics and the mass infections
scorned to contradict the fact that'tho disease is entirely non-in-
fectious under ordinary conditions. Juice from sick plants causes no
infeetion when it is rubbed into loavcs and stems of healthy tomato
plants. Nonetheless, the infoctiousnoss of the disease was proved in
1931 when its transmission to healthy plants was adcomplishod by
moans of grafting (Ryzhkov and Korachovskii0 1934). The only ex-
planation for the opiphytotics of the disease is tho hypothesis that it
is transmitted by some suctorial insect. For this reason Korachovskii
(1941) conducted tests with several varieties of aphids, cicada and
othor.insccts, but the carrier of "stolbur" was not found among them.
Since our basic task remained unsolvcd, various conjectures and
hypotheses began to surround this question. Tho wide-spread opinion
was that the pathogenic agent was not introduced into plant tissues
from without, but formed within thorn endogenously as a result of
physiological disturbances caused by unfavorable ecological con-
ditions. This case was a repotition of instances that have arisen more
than once in the history 'of medicine and phytopathology. It suf-
fices to recall, malaria - the very namo of this disease indicates a.
purely ecological interpretation of its nature dt a time when nothing
yet was known about its causal agent or carrier. In this category
,belong also hydrophobia, yellow fever, and - in phytopathology
pupation of oats, dwarfing of rico and d series of other discatos.
Inasmuch as stolbur is characterized by a southern realm of
distribution, the caugo of this disease was sought in high tempera-
tures of air and goil. An opinion to this effect has boon olpfossed
more than once Sorbinov, 1941; P.I. Dvornikov, 1941; A.I.
Sorcbriakov, 1941). Some saw the cause of disease in wator starvation
of plants (Modish, 1938). Those conclusions, not verified under
sttiet experimental conditiOng, were refuted by other investigatots
(I.K. Korachovskir, 1941; 0.N. Vortogradova, 1941). [Bogin p. 56.)
Tho contradictions found in all those hypotheses clearly indicated
that the nature of the disease had not been rovealod; as a rosult,
practicing'ontorprises could not obtain sufficient funds for its
control.
Investigations conducted for several years by the Laboratory of
Gonotics of Microorganisms and Variability of Virusos, Institut of
Gonotics, Akadomy of Sciences USSR, revealed fully the opiphytotics of
stolbur, and the method for its control proposed by the laboratory
passed succOssfully industrial tests and was accepted for adaptation
in industry.
? Epiphytotics develop on the basis of the. following principles.
In southern regions infection nidi are concentrated in perennial
woods of Which European bindweod [Convolvulus arvonsis) and cross
Ilopidium) arc the most significant. Those plants are attackod-by
stolbur on mass regardless as to whothor or not susceptible culti-
?
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vated plants are near-by. The great susceptibility ofthese weeds is
determined by their being the principal subsistance plants of a '
carrier insect which proved to be the cicada, Hyalesthes obsoletus.
The larval phase in the life of this cicada proceeds on the
rhizome of European bindweed and cress. If larvae develop on
diseased plants, they become infOcted by the virus which remains
within them for an extended time. In early June, after the for-
mation of wings, the cicadas appear on the surface of the ;round and
continue to live *there for some time in the places of brooding [vyplod]
Soon after, they migrate to the plantings of cultivated Solonaceao.
The greatest mass migration occurs between Juno 20 and July 5.
This period determines tho spreading of the virug to plantings of
tomatoes, pepper, eggplant, tobacco and potatoes.
After finishing migration, the cicadas load a stationary mode of
life within the limits of a given plantation where they hatch eggs in
the soil near the root neck of European bindweed, cress and some
other perennial weeds on tho planting. While feeding on those plants,
they cause their infection. Mature individuals begin to die off in
July, and adult, cicadas are found rardlyilwAugtAt. Larvae developed
from the egg still in-summer settle in the rhisomo of woods and
there pass the winter.
Since the incubation period of "stolbur" lasts in tomato plants
from 30 to 45 days, the manifestation of the disease occurs at the
end of July and lasts until the middle of August.
Epiphytotics reach their maximum development in August.
The period of mass migration of the carrier is fairly short -
two weeks altogether; this makes it posible to safeguard plantings
effectively by application Of-DDT dust. This method was worked out
jointly by the anther and A.M. Vovk in the years of 1947-1949. The
results were good, and in 1950 the method passed industrial tests
organized by the USSR Ministry of Agriculturc,-and later was accepted
for adoptation in a number of southern regions.
Four-time dusting carried out on Juno 20, 25, 30, and on July 5,
with each dusting consuming 20 kg of dust (dusting was not unbroken, but
was done over rows) reduced the percentage of infected plants by '
? 3-4 times, as established on the "Krasnyi Tamancts" kolkhoz in 1950.
This method combined with superior agrotechnics produces good
results. An important factor is also the wide-spread use of stem
varieties distinguished by relative resistance to "stolbur."
Weakening of plants under conditions of poor agrotochnics indroases the
losses inflicted by "stolbur."
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Solution of the problem concerning mass *ilt of popper, eggplant
and potatoes in southern regions was also now. It was demonstrated
under rigid experimental conditions that this disease is also caused
by the ustolburn'virus, although its symptoms (wilt) on the crops.
rOfOrrod to aro diffcront fftm those on tomatoes (K.S. Sukhov and
Vovk, 1949). [Begin p. 571
? Exporimonts have shown thnt an efficient method of safeguarding
sood-potatO 151antings against stolbur is the one proposed by Aca-
demician T.D. Lysenko, whereby-the planting is carried out in mid
.July or about the 10th of July. "Stolbur" is not found on such
plantings, because the disease vector loads of stationary mode of
life during this period and does not invade now areas. Yet, spring
planting of consumer potatoes must be carried out as early as
possible so as to give them enough time to accumulate a sufficient
yiold before"stolbur" wilt appears..
The work indicated was used in the People's Republic of Bul-
garia whore opiphytology of "stolbur" dovolops fundamentally along the
same principles. "Stolburt"wilt of potatoes has been identified in
Armenia (Khachatrian, 1950). According .to Samundzheva's data
(1949-1951), it has been established that transmission of "stolbur"
under Gruzian conditions is duo to the cicada Hyalosthos mloko-'
siowiczi, related to H. obsolotus.
Another harmful disease of tomatoes is mosaic; its infections
are paitticularly sovore under conditions of covered [greonhousc etc.]
ground. Even in the usual course of the disease, in which the main
symptoms are a mosaic coloring of loaves and some depression of
growth, yield is reduced by 10-15%.
Under certain ecological conditions more serious infections,
characterized by the dying off of tissue sOctions on stems and loaves,
arc likely to form on tomatoes with mosaic. In this phase it is
called the streak disease, Or striped mosaic, since the areas of dead
tissues on the stem show elongated dark stripos.
The noxiousnos of the streak disease is determined not only by a
general deterioration of the condition of the plant, but also by
the spoilage of fruit sets which develop necrotic spots and ulcerations.
Conditions conducive to retention of infection and to its spread-
ing are the same as those for tobacco mosaic, since the causal agOnt of
the disease in tomatoes and in tobacco is one and the same virus.
Distinct from the tobaaco disease, a small percontage of tomato mosaic
is transmitted by seed. This creates a supp1othontr9- and more
permanent source of infection on the plantings. M.I. Gol'din (1948)
proposed disinfecting 'tomato seed with a l% solution of postassium
permanganate for 5 minutes with subsequent rinsing of the seed in
water. Such disinfection rodacos initial cdsos cf infection, but
docs not eliminate them fully. Therefore A.M. Vovk's suggestion,
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that in covered ground farming only tomato seed gathOrod from healthy .
plants be uscd, must bc considered as more expedient. At present
there has arisen a need for establishing seed growing tomato nurseries
with covered ground and with open ground whore healthy plants, caf.o-
fully inspected as to virus transmission, would be grown for sood.
Testing of juice pressed from tomato leaves on the leaves of .Nico-
tiana glutinosa holps to datormino unmistakenly concealed virus vection
and to sort out the plants. Inoculation of the jico into glutinosa
loavos in which the virus is prosont causes easily Porcoptiblo
necrotic spots to devolop on them within three days. Maintenancu of
a 'certain number of glutinosa plants for this purpose presents no
difficulties on greenhouse farms, and the additional work connected
oaith-tho selection of healthy plants pays for itself in a very short
time.
Nonetheless, the sOoding of healthy seed for seedlings does not
solve the wholt problem. External sources of infection may.continuo
to exist in greenhouses. on different articles of equipment, such as
shelving boards etc. Therefore, the prerequisite for growing healthy
tomatoos'in'coverod ground is thc elimination of infection contacts
which occur during the thinning of 600d1ings, during their trans-
planting into ground etc. [Basin p. 581 The current practice of
sowing seedling tomatoes in peat-humus Tots eliminates such contacts
during the seedling period, which is very important since-tho first
infections occur already during the thinning of seedlings. Tomatoes
grown by this method are not subjcat to thinning and aro transplanted
into ground together with the pots. Transplantation, can be: accom-
plishcd without touching the stem or loaves with one's hands, which
cxcludos the possibility cf infection. After this, new contacts
with plants take place during the rOmoval of suckers.. Infection
occurs oasily during this operation. It can be fully .avoided by
removing suckers with pincers and soissors, and not by hand. After
removing suckers from 3-5 plants, the instruments are disinfected by
instantaneous immersion of their working parts in a 10% solution of
trisubstitutod potassium or sodium phosphates, or in a 10% solution ?
of tannin. Such disinfection docs not takc-too much time and insures
fully against the transmission of infection. Tho alkaline solution is
kept in a shallow glass flask which the worker carries on a strap.'
The pincers must be disinfoctod the same as the scissors. '
This simple method involving minor expenditures eliminates full*
the possibility of spreading infection during the removal of suckers.
Tomatoes cln be tied up only with a cord that has not been in use
previously. At the time fruit is being harvested, infection can bc
avoided by holding the cluster with pincers which arc being dis--
infected frop timo to time, and by tearing the fruit off by hand.
We have cited all those details for the purposo of showing how
siraplo motthods practiced to eliminate infection can be when they aro
known. At the same time, it must be emphasized that in covered
ground tonatt agrotochnics,? thoso methods have not boon taken into
consideration up to now. It?is Obvious that the time is ripe for the
initiation of appropriate scientific and organizational moasurcs
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bound to raise the level of covered ground fexming as a whole and to
create agrotoChnics answering modern demands.
' An interesting solution of a problem was obtained by developing
means for the control of white mosaic of cucuMbors. Cf1cumbors of
covered ground arc often attacked by groanmosaic NoI2. The bio-
logy of the virus-agent of this mosaic-has much in common with the
biology of the virus Of tobacco Mosaic. Tho virus is strictly
specific for tho .curd family and plants of other families are not
susceptible to it. Infection is spread by sood and is easily trans-
mitted through centact. Green mosaic is distinguished by moderato
noxiousness which decreases yields by 10-15%. On some farms, how-
ever, heavy opiphytotias f white Mosaic flare up on cucumber plant-
ings from time to time. In such a case the drop in yield reaches
70 and even 9. Indpoctions of covered ground farms conducted by
our follow werkor, A.M. Vovk, have induced him to draw the con-
clusion that white mosaic develops as a result of directed variation
of the vird8 ?i.*.grocn mosaic under the influence of extremely thigh
temperature.
Significant spreading -f green mosaic was observed on cucumber
plantings in greenhouses under the customary conditions, white mosaic,
however, was not found. Heavy attacks by white mosaic were noted only
in those greenhouses where temperature was maintained-high, at times
exceeding 35? C. To inquire into this situation, A.M. Vovk conducted
special experiments at the Institute of Genetics, Academy of Sciences,
USSR. Cucumber plants infected by the virus of green mosaic were
placed in a glass-inclosed compartment where temperature was main-
tained at 37-40? C.. Under these conditions, sifigle white spots
appeared on developed upper leaves in 7-10 days. Their number and
dithensions-grew gradually and in a short time white mosaic dominated
the plants. The same result was obtained'in a series of replications,
in accordance with natural law. [Begin p. 593 Thus, the virus
of green mosaic, under the influence of increased temperature,"
underwent directed modification into the virus of white mosiac. A
special feature of the latter.was its boin&adopted to now conditions
of temperature and being able to diselaco easily the virus of green
mosaid unadapted to these conditions.
Separate isolates of the virus of white mosaic infeeted cucumber
plants growing in a groonhouso-under ordinary Conditions. White
mosaic developed on the plant-s. The same result was obtained in
the next transition Epassazh].- Consequently, the new property of the
virus became fixed by heredity. It must be pointed out that during
two transitions the virus had. multiplied by hundreds of thousands and,
consequently, the retention of acquired characteristics in a series of
such multiple generations testified the high degree of Constancy of
the new form. Yet, under ordinary conditions, the virus of white mosaic
could 8urvive only man isolated state, in the-absence of green
mosaic. In mixed inoculation of cucumbers, i.e. with both viruses,
the original form, as the one more-adapted to ordinary temperatdro
conditions, displaced the new form. In proportion to the growth
of new loaves in a cucumber infected by both viruses, green mosaic
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was becoming more prodaninant on them while white mosaic disappeared
completely as timo,wont on. ?
Tho experiments described helped to establish fully and clearly
the cause for tho omorgonco of white mosaic. On the basis of this re-
search definite recommendations can be made to covered grouna farms
for the prevention of epiphytotics af a very noxious disease.
Antagonism between viruses of green and whito mosaics, their in-
compatibility within the organism of one and the same host plant in-
dicates that thoy belong to different species. In the given case,
as well as in a series of other cases, we witness the formation of
species in viruses under control conditions.
We doom it important to emphasize that a study of the regu-
larities of directed variation in phytopathogonic viruses is of
groat theoretical importance. Some aspects of this problem can be
solved more oasily'on the simplest nuclooprotcid viruses than on
cellular organisms. A contributing factor in this respect is the
rapid tempo of virus multiplication, their high plasticity and
their capacity to maintain activity under deviating conditions. Tho
relatively simple structure of viruses makes them a particularly
convenient object for theoretical in-ftstigations concerning their
variation and specific formation.
Tho theoretical significance of research on directed virus
variation becomes the more important as --no identification of viruses
with'so-callod "free genes" persists in Morganist virological litera-
ture. Variability of viruses i?eing treated from the position of
the occasional mutations theory. Many facts of virus variation are
under dispute and the appearance of new forms is being explained by
their separation from populations whore they, so to say, had existed all
along in an undisclosed state.
Work conducted at the Laboratory of Genetics of Microorganisms
and Variability of Viruses, Institute of Genetics, Academy of Scionces,
USSR, has disclosed a series of factors under the influence of which
rogular, directed virus variation develops. Cultivation of viruses
in plants belonging to different species, or in plants of one spOcios
grown under changed ecological conditions furnished such factors.
Directed changes in the form of the tobacco mosaic virus were ob-
tained,,for instance, as a result of cultivation of an infected host
plant at incroased temperatures. Tho-*roality of,,the process of
directod virus variation was demonstrated irrefutably in those in-
vestigations, and the hypothesis of a separation of ready forms from
populations was repudiated. The very fact that virus forms adapted
to life in plant tissues with a sharply disturbed metabolism [Begin
p. 60], ch,ractorizod by degradation and decomposition of proteins
obtained in those experiments, furnishes important material that will
help to understand virus nature.
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A study of the rogularities of directed virus variation opens up
wddc porspOctives for the study of the processes of species formation
among thorn,. Bacteria and. viruses can and arc already produOing ma-
terial most accossiblo to the investigation of this problem. A wido
field for the investigation of directed virus variability, is presented
by the problem of protective vaccination of plants. For the time
being, this question remains unsolved, with only the first steps
having been takon in this direction in our laboratory. The ?stab-
lishmont of numerous facts of antagonism and incompatibility of related
virus species within one and the same host plant offers a,good basis
for directed production of mildly virulent but harmless forms of
viruses, which might serve as live vaccine safeguarding plants ino-
culated with it against sOvore infoction causod on the part of related,
strongly virulent viruses.
The development of this mchtod encounters groat difficultios, but
its perspective for certain agricultural crops is clear, and it
deserves much attention.
Control of virus -diseases of plants should be expanded in
other directions also. Chomother'apy of virus discasos has remained
until now a neglected department. Systematic work for the dis-
covcry of antibiotics which inactivate viruscs is being done nowhere.
The initial phase of such,work must of necessity be included right -
now in the complex investigations by virologists and microbiologists.
But even in those spheres of agricultural virology where re-
search has been conducted long ago, and whoro many achievements have
boon attained, tho work must be strengthened.' This, to begin with,
concerns development of agricultural conditions that will increase
the resistance of plants to virus diseases. Means of crop pro-
tection such as chemicals must be considoi-ed as strictly extra-
ordinary measures required in acute cases. They must be replaced
with special moans of agrotechnics in which primary importance will
always be attached to factors of plant nutrition, time and methods of
seeding,-structuro of crop rotations, weed control, and prophylactic
measures.
Efforts must be made to expand selection-work for the develop-
ment of varieties resistant to virus diseases. The roults of
selection work obtained in cotton and tobacco culture, whore virus
infection can literally by eliminatod by high varietal resistance,
indicate a cloar prospect [of success].
Finally, it is. very important to expand and intensify the
theoretical work. in the spherc'ef virology which will always be of
great signigicance in practice.
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Samundzhova, E.M. 1949: &if:1g data on tho study of the tomato"stolbur
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Serobriakov, A.I. 1941. Tomatc stolbdr in'tho zones-of canning'
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Sukhov, K.S. and Vovk, A.M. 1940." Pupation of cultivated cereals '
and it spread in nature. M.- 1949. Stolbur of Solanaccac. M.
Sukhov, K.S:. Charactcristics of two ccroal viruses. Mikrobiologiia,
It, No.5 - 1952. Virus of top chlorosis of makhork.a: DAN
S8SR;.85? No.6.
Sukhov, K.S. and Kozlova, V.I. 1951. Methodical instructions in
inspation of citrus crops as to psorosis infection (pooling of
bark).-
Sukhov, K.S., Vovk, A.M. and MUshnikova, K.S. 1952. Industrial tosts'of
DDT dugt in the control of t6mato gtolbur. Dokl. VASKHNIL, No.8.
Sukhov, K.S. and Razviazkina, G.M. 1953. Top chlorosis of makhorka,
its causal...agent-and vector. Tr. In-ta GonOtiki AN SSSR, 20. ,
Ternovskii, M.F. 1950." Ithmuno tobacco varieties. Solokts.
.Semonovodstvo, No.10. - 1951. Resistance variation in tobacco
mogaic as a result of vegetative hybridization. DAN ,SSSR, 29,
No.3.
Fedoronko, Mall. 1952. 85.4 contners of makhorka per hectaro.,
Poltava. 7 -
Khachatrian, G.A. 1950. Stolbur potato wilt in Armenia and its
tranSthigsion in grafting. Izvost. AN Arm. SSR, 3, No:12.
Khudina, IP. 1936: Virus diseases of tobacco in the USSR. Kras-
nodar. - 1941. Virus disoages of tobdcco and Makhorka in the
USSR and thOir control. Tr. Sovoshch. po Virusn. Bolezniam
Rastonii, M.
Institut Genctiki
. Akademii Nauk SSSR
Article received at
Editor's Office '
February 12, 1954
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? .1
Brezhnev, D. D. 'and Aizenshtat, IA.. S.
Novoe v selektsii rastenii.
MoskIa
144p. 1954
463.62 B75
Leningrad
[Something new in plant breeding].
Gosudarstvennoe Izdatelistvo,
Seliskokhoziaistvennoi Literatury
SONETHING 1JE7 IN PLANT BREEDIFG
CONTENTS
Trans. A. 532
Contant and Conclusion
By R. Adelman
FOreword 3
Influence of cultivation factors upon the-foi'Mation of properties
and characteristics of tomato plants 5
Inheritance of acquired Characteristics 29
Means of increasing the vitality of organisms 49
Influence-or-f)reliminary grafting upon thO-inhei-itenCe - af-ohaaCte- '
tics-----7 64
Role of age -iiafiabilitSr-of - anndal-herbadeads-plant6 in-hereditary trang-
mission 76
Role of age variations of sex elements in inheritance of characteristics 88
Role 0 amount of pollen'and-ffiltiplioitSr-Of"pollination in-inheritance-
of characteristics 104
Influence of extraneous added-pollinatiein upon - .critality dheditaitransmission 117
117
Influence of ocologo-Lgeographical-factors upon the-fottation af-dhasra8=-
tcristics in tomato hybrids 129
Conclusion 136
References 141
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CONCLUSION
Problems analyzed in the present work deal with the control of
vitality and heredity in annual herbaceous plants and in their hybrids.
The influence of cultivation conditions upon the formation of various
characteristics has beon demonstrated on a wide assortment of material.
A special foature oftheso investigations was the comparative study of
plant behavior in different agro-climatic zones'. The large varietal and
specific stock utilized in the work speaks very convincingly of the in-
fluence of conditions exerted upon the formation of a whole series of
agriculturally importmnt characteristics. Morphological characteristics as
well as biological (and biochemical)-proportios of plants change under
the influence of cultivation factors. Dynamics of growth and the incep-
tion of generative organs proccbd differently; time of flowering, ripen-
ing of fruit and their chemical composition, vary. The variations which
crop up may,beso cebsiderable as 4o?.exceddAhe1.limitsliahtrynt4nthe
indicators of a species of varioty.
A varioty changos its nature When transferred to now conditions of
cultivation. ,qAance the clarification of the reaction of different varie-
ties to the influonco-of external environment is of groat importance in
the work of selection. It is well known that varieties developed and
rogionalized.for:a oortain locality can' be cultivated successfully in
other regions and other climatic zones. In clarifying the possibilities
of rogionalizing a variety, it scorns to us that its testing by the method
adopted by the state variety testing network only is insufficient.
It is important to show to what extenta variety is capable of assimi-
lating factors of cultivation and of rcSponding to them with changes of
separate properties and characteristics. Only the knowledge of the
nature of behavior chafactoristics under appropriate conditions makes
their control possible. Tho work cites data which demonstrate the moans
of Modifying the characteristics of the length. of the vogotatilio period,
the chemical properties of fruit, their form, productivity etc. Special
features meriting attcntion are those involved in the formation of cha-
racteristics under conditions of irrigation) this must be taken into
account when now varictios are being-doveloped and those existing in
irrigated regions are being utilized.
All methods developed for control of the formation of characteris-
tics assume importance in the event they can be fixed in the, progeny.
Quite logically, the following chapter of tho'work is devoted to the
problem of inheritancc of acquired variations. Thus far, Nichurinist
biological science has at its disposal cotvarativety lit1453 fwd.:U.41kt
material corraborating inheritance of characteristics under the in-
fluence of environmental conditions, (Begin p. 157) and the accumu-
lation of now facts in this sphere is of groat importance. The book
cites data which demonstrate that acquired variations become a re-
quirement of the organism and are preserved in progeny cultivated under
now conditions. The capacity of plant organisms-to fix acquired
variations is extremely significant in selection. WI= the factors
facilitating the formation of a definite characteristic and its fixation
in progeny will be known, then the breeder will be able to control its
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development. The book cites data indicating the possibility of fixing
modified properties and characteristics of tomato plants in the progeny:
length of the vegotatile period, form of fruits, their chemical composition,
and a series of others. The basic method used to modify properties is
rearing plants under conditions deviating from the customary (intrinsic)
requirements of a crop. A tomato crop that thrives in a warm climate
changes its nature (heredity) and acquires the property of enduring low
temperatures, if its subsequent seed generations are roared under con-
ditions of low tamporatures. Directed selection aimed at acquiring cold-
resistant properties is pirticularly important when a crop is being moved
to more northerly regions. Increased cold-resistance in tomatoes is -
important also for the central bolt, the Baltic Republics, White Russia.
The book contains methods for increasing cold-rosistan6c of this parti-
cular crop. Results obtained by one of the autly.-rs (D. D. Bfezhnev)
arc indicative of the effectiveness of the methods suggested. With the
cooperation of P. P. Gusov, a science fellow of the Polar Station of VIR
(All-Union Institute of Plant Industry); ho succeeded in resolving the
problem of'growing tomatoes in open ground [outdoorsi under trans-polar
conditions. The book reveals conditions for the inheritance of mor-
phological, biological, physiological and other properties in tomato
plants. As in the case of influence exerted by cultivation conditions
upon the formation of characteristics, there is data on the hereditary
diffOrences in plants cultivated in arid areas and in those under irriga-
tion.
A separate section of the book (chaptor) is devoted to factors in-
creasing the vitality of hybrid organisms. Now methods of intra-varie-
tal crossing are also included. A distinctive characteristic of these
methods is the growing of parent plants under different conditions.'
The additional differentiation of sox gametes thus created loads to a
considerable ificreaso in the vitality of progeny, its productivity and
early ripening. There are indications that the opinion established in
selection science, that utmost heterosis can be obtained by selecting
parent pairs which differ in morphological characteristics, is not
always correct. Obviously, the biological differentes-An,the-plants-to
be crossed, created by dissimilar conditions of cultivation, perform
an important role. Works on crossing plants reared under conditions
as sharply diverse as hothouse and-open air culture, and a series of
others, are of interest. [Begin p. 138] Methods connected with regu-
lating the state of the plant and its sexual elements in terms of age
are completely original. At different stages of its development?the
plant and its reproductive organs produce a progeny with dissimilar
vitality. The most productive seed produce plants in the period when "
their development proceeds most vigorously,"at 146 called 'sage: Of puberty".
The same applies to the state of the flower. Considerable increase in
the yield of hybrid progeny is obtained by pollination coinciding with
the greatest vitality in the state of the plant (the flower). The
method of preliminary grafting is also effective. In a Series of cases
it is instrumental in nearly doubling tho yield of fruit as compared
with ordinary hybridization (crossing). Finally, it demonstrates the
possibility of using the method of added alien pollination for the
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purpose of increasing the vitality of hybrid progeny.
A number of'the following chapters are devoted to Control of hero-
dity in crossing. The methods suggested aro founded on intorfcronco
(control) in the sexual process and on nurturing the embryo at the
earliest possible Stages of dovolopmont.
The -method of preliminary mentor which combines soxual and.
vegetative hybridization is aimed at influencing the maternal organism
prior to crossing, and the developing hybrid embryo after fertilization
has taken place. Preliminary grafting makes it possible to obtain
hybrids with paternal heredity even in cases in which the paternal
plants aro represented by more "persistently" recessive forms, as, for
instance, a leaf of the potato typo and the yellow color of tomatoes.
Particulars of preliminary grafting aro explained as applied to tomatoes
under southern and northern conditions. To Obtain variations in more
consorvdtivo characteristics in the South, a one-time grafting is suf-
ficient. In the North in such cases it is necessary to resort to grafting
F1 plants on the original parent forms. According to the characteristics
of the fruit, in the work of selection a one-time grafting is sufficient
even in the North. Tho role of aEo variation in annual plants' in the
inheritance of characteristics is also demonstrated. In different phases of
age, plants transmit their properties to the hybrid generation with
different vigor. The maternal tyke of heredity which responds to a
more vigorous plant development is transmitted more fully by pollina-
tion of clusters of the central formations. Characteristics of ma-
ternal plants aro transmitted loss vigorously in pollinations of the
first flowering clusters and blossoms of aging plants. As applied to
tomatoes, the cultural practices which assume importance aro:
Greenhouse and -open air cultivation. In the South this period
(transplantationin the open) is loss severe than under northern con-
ditions. Hence, somewhat different results arc observed in the pollina-
tion of the fii-st flowering clusters which usually form while still in
the greenhouse: In the case of pea culturO, ago variations appear in
a "purer form". In crossing peas ['Begin p. 1391 (ehis, obviously,
applies to a certain extent to all 'annual crops) one =1st keep in mind
the early ripening of the plants - "ago within itself". In cases in
which both parents ripen equally early, ago differences of the mother
plant suffice to obtain certain variations; if, however, development
(early-ripening) varies, one 'should resort to additional ago differ-
entiation by planting the parents at different dates.
A very effective method is"tho roulation of the ago state of sex-
ual elements (ovicolls, pollen). Pollination of a flower (ovicell Whorl
Its flowering period is ending strengthens considerably paternal here-
dity in the progeny. Hero, a change of domination can take place, even
cases in which the parent plants arc represented by clearly recessive forms.
It must be remembered that the ago of a flower has meanings other than
the calondar idea, and is charactoripd by d physiological condition due
to many factors, chiefly weather conditions.
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Trans. A. 532
Tho application of "highly mature" [listarovozrastnoin] pollen makos
it possible to strengthen the maternal typo of heredity and may provo to
bo an efficient method in selection, particularly in casos in which it
is desirable to proscrvo the cultivated typo of plant in crossing with
wild forms.
Apart from utilizing pollen of various ages for the purpose of con-
trolling heredity, knowlodgo of the characteristics of progeny behavior
during it application has still another significance in the work of -
selection. It is known that it often becomes necessary to store pollen.
It is enough to point out such a necessity in crossing forms with non-co-
incident dates of blossoming, forms geographically, remote, and varieties
grown in different localities (duo to transfer of pollen) etc. The
data *Filch we have cited can servo as certain orientation in all these
cases. Up to now, the problem Under consideration has found no clari-
fication whatever in literature. The book demonstrates the possibility
of controlling the force of hereditary transmission by regulating the
amount of pollen dropped on the flower stigma, and by multiplicity of
pollination. Reduction in the amount of pollen applied (limited pol-
lination) strengthens the transmission of maternal properties. The
opinion established in literature that offspring of limited pollina-
tion must possess certain depressiveness is not justified in cases of
intorvariotal crossings. Two contrasting phenomena are encountered in
fertilization duo to a limited amount of pollcn: weakened activity of the
sex process, and bettor conditions of development (nutrition) of the
hybrid embryo as a result of tho small amount of seed formed. The -
latter circumstance may lead to a progeny possessed of high vitality.
Miltiplc pollinations strengthen transmission of paternal properties
to hybrid progeny. -Investigations of this sort were prompted by the
hypothesis [Begin p. 140] that pollen is capable of germinating and of
reciprocal action with the vogotativo parts of the ovary and developing
embryo in the course of extended time (at least 4-5 days) after ferti-
lization has occurred. Tho high percentage of sets in instances of
multiple pollinations .(up to logg of crossing success) anethe effi-
ciency of the method make it acceptable for selection work. A sepa-
rate chapter is devoted to the role ar supplementary alien pollination
during hybridization. The breeder quite frequently encounters pheno-
mena of insufficient pollination, or has to deal witn pollen that is
inefficient, partly sterile etc. In all such cases the normal course of
fcrtiligation is being disturbed and the character of progeny behavior '
changes., We have explained in our work the foasibility of compensating for
the quantitAtive and qualitative deficiency of pollen by alien pre-
pollination. Application of pollen of distant species (which takes no
direct part in fertilization) increases the percentage of sots, and, in
many rospocts, restores the usual (control) picture of hereditary charac-
teristics. Addod alion-pollination with highly mature pollen takes on
particular significance. Data cited in the book show that oven pollen
which has practically lost its capacity to fertilize, restores it to a
considerable degree by added alien pollination.
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Tho last chapter is devoted to the problem of ccologo-gcogrdphical
J influonco upon tho formation of characteristics in hybrid plants. In-
vestigations conducted up to now on a small. scalc explain definito re-
gularitios of the development of ch6ractoristics in hybrids reared at
certain ocologo-geographical,points. The availability of a wide notwork
of state soloction stations and other oXperimont-1 institutions located in
quito different agroclimatic zones offers tin opportunity to dcvelop-this,
substantially now, trend in selection work. This still is the tomorrow r
of Sovict .soloction, but-wo deem it timely to Call -it to the attention of
wide circles of brooders.
On the wholo, the methods which we propose aro tochniqally simple
and entirely acccssiblo to any experimental institution.
-
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Avdonin, N. S. and Khudiakova, Iu. A.
0 sposobakh effektivnogo ispoP.zovaniia
ezotogena
[Methods for effective utilization of
? ,azotogen].
Voprosy Povysheniia Plodorodiia Pochv
Nechernozemnoi Polocy
Iidatelistvo Moskovskogo Universiteta; 1954
p.83-103 56.6 M85
CONCLUSIONS
Trans. A. 533
Conclusions only
By R. Adelman
1. Azotogen applied-on saline sod-podzolized soils by methods now
available is ineffective. This can be explained by the fact that
azotobacter introduced in sod-podzolized soil finds itself in an environment
deficient of phosphorus and organic substances, with an excess of motile
forms of aluminum, in the presence of rivals of azotobacter adapted to
conditions of podzolized soils. The destruction of azotobacter'in saline
podzolized soils is aided by its introduction in pulverized fofm after
having been sensitized through maintenance on artificial media.
2. To apply azotogen successfully, it is necessary to create for, it
favorable living conditions in soil and to change the methods of its
introduction in soil, along with the development of stable azotobacter
races.
3. Special "azotobactor granules" repregenting nidi with conditions
favorable to azotobacter have been suggested. These granule g are prepared
from neutralized soil, humus, and neutralized superphosphate. Tho mix-
ture of substances indicated is infected with pure azOtobacter cultures,
converted into granules, and introduced into the soil.
4. Whenever azotogon was introduced into specific "granules", azoto-.
bactor lived, multiplied and fixed atmospheric nitrogen upon finding these
"granules" in acid sod-podzolized soil (p11.4.6) in the course of two -
vegetative periods, while it perished if introduced by the old method.
Azotobacter "tranulesn'introduced under potatoes increased their yield
substantially.
5. A pfoPorty of "granules" important to azotobacter is their buffer
capacity, i.e. the capacity to keep environmental reaction on a certain level.
This was achieved by including neutralized soil in the "granules" and by
creating a limp coating about them. Thus the cfivironmental reaction in the
"granules" upon finding them in'acid-sOil (pH 4.6) kept up for two vegeta-
tive periods 'on a level of pH 6.2--6.8.
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6. Preliminary propagation of azotobactor on cultivated soil with a
,mildly acid or neutral reaction of environment increases the reAistance of
azotobactor to subsequent unfavorable conditions of subsistence.
7. Better devOlopmont of azotobactor was observed during the first
,vegetativo period whon suporphosphate content in the granules was 0.5%,
and daring tho second vegotative pOriod when suporphosphato was maintained
at 1%.
8. An addition of 5% of straw to the granulo compositioristimulated the
development of azotobactor during the first vogotativo period, but depressed
it in the course of the second vegetative period. ,
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Conclusions only
R. Adelman
Kalinkevich, A. F.
Formy asotnykh udobrenii pri
vnekornevom pitanii rastenii
(Forms of nitrogenous fertilizers
in external feeding of plants].
Zemledelia. 2(6)45-49
June, 1954 20 Z44
(In Russiam)
CONCLUSIONS
Experiments in the use of various forms of nitrogen fertilizers as
nutrition for maize and buckwheat have shown that urea, an organic
form of nitrogen, ii the best for external nutrition, and ammonium
sulfate the poorest.
The advantage of urea over other forms of fertilizers lies in its
considerable content of nitrogen which facilitates its introduction
in small concentrations in fairly large doses through the leaves.
Nitrogen introduced in the form of a urea solution converts more
readily to albumin substances in the leaves. A 0.5% urea concen-
tration,produced no external changes in the leaves., while ammonium
sulfate caused decomposition of the-green part of chlorophyll and
thus decreased their photosynthesis.
Such diverse action of mineral fertilizers (ammonium sulfate and
ammonium nitrate) and organic fertilisers (urea) in external
rnitri-
tion is due to the characteristics of foliar nutrition. In ordinary
root nutrition, nitrogenous mineral fertilizers undergo synthesis into
primary organic compounds already in th& roots and, as a rUle, enter the
leaves in the form of organic compounds. In ordinary root nutrition,
miner41 nitrogenous compounds enter the leaves only in exceptional
oases. This is observed in increased nutrition or plants with
nitrogen When a part of mineral nitrogen reaches the leaves, and also
in cases of protracted weakened photsynthesis and carbohydrate
deficiency in plants.
Urea, upon landing on the tissue surface of a leaf covered with
fatty substances, disolves in these substances and thus penetrates inside
the tissue more rapidly and, being an organic substance, enters
into reciprocal action with the plant tissue more readily than mineral
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substances (Gunar, 1952). It ex6els all available nitrogen
mineral fertilizers in external feeding.
All this, plus the experimental application of herbicides and
physiologically active substances, demonstrates that organic,
soluble forms of nutrive substances, obviou?sly, have advantages
over mineral fertilizers in external nutrition.
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Borisova, N. I. and Zagainova, 0. A.
Isollzovanie izotopa p32 dlia otsenki postupleniia
v rasteniia fosfatov iz pochvy i iz udobrenii pri
posloinom i ravnomernom raspredelenii
[Utilization of isotope p32 in evaluating intake
of phosphates by plants from the soil and from
fertilizers distributed in-localized layers and
spread evenly [throughout].
Izvestiid Akademii Nauk SR, no. 6, pp.111-116
Nov./Dec. 1953 (Ser. Biol.) 511 Sa2B
(In Russian)
CONCLUSIONS
Trans. A. 535
Conclasions only
By R. Adelman
1. Localized introduction of fertilizers into soil layers leads
to more intensive utilization of fertilizers in the first period of
plant development than does their even distribution.
2. Alecording to the difference between phosphate intake in
unfertilized and in fertilized experimental varients, determination
of plant intake of phosphates from fertilizers introduced in the soil
gives an impression of a minimal use of fertilizer phosphates in
localized application.
3. In localized application of fertilizers, the consumption of
soil phosphates during the first period of plant growth is less than
in an even distribution of fertilizers.
ea ?
4. In an experiment conducted on
localized introduction of phosphates,
phosphates was relatively higher than
per comparative experiments conducted
podzolized &layoff- soil with
utilization of fertilizer
that of soil phosphates, as
on chornozem and sierozem soils.
5. The advantage of introducing fertilizer in the soil layer
persists up to the time of harvest, he regards utilization of phos-
phates from fertilizers, but'it is considerably less than in the
first period of plant growth.
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6. In the second period of plant growth consumption of soil
phosphates was more intensive than in the first period of their
growth, when fertilizers were localized.
7. Aided by the method of tagged atoms, it was possible to
obtain data allowing the hypothesis that the fundamental advantage of
localized introduction of fertilizers (into a layer of soil) lies in the
increased utilization of fertilizOr phosphates by plants in the
first period of their development.
PochIonnyi Institut
im V. V. Dokuchaova
Akadomii Nauk SSSR
Article received
at oditbrial office
Doc. 1, 1952
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Bushinskii, G. I.
Agronomicheskie rudy
[Agronomic ores].
Vestnik Akadanii Nauk SSSR
Vol: 24, no. 12, pp.30-34
Dec. 1954 511 Ak14V
(In Russian)
(i7) Trans: A. 536
By: R. Adelman
AGRONOMIC ORES
Artificial fertilizer 6 have been utilized extensively-in ;-
agriculture for many years. If used skilfully, taking into account the
characteristics of the soil and the crop to be grown, they increase -
the yield by one and a half to two times, and frequently by even more.
Their application is economical. The cost of the fertilizefs, compared
with the increase in yield which they produce, is not great. Thus,
fertilizers are a powerful medium in raising the productive capacity of
farm labor.
Various agronomic ores are used in the preparation of artificial
fertilizers. The most significant among them are the potassium and
phosphate ores, and also carbonate rock (limestone and dolomites).
Gypsum arid various minor elements are used in smaller amounts - to
begin with, boron, copper,"cobalt, manganese, iodine, molybdenum,
uranium, vanadium and zinc.
The most-prominent of the phosphate ores are apatite, phosphorite
and vivianite,
Apatite ores are composed of fluorapatite-mineral and different
admixtures, principally nepheline and pyroxene. They constitute the
Ehibinsk apatite deposits onthe Kolisk peninsula. These deposits
formed during the solidification of alkaline magrha, apparently in
the Devonian period, about 350 million years ago. The concentrate of
these ores obtained by their granulation and enrichment consists
almost entirely of fluorapatite and has proved itself an excellent
product fcr chemical pfocessing into soluble phosphate fertilizers,
chiefly superphosphate.
The large apatite crystals found in mica veins in the vicinity of
the city of Sliudianka on the southern shore of the Baikal lake have
been know for over a hundfed years and have boon described in ,
literature more than once. The quantity of these crystals, however,
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is not largo. In recent years, Soviet geologists have found an
accumulation of fine apatite grain the deposits of which are signi-
ficant. The apatite in these deposits is accompanied by a large
amount of admixtures and, therefore, it has not as yet been adapted
to practical use. These deposits require further geological re-
search for the purpose of finding ores of a higher grade, and a
technological investigation aimed at defining a more efficient method
far their processing.
Phosphorites have a fairly complex mineral composition. Their
phosphate substances occur in the form of throe *idely distributed
minerals - fluorapatite, francolite and kurskite. The catont of
P205 is. the main qualitative indicator of raw phosphorus,
Fluorapatite Cont6ins 42.23gof P205, franoolite - 37.10, and
kurskite - 34.52%. [Begin p.311
Tho minerological nature of francolite and kurskite has thus far
only been studied superficially. There is no unanimous opinion
even On as important a problem as the state of carbon and phosphate.
Some insist that all phosphorites have been.formed by fluorapatite and
that kurskite is non-existent among than; others demonstrate the
existence of these two minerals on the premise that their properties
differ essentially from fluorapatite. The verdict of this hypothetical
dispute is of substantial practical importance. If the voracity of the
supporters of the first point of view will be established, then there
will be hope far increasing artificially the P205 content in phos-
phorites up to 40-42g; if, however, the opposite opinion will
triumph, then the attempts-to raise the quality of franOolite phosft
phorites above'37% of P205, and of kurskitcs - above 34.. of P205
will be futile.
Minerologists must pursue the study of the minOrological nature of
the phosphate substances of phosphorites in earnest.
The two known tost:important types among phosphorites arc the
scam [plastovyi] and the nodular [zhelvakovyi] types. With such
distinctions in form arc connected distinction of a.mincralogical
nature - scam phosphorites are deposited by francolite and partly by
fluorapatite, and nodular by kurskite. Kurskite phosphorites are
assimilated by plants more readily than.francolites and fluorapatite:
hence they are being utilized as phosphorite fertilizers without
chemical processing. As separate entities the podollsk phosphorite
nodules stand out. They'are deposited principally by fluorapatite and
in part by carbonapatite.
Seam and nodular phosphorites had formed in shallow areas of
ancient seas as biochemical precipitates and they occur among-sea de-
posits.
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The P205 content in seam phosphorites usually is eqUals to
25-30%, and in nodular, to 15-20 and infrequently to 25%. The
magnitudes of seats in the first reach 10-15 in, and in the
second 1-2 meters. Consequently, seam phosphorites have.a sub-
stantial advantage over the nodular type with respect to 6 higher
P205 content as well as the great magnitudes of the seams.
The seat type includes the Khoper, Seleuksk and Kara Tau
phosphorites.. The latter are of great industrial importance: they
are being shipped to the superphosphate factories in Central Asia.
The Kara Tau deposits are distinguished by a high content of
dolomite, which hinders the production of granular superphosphate and
complicates the technological process phosphorite treatment.
Deposits of nodular phosphorites are widely distributed over the
Russian plains'and occur in the Urals, the Caucasus, in Central Asia
and in Siberia. By the character of admixtures, quartz-sand and
glauconito sand phosphorites stand out.
Deposits of quartz-sand phosphorites arc known chiefly on the
northern outskirts of the Dnepro-Donots depressioh (Briansk and
Kursk Regions), and also in the Aktiubinsk Regf,on. These are not
high-grado phosphorites: their P205 content is equivalent to 15-
19%. Their reserves, however, arc very considerable and thanks to
their occurrence at the surface, they can be extracted by open-cut,
mining.
Quartz-sand phosphorites uscd-as phosphorite fertilizer wore
until recently of local importance. Modern methods of these phos-
phorites by flotation and producing a concentrate containing up to
25-27% of 1,205 hai-e increased to a largo extort the possibilities of
their application.
Glauconite sandy phosphorites are distributed principally in the
Moscow and Riazansk Regions, in the' Chuvash Alltonomous SSR, and in the
vicinity of the upper Kama [river]. [Begin p:32] These phosphorites
contain 20-25% of P205 and are easily crushed. As a result of their
high content of ferric and aluminium oxides, they are not well suited for
chemical treatment, and similar to quartz-sand Phosphorites, are being
utilized principally as phosphorite fertilizers. Their-deposits are
also considerable and are available for open-cut mining.
Until recent times, only phosphorite ores containing no less than
10-12% of P205 were considered eoonomically valuable. For this reason
no attention was paid, for instance, to obel4s [obolovye] phosphorites
long known in Egtonia and the Leningrad RegiOn (5-8 and accazionally
up to la% P205). Investigations have shown that these ores were com-
posed of phosphate shells of the simplest Brachiopoda of the marine
animals of obelus [obolius] and lingula, and an admixture of a large
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quantity of quartz sand grains. It was found that by crushing obolus
phosphorito, it is possible to produce phosphate and quartz grain with
almost no adhesions. Further flotation of this material permit S the
production of a high-grade product with a P205 content over 30%. Thus
it became possible to expand-considerably the possibilities for
utilizing ?bolus phosphorite.
in"Siboria, until recent times, deposits of phosphate ore were
unknown. Formations of apatites and phosphorites found here in the last
few years a re as yet scarcely adequate.
Vivianite is an aqueous phosphate of ferrous oxide. Its deposits
occur in lowland peat moss. Thoy are found in large quantitios in the
swamps of-Vhito Russia where vivianito is used as atortilizer togothor
with peat. Usually vivianite deposits are not largo, but thanks to
their distribution over a. large territory, their occurance at the
very surface, and the simple mannor-of extraction, they are being
exploited successfully by kolkhozos.
Of groat oconOmic importance aro potassium fertilizers for the
production of which the deposits of chlorous and sulfate salts of
potassium or of potassium with sodium and magnesium arc utilized.
Those salts had precipitated from the salt-water in lagoons of early
geological periods under arid and hot climatic conditions.
Tho USSR-is mining two potassium formations - the Solikamsk and
cis-Carpatian. "The first had formed in the Permian period; the second,
in the tertiary. Lately now, very promising deposits of potaSsium
salts which had formed in the Devonian period were discovered.
Potassium deposits aro know- also in the Ural-Embonskii Region and in the
South of Central Asia.
All of those deposits have enough reserves to supply the countryts
demands for potassium fertilizers. The problem simmors dawn mainly to
the pro-exploration of the old, and exploration of new localities for
laying mines for the purpose of more efficient exploitation of all
salt doposits.
Liming of acid soils is accomplished with various carbonic mountain
rock dePoSitod by calcium or dolomite - limestone, dolomitic limestone,
dolomitos, marble, chalk, marl and limo tuffs.
The exploitation of deposits of fresh-water limo tuff and swamp limo,
or lake chulk, is more simple. Their formations usually are not largo,
but they are very numerous. Thanks to thoir friability those rocks
need not always be crushed and, therefore, can be delivered directly
to the field. They usually occur at the very surface which makos their
exploitation much easier.
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The USSR-is known to have a largo quantity of deposits of
carbonic rock. Many of them have been diluted in detail and are being
processed for various purposes: In so doing, frequently a considerable
amount of the waste [Begin p.33] of tpoor cross-section' [malogabar-*
tnyi] gravel which can be utilized successfully for liming of soils is
obtained.
In conjunction with carbonic rock, good results a re produced on
acid soils by manganese. Tho waste of manganese concentration factories
serves as manganese fertilizer; law-grade manganese ore is used for the
same purposc (in the Caucasus, the Urals and a.-number of localities in
Siberia).
Gypsum is being used successfully for the improvement of alkaline
soils in'arid regions. Considerable deposits of gypsum are found in
the Ural-EMbensk Region, in Central Asia - precisely in the places
where alkaline soils cover largo areas.
Recently the application of magnesia fertilizers was begun; tho,
proved to be particularly effective on sandy loam and podzolic soils.
Dolomites and dolomitic limestone arc the principle sources of magnesium.
Deposits of those rocks are distributed extensively in USSR territory, and
in many placcs they arc diluted in detail and are easily available for
extraction.
Magnesia silicato dunito, py'roxenito and sorpontinito can serve as
supplementary .sources of magnesium. Although, as reards assimilation
of magnesium, they do not comes up to such readily soluble forms of
magnesia fertilizers as sulfuric acid or potassium and magnesium
chlorides [khloristyc soli kaliia i magniia], or even to dolomitcs,
thoir worth-is enhanced by the frequent presence of cobalt and nickel
within thorn. Hugo dopoits of magnesia silicates are known to be in
tho Urals, the Caucasus, in Central Asia and in many localities in
Siberia. In some Ural mines large quantities of magnesia silicates
are treated as t'lasto products in the oxtraction of asbestos and other,
useful minerals.
The agrochemical effect of magnesia silicates has not as yet
been sufficiently invostigatod; hence their geologic reserves arc not
considered as agronomic ore.
The work of many investigators has demonstrated the high offective-
ness of boron fertilizers, especially on podzolic soils and forest
siorozoms. In the production of these fertilizers, the calcium and
magnosiuft salts of boric acid - hydroboracitc and ascharitc are
utilized. Their doposits arc being mined on the Indcrsk formation in
thc Gurcvsk Region. Other possible boron sources are the complex
borosilicatos datolito and tourmaline. Boron occurs also in the
waters of some lakes in-Contral Asia and in volcanic mud in the
Crimea and the Caucasus. Tho boron content of those sources is law,
and the expediency of'extracting it from those waters and muds has not
yet been investigated.
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(6) Trans. A. 536
For.tho time being, the pure 8a1ts of corresponding metals arc
being utilized as micrefortilizers. Considering, however, the vast
cropping areas in the USSR, the demand for these salts is very great and
their supply to agriculture is becoming a serious geological problem'.
It is essential to ascertain which natural minerals or rocks containing
microelements Could be utilized directly for fertilizing, and to estimate
their deposits. In many eases it is possible to use industrial waste
with microelement content.
Due to the absence of approved technical specifications for
microelements as a raw material for feftilizers, geologists are unable
to evaluate or estimate their deposits. The Ministries concerned must
eliminate these inconsistencies.
At the conference concerning the geology of agronomic ores, held
in May of this year at the Department of Geologic and Geographical
Sciences of the Academy of Sciences USSR, the situation of the raw
material base of the most important agronomic ores phosphorite,
apatites, potassium salts and raw boron - was-examined, [Begin p.34]
and moans for its future expansion were noted. The conference resolved that
it was necessary to continue the search for phosphorites in the Ukraine,
the Caucasus, the Urals, in Siberia and the Far East. Recent now
finds of phosphorites indicate tic prospects of the search for agronomic
ores in many regions of the country.
The conference noted that it was necessary to study the genesis of
phosphorites and their minerology. The mincrology of calcium phos-
phates has; by the way, boon studied up to now only by separate
geologists. Tho problem, however, proved very complex - the molecular
structure of calcium phosphates which form phosphorites has not as yet
been sufficiently ascertained. The hypothesis that carbon enters the
phosphate molecule is particularly contravefsial. The study of those
problems must assume a systematic character.
Discussing the moans for expanding the raw material base, of the
potassium industry, the delegates to the conference noted that geo-
logical petroleum organizations drilling wells fail to acquire complete
selections of core [samples] from salt scams.
A potassium variety with the atomic weight of 40 possesses
radio activity and occurs in potassium salt in the form of a constant
admixture. Gamma-rays omitted by radioactive potassium penetrate
easily the drive pipes of dfill holes and, therefore, can be measured with
the appropriate instruments. By measuring gamma-radiation in drill
holes, it is possible to locate the petassium scam and to determine
its magnitude and depth of occurrance. The conference resolved that
it was necessary to accomplish such measurements in drill holes
from which no samples had been taken during the time the drill stem
was passing through the vein or scam of potassium salt.
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Trans. A. 536
Thus, to solve the problems confronting our country's agriculture,
an extensive geological investigation of the depo8its of various
agronomic ores assumes an ever greater importance.
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Kodrov-Zikhman, O. K.
Khimizatsiia zemledoliia
[Chemicals applied to agriculture].
Prir oda
43(3):11-20
(I) Trans. A. 537
By R. Adelman
March 1954
410 P933 .
CHEMICALS APPLIED TO AGRICULTURE
Tho September [1953] Plenum of the Central Committee, Communist
Party of the Soviet Union, has charged our countryls agriculture with the
imposing task of raising the agricultural standard to a new, still
higher level.
Successful implamentation of this task will ensure a sharp rise
in the productivity of agricultural crop, and will create an abundance
of popular consumer goods in our country.
An increase in productivity is connected with a widespread
adaptation of a series of agrotochnieal measures of which the use
of chemicals in agriculture ranks among the most important.
' The application of chemicals to agriculture is accomplished
fundamentally by the extensive use of fertilizers, and primarily
mineral fertilizers manufactured by the chemical industry.
The production, and use of mineral fertilizers expanded in our
country only after the Groat October Socialist Revolution. In Czarist
Russia manure was the main fertilizer used, but oven that was applied
in insufficient amounts and, in most cases; without observance of the
most elementary agrotochnical requirements. Mineral fertilizers were
imported from abroad in small quantities and Used only on a few
large manorial ?States; to peasant farms, however, those fertilizers
mere unavailable.
After the revolution the situation was radically changed - produc-
tion'and use of mineral fertilizers began to develop at an unprecedented
pacG. As a result of investigations conducted under the guidance of
ofitstanding Soviet scientist - the academicians N. S. Kurnakova,
A, E. Forsman and Prof. P. I. Preobrazhenskii - rich deposits of
phosphorus and pOtassium ore Were discovered in the first years of
Soviet authority. This permitted the organization and production of
phosphorus.-and potassium fertilizers.
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(2) Trans. A. 537
In the Soviet Union, the basic phosphoric ores aro found in the
rich deposits of apatite on the Kolisk peninsula and in the formations of
phosphorite in different regions of the country including the large
deposits discovered shortly beforo-the Groat Patriotic War in the
Kara-Tau mountains in Central Asia.
The main sources of potassium fertilizers are the richest potassium
salt deposits found at Solikamsk, and also in Kalusha and Stebniki
(Western Ukraine).
Along with the production of phosphorus and potassium fertilizers
developed also the production of nitrogen fertilizers based on the
fixation of atmospheric nitrogen into ammonia. In selecting the
technological process for this industry; the investigations of the
eminent Soviet-scientist,-a.cademician D. N. Prianishnikov played an,
important role. [Begin p.12) He demonstrated that not only nitrate
[nitratnyil, as held prior to his works, but also ammonia nitrogen-
[ammiachnyi azet] can be utilized successfully in plant nutritionl.
Organization of production and vide-spread use of mineral ferti-
lizers in the USSR were closely connected with the realization of
socialist industrialization of the country and collectivization of
agriculture, The fertilizer industry Was established it the USSR
during the first five-year-plan. At the beginning of the second
five-year-plan, our agriculture began to receive considerable amounts
of mineral fertilizers and to use them under the basic technical
crops - cotton, sugar beets, tea, citrus, flax, hemp. As a result,
the yield of thee crops increased considerably during the sedond
five-year period. Thus, for instance, the average yield of raw
cotton in Uzbekistan was 8.9 oontn6rd per ha in 1933; yet by-1939
it had increased to 17 c per ha; i.e., it had almost doubled. The
yields of other technical crops which were receiving mineral ferti-
liprs also increased considerably.
During thc third five- year plan, up to the beginning of the
Great Patriotic War, production and use of mineral fertilizers in
the USSR continued to expand rapidly. In war time the production of
mineral fertilizers was greatly impaired; their application on the
fields of kolkhozos and sovkhozos was sharply curtailed. After the
victorious termination of tho Groat Patriotic war, mineral ferti-
lizers played a very important role in the reconstruction of agri-
culture, and by decision of the Party and the Government, they
have been applied in recant years not only under technical crops;
but also under subsistence crops - cereals, potatoes, vegetables.
Tho September Plenum of the Central Committee, Communist Party of
the Soviet Union resolved on a further sharp mtpansion of production
capacity and of the use orminoral fertilizers. It says in the
Plenum decree - HT? instruct the Ministry of the Chemical Industry and
the Ministry of the Metallurgical Industry to increase from 1954-1963
the production capacit for mineral fertilizers (cbriVorted to
For details concerning-the works of Aeadomician D.N. Prianishnikov
see "Prirode, 1954, no.l.
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conventional units) up to approximately 16.5-17.5 million tons in the year
1959, and up to 28-30 million tons in the year 1964. To bring the output
of suporphosphate in granular form in the year 1955 to 45 percent,
and in tha yoar 1958 to 60 percent of the total production of super-
phosphate. To supply agriculture, beginning with the year 1956. with
ammonium nitrate in granular form only. To work out a plan for wide-
spread utilization of Khibinsk apatitos as well as local phosphorites
to meet the needs of agriculturc"1.
The manufacture and application of so large a quantity of mineral
fertilizers will play a particularly important role in the sharp in-
crease in Yields of cultivated plants and inthe growth of agricultural
production.
The principal typos of mineral fertilizers manufactured by
our chemical industry aro nitrogon, phosphorus and potassium fertilizers.
At present, the bulk'of nitrogen fcrtilizors is being utilizod to
cultivate technical crops. Their application 'undor grain crops is
limited to small amounts of nitrogen used as spring sidc-drossings,
and the portion of vogotablos and forago crops receiving nitrogen
fertilizer is very small. In future the standards for the nso of
those fortilizors for subsistonco crops will .widen markedly.
The instruction of the September Plenum of the September Plenum
of the Central Committoo, KPSS, on supplying agriculture, beginning
1956, with ammonium nitrate - the principal nitrogen fertilizer -
in granular form only, is of grat importance. Ammonium nitrate is a
very offoctivo fertilizer, but it deteriorates in storage and very
often forms lumps; hence it must be ground a gain before it can be
used. This groat shortcoming of ammonium nitrate can be eliminated
by granulation, sinco granular nitrate does not deteriorate.
In the future phosphorus fertilizers will also be used to a .
much groaterextent than at present. Furthermore, as the amount of
suporphosphato increases; the manufacture of phosphorite fertilizer will
also increase; (Begin p.13] the wide possibilities for utilizing
this good phosphorus fertilizer are not by far being fully exploited.
To raise the effectiveness of this frtilizcr, composting'of phosphorite
fertilizer with manure must be practiced more extensively. Composting
increases tho offativonoss of the manuro as well as of the phos-
phorito fertilizer. Besides, phosphorite fertilizer in composts with
manure produces considerable effect not only on highly and moderately
acid soils, but also on mildly acid and neutral soils.
IConcofning measures for the future development of agriculture
in the USSR. Decree of Plenum of Central Cothmitto'KPSS adopted
September 7 , 1953 purduant to report of toy. N. S. Khrushehov,
Gospolitizdat, 1953, p.38.
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(4) Trans. A. 537
The output of superphosphate in granular form is also highly
significant. If superphosphate is introduced in soil in pulverized form,
the availability of its phosphoric acid to plants decreases considerably,
which is due chiefly-to the formation of slightly .soluble iron and
aluminum phospIhntely. If, however, granular superphosphate is in-
troduced in soil, fixation of phosphoric acid by ferric and aluminum
oxides decreases sharply, since granular superphosphate has a much
smaller iii4ea of interaction with the soil than pulverized super-
phosphate. Therefore the effectiveness of granular superphosphate is
considerably higher inmost cases than that of pulveriied super-
phosphate, particularly when applied on red soils and podzolic soils
rich in ferric and aluminum.oxides. Granular superphosphate-is expecially
effective when it is introduced into rows together with seed.
Heretofore, it was held that our [Soviet] agriculture is considerably
less in need of potassium fertilizers than of the nitrogenous and the
phosphoric types. Scientific investigations and industrial experience,
however, show that, as the yield increases, the need for potassium
fertilizers grows. In addition, the application of potassium ferti-
lizers is even now most important for a series of agricultural plants
(legumes, plants with edible roots, potatoes).
It must also be noted that plant requirements for potasdium ferti-
lizers depend to a high degree on the properties of the soil. The
application of potassium fertilizers is particularly important for light
sands and sandy loam soils stretching in our Union over wide expanses.
It is entirely impossible to manage a farm without the application of
potassium fertilizers on peat-bog soils. Here it is necessary to apply
potadsium fertilizers it considerably larger doses, and every year at
that. Hence, in connection with the anticipated drainage and adaptation
Illustration
Mechanized placement Of'mineral fertilizers.
Kolkhoz named after I.V. Stalin, Novo-Ukrain
District, Kirovograd Region, lokranian SSR
Photo by G. Verushkin
of vast areas of peat soils within the near future, agriculture's
demand for potassium fertilizers will increase sharply. At present,
the main potassium fertilizer is potassium chloride manufactured by
our chemical industry, at the base Where Solikamsk potassium salts are
being processed. Potassium chloride is an effective fertilizer with
a high potassium content (about 55g), but it has an essential short-
coming; namely, chloride fixed with potassium has a negative effect on
a number of agricultural plants including such important and widely
distributed crops as potatoes, tobacco and grapes.
? Therefore, our chemical industry louts out for fertilizing pur-
poses also some potassium sulfate the amount of which will have to be
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Trans. A. 537
increased in the future. Besides, there will be a marked expansion in
the production of potassium fertilizers from salt deposits inthe
Western Ukraine. They offer the possibility of manufacturing potassium -
fertilizers in which a part of the potassium is fixed with sulfuric acid.
Along with potassium, these fertilizers contain a considerable mount
of magnesium one of'tho most iziaportant elements in the nutrition of
agricultural plants. [Begin p.14]
ILLUSTRATION
Broadcasting of mineral fertilizers by airplane.
Kolkhoz "Chervonyi stop", Trestianots District,
Vinnets Region, Ukranian SSR.
Photo by E. Kopyt
Many crops quite frequently experience a lack of magnesium, particularly
on light soils, and under conditions of liming, not only on light
but also on heavier soils if the lithe fertilizers used do not contain a
considerable amount of this element. From the salts of the West
Ukranian deposits potassium magnesium [kali-magnezia] is being-produped
-a splendid fertilizer containing about 26-287 potassium oxide. In
the futrue, the production of this fertilizer should be expanded
considerably. Tho unit weight of law-percentage potassium fertilizers
(kainite) manufactufed also by industry on the basis of West Ukrainian
salts must decrease.
Tho assortment of mineral fertilizers must be extended chiefly on
account of tho output of fertilizers with a higher content of active
substances, particularly on account of compound fertilizers containing
two or three nutrient substances.
The use of organic fertilizers, first of all manure and peat, is
of utmost importance. In the Decree of the September Plenum of the
Central Committee, KPSS, it is indicated that it is necessary "to pro-
vide for a censideraba6 increase in the accumulation and application Of
local fertilizers (manure, peat, liquid manure, various composts etc).
In regions with peat deposits, to increase the preparation of peat
for fertilizing and litter. To organize on a large scale the pre-
paration and application of composts and organo-mineral mixtures so
as to increase within the next 2-3 years the use of organic ferti-
lizers on kOlkhozes by one an a half 2 times to compared with the
year 1952'11.
Up to now, there still is the widespread opinion that the application
of local fertilizers cannot be considered as a chemical means applied
to agrieulture; that, for example, the, effect of manure or other organic
fertilizers is not due to apy essential chemical action upon the-soil.
I-Msasures for the future developmet4 of agriculture in the USSR.,
Decree of Plenum of Central Comraittee, KPSS, adopted September 7,
1953, pursuant to report of toy, N,S. Khrushchev, p.38.
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(6) Trans. A. 537
The inaccuracy of this opinion is visible from an examination of the
processes caused by the introduction of manure tn the soil. Not only
large amounts of nitrogen, phosphorus and potassium are introduced into
soil together with manure, but also a great deal of calcium, magnesium
and other cations which, following the decompositien of manure in the
soil, displace hydrogen-ions and enter the absorption complex. As a
result, soil acidity decreases under the influence of manure, although
to a'lesser degree than through the introduction of normal doses of
limo. Manure causes also fixation of motile soil aluainum, thus re-
ducing the injurious action of the latter upon plahts. Manure contains
different microelements including boron, manganese. The Positive effect
of manure is in a large measure duo to the fact that it adds to the
soil a largo quantity of useful microOrganisms and energy material
essential to the activity of microbes.
As manure decomposes, a large mount of carbon dioxide bocomes
isolated in the soil; first itzaturatos'tho soil solution; second,
while separating from tho soil, [Begin p.151 it increases the carbon
acid content in the lower layers of the air. Saturation of the soil
solution by carbon dioxide makes it a more onorgotic reagent which
facilitats the solution of nutrient elements contained in the soil
(for instance, phosphorus compounds) and their conversion into.a stato of
greater availability to plants. An increased content of carbonic acid
in the lowor'atmosphoric strata produced a positive effect on agric 1-
tural plants. Besides, as demonstrated by investigations including
those conducted with tho radioactivo carbon isotope (014), carbon N.
dioxido cdn enter the plant not only through the leaves, but also through
tho roots. As a result, tho increased concentration of carbon dioxide
in tho solution of soil helps to'increaso its content in the plant
and to strengthen photosynthesis.
Manure produces a strong positive effoct upon tho yield of agri-
cultural plants not only in the first year of its introduction in tho soil,
but also in subsoquent years. An experimont conducted by tho Institute of
9ocialist Agriculturo of the Academy of Sciences BSSR, [Mite Russian
SSRI, at its central experimental base in "Ustion, Orsha area, Vitebsk
Region, can be cited as a brilli6nt example of the high effectiveness
of manure on sod-podzolizod loam. In this experiment, after an appli-
cation Of 40 taxa of manure, the yield of winter whoat grain rose
from 17.9 to 32.3 contners por hectare, and tho hay yield of mixed
grasses seeded undor rye and composed of rod clover and timothy in-
ordased the following year after tho introduction of manure from
66.5 to 111.2 centnors per hoctaro.0
Duo to inadequate storage, frequently more than half of tho nit-
rogen content of manurO and a considerable portion of its phosphorus
and potassium are' lost. Yet those losses could be sharply decreased
by implamonting certain measures that can be easily realized on con-
solidated kolkhozos and sovkhozcs. We have in mind the construction
of manure storages and liquid manure receptacles, an increaso in the
amount and improvement of the quality of littorm plowing undpr of
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Trans, A. 537
manure in good time after it has been spread over the field. These
measures will conserve a larger amount of nutritive subs-mnees for
agricultural plants than our farms can obtain.from industrial fertilizers.
The fertilizifig action of manure depends largely on the method
of its preparation. The basic one'should be the currently recommended
compact method for stacking manure. Stacking it solidly in manure
storages as well as in stock-piles built near the farm or In the
field will ensure a fertilizer of good quality, with lossev7of
nutrients falling considerably belaw those in loose storage.
Our country's rich reserves aC peat can be utilized wily as
fertilizers. AS a rule, peat manure iSas.good ag the customary straw
manure and very often exceeds it in effectiveness. Thus, for example,
according to data of the Institute cf Socialist agriculture of the
Academy of Sciences, BSSR, one of the experiments conducted on sod-
podzilized loam at the Atolino-Priluki sovkhoz, Minsk Region, produeed
the following potato yield: without the application of manure - 121.0
centners pr ha, after introduction of 36 tons of straw manure on
1 ha - 179.0 6entners per ha, and after introducing 36 tons at .?eat manure
on 1 ha - 186.4 centners per ha.
To utilize peat manure and other peat fertilizers to a greater"
extent, it is essential to adapt the application of a special factory-
made peat litter to agricultural production, along with peat prepared
by kolkhozes and sovkhozes themselves. In pressed form, this litter
can stand long distance transportation and, consequently, it will find
application also on farms which have no peat moss near their fields.
Peat litter might play a particularly important role On suburban vege-
table farms which usually have a poor supply of straw. Besides, it muat
be taken into account that each-ton of factory-made peat litter will
produce 6-7 tons of good manure.
Peat ought to find widespread Use also in the form of composts
and mixtures with various materials. According to data of White Russian
scientific-research institutions, the effectiveness of a peat-manure
mixture (one part of manure to one-two parts of peat) is equivalent
or almost as good as that of manure. This can be :judged by the results
of an experiment conducted at HUrst'en. In this exPeriment the yield
of 'winter wheat grain constituted: (Begin p.16118.1 centnors per ha
without introduction of organic fertilizers, 31.7 centners per ha after
adding 40 tons of manure, and 30.3 centncrs per ha after adding 40
tons of a peat-manure mixture (owe Apart of manure to one part_of peat);
the yield Of hay of perenni61 grasses (clover timothy) seeded under
rye was 91.6, 123.9 and 117.0 centners per ha respectively.
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(8) Trans. A. 537
ILLUSTRATION
Preparation of peat composts.
Unloading of liquid manure from
motorized cistern
At present peat-manure mixtures are being applied not only in the
Mite Russian SSR, but also'in various regions of the RSFSR - Moscow,
Leningrad, Kirov and others. The most importaht peat ccanposts are
peat-liquid manure and peat-excrement composts.
Along with the intensive peat fertilizers listed above, the dried-
off dnd wind-aired lowland peat must find application in pure form
also. Lowland peat produces a smaller increase in yield than do
intensive peat fertilizers, such as peat manure, peat composts or
mixtures. Peat in pure form can, however, be utilized in instances
when there is more peat available for fortiliging than is needed for the
preparation of mixtures, composts and manures. If applied correctly
and in sufficient amounts (not less than 40 tons pr ha), then dried
and wind-aired: peat aids considerably in raising the yield of agri-
cultural crops. Thus, for instance, in the experiment conducted by
the Institute of Socialist Agriculture of the Academy of Sciences,
BSSR? at "Ustiell,"the yield of hay -af perennial grasses (clover It timothy)
increased from 24.1 centners to 93.7 centners per ha, because the
soil was fertilized with 40 tons of peat.
A highly effective method of utilizing peat is to employ peat
humus pots to grow vegetable seedlings.
In addition to manure and peat, it is essential that other local
fertilizers, such as excrements, eity refuse, barnyard manure, pond
silt be utilized moro extensively. A considerable increase in produci,
tivity can be achieved if these fertilizers are applied corrcctly..
The high effectiveness of excrement fertilizer can be estimated by
data of an experiment conducted by the Institute of Socialist Agriculture
of the Academy of-Sciences, BSSR; the yield of forage beets on a control
test plot was 181.9 per ha, but after application of 36 tong of peat-
excrement compost on 1 ha, itredt-tcc745A. tentners7petnha.
City refuse can be utilized widely as a fertilizer on suburban
farms; it is best to let it first seep throught a compost pile. In
addition, city refuse is used on truck-farms as a biological fuel.
In many regions conditions-favor the application of green manure,
primarily in the form of lupine. Lupine is particularly important in
the reclamation of sandy soils. Nonethologs, it can be used as green
manure also on clayey sad-podzolized ;oils. Thus, for instance, at
"Ustle? the yield of winter rye grain obtained on a control test plot
on clayey sod-podzolized soil was 11.2-contners per ha, yet on the
test plot fertilized w ith lupine - 21.2 c per ha.
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(9) Trans. A. 537
Results of investigations conducted in recent years show the high
effectiveness of green manure combined with peat. Thus, on the ox-
perimontal field fertilized with green mature combined with peat
(40 tons per ha at the "UWe" oxperimontal base,'tho hay"yield of
perennial grasses (Clover timothy) rose from 22.0 to 78.2 contnors
per ha.
The principle plant cultivated for groon manure is the blue
angustifolium lupine. However, in'various localities of our country,
particdlarly in the 'Jost, [Begin p.17] it is oxpodiont to sow yellow
lupine. Tho crop of perennial lupine should also find application,
particularly in the northern part of the USSR.
Special attention ought to be paid to a mote extensive distribution
of the crop of mildly alkaloid (forage) lupinos. These lupincs mako
splendid forage plants, while their root mass, harvogt waste as well as
their aftermath porform the function of green manure. Tho aftermath
of serradella could also be used as green manure to good advantage.
In the Soviet Union, especially in the non-black soil bolt, are
found vast areas of land with heightened acidity detrimental to most
cultivated plants. Such soils contain little organic matter or nutri-
tive elements, possess poor'physical properties, poor structure, a nd
hence are littlo productive. On acid soils, various.agrotechnical
measures including the use of fertilizers very often not only fail to
produce a full effect, but sometimes oven exert negative action.
To improve the agronomic properties of those soils it is necessary,
first of all, to reduce their oxcossivo acidity by liming. In liming
soil in grass-ficld crop rotations, one must strive to reduce ex-
cessive soil acidity to a mildly acid reaction which is best for the
development of perennial grasses used-in grass-field crop rotations
and for most other agricultural crops.
Liming exerts on soil multiple action. Its positive effect is
manifest in the following: it eliminates excossivo acidity and the
action of motile aluminum compounds harmful to plants, *hich is
connected with the acidity, as well as excess manganese. Liming creates
favorable Conditions for the activity of useful microorganisms.
Under the influenco of chemical, phsicochamical and biological
processes which originate in the soil duo to liming, there occurs
an increase in the contont of nitrogen, phosphorus, potassium, calcium,
magnesium, and molybdenum compounds, end of other nutritive substances
available to plants. The physical propertios of the soil improve,
particularly soil structuro, as: do the water, air and nutritive
regimes of the soil. Together with lime the soil is fed calcium - one
of the basic nutritive substances of plants, the lack'of which is often
felt by different crops grown on sod-podzolizod soils.
Liming of acid sod-podzolizod soils is a vell effective method of
incroasing the productivity of agricultural crops. Tho general
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(10) Trans. A. 537
inference of data on field experiments conducted in the Soviet Union
is that average gains in yield from soil liming fluctuate for grain
crops (grain) from 2 to 6 centnsrs per ha; for hay of perennial
grasses (clover timothy) from 10 to 15 centners per ha; for fodder
beets (roots) and fodder cabbage from 40 to 100 centners per ha; fof
table beets (roots) and table cabbage from 30 to 80 centners per ha.
Red clover and mixed perennial grasses, in the composition of
which clover belongs, respond to liming particularly well. The in-
crease in hay of mixed grasses, consisting of clover and timothy,
frequently reach 30-40 centners .per ha. and above. Furhtermore, under
the influende of liming, hay of mixed grasses experiences an increase
in albumin and vitamin content, as well as in phosphorus.and calcium,
which is most significant for the quality of forage; clover which
very often fails on unlimed soils shows increased winter hardiness.
ILLUSTRATION
Transporter broadcasts fertilizers
."TUR 7" on the field
(Begin p.18] In view of the high responsiveness of clover as
well as of other perennial grasses to liming, this measure is most im-
portant factor in intfoducing correct crop rotations with seeding of
grasses.,on acid sails. Soil liming is the medium for obtaining high
and stable crops of perennial grasses with a powerful root system..
Liming produces high increases in the yield of v6rious other crops:
plants with edible roots, cabbage, wheat, barley.
If lime is addOd in doses reducing soil acidity to a mildly
acid reaction; i.e., in normal (full) doses, tAe action of liming
will manifest itself for 10-15 years and longer. Thus, for the period
of its effect on sod-podzolized soils with heightened acidity, each
ton of lime produces a total increase in the yield of all plants grown
during that pefiod equivalent in terms of grain to 12-15 centners of
the production.
Liming influences not only the magnitude of the yield but also
its quality; wlth an adequate magnesium and boron content in the
soil, the influence of liming increases the sugar content in the roots of
plants-with-edible roots, of starch in potatoes, fats in seed, vitamins
in different plants, and also improves the properties of seed grain
and increases their productivity. In addition, the improved seed
grain (yield) properties can be transmitted though heredity from
generation to generation for a number of years.
Tho effectiveness Of liming depends to a high degree on the
doses of lime and methods of placement. In determining lime dosages,
one should proceed from the above cited basic hypothetical principle of
soil liming pursuant to wtich heightened soil acidity must bc reduced
to a mildly acid reaction. The amount of lime required for it del5ends
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(1/) Trans. A. 537
on the soil properties, particularly on soil acidity and the me-
chanical composition of soils, as well as on thoir content of organic ,
substances. If the content of organic substances in sod-podzolizod soils
is not over 3%, then it comes to 4-6 tons per ha far medium axid heavy
barns, and to 2-4 tons per ha foraight looms and sandy soils. Tho
application of such quantities of lime ensures considerable growth
in the yield of most agricultural plants during one or even two ro-
tations of a seven-ten field crop rotation.
Tho amounts of lime iAdicated above can be applied not only by
one method, but by several. This constitutes another important hy-
pothesis of modorn soil liming which proves that it is possible to
repeat tho application of -reduced doses of limo more frequently than
in liming with full doses.
Soil can be limed with different typos of lime rook?end-induttrial
waste and, in the first place, with materials which need not be -
ground or calcinatod; namely; loose lime reek and Iime.waSte of-in-
dustry of which the Soviet Union has huge reserves. Tho basic loose
limo rock constitutes limo tuffs, ground dolomite [dolomitovaia
mukal? lake limo, chalk-marl and marl. Tho main limo wastes are:
ash of igneous-shales, waste of tho sugar industry (defecation),
vast() of the limo, cement and Silicate, metallurgical industries,
waste accumulated in pits at. tho placos whore lime is '001.4 minod for
tho needs of .industry and road construction, and peat ash.
Soil can be limed with hard lime rock, with ordinary and
dolomitic limestone and dolomite, as woll as with solid chalk. To
do this, it is necessary to organize tho grinding of of those rocks
and in areas rich in fuel their calcination.
Some of the indicated materials contain magnasium along with
calcium; for instance, ground dolomite, different products of aerated
dolomitic limestones and dolomites, as well as ground limo [izvest-
kovaia mukai pfeparod form those rocks, slaked limo, dolomitic dust,
and some marls. Lime fortilizers containing magnesium produce a larger
increase in tho yield of many agricultural plants (clover, alfalfa,
lupine, scfrolla and other legumes, plants with odiblo roots, potatoes,
onions eta%) than foftilizers with an inconsiderable admixturo of
magnesium. [Begin p. 19]
.I is extremely important to combine the liming of soil with tho
application of manure and ether organic as well as mineral fertilizorsi
tho highest yield of agricultural crops can be obtained by this medium.
If soil is limed in correct crop rotations with seeding of
grassed, one to bogin with must strive to create cofiditions favorable
far the growth and dovelopment of perennial grasses. This embodies
the third fundamental hypothesis of modern soil liming. 'Proceeding
from this hypothesis, it is necessary to introduce lime under a winter
or spring crop serving as 4 cover crop for grasses, if good yiolds
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(12) Trans. A. 537
of poronnial grasses are to bo obtained.
Liming of sod-podzolized soils with heightened acidity is also
highly significant in solving othcr agricultural problems - in the
dultivation of sod-podzolizod soils, dooponing of the plowing layer,
creation of a forage base, moving different plants from more southerly
regions onto the non-black soil bolt, creating conditions Tfcr better
utilization of commorcial mineral fertilizers.
Tho prime importance of liming was noted bSi comrade G. M. Malonkev
in a report to the 19th conference of the Party. Posirg on the problem
of how to increase productivity in the non-black soil bolt of the
Soviet Union, ho said: "To obtain high and stable yields. Of
agri-
cultural crops hero, it is necessary, first of all, to organize largo
scale liming of acid soils with a simultaneous application of sufficicnts
amount of organic and mineral fdrtilizors, to dbvelop grass culture to
the utmost, to inprovo tillage"'. Our country has vast areas of saline
soils whiCh innmost cases contain much sodium and sOmotimos magnesium
and other cations, yet its calcium content is small. Those soils
are digtinguishod by poor agronomic properties and they produce low
yields. Tho proportios of those soils can be radically improved by
substituting sodium found in a state of absorption (or some cation)
for calcium. This can be accomplished by treating the soils with
gypsum. In localities with saline soils gypsum treatments are of prime
importance.
Striving to fulfill the decisions of the Soptombor Plenum of the
Central Committee, ISS, on raising the agricultural lovol, in addition
to fertilizers already adapted to agricultural production, it is necessary
to utilize also fertilizers which arc not as yet in uto, or have boon
applied on a very small scale. This pertains primarily to microforti-
lizors containing elements required by plants in negligible quantities
(microolements)1.
Microfortilizors currently in use arc the copper typo applied on
peat soils on which for lack of thorn, the grain in a series of grain
crops fails to develop altogether, or develops abnormally, while many
other crops produce reduced yields. The basic copper fcrtilizer is
pyritc cindorg containing, besides copper, different other elements in
largo c4mounts.
In the Soviet Union boron fertilizers are also applied, although
in small quantitics. These fertilizers produco an espocially high effect
undor conditions of limed sod-podzolizod soils. Boron fortilizors not
only increase the yield of flax, but they also improvo thd quality of
its'fibor and'raisc the plant's resistance to bactoriosis. Tho uso of
1 G. Malonkov. Complete roport on the work of thO Central Committee of
the VKP(B) submitted to tho 19th Party conference, Gospolitizdat,
1952, pp. 54-55. .
I Concerning this item see also J; V: Poive's article "Microolomonts in
agriculture", "Priroda", 1953, no.11.
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(13) Trans. A. 537
boron"fertilizers is also very significant in the culture of sugar
beets. Liming acid sod-podzolized soils facilitates considerably
the increase in sugar beet yields; yet at the same time infection
of the beet roots by heart-rot'and the dying-off of the point of
growth are frequently observed. Boron fertilizers are a good medium
in safeguarding sugar beets from these diseases. Everthing said
about the sugar beet concerned the forage and table varieties, and
also, although to a lesser degree, different other plants with edible
roots.
Boron fertilizers have a particularly good influence upon the
generative organs of plants; not only the yield of seed increases,
but the properties of seed grain improve as well, and productivity
rises. [Begin p.20] Hence it is essential that these fertilizers
be utilized extensively in growing various plants for seed. This
applies particularly to clover and to different vegetable cropss also
to flax.
Manganese fertilizers are used'in-our country chiefly on chernozem
soils in the culture of sugar beets. Nonetheless, results of in-
vestigations conducted in recent years have shown that they produce a
considerable effect also in liming of sod-podzolized soils With
heightened acidity. Thus arises the question concerning the appli-
cation of manganese fertilizers also in the Aon-black soil belt of the
?oviet Union under conditions of soil liming.
Our agriculture is making little use of bacterial fertilizers
which represent the cultures of various bacteria facilitating the
accumulation of nutrient elements in the soil or the conversion of, 4
substances contained in these elements to a state available to plants.
Of bacterial fertilizers, only nitragin is being applied to a consider-
able extionicondomitantly with nitragin, the culture of nodular,
atmospheric nitrogen-fixing bacteria is being introduced in the soil.
Those bacteria live on the roots of legumes forming special nodules.
The action of nitragin upon agricultural plants and the conditions
of application of this fertilizer in relation to the biological pro-
pertiee of the plants under cultivation have been studied in great
'detail. This circumstance permits its effedtive use in increasing the
productivity of'various legumes in relation to the specific characteristics
of these plants.
Azotobacterin is used much loss than nitragin; it represents the
culture of the azotobacter microorganism which also fixes atmospheric
nitrogen, but, contrary to nodillar bacteria, lives free in the soil
and not on the roots of plants.
In recent years phosphobacteiin has been adapted to agriculture;
it represents the culture of bacteria the activity of Which is re-
sponsible for increasing in the soil the content of phosphorus compounds
available to plants. The considerable increase in the yield of
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(14) Trans. A. 537
agricultural plants observed in many,experiMonts with phosphobactorin
justified the recommendation of this fertilizer for use in agricultural
production. It =1st3 however, be noted that conditions of offodtive
application of azotobactollin and phosphobacterin have not as yet boon
sufficiently investigated.
For a number of years now, agricultural production is utilizing
the bacterial preparation AMB tagar meat bouillon (?)] which very often
produces a considerable effect. Nonotholoss, the conditions for its
use must also be defined more precisely.
At present a large part of the wofk with fertilizers and their
use is little or not at all mechanized. This pertdins particularly
to manure, peat and lime fertilizer., to gypsum otc. It is sufficient
to point out that hundreds of milIons of tons of organic fortilizors
are loaded on trucks,'unloadod and spread over the field only with
the aid of pitchforks. In connection with the extensive uso of chemi-
cals in agriculture to be adapted within the next few years, it is
essential to increase sharply the mechanization of all processes
connected with the preparation, transportation and placement of
fertilizers.
Largo scale application of fertilizers is a powerful factor
in incroasifig the fertility of soil and the productivity of agricul-
tural crops.
The natural riches of oUr great motherland and the huge successes
of socialist industry permit the utilization of this factor to the
fullest extent in obtaining an abundanco of products and in unceasing
the prosperity of the Soviet people.
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(1) Trans: A. 538
By: 2. Adelman
Gorbachev, I. D.
Resul'taty Ispytanii novogo fosfornogo
udobreniia termofosfatal
[Results of testing Thermophosphatel -
a new phosphorus fertilizer).
Doklady Vsesouznoi Ordena Lenina Akademii
SellskokhoziaistvenAykh Nauk imini V. I.
Lenina, vol. 15, no. 10, pp.35-38
1950 20 Akl
(In Russian)
RESULTS OF TESTING TBERMOPHOSPBATE -
A NEU PHOSPHORUS FERTILIZER
Submitted by Academician P. A. Baranov
It is well known that the best fertilizers for acid mifieral pod-
zolic soils and for peat-bog soils are alkaline fertilizers. These
include thermophosphate2, the new type of phosphorus fertilizer obtained.
by the method developed by B. N. Melentlev (Academy of Sciences, USSR)
by mixing apatite concentrate (or apatite rock), olivenite, celskite
[kol'skit3 and quartz in proportions determined by weight and by
appropriate subsequent treatment.
This fertilizer contains up to 26% P205 (including citrate-
soluble up to 24%), Mg0--8-10%, Ca0--18-20%, S102--43-45%, Fe203--A1203--
2-3%.
A study of the fertilizing action of the present thermophosphate
was conducted by the Author from 1946 through 1949 with different
agricultural crops and on different types of soil (Begin p. 363 '
characteriStid? of the northern part of the Siberian forest zone.
The agrochemical soil characteridtics of the experimental plot
(arable horizon) are cited in table 1.
I In articles and reports published earlier, the given fertilizer was
known as magnesium phosphate, but in the present article it was given '
the name of thermophosphate which is mofe'accurate and fits its nature.
2 The pH of its liquid extract equals 8.0.
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(2) Trans. k. 538
Table 1.
Type of soil
Salt
_
Hydrolytic acidity
m.-ed. (m.ek.) per
100 gr. of soil
Sum of absorbed
Iek.) Der 100 gr of ,
soil
Degree of soil
Isaturation in %
p20q mg (according
to kirsanov)
Experi-
mental
crop
Sandy-
4;62
745
1:44 16:19
20:55
Potatoes (exp:
It
475
8:28
554 40;09
4375
Potatoes (exp.:
Sandy loaa-.;
r4:29
8:57
380 ,30:70
4:50
Oats & barley
Clay soil...
494
? 5:62
3:38 13755
750
Oats
Lowland bogs
418
72:00
49:20 i4059
79;30
Potatoes
n n
4:64
6595
33.:90- 34:00
23:55
Oats
Upland bogs
4:24
104:10
3015 22:46
7;70
Oats
Lowland '
4:24
65:95
30:15 31:37
23:55
Barley
tt tt
4;24
60:40
4560 3:42
23:10
Turnips
n n
4.50
63.63
.65.45 150.70
91.90
Perennial gras
On mineral podzolic soil the investigation of thermophosphttte
action was conducted with potatoes, and oats and barley for hay.
The experiments with potatoes were conducted for two years on a
well tilled plot, on the base of organic fertilizers (peat-manure
composts) in doses of 80 tons per hectare. The soil was a sandy
ferrous podzol.
The experimental results are cited in table 2.
T a b 1 e 2,
1)
es
Experimental scheme
Potatoes
(1947-48)
yield of
tubers
c/ha.
Potatoes
(1949)
yield of
tubers
c/ha
Oats, yield
of hay
c/ha
Barley, yield
of hay
c/ha
NK
NK 7LP in superpho
phate
NK ill? in theimophos-
phate
90.9
.
149.8
?
131.6
73.9
129.5
.
132.5
18.2
23.0
_
24.2
9.3
14.3
_.
22.8
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Trans. A. 538
In the given experiment phosphorus fertilizers were applied with
a disk-harrow after plowing, which is less expedient than deep appli-
cation.
Hence, in 1949, the experiment with potatoes wasrepeated. In
this experiment the fertilizers were plowed under.
The results of this experiment cited in table 2 show that this
time the potato yield derived with thermophosphate was as good as the
yield achieved with superphosphate; this fact corroborates the above
statement concerning the method of thermophosphate application.
The industrial experiments conducted on kolkhozes and eovkhozes
also demonstrated the high effectiveness of thermophosphate. In these
experiments, the application of thermophosphate prodaced a yield that
was higher than the one obtained with superphosphate.
Experiments with oats were conducted for 2 years, with barley for
one year. The experimental plot represented reclaimed virgin land
from a spruce-birch forest. The soil of the plot was sandy loam and
humic ferrous podzol. In the experiment concerned, phosphorus ferti-
lizers'were plowed under.
Table 2 shows also the advantage of thermophosphate over super-
phosphate when applied under the crops of oats and barley planted
for hay on podzolic soils. This was particularly evident in the case
of barley in which thermophosphate applied in the same dose as super-
phosphate'increased the yield of hay by 8.5 cAa as compared to super-
phosphate.
Analogous results were obtained also in vegetative experiments
with oats and barley crops on pOdiolic (virgin as well as cultivated)
sandy, sandy loam, and clay soils, Barley, for instance, gave the
following yield of an airdried mass per container: - on virgin and
sandy soil with superphosphate - 33,5 gr, with the same dose P20.6
in thermophosphate - 39.5 gi, and-with d one and a half doses 50.1
gr; on cultivated soil --52.9; 70.2, 73.1 gr respectively; on virgin
clay.- 36.71 42.9 and 50.1 gr.
On peat-bog soil, experiments were conducted with potatoes, with
oats for hays barley for grain, with turnips and perennial grasses.
it
The experiments with potatoes were conducted for 3 years, ex-
periments with barley, turnips and perennial grasses (mixture of
cereal and legume .grasses of 1935 crop) for 2 years on well cultivated
lowland bog. The results of these experiments are cited in table 3.
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(4) Trans. A. 538
T a b 1 e 3.
Experimental scheme
NK
NK1ZP in superpho6-
phate
NKXP in thermo=- -
phosphate....
Lowland bog
Upland bog
Perennial
CH
'Oats, yield
183.5
225.0
261.4
4.7
5.2
7.8
115.2
165.6
208.5
42.7
43.8
46.9
f hay, c/ha
57.8 0
0
166.3 25.5
for the
66.7
Notes: 1. The experimental results in tables 2 and.3 are cited
of years equalling the time in which the experiment was
conducted.
2. Fertilizer was applied under experimental crops in the
following doses; potatoes - 1480K 1201)80; oats and barley for
hay - N6K6 oP60; barley for grain - N3K7oP6; turnips -
9 80
N80K0 P ; perennial grasses N60K80P6*
3. The good yield obtained with NK in the experiments on
peat-bog soil can be explained by the fact that in the
years preceding the experiment, large doses of phosphorus
fertilizers were applied on the plot.
Data in table 3 show that the yield of potatoes, barley (grain),
turnips and perennial grasses on boewith thormophosphate is higher
than the yield gained superphosphate.
In addition, it must be noted that thermophosphatc, concomitantly
with increasing the productivity of perennial grasses, exerted an in-
fluence also on the composition of the grass mixture; on test plots
whore this fertilizer was applied, the amount ofgrasses valuable as
forage cilops had increased (timothy, foxtail).
[Trans. note: fertilizer dosages in note 2 are practically illegible
in original text.)
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Trans. A. 538
Industrial experiments in which thermaphosphato was:applied-under
perennial grasses on lowland bog produced the same results.
The data show that with the crop of oats planted hay on lowland
bog, thermophosphate equals supprphosphato in effectiveness. Analogous
results were obtained also in vegetative experiments with virgin peat
of lowland bog.
Tho situation is entirely different on upland beg where we were
unable to Obtain a yield of oats with'a selection of the usual mineral
fertilizers containing superphosphate. Sprouts -advanced in height
only 5-6 cm, had a stunted appearance and a ydllow color, and they
remained like that for the rest of the summer. In contrast to this,
the yield of hay' from oats ebtainedlwith thermophosphatc. was 25.5
contners per ha.
Similar results were obtained in vegetative experiments with p.-)at
of upland bog in which, upon application of superphosphate, oats
perished completely after reaching barely 5-6 cm in height; yet with
the use of thermophosphatc it showed powerful development, had a normal
dark-green Coloring, reached a height of 70 cm and produced-the high
yield of 61.5 gr of air-dried mass per container. LBegin p. 38]
Besides the experiments conducted to investigate the effect of
thermophosphate upon the productivity of agricultural crops in the
year it is applied, a study was also made of the action it exerted
in subsequent 1-ears, and of the influence of thermophosphate of
various grists.
A study of the aftereffect of tnermophosphate on peat-bog soils
was conducted with perennial grasses. The experiments demonstrated
that the aftereffect of thermop'hosphate on these soils is more pro-
nounced than its direct action. For example, the yield of hay of
perennial grasses Obtained a year After the application of super- '
phosphate-amounted to 44.2 centners per ha; yet after thermophosphate
it was 51.5 cAla.
Studies of the aftereffect of thermophosphate on mineral soils
are still incomplete and more work must be done in this direction.
A study of the effectivenessof.thermophogphate of different
grists was conducted in vegetative experiments. They showed that to
obtain good regults, the grist of thermophosphate must compare to
Thomas slag; i.e., it must not exceed 80 mesh in coarseness; the use
of coarse grist thermophogphate reduces strongly its effectiveness
in the year it is applied. This ig particularly apparent on mineral
soils, but less pronounced on peat. For example, on sandy ferrous
podzol, in the year of application, the yield of the air-dried mass
obtained from one container was: oats, after coarse thermophosphate
(30 mesh)'- 17.34 gr, after fine (80 mesh) - 31.29 gr; barley - 11.52
gr and 41.56 gr respectively.
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(6) Trans. A. 538
CONCLUSIONS
1. As a phosphorus fertilizer, thermophosphate-is fully adaptable
to mineral podzolic and peat-bog soils of the North. With respect to
its effeativeness on mit6ral soils, it equals superphosphate, and on
peat soils it exce116 it.
2. To obtain greater effedtiveness of thermophosphate on mineral
soils, it must be plowed under. This method of application increases -
its availability to plants.
3. The grist is of decisive importance-in the adaptation o
thermophosphate, it must not exceed 80 mesh.
4. The application of thermophosphate'produces satisfactory
yields on acid'upland bogs without the use of neutralizers (lime,
nepheline etc).
5. Thormophosphate is a very valuable fertilizer for the northern
part of the podzolic zone gnd for peat-bog soils; it oxceld super-
phosphate in effectiveness.
Vsesoiuznyi N.-I. Institut Received at Editor's Office
Rastenievodstva March 31, 1950
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Trans: A. 539
--Byl R. Adelman
Baranov, P. A. and Korenllov D. A.
0 nevem vide slozhnykh kontsentriro-
vannykh beschlornvkh udobrenii
[New type of -compound conb'entrated
non-chlorous fertilizers].
Zemledelie, vol. ?4
Aug. 1954
(In Russian]
no 8, pp.60-,67
aD Z44
NE7 TYPE OF COMPOUND CONCENTRATED
NON-CHLOROUS FERTILIZERS
The Institute of General InorgafiiC Chemistry of the Academy of
Sciences, USSR (A.Ia. Zvorykin and F.M. Perel'man), 'has developed a -
method for obtaining a compound concentrated non-chlorous :fertilizer.
It is composed of potassium and ammonium phosphates which, crystallize
themselves in the process of their production in so. close a-Contact,
as if each of the salts had been "dissolved" into the other. Because
of this, the autHors gave the fertilizer the additional name of
"solid solution". This fertilizer contains about 5% of nitrogen,
about 50% of phosphorus, and about 22-23% of potassium.
In recommending this fertilizer for production, the authors
point at a series of its advantages over ordinary fertilizers.
In the opinion of the authors, this fertilizer is more effective
than acorresponding mixture of simple fertilizers and, in addition,
it eliminates the need of mixing simple fertilizers. The other ad-
vantage of this fertilizer, the authors point out, lies in its high
content of nutrient substances and in the absence eff ballast matter,
particularly chlorine.
Regardless of the fact that the matter had not been sufficiently
investgated, the above fertilizer was widely advertised by the authors.
Particularly A.Ia. Zvorykin, in his article "New Type of Mineral,
Concentrated Fertilizers", published in No. 273 (6379) of the news-
paper "Socialist Agriculture [SOtsialisticheskoe zemledelie]" of
November 19, 1952, cited a line of figures obtained from results of
exp3riments conducted with different crops which indicated the ad-
vantages of the above fertilizer over ordinary types.
Upon the suggestion of the Ministry of Agriculture, USSR, the
All-Union Scientific Research Institute for Fertilizers, Agrotechnics
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(2) Trans. A. 539
and Agro-Soil Science, and a number of other scientific research
institutions conducted in 1953field and vegetative experiments-to
test the proposed fertilizer (hincefarth-to be referred toas potassium-.
ammonium-phesphate) on a series of agricultural crops under differont
soil and climatic conditions.
During the testing this fertilizer was compared with concentrated
fertilizers containing no chlorine (ammonium nitrate, double super-
phosphate or ammophos, potassium sulfate).
The varied content of nitrogen, phosphorus and potassium within
potassium-atiamoniuM-phosphate was taken into consideration during the
experiments. H once this fertilizer was tested with and without
additive of other concentrated fertilizers, bringing the nitrogen,
phosphorus and potassium'doses up to the proportions adopted in practice.
To bring out the advantages of potassium-ammonium-phosphate more.
fully, the experiments were conducted under conditions favorable to
the action of this fertilizer: tho choice of crops was limited to those
sensitive to chlorine and to saline concentrations in general, and of
soils at light as possible. Along with broadcast placement, [Begin
p.611 the fertilizer was tested also under conditions of localized
application at planting time.
Vegetative experiments. The experiments conducted by VIUAA
.A.11-Union Institute of Fertilizers, Soil Science, and Agricultural
Engineering' .(Candidate for Agricultural Sciences, A.M. Shchepetile
nikova) with flax on podzolized sandy loam and clay soil have shown
(table 1) that the compound fertilizer potassium-ammoniuth-phosphate,
applied without an additive of nitrogen fertilizer, (var. 2) produces
the same effect'as an equivalent amount of simple concentrated ferti-
lizers (var. 3).
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Trans. A. 539
Tablel.
No. ill
conc.
order
Expermental variant
IYield of flax (in gr per container)
on:
Sandy loam Clay soil
Fiber Seed
[solomIca]
Fiber
.Seed
No fertilizer 4.21
2 Pbtassium-ammonium-phosphate
(in dose of'0;2-gf-PO-lief'
container)
Binary superphosphate (0.2 gr
P205 per container)/ammonium
nitrate and potAssium sulfate
(in dose of var.2)
Ammonium nitrate. itpotast'iUM-
sulfate .(base)
5 -Base 71 potaddiUMamffionitim--
phosphate
7
Base binary superphosphate
Base 71 simple superphosphate
8 Ammonium nitrate, simple super
phosphate, potassium chloride
5.48
0.72
0.96
5.73 1.15
11.36 3.11
14:18 1 3.67
13.96
3.69
11.62 3.23
10.88 3.29
18.69 k 4.78
17.90 4.92
15.37 4.55
Only by-adding a sufficient amount of nitrogen fertilizer?and potassium
sulfate (var.5) a marked increase in fiber yield (b9- 2-2.5 times) as
well as in seed yield (by 3-3.5 times) Was obtained.
A 2-3 time increase in yield was obtained from potassium-ammonium-
phosphate and from the comparable potassium phosphate [fosforno-
kaliinogo] fertilizer on a-nitrogen base on light sandy loam as well
as on heavy clay soil (Var. 1 and 2, 4 and 5). However, the simple
concentrated fertilizers - ammonium nitrate, binary superphosphate
and potassium sulphate - produce under analogous conditiona practi-
cally the same effect as the fertilizer under investigation (var,6).
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(4) Trans.- A. 539
On a base of an adequate dosage of nitrogen and potassium ferti-
lizer (in the fm of potassium Sulfate), the fertilizer under investi-
gation produces a. large increase in yield as compared to a selection cf
simple fertilizers only if binarl superphosphate is substituted with
an ordinary pulverized type (var.7), and potassium sulphate is sub-
stituted with a chlorous type (var.8).
The effectiveness of potassium-ammonium-phosphate in localized
application at plahting time was studied in another eicperiment with
flax on sandy soil. The results are cited in table 2.
Data in table 2 show that neither the absolute yield nor the
increase produced by the fertilizer was large (war.2), when the
application of potassium-ammonium-phosphate was localized (under the
seed) (war.2). An equivalent dose of phosphorus and potassium
(in the form of binary superphosphate and potassium sulfate) ptoduces
the same effect as the fertilizer under investigation (war. 4).
Yet, by applying an adequate dose of nitrogen fertilizer (in
the form of a side-dressing at the bottom of the container), the
effect derived from applying the fertilizer undei- iaestigation
under the seed increased by more thah twice (var. 6).
T a b 1 e 2.
No. in
cons.
order
Experimental variant Yield of flax
(in gr per container)
Fiber
Seed
1
2
No fertilizer
Potassium-ammonium-phosphate (0 1
gr P205)
4.21
5.1
0.72
0.83
3
Binary superphosphate (0.1 gr P205)
4.23
0:69
4
Same 7? K2304 (equivalent to var.2)
5.04
0.93
Base (ammonium-nitrate 7? potassium
sulfate) (N--0.25 gr, gr)
7.84
2.70
6
Base 71 potassiuMatMoniuM=phaSphate
(0.1 gr P205)
9.95
4.07
7
Base 71'binary superphoSphate-(0.1-gr
P205)
9.54
3.84
N o t e: phosphates were introduced 2 cm bulaw seed-(in rows).
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Trans. A. 539
Under these conditions, the effect Of simple concentrated fertilizers
also rose sharply, an4 4 certain decrease of it, as compared with .
the fertilizer under investigation, can obviously be explained by the
fact that no potassium sulfato was introduced in the nrows" in these
experimental variants (var.7).
The experiment with flax was conducted also by the All-Union
Scientific Research Institute for Agricultural Microbiology on sod
podzolized moderdtely clayey soils. The-fertilizers were introduced
at the rate of 0.1 gr K20 per kg of soil. The amount of the other
fertilizers was calculated in 6onformity with this dose. Data of
yield are presented in table.3.
T a b 1 e 3.
NO. in
cons.
order
Experimental variant
Yield of flax
gr per container)
Fiber Seed
1
Control (to fertilizer) 21.9 6.8
2 Totassium-amnonium-phosphate
3. Ammonium nitrate, binary superphogphate?
potassium sulfate (in dose of var.2)...
4 Potassium-ammonium-phosphate with addi-
tive of ammonium nitrate andipotassium
sulfate as a side-dressing .4.
5
6
Ammonium nitrate, binary superphosphate
and potassium sulfate (doses and p1666.1
ment methods analogous with var. 4)....
Ammonium nitrate, simple superphosphate
and potassium sulfate (doses and-pldde=
ment methods analogous with var.4)
Ammonium nitrate, simple superphosphate,
potassium chloride (doses and-pladeMent
methods analogous with var.4)
31.5
32.6
6.8
8.44
49.8 14.3
53.6
53.9
52.9
15.4
15.5'
17.2
The data cited confirm that potassium-ammonium-phosphate (var.2)
and simple concentrated fertilizers (var. 3) added to a dose of
potassium-ammonium-phosphate produced poor results, while an addi-
tive of nitrogen and potassium-fertilizers mixed with the ferti-
lizer under investigation (var.4) contributed toward a donsiderable-
increase in the yield of the fiber and the seed of flax. At the
same time a mixture of ordinary fertilizers, even in the form of
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Trans. A. 539,
simpie superphosphate and potassium chloride, (var. 6,7) showed better
action than the fertilizer in question (with an additive of nitrogen
and potassium up to a normal ratio).
Experiment with maize. A vegetative experiment liad conducted by
the Soil Institute of the Academy of Sciences USSR (D.M. Kheifets-
Shtrausberg) on cis-Caucasian chernozem soil to study the influence
of potassium-ammoniuth-phosphate upon the development of the vege-
tative mass of maize. Data on this experiment are given in table 4.
Table 4.
No. in
cons.
order
Experimental scheme
2
4
5
IPotassium4ammonium-phosphate (at
rate of 0.5 gr P205 per container)
No fertilizer
. ?
NB003, KCl; Ca(Bg04)2;H0-(in
dose of-var. 2)
Potassium-adtmonium-phosphate (in
dose of var. 2) 7.(NRINO3 (up to 1
gr N) . pOr' - 66n-
tainer)
NB4NO3, KC1; Ca(W04)2-Hi0-(in
dose of var. 4).....
Weight of dry plants
(in gr per container
for 30 days of
vegetation
1
far 51 days of
vegetation
2.17 6.2.
3.67
10.32
4.00
10.60
6.12
31.75
7.40
33.50
Table 4 shows that potassium-ammonium-phogphato (var.2) and an
equivalent quantity of simple fertilizers (var.3) produce almost the same
action exerting a mild influence upon the development of the vegetative
mass of maize. By supplementing potassium-ammonium-phosphate with
potassium nitrogenous.fertilizers up to a nofmal'ratio, the develop-
ment of maize was considerably improved (var. 4). A mixture of simple
fertilizers, however, produced a somewhat bettor effect than ,an equivalent
amount of the fertilizer under investigation (var.5).
Field' experiments. To test potassium-ammonium-phosphate, a
series of experiments was conducted with potatoes on the podzolic
soils of TSCS [Central Insurance Department of the People's Commis-
sariat for Rail Transport (?)] VIUAA0 the Novozybkovsk Experimental
Station of VIUAA, at the Institute of Potato Economy, and at the
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Trans. A. 539
Institute of Grain Economy of the Non-BlaCk Soil Belt. The results
of these experiments are cited in table 5.
Table 5 shows that the increase in the yield of tubers and starch
derived from the application of potassium-ammonium-phosphate is in- ;
signifi6ant in localized (var.6) as well as in basic application
('var.2). High yields of potato tubers and starch were Obtained on all
types of podzolic soils, including light sands and sandy loams, when
potassium-ammonium-phosphate was supplemented with an adequate amount
of nitrogen and potassium fertilizer (var.4). In addition, simple
concentrated fertilizers were in every case as good as potassium- '
ammonium-phosphate, and in a number of cases their action was better.
Concurrently with the above, the eXperiments of the Institute of
Grain Economy of the Non-Black Soil Belt, conducted on heavy clay soil
and sandy loam (table 6) at the "Put' Oktiabria" kolkhoz, Kuntsev
District, Moskaw Region; demonstrated that the effect of potassium-
ammonium-phosphate (var.2) upon the yield of potato tubers was inferior
to that of an equivalent amount of ammoniu6 nitrate, simple super-
phosphates and potassium chloride (var. 3).
Only by supplementing the fertilizer being tested ammonium nitrate
and potassium chloride (var.4), the yield of potato tubers was con-
siderably increased, particularly on light soil. At the same times
simple fertilizers in normal proportions (var.5) produced better action
on light sandy soil than on heavy clay, as compared withthe potassium-
ammonium-phospate variant plus an additive of potassium nitrate ferti-
lizers (Tar.4). [Begin p. 651
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No.
in
cons.
order
0-1-0001-000Z01-09Z171-0108dCl-V10
Experimental variant
TSOS VIUAA
?
( )
NovOzybkovsk Experimental
Station
Sandy soil
No manure
Heavy clay mo-
derately pod7
zolized -soil
No fertilizer
Potassium-ammonium-
phosphate
Simple concentrated
fertilizers (in dose
of var.2)
4 Potassium-ammonium-
phosphate with addi-
tive of potassium
nitrate fertilizer
5 Simple concentrated
fertilizers (in dose
var.4)
6 Potassium-ammonium-
phosphate under tuber
Simple fertilizers
(in dose var.6) under
tuber
Tabers tStarch
,L21.4.7.,6 40,1
260.7 42.1
264.7 43.5
322.0
318.1
250.4
51.8
53.0
40.2
Table 5.
-I Institute of Potato Economy
Midly podzolized soil
0
(D
0
Institut g;
of Crain gi
Economy, a
Non-Blac"Sa
Soil Bel a>
avy 33
Clay mo-E3
derately
podzoliz
11
0
Tubers -'7J
274.9 E.
n.)
45.5 285.5
co
48.2 286.5
ubers
154.8
169.8
164.4
231.1
249.6
157.g
Starch
1 20.2
22.6
22.7
31.0
33.8
23.1
With manure
Yield (in
Tubers
212.2
228.9
220,7
253.0
260.0
199.6
No manure
'With manure
centners per ha)
Starch Tubers StarchLTubers Starch
26.9 217.8 38.1 1 275.0 43.7
33,9 1 249.5
29.3 257.9
29.3
263.8
32.9 I 291.9
24,1 j 229.3
43.2 286.4
45.9
45.6
50.5
4O.1
299.6
279.3
281.1
287.6
43.0 292.9
4-3.9
)46.9
293.1
281.3
0
-0
co
0
0
n.)
251.8 41.9 1 t 50.4 289.5 n.)
Note. Second and third experimental variants had the following doses of fertilizers: N--4.8, P205--50, Kz0--23.51
fourth and fifth variants had N--60, P205--50,.K2&-60,_sixth and saventh_had W--3.3, P205-33.3, ,K20--14 kgg per ha. 8
0
In the first three experiments potassium-ammonium-phosphate was compared with simple concentrated fertili:zzers:
185.2 25.8
212.6
28.0 I 226.4
39.9
298.1
ammonium nitrate, double superphosphate and potassium sulfate, and in the experiment of the-Institute of Graain Economo
of the Non-Black Soil Belt, with ammonium nitrate, simple superphosphate and potassium chloride.
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(9) Trans. A. 539
T a b 1 e 6.
No. in 1
Potato-yield
cons.
Experimental variant
(in cent. per ha) on:
order
Heavy -
clay
Light
sandy soil
J J
1.
No fertilizer
109.5
60.0
2
Potad - 0 : 66
c/ha
170.2
110.0
NPK (in dose of var. 2)
192.0
130.0
4
Potassium-ammonium-phosphate 0.66
c/ha NK (N--33, K20--33 kg/ha
223.5
188.0
5
NPK (in dose of var.4)
207.7
192.0
6
Potassium-atiamonium-photphate 0-.66-
c/ha N 29.7 kg/ha
210.0
152.0
An additive merely of a nitrogenous fertilizer increased sharply
the yield of.potato tubdrs (var.6) ensuring an additional increase in :
yield up to 40 c per ha.
Experiment with tobacco. Tests of potassium7ammonigm-phosphate
on tobacco were carried out at the experimental stations of he All-
Union Scientific Research Institute for Tobacco and Makhorka. These
investigations. also confirmed that potassium-ammonium-phosphate fails
to produce significant increases in tobacco yield without a supple-
mentary application of potassium nitrate fertilizers; yet an additive of
nitrogen and potassium to the fertilizer up to a-normal ratio of
the substances increased the yield; and the output of high-grade
varienties of tobacco. At the same time, the action of an equivalent
amount of simple concentrated fertilizers was in most cases approxi-
mately the same, and in individual cases better. At the Crimean
Experimental Station, in 19523 the application of ammonium nitrate
(N30), superphosphate (P205-100) and potassium sulfate (K20--100)
producod a total tobacco yield of-14.1 c per ha, with the yield of
its high-grade varienties being 5.5 c:per ha; yet with potassium:.
ammonium-phosphate (P205--100) the total yield 6f tobacco *as 11.4
c per ha, and the yield of superior varieties 5.1 c per ha. By
supplementing potassium-ammonium-phosphate with a-potassium nitrate
fertilizer, the total yield of tobac6o came to 13.3 contners including
the yield of superior varieties of 5.6 c per. ha. At tho same staltion,
in 1953, the tobacco harvest in the N30 P100 K100 Variant was l0.1
contners per ha, with the yield of superior varieties being 5.7 c per
ha, but in the potassium-ammonium-phosphate variants supplemented
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(10) Trans. A. 539
with ammonium nitrate and potassium sulfate up to the proportions in-
dicated in the above) variant, tho tobacco yield cathe to 10.8 a per ha,
and the yield of high-grade tobacco varietios? to 5.6 c per ha.
At the Kotaisk Experimental Station, the following tobacco yields
were obtained in 1953 (table 7).
Table 7.
No. In
Cons.
Order
Experimental variant
Total fIneluding yield
yield fof superior va-
ieties as of 3rd
IbrOaking
in cent. per ha
1
2
No fertilizer
Totassium-ammoniuthho?phat6-%-,'
1390K40,21 .kg/ha
22.86
25.75
Ammonium nitrate, superphosphate
[potdssium sulfdte-(in-de6e-of-id
2) 25.63
Potassium-ammonium-phosphate-/
KNO3 NNNO3 (N45P90K100) 28.30
Ammonium nitrate, superphosphate,
potassium sulfate-(ih doso'Of-Vaf
4) .
24.95
7.0
8.1
7.7
11.3
'[Begin p.661 Table 7 shows that potassium-ammonium-phosphate
(var:2) and an equivalent amount of single concentrated fertilizers
(var.3) exerted similar action upon the total yield of tobacco. But
the yield of high-grade varieties fertilized with simple concentrated
fertilizers was by-3 c per ha higher than in the case of potassium-
ammonium-phosphate. By' supplementing it with ammonium nitrate and '
potassium nitrate -(var.4), the total tobacco yield was 3.35 c per ha
more than *ith simple concentrated fertilizers used in the same propor-
tions (var.5). The yield of high-grade tobacco varieties, however, in this
experimental variant with simple concentrated fertilizers was also higher
by 3-6 centners than with potassium-ammonium-phosphate combined with '
potassium nitrate fertilizers.
Experiments of the same stations have demonstrated that the action
of a mixture of simple fertilizers is more effective than that of pot-
assium-ammonium-phosphdte, if its potassium sulfate is substituted
by potassium magnesium.
In addition, we shall cite the results of testing potassium-
ammonium-phosphate on sunflowers and castor plants obtained in the 1953
experiments of the All-Union Scientific Research Institute for Oil
Crops (table 8),
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.11
No.
in
cons.
order
Experimental variant
1
2
3
4
No fertilizer
Potassium-ammonium-phos-
phate N0163P6,25K2,75
kg/ha
Organo-mineral granules
NPK (in dose of var.2)
Granular factory super-
phosphate P6,25 kg/ha
Potassium-ammonium-phost
phate ,(in dose of var.2)
Trans. A. 539
T a b 1 e 8.
Method
of
Sunflower Castor plant
in c/ha
place-
merit
yield I in-
'crease
yield
in-
crease
26.6
-
9.9
on one
28.2
1.6
10.8
0.9
side of
hill
same
27.9
1.3
10.9
1.0
28.6
2.0
11.1
1.2
in
hill
26.9
0.3
11.0
1.1
These data shauthat potassium-ammonium-phosphate (var.2)'is
equivalent in Offeetiveness to organo-mineral granules applied in the
same dose (var.3), and is somewhat less effective than granular super-
phosphate applied in doses identical with the second and thirds variants.
Thus, the results of field and vegetative experiments indicate
that potassium-ammonium-phosphate has no advantage over an appropriate
selection of plain concentrated fertilizers the action of which, in a
nuMber of cases, proved to be better.
The advantage of potassium-ammonium--phosphate simmers down to nothing
if the chlorous potassic salts [khloristykh kaliinykh solei] are "sub-
stituted filth potassium sulfate or even with the cheaper potassium
magnesium. This substitution incorporates in the fertilizer not only
potassium, but also other valuable nutritive elements, such as sulfur
and magnesium. ?
The experiments have also shown that a simultaneous combination of
nitrogen, phosphorus and'potassium in one crystal of a compound ferti-
lizer is not a necessity.
The difference in the motility of the ions NH4, K, PO4 in the soil,
the series of conversions and variations which fertilizers undergo in the
soil under the influence of physico-chemical and biological processes--
all these phenomena and facts practically void the importance that once -
was attached to the close union of nutritive elements within one crystal.
They remain in Such a state only until they are placed in the soil, and
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.1 .1
(12) Trans. A. 539
theAT joint locale (focus) in the soil is very Conditional for potassium
and particularly for nitrogen which afe distinguished by great motility
in the soil as compared to phosphorus. Furthermore, the simultaneous
introduction of nitrogen, phosphorus and potassiuM fertilizers into the
soil is not prompted by therequirements of plants. As the plant de-
velops, its requirement for nitrogen, phosphorus and potassium varies,
In this connection, already D. N. Prianishnikov pointed out that the need
fur fertilizers depends not only on the species of the plant, but also
on the properties of the soil. Potatoes on light soils, for instance,
may require most of all potassium, yet on chernozem soils, phosphorus;
in additiqn, potatoes on the same soil, thriving on manure, will sooner
require a supplement of phosphorus and nitrogen-than of potassium, in
contrast to potatoes not fertilized with manure. The degree of Clover
and alfalfa participation in crop rotations also leaves its mark.
Hence the simultaneous introduction of nitrogen, phosphorus and potassium
fertilizers into the soil is not an absolute precondition. Besides,
the results of the investigation have shown that potassium-ammonium-
phosphate, thanks.to the disproportion in the content of its nutritive
substances, produced a high effect only when it was mixed with simple
mineral fertilizers, bringing its nitrogen, phosphorus and potassium
content into more effective proportions. As a result, the authors
argument that tho application of the compound fertilizer proposed by
them will relieve agricultural production of the task of mixing fertilizers
lacks foundation.
It follows from the above statement that, in view of the absence of
any advantages of potassium-ammonium-phosphate over simple concentrated
fertilizers, the question of expediency of its production cannot be
settled by agronomic evaluation, bdt solely by the technical and
economic factors of its production.
?
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(1)
Genkell, P.A. arid Oknina,
Diagnostika morozoustoichivosti
rastenii po glubine pokoia ikh
tkanei i kletok
[Diagnosis of frost-resistance in
plants by the depth of dormancy of
their tissues and cells.]
lzdatelLstvo Akadsmii Nauk SSR,
Moskva, 1954, Not in USDA
(Instruction in methods).
(In Russian)
Trans. A. 540
Abstract and trans.
of Conclusions
By: R. Adelman
DIAGNOSIS OF FROST-RESISTANC7. TN PLANTS BY TEE
DEPTH OF DORLAITCY OF THEIR TISSUES AND CELLS.
Chapter I contains the following list of indicators of the depth of
dormandy and frost-resistance in plants:.
1. Conversion dynamics ofreserve substances;
2. Process of the separation of protoplasm;
3. Character of plasmolysis in a saccharose solution;
4. Time of advance of cap plasmolysis;
5. Resistance of lipoids to temperature influendes ?
(in plants in whose tissues lipoids develop).
In most grads crops starch converts into sugar, in woody species into
fats :end lipoids. Observations of conversions are conducted with the aid of
microreactions.
Plasmodesma are observed in a plant while it is in a state of vegetation.
It connects adjacent plant cells, but separates itself from the cell walls
as the plant enters a state of dormancy.
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(2) Trans. A. 540
If the process of protoplasm separation is not discovered directly
microscopically, then the state of the cell (dormancy) can-be determined by
the shape of plasmolysis in a molar solution of saccharose.
The time of approach of cap plasmolysis indicates the degree of swell-
ing of protoplasm colloids; this circumstance is one of the best indicators
of the depth of dormancy.
Frost-resistant varieties of woody species are characterized by a deeper
conversion of substances than varieties of poor resistance. Conversion of
substances in frost-resistant plants goes far enough to form fats and lipoids;
in poorly frost-resistant varieties it ends with fat-like substances or sugar.
In the first, decomposition of lipoids occurs at a temperature of 70?C, in
the second at 40-50?C - a fadtor of utmost importance in the survival of a
plant.
Chapter II deals with the comparative evaluation of diagnostic methods;
chapter II, with evaluation of the degree of frost-resistance in fruit
crops obtained on the basis of microchemical reactions, and chapter IV,
with thc depth of dormancy and frost-resistance in cereals and other grass
crops.
CONCLUSIOMS?
In summing up the above account, we are ableto submit the following
diagnostic scheme for frost-resistant plants.
1. Observation of dynamics of the conversion of reserve substances
(the disappearance of starch and accumulation of sugar ifi grass crops, and
fats in tree species characterize the state of dormancy).
2. Microscopic detection of the separation process of protoplasm (in
a state of dormancy protoplasm separates to some degree from the cell walls)
and the color of plasmodesma (the absence of plasmodesma characterizes the
state of dormancy of woody and grass crops).
3. Investigation of the character of plasmolysis in a molar
saccharose (bulging plasmolysis - state of dormancy or emergence
4. Investigation of the rate of approach of cap plasmolysis
nate potassium (the slower the .approach of cap plasthelysis, the
is protoplasm, the more sound the state-of dormancy). The given
particularly pronounced in tree species.
solution-of
from it).
in thiocya-
less bulging
property is
5. Investigation of the magnitude and stability of lipoidic layers
(the greater the depth of dormancy, the sharper is the reaction to lipoids
in tree species, and the temperature dausing the destruction of lipoids must
be the higher).
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(3) Trans. A. 540
The above described methods determining the depth of dormancy and itg
relation to frost-resistance are fully within the reach of any laboratory.
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