JPRS ID: 8317 PERISHING OF WINTER CROPS
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JP1tS L/8317
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7 March 1979
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PERISHING OF WINTER CROPS
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REPORI' DOCUM~NTATION I~t.pE1'ONf N0. I~. q~ain~~m~~ ~eat~~on No.
PAGE JPI2S L/$317
Till~ ~nd Subfdt� b. R~port D~1~
I'IsRI5HINC OA' WINTCIt CItOP5 7 March 1979
~
7. Aulho~t~) P~HorminQ Or~~nu~tion p~p1. No,
F. M. Ku erman und V. A. Moise ch k
f. V~rlorminr OrtanitaUon Nam� and Addra~ l0. Pro~~et/T~+N/WOrw Urot Nu.
JoinC Publications Ftesearch 5ervice
1000 North Gl~be R08c~ 11. Co~tne~(C1 0~ arant(o) No.
Arl~ngton, Virginia 22201
tc~
12. Spon~o~ln~ Or~anft~llon N~m~ and Addns~ 13. TyD~ a~ p~DOrt 8 P~ilod Cov~nd
As above
- u.
13. SuOD~~m~nbry Not~f
, VYPl2I:VANIYC 0'LIMYKH KUL' TUit, Leningrad, 1977
1R. Ae~trset tUm~L� 200 ~ord+l
'I'his report contains a summary o~ the many years of experimental sCudies on -
ti~e physiological processes resulting in the loss of winter crops by rotting
during t{ie wintcr and early spring period. Data is analyzed concc~rning the
- growth and dcvelopment of pl~nts in ~he f:ill, winter and spring as a facCOr
affecting the wintering ability pf winter wheat and rye. Agr~me~eorological
Eactors causing the rotting of winter crops and laws effecting their seasonal
ci~anbes are also examined. MeChods are described for long-term forecasting
in various zones of the USSR.
:
17. Doeum~nt Andys~s O~u?Ipton
USSIt Crop losses Snow
Winter wl~eat, Agrometeorological factors Rotting
Winter rye Forecasting Mold
Q I~Mllfi~n/Op~n End~d Tenns
cos~n n.iaic~o~o 02D 06U 06F
_ 1~. AvaJ~Whty Sbt~m~nf 1!. S~CUntp Clau Rhif R~port) 21. No. of Ia{~s
For ~?fficial Use Only. Limited UNCLASSIFIED 1 82
Vu.~nber of Copies Availabte From JPRS. 20..~turiryClastfiA~sPiR~) 22. Pnc�
UNCLASSIFIED I
(SN AtISW9.lt) S~~ In~truetlen~ on R~v~n� OIRIONAL fdRM 272 1~-171 -
(ionn~rty NTI~3S! -
O~p~Ren~nt o/ Comm~?t� -
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JP RS L/ 8 31. 7
_ 7 March 1.9 79
PERISHING OF WINTER CRnPS -
Leningrad VYPREVANIYE OZIMYKH KUL'TUR in Russian 1977 signed _
_ to press 22 Apr 77 pp 1-168 -
[Book by F.M. Kuperman and V.A. Moiseychik, (7idrometeoizdat, -
_ 3,2~~ copies] _
COhTENTS PAGE _
- Experimental Researct~ on Physiological Processes That Cause
- Perishing of Winter Crops Under Snow 1 ~
Research on the Processes of Growth and Development o� Plants
in the Autumn-Winter-Spring Period as Factors in the Formation
of Winter-Hardy Winter Crops 31
Main Agrome*_eorological Factors Causing Perishing of Winter
Crops Under Snow and Patterns of Their Seasonal Change 87
Quantitative Evuluation of Agrometeorological Conditions cf
Winter~ng and the Condition of Winter Crops in Rer~ions With
a Deep Snow Cover 123 -
Tt~e Effects of Agrometeorological Conditions on the Gross Yiel,i
of Crain and the Area on Which Winter P1a~1ts Perish Under the
Snow 137
Methods of Comparing Long-Term Predictions of Perishing of Winter
Crops Under Snow 148
The ProbabiliCy of Perishing of Winter Crops Under the Snow in
Various Zones of the USSR.......e 166
In T~ieu of a Concl~ision 173
Bibliography 176
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pUBLICATION DATA
English title ~ PERISHTNG OF WINTER CROPS
R~ssian title � VXpREVANIYE OZIMYKH KUL'TUR
AuChor (s) � F. M. Kuperman and V. A. Moiseychik
. Editor (s) ;
:i
Publishing House � Gidrometeoizdat
Place of Publication � Leningrad
Date of Publication � 1977
Signed to press . 22 Apr 71
= Copies . 3,200
COPYRICHT . Gidrometeoizdat, 1977
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� FOK UF'~'ICIAL US~ OIJI.Y
~XPERIMENTAL RE5EARCEi ON Yi~YSIOLOGICAL PROCESSES THAT CAUSE PERISHING Or
WINTER CROPS ill~?DER SNOW
[Text] The perishing af winter crops under snow fs an eactr~~mely complex process.
It is observel when the plant3 remain for an extended time at a temperature
close to 0�C (0, �3�C) and when the soil, under a heavy snow cover, is
withouC light and is frozen to a slight depth. Under these conditions the
plants intensely expend supplies of nutritive subst~nces in the leaves
, and tillering nodes and Chey become exhausted. With a shortage of oxygen
and an excess of carbon dioxide in the plant tissues there is destruction
of Che cells, undifferenCiated, anomalous growth of the vegetative cones,
decomposition of pigments and a number of othar ~.lienomena that lead to
damage and destruction both of individual shoots and of whole plants. Plants
like these that are weakened in the winter, after the snow is gone and
below-zero temperatures return in the spring, are damaged by less severe -
frosts than they are in the autumn and winter. They are easily susceptible
to fungal diseases, the most widespread of which c-~hen the plants have been
damaged under the snow are various forms of snow mold and collar rot.
_ Damage to plants under snow is obszrved primarily in Che nonchernozetn zone -
of the RSFSR--on heavy soils with poor water penetrability. But in a number _
of years it covers large territories, including the western regions of the
USSR.
A considerablE thinning of winter plantings under an abundant snow cover
was observed as early as 1905 by V. V. Viner at the Shatilovskaya Experimental
_ Station (35j. According to his data, the deeper the snow cover, the more ~
severe the thinning.
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In the winter of 1927/28 when there was mass freezing of winter crops in the
souChern regions, in Moscow and Ivanovskaya OblasCs,'in Che Belorussian
SSIt and in a number of northern regions of the European terrieor.v of the
5ovieC lJnion, winCer plantings were severely harmed because of damage under
- snow. During this winter the snow fell on soil in which only the surface
' layer was frozen and then, under a layer of 40-50 cenCimeters of snow, the
' soil compleCely thawed ouC. The.snow remained for more than 5.5 months
Chat winrer.
- A considerable amount of destruction under an excessive anow cover ts also
' observed under mountainous conditions [46].
The ma~ority of researchers thought that the destruction of winter crops
iinder a deep layer of snow takes place either because of a shortage of
oxyg~n or because of suffocation with carbon dioxide gas that is discharged
by Che planCs.
_ In 1930/31 I. I. Tumanov and in the winters of 1931/32 and 1932/33 he in
con~unction with I. N. Eorodina and T. V. Oleynikova [34] in Pushkin
(Detskaya Selo) conducCed a large series of experiments. On the basis of
certain data concerning the content of oxygen in the dense firn snow
_ Tumanov expresse~ doubts about the possibility of the plants' destruction
because of a shortage of oxygen under the snow.
I. I. Tumanov formulated the problems in setting up his experiments in the
. following way: "First, to test the generally accepted opinion about the
destruction of winter plantings under a deep snow cover from suffocation.
Along with this, we explained the conditions for the destruction of winter
plants under snow, established the causes of the desCruction and the
phys~.ological bases of of the resistance of winter crops to the snow cover
and developed methods of evaluaCing the resistance of winter crops to -
damage under snow. In addition to past analyses, we periodically checked
the amount of damage and destruction of winter plantings under snow and
determined changes in the frost resistance of winter crops as well as the
_ content of carbohydrates." [34] `
_ Thus I. I. Tumanov outlined and implemented a large program of special
experiments for studying the perishing of plants under snow. The results -
- he obtained were verified under various conditions and augmented by many
physiologists and agrometeorologists and although by now a comparatively
large amount of new material has been accumulated, the classical experiments
of I. I. Tumanov are still of great interest. _
~ Without discussing the methodology of gas analysis of the air adopted by
I. I. Tumanov, let us note that when the oxygen content in the air above `
_ the snow is about 90.2 percent, under a snow cover of 50 centimeters three
months afCer it has fallen the oxygen content at the surface of the soii
ranged between 20.8 and 22 percent and the COZ content did not exceed 1.85
~ercent. With mor~ precise analyses the figures obtained were 02--21.6
percent and C~~--0.91 percent.
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In the winter of 1931/32 r.he e:cperiments were conducted when Che anow cover `
- remained constant at 100 cenCimeCers throughout the winter. The content of
02 grAdunlly decreased but after four months it had decreased only by 1.5
perce~;;: and was no less thun 20-25 percent. The Cemperature under the
100-centimeter snow cover during the course of four months ranged from
- 0 to f 2�C, the planCs breathed intensively and rhey consumed oxygen.
The insi~~nificant change in the (l2 conCent show that even with a 1-meter
layer of snow tihe gas eacchanfie provided ~or the vital activity of Che winter
- plnnts.
'Thus the gas analogies showed ttiat there was a sufficient quantity of
oxygen, but ~he winter plants died under these condiCions. Consequently
it was proved w3th precise experiments that the destruction of winter plants -
aC a temperature close to 0�C rakes place not from suffocation as was
previously suppc~sed before the experiments of I. I. Tumanov.
In order to exp~ain the degree to wh:ich a 50-centimeCer cover of snow
warms winter plants, the minimum tem~~eraCure of rhe air under the snow
was measured for three years (Figure 1). -
t'C -
4 -
3~ _
~ ~ r .,,i
-4 \V~'~ f ~ r
-B ~ F / v � 1V/ \ ~ ,
~I I
hj
-~z II
-~s ~
zo
i I i
-2420 30 10 ZO f !0 20 30 ,'0
l Il lll lV
Figure 1. Change of temperature under snow cover of 50 cm (1)
and 10 cm (2) and air temperature at time of observa-
Cions (3) [34]
As one can see from Figure 1, in 1931/32 the air temperature under a "
50-centimeter snow cover did not drop below -4.5�C while the temperature
under a 10-centimeter snow cover, wiCh severe frosts, dropped to -10 --16�C,
even though it was higher than the air temperature. The minimum air
temperature in the winter of 1932/33 dropped to -20 --21�C, but under a
10-15-centimeter snow cover it did not drop below -8 --10�C and under a
50-centimeter snow cover it was not lower ~han -4�C. An increase in the
layer of snow from 50 to 100 centimeters had relatively little effect on the
- . temperature conditions for the plants that were under the snow. Under
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' ~~roii c~rrrc:lAL us~ c~Nt,Y
such n layer of snow it is determined more by the freezing of the soil
before the snow cover was formed.
Very frequently when snow falls on Chawed soil Che plants spend a long
= Cime in the dar.k in an un�rozen condition gt 0�C. According Co
observaCions of N. N. Kalitin, ciCed by N. N. Yakovlev [44), 20 percent
of th~ radiation on the surface of the snow penetraCes to a depth of
10 centimeters in dry snow and about 1 percent of the radiaCion peneCrates _
to a depth of 50 cenCimeters.
K. M. Pyyklik [27j, in connecCion with reaearch of the phenomena of plant _
- destruction under snow, also studied the total radiation that passed
through the snow to various depChs. He established that while the snow
allows from 10 to 15.8 percenC of the total radiation falling o~ the ,
surface to pass through to a depth of 3 centimeters, half as much passes
through to a depth of 8 centimeters and no more than 2.3-3.4 percent to
a depth of 15 cenCimeters (Table 1). According to the data of P. P. _
- Kuz'min, as N. N. Yakovlev notes [44], even in the first 5-centimeter
layer of snow, from 34 to 88�percen~ of all the radiation penetrating
the snow cover is absorbed, depending on its density and transparency.
Table 1. Total Radiation at Various Depths Under Snow Cover
~l~ CyMM7PH~A p~aN~ui~a
('ny6NNO or nueepx� 2'' ~ ~9 18 I11
iiocti+ cHera ~
_ / r~ \ Ki 1)4 M� � YHH) I K~11 ~CM~ � 11NH~ I ~ ~A/~C4t � MNH~ qp
\L/
!la nosepx~iocr?t
~5~ ci~cra 13,0 100 38,6 100 51,5 100 -
(6)H~ ri~�6une �
- 3 c~i 1,4 10,8 6,1 15,8 6,5 12,6
15 cni 0,3 2,3 1,3 3,4 1,5 2,9
_ Key:
1. Total radiation
2. Depth from surface of snow
3. Cal/(cm2 � min)
4. On surface of snow
5. At depth of:
Autumn overmoistening of the upper layers of soil in a number of regions
oE the USSR also affects the wintering of winter plantings under a deep
snow cover--the probability of damage increases. Thus K. M. Pyyklik
- [27] notes that under the conditions of the Estonian SSR, in order to
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~ I~OIt (11~ f~ I C 1 ~I~ II~+1? c1NI,Y
dingnose Che destrucCion o~ winter plants under snow it is necessary to
- take into account the condition of Che autumn moisture co ntent of the
soil.
= N. N. Yakovlev [44] in 1937/38 beginning on 25 November, created an
excessive~snow cover to a height of 60-70 cenCimeters; before thaC the
layer of snow remained nt 7.0-25 centimeCers. Under these conditions the
' destruction of Moskovskaya 2411 winter wheat, determined at various times
from 25 November through 25 March on plots with various degrees of moisture,
= in February wiCh excessive moisture was twice as great as with normal
moisCure content. The longer Che excessive sr~ow cover remained, the
greater the destrucCion of the plants. When Che planCs remained under
- a deep snow cover for an extended pe.riod of Cime (up to 150 days) Che
destrt!ction on excessively moist soil reached 57 percenC and on naturally
_ moist soil--24 percent, when in all cases the soil was frozen by the time
the snow fell (down eo 34-38 centimeters).
Consequently, under a snow cover of 50-100 ceciCimeters with a temperaCure
t}iat does not drop below -5�C, the freezing of the plants, like suffo~ating
from a shortage of air, was ruled out in the exper:tment of N. N.
Yakovlev [44]. _
_ In order to explain the causes of the destruction af winter crops when they -
- remain under snow, it was.necessary to turn to a study of physiological
processes that take place in winter plants. It was necessary to F
investigaee the dynamics of the sugars in the first and secoz~d halves of _
= the winter, to explain the respiration energy of the plants under a snow -
cover and to take into account the importance of th~e age, stages of
development, bushiness, depth of tillering nodes and a number of other -
indicatars of the physiological conditions of the plants during the _
autumn, winter and spring periods. It was necessary to explain the con-
diCion of rest at temperatures close to 0�C and also to investi~ate the
growth of leaves under the snow, which was repeatedly noted by- many -
writers, especially the growth and differentiaCion of the vegetative -
cone. _
- After it was established that the destruction of winter crops under an -
excessive snow cover is not the result of suffocation of the plants from
~ a shortage oF oxygen and that plants perish under snow with adequate
gas exchange, I. I. Tumanov [34,35] began research on the resistance of
plants to destruction under snow depending on their condition before the
beginning of the winter and also those processes that take place in winter
' plants during the winter period. Here special attention was devot~d to the
dynamlcs of the carbohydrate content during the autumn, winter and early
spring periods. He investigated plants that were planted at various
' times, plants of various ages, plants that were well tempered and plants
that had not been tempered before the beginning of winter under the
conditions of various snow covers.
5
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AS a result of Chree years o� experiments it was esrablished thnt the
intensiveness of the expendiCure of sugars under a deep snow cover ig more
significant at a temperaCure close to 0�C than at temperatures of -5,
-8�C and lower, it also became clear Chat we11 tempered plants expend -
sugar under a deep snow cov~er considerably more "economically" than untem-
pered plants do. Under these conditions th e content of sugars in tempered
plants decreased in Che tillering nodes from 25 to 7.5 percent and in the
leaves, from 17.6 to 5 percent. while in unCempered planCs it decreased
to 2-2.5 percenC. Moreover in tempered planCs that have accumulated more
s usars since auCumn, the "critical minimum" of Che content of sugars _
(about 2-5 percent) comes later than in untempered plants. Entering the
~ wintc?r with a smaller supply of sugars, poorly tempered plants begin to -
starve under deep sn~w considerably earlier than well-tempered plants do.
In a special experiment plants that were well tempered under natural
conditions were compared with plants that were preliminarily kept for
10-12 days in a warm greenhouse where tempering decreased significantly.
Almost all of the untempered plants died (only 6-14 percent of Che plants
_ survived) while 41-52 percent of the well. tempered plants remained
unt~' spring under these cond3tions. -
The number of surviving tempered plant.s fluctuated somewhat in various
years but in all years there were significant differences in the
resistance to damage under snow between tempered and unCempered plants. -
Three years of exp~rin?ents confirm that the length of Cime the plants -
spend under the sno�:,r has a great influence on their damage. The longer ~
the period a deep snow cover remains and the later the snow disappears
f rom the plantings, the more sugars they expend and the more severely
they are exhausted anc~ thinned.
A systematic analysis of specimens of plants that hava been under a
constant snow cover of 50 and 90 centimeters showed that winCer rye did
not die off for app~oximately 2 months, but in the third month the
destruction accelerated. With untempered winter wheat plants the
destruction proceeded throughout the entire winter and a typical picture
of perishing under snow was observed.
Thus even the early experiments of Tumanov confirmed observations of
~ agrometeorologists and agronomists that perishing under snow takes place
only when the plants spend an extended period of time under deep snow
cover.
Experimental investigations of the links between the dynamics of sugars
and the perishing of winter crops under a deep snow cover have been _
conducted by many writers. K. M. Pyyklik [27J, for example, reports on
- his experiments in connection with expenditures of sugars by plants during
the winter. His data are presented in tabYe 2. From these data it is
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clrar th~tt a connec~ion existe between expendirures ok sugars and the
temperarure of the soil 1t Che depCh of the tillering nodes of rhe planrs.
'I'he mosr rapid expenditure ot sugar Cook place in Che winCers of 1955/56
and 1958/59 when [he temperaCure at the depth of the Cillering node was
almost cons~antly elevated. The ex~enditure of sugars was much less in
1956/57 and 1957/58 and 1easC of a1~. in the winter of 1959/60 when the
temperature was the lowesC.
Table 2. Expenditure of Sugars in Winter Rye in the Winter Period
2 Ni~rc eon~ ~Pe~~iuit cyta~m~n pacso.t
~ P casapon ~uMOO (~~r ea t r
~1~ npeobneaa~n. 3 cyxoro oeutecrea)~
fOAM 41H:f TCMIICPB� -
. ryp ~~a r.~y0~mc
yane (~y~jtc~u~n o.i~~cr~nx B Y3.70X KyLLIC� -
~ C ~ HHq
.J
1955~Sfi 0, - 1 1,65 2,16
, 195t~57 -O,S, - 2 1,10 1,53
1957/58 D, - 3 1,01 1,60
~ 19SSJ59 0, - 1 1,G6 ?,58 -
_ 1959~60 --2, -5 0,90 O,~JB -
Key :
1. Years
2. Intervals of predominant temper~tures at the depth of the tillering `
node (�C)
3. Average daily expendiCure of sugars in the winter (mg per 1 g
- of dry substance)
4. In the leaves
5. In the tillering nodes
~ 1From the beginning of wintering until the period of intensive snow melting
in the spring.
Taking into account the fact that one cannot create a snow cover of the ~
necessary depth everywhere or every year, in 1935 T. I. Tumanov [34]
suggested that when evaluating various strains of wiiiter crops for
resistance to destruction under snow, experiments be conducted in dark
thermostatic rooms and that the boxes of plants be covered with a layer of.
wet sawdust instead of snow. This way a temperature close to Q�C is
easily maintained for a long period of ~ime at the level of the plants
and if necessary it car~ be regulated. Wet sawdust makes it possible to
maintain the moisture of the air at the point close to saturation as it
is in natural conditions under a deep snow cover. The results obtained -
in the experiments with wet sawdust corresponded to the results of field
experiments under snow. _
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_ t
'Che condi~ion of the soiZ during ehe winCer--thawed or frozen--heg n
gr~gt influence on the expenditure of sugars. ihawed eoil ghowe thn~
under the gnow th~ CemperaCure was 0�C or ~ lit�tl~ higher a11 the Cin~.
Th~ exp~ndit�ure of srored eubetances in Che planCs ig especially greaC
under n deep ~now cover when Che soil is th~wed.
In Che winter of 1955/56 aC th~ experimenCal basis in Kuuziku on one -
plot of winter ryp n eh~wed conditian of Che ~oil was arCifically creaC~d
~ (by regclation of Che snow cover) and on another--a frozen cottdition. The
difference in the CenperaCure of the soil on the p1oCs was insignificant--
an average of 0.5-1�C. The difference in ehe expenditure of gugars by the ~
pl~nts th~t apene the winCer in thawed and frnzen aoil, both in the 1F,~ves
and in ~:he tillering nodes, reach~d 40 percene. Considerably more planCg
died in the thgwed soi1.
Moreover, during the course of three wintera (from 1959 through 1962)
oth~r experiments were conducted in Kuuziku. In order not to allow
fre~r.ing of the soil ~n the winCer rye field, at the beginning of wintering
o,~e plot was covered with a layer of sawdust 20 centimeters thick.
In the winter of 1959/1960, as a result of an exceptionally severe freeze
in Uecember, the soil even under sawdust froze slightly (3-4 centimetera),
but in th~ secon~ half of Che winter it thawed. At Che e.nd of the anow
thaw (20 Aprll) one-fourth of the plants under the sawdust had died and _
in Che le:~ves of the reroaining planta there was 2-5 percent sugars and
in the tillering nodes--7-9 percent.
In 1960/61 in the winter rye under sawduat even by 16 January the supply
of sugars had decreased to 7 percent in the l.eaves and to 12 percent in
the tillering nodes. A significant decline ot the plants under sawdust
began at the end of February. In the beginning of March in the leaves of
the live plants there was 3-4 percent sugars and in the tillering nodes--
7-9 percent. By Che middle cf April all Che plants had died.
Thus in thawed soil, that is, at a temperature of 0�C and higher, the
_ expenditure of sugars takes place 2-4 times more rapidly than in frozen
soil and after winter crops have been in thawed soil for 2-3 months tl~ey ~
begin to die--slowly at first and in the second l~alf of March and the first
halE of April, in large numbers.
One of the main reasons for the intensive expenditure of sugars under a _
deep snow cover is the accelerated expenditure of sugars for respiration.
V. A. Moiseychik [Z2], utilizing the dzta of N. V. Savinskiy, obtained
at the Sobakino Agrometeorological Station, established that the expenditures
in winter rye wit}i a reduced temper~ture at the depth of the tillering
node from 6.0 to 1.5�C increased 1.6-fold when the initial supplies of
sugar were 10-18 per;.ent. In plants with supplies of sugars of 26-28
percent in the middle of the winter the same increase in the expenditure
of sugars was observed with a considerably lesser increase in the temperature
at the depth of the tillering node (from 7 to 5�C).
8
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F'Ok UI~F'ICIAL USI: ONLY
- V, A. rtoi~eychik [22~, like other rese~rcher~ l,tttez', e~t~nbliahed rh.~C
~ large ~upply nf gugnry in wintex crops wiCh ~ more developed gr.~s:~
~ ~tand ig E~xpenci~d nn ~egpir~Cion more r~~pidly (subgequently it bec~me
ciear thnt thig ig ~71so true df ~rowth proceK~e~). 11~is i~ on~ o~ the
re~r~on:~ tor the gre~ht thinning when overgrown p1~nCed ~~r~hs 5pend Che
wintcr under snow~ I. L. 'Tumanov thinks that we~~kly tempered pt~nts
expend gugnrs congiderably more rapidly ae th~ snme temp~r~eure~ (2-~
times). I. M. perunin [~3] using the formul~ for respir~tion
C~H120~ + COZ 6C02 6N20 (1) -
~7nd the ~Comic weighCa of elerneties tti~t comprige thi~ frrmula, determined
the ~nmount df Chc ratio between disch~rged cnrbnn dioxide and expended
~ugnr (pluCOSe). By dividing ttie weights of glucose molecules by the -
weight oL the carbon dioxide digcharged duri.ng reepiration, he esCablished
that i?~ tc~rms oi' weight the sugnr expended during respir~~Cion is 0.682 -
the weight of the discharged cttrbon dioxide. Tak.ing into account the
dependen~y oF the c{u~ntity of carbon d,~oxide discharged during the process
of respir~~tion on the temperature, he calcul~Ced the amount of expenditure
oE sgunrs by the plant on respiraCion during a day (in milligrams) per ,
1~rnm of dry substance (Table 3).
Tab1e 3. ~xpendieure of Sugar~ by Well-Developed and Tempered Plants of
Winter Crops on Respiration Under a 5now Cnver During Qne Day,
Depending on the TemperaCure
- (1) ?eWnepat~�pa, �C ? 6 5 4 3 2 1 0 --1
~2~ F~arxoa ca~apoo,
~~r . . . , . . 9,86 ~J,O7 $,30 ?,56 6,91 G,31 5,74 5,23 ~,73
(1) Tc~inepaT}�pa. �C -2 -3 -5 -6 -7 -8 -9 -10
_ (2~ Par~oa caxapoe,
- Mr 4,?7 3,84 3,43 3,05 2,66 2,30 1,97 1,66 1,37
Key:
1. 'Cemperature, �C
'l. ~xpenditure oE sugars, mg
'I'he perisliing of plants does not begin immediately upon the expenditure
of supplies of sugar, but considerably later, since the plants are -
~apable oC augmentinq them to a certain degree by transforming a large
supply of starch into sugars. But then plant starvation takes place
(expenditure of proteins and decomposition of the tissu~s of zhe plant
cells) and a second phase of perishing of winter crops under snow begins.
'Ct~e pl.7nts begin to expend protein when they have only 2-4 percent sugars
9
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~oK d~rt~rnt, ~isc drri,v
left. 'Thig tnkeg place usu~lly ~t th~ end of ehe wint~r nnd during rhe
period of inelCing of thQ snow. The expenditure of proteins ig d~ngeroue _
~or ehe liLe of ehe planes (i19b becnuee Che he~t dtgch~rged in thig proe~sg
r_rpaCp~ fnvor~ble ~ondiCions for ehe development of miCroorggnismg gnd
the growth of mycelin of various fungi. Th~ laCt~r, growin$ rgpidly attd
powprfully on ehe sCgrv3ng planCg, shnrply accelerate ehe exp~ndieur~ of
prntein, whi~h leads eo degeh firge of ehe leaveg eh~e eouch rh~ goil and
ehen the boCCnm9 0~ the shnotg gnd sub~equpnely ~1ec~ ehe eillering node~
of winter Crops. The longer. Che period of starvation rhe more the pl~nts
are dam~ged from being under the Anow. The resulte of an gnelyeis nf
� m~terigls from mgny years of observaeions hgve made it po~gible for V. A. -
Mnigeychik Co come Co Che conclusion that all or Che ma~ority of planCs _
die wh~n ehe period of ~tarvation is no leas rhgn 30-40 days gnd Che
~ meltin~ nf snow e~kes a long time (22].
f3ut, as I. I. Tumanov made Clear as enrly es 1935, the exhnugtion of
_ plants doeg not directZy CgUBe their death. Thus in special experir.~enl�s
in cert~in variants the leaveg of winCer planCs were cut and removed
before they were covered with ~now--onty nne tillering node and the rooe
system went through the winter. In ehis variane 85 percent of the plants
survived ~nd produced aftergrok*_h. In the second variant when the above-
ground part was Cut o~f at the same ticne but nc~ removed and left lying
on the plot, only 33 percent of the planCs survived. The exhaustiott of
the winter plants was the same but in places where Che leaves were not cut
and also in places where the cut leaves remained on the plot, snow mold ~
developed. On the plots wher~e Che leaves were removed there was no snow
mold.
It is probably no accident that when there has been abundant growing of
cain[~r crops in the autumn farmers have grazed animals on areas plan[ed
in winter crops before the beginning of winter. And although the grazing
of animals on winter plantings also caused damage to the plants, it was
apparently les~ than the damage from apending the winter under the snow.
There nre interesting experimenta with cutting back the plants in the autumn
under the conditions of three backgrounds--a natural snow cover, an
excessive snow cover and without snow [44~. The plants were cut back at -
the end oE 5eptember to a height of 10 centimeters and the mass that was
cut was removed. On the background of a naGura~ snow cover there were
no essential differences in the wint~ring; when there was no snow the
tillering nodes not ~overed by leaves suffered more; and under conditions
of an excessive snow cover the removal of the cut mass--the nutritive
environment for the development of fungi--led to a reduction of the death
of plants from spending the winter under snow (Table 4).
The mas~ appearance of snow mold on the planted areas usually coincides
with the end of winter when the plan[s are already weak and exhausted and
When, under the influence of starvation, chere is an intet~sive decomposition
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.
~OE~ O~~ICIAL USL ONL~'
of p;nCein~ end the qunnCiey of nitrogen compo~irid~ necessnry for Che
Jc~velopm~ene of gnow mold inrreases. Moreover the amaller Che mass df
. leaves, the le~g the fungi develop, which is clear from experimeneg wieh
plgnGg ChnC hnve be~n cuC bnck.
Tgble 4. Deneh of Plgr~ee that were CuC Bnck and Noe Cue gack in the
~n1L (9~)
_~2~ CneMnWn nacpoe I
~1~ - I~etene~be
_ ~3) ecreerreHaaa ~ ~~3)iuauTOV~~dn (5)
l~1ltT0 II~A,iN~a!lINII (1~CTQHNM
N 1'OS of161f~
noapraur neno~pe� noape�u� neno.tpe� noapcaau� ueno~pc�
NyC~~ 1HNC ur r~,iNbe I NY~ uuue
l~l ~
~9~Korrpo~ia, 1938 ~H,1 17,R 14,5 42,G fiG,4 40,8
~ld)Oniunio, Inan 74,~'i 24,8 30,0 36,4 ?2,? 64,G
] t)fl~�iuknN, I'.~54 3'l,8 Y6,fl 22,5 '~T,e 55,5 30,9
Key:
1. Place of observuCions and year of experimenC
2. Snow cover
3. Nururxl 8. NAt cut back
4. ~xcessive 9. Kostroma, 1938
~ 5. No snow 10. Ovtsino, 1940
6. P1anCe 11. Pushkin, 1959
_ 7. Cut back
'I'hus it has been established that starving creates the necessary
preconditions and ~now mold kills the plants under these conditions. Under
conditions favorable for the development of mycelia of fungi (temperature
under the snow--about 0�C and higher, air humidity--about 90 percent), -
the destruction of the plants begins within several days.
The experimental data obtained by I. I. Tumanov [34~ enabled him to eingle
out tl~ree qualitatively different phases depending on the physiological
processes predominating during the course of perishing of the plants:
' first--c~~rbohydrate exhaustion, second--starvation and disintegration of
organic substances, and third--Che death of the plants with the development
of fungal diseases. _
As has already been confirmed by other researchers, the first phase of
perishing is characterizE~d by hydrocarbon exhaustion of the plants. When
the temperature under the snow is close to 0�C the winter plants maintain
a marked respiration energy and the process of growth of the vegetative
cones and root system continues almost continuously, although weakly.
Each day ~ven weak (especially with temperatures somewhat above freezing)
expenditures oE sugars on respiration and growth during the course of an
extended period under the snoW cover are accumulated and finally lead to _
an almost complete exhauation of the plants.
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~
I
~Oit nI~~ICIAL U5L ONLY
'fhe procese of exhaugCion of Che plgnts continue~ for 2-3 monChs. 'The
_ expendieure of ~ugars increae~s e~peci~lly aC Ch~ b~ginning of Mnrch, wiCh
ehe excepCion of ehose years when ehe Cempera~ure of March drops sharpLy
- and the soil freezee to a deeper level.
CxpendiCure of sugars in witteer crops incre~ses shgrply, which was
- repenredly observed by K. M. Pyyklik [27] during t~te period of inCeneiv~
~ndw melting, egpeci~lly ae the end of this period (up ro 5 milligrams and
more per 1 gram of dry gubstance). The periahing of planCg increases
sharply during Chis period 5-6 days before ehe snow is gone, which frequenely
h~s a decigive significance for the plants' survival ehrough the winCer.
Thus K. M. Pyyklik in 1951 compared ehe reeules of an analysis of apecimens
of wineer rye planted at an optimum time which were taken on 12 and 20 ~
Mnrch at the time of intensiue snow melting on the fields. During these
g days the quantiey of sugars in the tillering nodes decregsed from 20.1
to 13.2 percent and in the leaves--from 12.8 to 6.5 percent, which amounCed `
Co ~pproximately 8-9 milligrams a day. In Che s~bsequent days the content
of s~gars dropped to 3-4 percenC.
'The works of ather writers also confir~ned that the most active expenditure _
of eugars takes place under conditions of inelting snow where the temperature .
remains sCable at nbout 0�C and the high moisture content contributes to the
rapid hydroliais of sugars in the damaged plant Cissues. The quantity of
sugars in the leaves and especially in the tillering nodes before enCering
the winter is of great significance for Che aurvival of winter plants under
a deep snow cover.
The First experimenCs of I. I. Tumanov [34, 35] showed that tempered plants
that accumulate a high percentage of sugars in the autumn remain longer and
- the second phase, starvation, begins laCer in them. Therefore one should
discuss the factors that determine the degree of autumn tempering of plants
under the conditions of the nonchernozem zone.
As we know, according to the theory of tempering developed by phyaiologists,
a considerable increase in the content of sugars in the leaves and
especially in the tillering nodes is a result of the fact thaC in the
- autumn during sunny days the processes of photosynthesis proceed relatively
intensively in wheat plants and in the evening, night and morning hours when
the temperature is reduced their processes of respiration and growth slow
down, which leads to the accumulation of sugars. The longer the transition
period from high temperatures in sutumn to lower temperatures in the winter
and the more optimal these temperatures are for tempering, the more inten- `
sively the process of accumulation of sugars in the tillering nodes takes
place.
Table 5 gives figures from an ^xperiment during which, after remaining
' at a temperature of 5�C for 14 days, the plants withstood from 2 to 5 days
at temperatures oE -2, -5 and e7�C. From these data it is clear that the
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I~OIt i)I~i~'1:(;tAL U;;1. ONI~Y
m~ximum monosug~r~ were ~ccumul~Ced in Che vari~nC with 14 dayx et n
tempergCur~ n~ 5�C plus 2 days with -2�C ~nd the largest number of di-sugare
was ~ccumul~Ced in the variant with 14 days ~t 5�C plus 2 dnys aC -7�C.
Tuble 5. Accumulation af Sugarg in Vgrinu~ 5Crains of Winrer Wheat
of Dry SubgC~nce)
~1~ tl ineA npa b' C
~o~ OHipO.ib t~ lna npH ~}S AHCR IipN ~}7 AHp flpN
Copt ~2~ ~3~ -t C (4~__,,, (5)
NUIIO� ~II� ?IONO� dH� AI011o� AN� MOIIO- All�
Yy ~ I � p~ ~ ~p~ I~~ p~ e~~ ~ I uxop~ e~x~p~ az~p~
d s
~g ~ MHH%8p~11 22,4 28,3 25,8 Y6,9 19,8 3A,1 17,7 38,"
~10~ ~HCtCKe ~1~9 ~3~n ~s~~ ~5~~ 25~~ ~~~6 7$~i
11 pii~inav `12,u 4,1 17,1 13,5 11,7 22.0 15.'l 24,7
- (12)Wr)~Ge '.1i~KKOn~~ 12,1 3,a 20,7 4,7 11,6 18,4 17,4 15,5
Key :
1. 14 days at 5�C 7. Di-s~igar
2. Control 8. Strain
3. + 2 days at -2�C 9. Minkhardi _
4. + 5 days at -5�C 10. YaneCska
5. + 2 days at -7�C 11. Rimpan
6. Mono-sugar 12. Shtube Dikkopf _
One of the essential facCors thaC provide for good Cempering of the plants
and accumulation of sugars is optimal planting times that correspond to _
' the best agro-meteorological conditions for the growth of the plants and
Che strain peculiarities of winter crops [2, 7, 13, 16, 22, 27, 25, 29J.
K. M. Pyyklik [27] notes thaC winter crops planted at Che optimal time
accumulate in the tillering nodes an average of 10-15 percent more and
in certain years even 20-30 percent more than plants planted at laCer times. _
'Che period oE autumn tempering of winter plants and the accumulation of
sugars is usually observed when the average daily tempering of the air
steadily decreases to 8-5�C with a large daily temperaCure range [22J.
Observations of the dynamic accumulation of sugars in 1932/33 at the
Khar'kovskaya experimental station in such strains of winter wheat as
Dyurabl', Ferrugineum 1239, Gostianum 237 and Ukrainka showed that most of
the sugars were in the tillering nodes; up to 30-45 percent nf ~hem were
accumulated under conditions when the temperature for 13-16 hours reached
15-20�C and at r.ight dropped to 2-3�C [7j. Summary data are also given
~ for the Estonian SSR. Thus in the autumn of 1961 during the third ten
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~Olt 0~'F~'YCIAL US1. ONLY
dny~ df Sept~mber nnd the firsti five dnyy of OcCober Chere wag dry sonny
wenther, the ~verage d~ily Cpmperarure was $-9�C, Che muximum--13-17 C and
ehe minimuml-- -1 -*3�G. 13y Che end o� this period ehe winter planCs
' had nccumulated abouC ~0 percent of Ch~ sugar~.
tf ehe avergge dnily remperaCure rises Co above 9-10�C or Chere is
considerable cloudiness, Che accumul.~eion of sugars ia retnrded and in a
number of cases the quanCiCy of already gccumulated sugars, especially in _
ehe leaves, decregses considerably. With comparaCively low temperaeures
(average daily--3-4�C), Che accumulation of sugara is already slowing up =
even ehough Chis prncess conCinues right up to the Cime when the average -
daily eemperaCures are below 0. K. M. Pyyklik [27j tn his work gives a
description of the condiCions for the tiempering of winter rye and the
' cnurse of accumulation of sugars for a number of years (Table 6, 7).
When analyzing days Chat are favorable or unfavorabl~e for tempering, the
course of them and their distribution throughout the auCumn is significant.
If ~lear sunny days predominaCe over overcasC ones ChroughouC the enCire _
autumn, Che process of Cempering proceeds almost continually unCil the end
of the growing period. If after the days that are favorable for Cempering ~
- there ~ppear overcast days and the temperature increases, frequenely
the accumulated supplies of substar?ces are used by Che plants for growth
of the organs and the sugar content decreases.
It has been established that about 90 percenC of the sugars in the tillering
nodes in the tempered plants are localized by buriers that are difficult
to penetrate. This property of tempered plaaCs keeps the sugar firmly in
ur.damaged cells and is possibly one of the reasons for the reduced losa of
sugars under conditions where Che Cemperature is close to 0�C, which in
itself does not damage ehe cells, especially in the first half of the
winter. _
The dynnmics of hydrocarbons can show the degree of the plants' readiness
for wintering. But only after the temperaturea begin to drop below zero
can one determine the content of sugars wiCh which the winter crops will
enter the winter. When the temperatures drop below zero before the snow
falls in cold winters (especially in the first half of'the winCer), the
content of sucroses continues to increase and the more winter hardy the
strain, Ch~ higher it is. During warm winters the proportion of simple _
forms of sugars increasesr above all fructoses [31].
Many writers have engaged in the development of inethods for diagnosing
the condition of winter wheat in terms of the hydrocarbon content. Thus
IV, I. Goysa, R. V. Gatsenko and I. I. Kovtun in 1975 published the results ~
of comprehensive research during 1967-1971. They Cested strains of the
Mironavskaya selection--Mironovskaya 808, Mironovskaya Yubileynaya,
Lyutestsens 2488, 2917, 2877 and also BeZOStaya 1 and Odesskaya 16.
14
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Tnble 6. Connection I~etween the Conditions for Tempering SangnsCe Wineer
_ Kye P1nnCed ae the Optimal Time and thr QuanCity of iug~?r~ aC
Che ~nd of ehe Tempering p~riod
(1) c~~ ~ ~~~~ek- ~ ~ ~'r.~oeun aaKa.iNBONNp C~;,tn~~eC;~,cat~poo
T,+n~iNx reMne� 3 -
rux p�ryp or nocee~ C 4NCeo aHeR ~ rq
~o npeKp~ute� ~ (8 10 (11
unp eererwNU ` o ne~e. bb~HK~ ~ ~ e ya~t~ex
(-C) 6) 7) C4lt0 H~ NONLLB B JIHCTbNx g pllHXY
xo ottuHx+ c~eA(int xopowHe nepNO~i~ y
1955 312 4 A 8 3 I 76 35
1956 146 4 6 7 3 1`l 32
1957 192 5 H 9 3 21 31
195A ?5`l 7 11 13 4 29 45
1959 151 5 G 8 3 17 25
1~J(,~ I 9y 0 2 I 2 12 20 -
~ _ .
Key: -
1. Year
2. Total effective temperatures from planCing time unCil end of
~ growing period '
- 3. Tempering conditions
4. Quantity of sugars (9~)
5. Number of daysl
6. Good2
7. Average
� 8. F.xpressed in Cerms of good ones
9. Evaluation at end of period3
10. In leaves
11. In tillering nodes
lA "good" day is expressed as 1 and an "average" day, 1/2.
- � 2A good day--clear, sunny, not cloudy weather; average--slightly cloudy or
_ with variable cloudiness
3Evaluation made with 5-point system: 1--very poor conditions for tempering;
S--excellent conditions.
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I
~OFt 01~~~CTAL US~ ONLY
Table 7. ConnecCion Beeween Che Cottdition~ for 'Tempering Sangaste Winter
Rye Planted gC Che OpCim~1 Time and ehe QuanCiey of 5ugars in
Che Tillering Nodes
4NC.~o zopowur aueil OueuKO yctaoull Ko.tNVectao coxipos
/y `A.iq ~~K~.iNtllHNN JOIt/~1NB81111q 8 \'JABk N)'Illtlllip
\ii \ /
-
~2 (4)Xopowue ~25
8-12 (5}CPent~~~e 1~ 35
s ~6~ n,~ox~ic
Key :
1. Number of good days for tempering
2. EvaluaCion of tempering conditions
3. Quantity of sugars in Cillering nodea
4. Good
5. Average
6. Poor
It was established that Che maximum quantity of Cotal sugars accumulaCed in
the autumn period in the tillering nodes of winter whpat changes little
fr~m strain to strain and amounts to 38-44 percent of the weight of the
dry sub~tance. There was no connection between the quantity of plants that
' live through the winter and the total sugars (R = 0.06). Thus the maximum
_ content of sugars in the tillering nodes in the early autumn period does
- not serve as an indicator of the level of frost resistance of the plants.
'The authors revealed a close connection between the quantity of planCs Chat
wiChstood the winter and the sum of sugars remaining in the Cillering nodes
- at the end of wintering (R = 0.74). A reduction in the content of sugars ~
is accompanied by a marked drop in the number of planrs that withstood
the winter. If one can understand the connection between the minimum
cantent of sugars in the tillering nodes and the yield of grain of winCer
wheat (R = 0.65).
The writers think that the amo~mt of the minimum content of sugars in the
tillering nodes ~n the early spring can be recommended as a biologfcal
_ indicator for diagnosing the condition of the planted areas after wintering
and, perhaps, can be utilized as a prognisticatory indicator of the harvest.
In the authors` opinion, the advantage of this method of diagnosing the
condition of the plants in the spring as compared to other widely known ~
methods (sprouting the plants in core samples of soil, freezing the plants
- in refrigerated chamgers and so forth) is that it makes it possible to
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~OR Ori~'ICYAL USL ONI~Y
obtaitt in prnceice n simple (nvertt~e for a given field) And opernCional
idea oF,Clie relativ~ely poseible Chinning after winrering and also Co _
determind'the need fox replanting 1..5-2 weeks before Che beginning o� the
- spring growing period.
But what is the nature of the procESa of eempering and the accumulation of
sugars in pl~nts ChaC are planCed ~C varioua timea or, more precisely,
planCe of varioue ages?
A study o� the effecCs of Che planting rimes and winter warm periods on Che
dynamics of ehe sugaxs was made by F. M. Kuperman in 1935 under the
conditions of ehe Khar'kov experimental station for several years [7].
Analyses showed thaC when there was an overall drop in Che contenC of sugars
in r.he planCs under the snow, afCer the snow was gone during an extended
Ch~wing period when the temperatur~s were above zero during the day
~ (even 3-10�C) the planes are capable of carrying out photosynthesis. 'The
content os yugars in them increases.
Many researchers have obtained similar data in the spring of years when
winter plants are relieved of the snow cover early and conditions are
established that are favorable for photosynthesis. In these years snow _
mold does not manage to develop in the plants, despite the facC Chat they
are weak and exhausted from the lengthy expenditure of hydrocarbons under -
the deep snow cover, still survive and be~in to grow. _
K. M. Pyyklik [27J each year compared Che condition of winter plants
thaC came out from under Che snow and those that were still under it. In -
the planCs that came out from under the snow and were in the light for
from 2 to 5 days there was 80 percent more sugars in the leaves and 60
percent more in the tillering nodes than in plants that were still under
the snow [Table 8].
- Early in the spring when there is a considerable range of day and night
temperatures the plants can again be tempered and therefore withstand
the return of frosts. One can 3udge the content of sugars in the leaves
and tillering nodes in the early spring from the data in Table 9.
Cven in his early works I. I. Tumanov [34] published figures on strain
difEerences in winter crops in terms of their resistance to perishing
under snow [Table 10].
N. N. Yakovlev (44] established the various resistances to perishing under
snow of strains of winter crops with various origins. He noted a direct
connection between the resistances of plants and the ecological-geographical
conclitions of their origin.
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rok c~i~i~�~.c;rn~, crsi: c~N~.~Y =
Ts~ble 8. Qualiey of Sugars in Winter Rye in Kuuz~.ku in the Spring
oE Dry SubsCance)
~ 2 ~ perw onpeAeneeNn c~a~poe
M ecrouo.~o*cuee p~crenHp
_ ~ Z~ 12 I11 I 11 !11 I 20 III I 41 111
~ ~ CIOA CNCPODI _
_ ) B !11{CTbAX 11,9 ` 6,5 6,2
S) n~�3nax Kyu~etiNa 13,2 12,6
( ~ Ha ceery
B lIIICTbqX 20,5 21,0 17,3 20,7
5 o y3nax KpLLICNNA - - 16,5 16,9
Key:
1. I,ocaCion of plants 4. I~ leaves
2. Dates o.f ~:easuring of sugars 5. In tillering nodes -
3. Under snow 6. In the light
Tab 1e 9. Content of ~ugars in WinCer Plants in the State of Stem Extension
After Early llisappearance of Snow
CatepMaHHe c~xapos
(1) (2) 3~96 ot cyau~o eec~?
C poK eee~ COCTONNN! PICTCNNR Q~HH!(1 eecNOR .
s aNCreak � Y~~~=
r 1jxJ1(lHNA
~
(6) OnrHn~anbHwti i'naetiwe noGeri~ (Tp)'6tca) 33,0 35,9
_ ~ 36oKOdae noGeni (rpy6K?i Her) 31,5
~1 1P8CTCHIIA, KOTOPW7~ He Gd10 .
~ 7P}'GK002HNA 32,7 ~,5
I~I03AHNF1 ~j CTEHNA B ~183e KYUleHNA
ly~TP)'GK088HNA HfT~ ~~2 28r3
Key:
1. Planting time
2. Condition of plants in early spring
- 3. Content of sugars of dry substance)
4. In leaves
5. In tillering nodes
6. Optimal
- 7. Late
8. Main shoots (tubule)
- 9. Side shoots (no tubule)
10. Plants in which there has been no stem Qxtension
11. Plants in the tillering stage (no stem extension)
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~ roa c~t~T~'iCTAL USL ONi~v
T~ble 10. Description of WinCering of Plants Under Deep Snow
CpoKU ceaa
~Incao AHe11 aponeaa� 2, VIII 5 IX 25 IR
UNN CeMnll 0 ~0 13 0 10 "0 ~0 43 0 10 20 10 ~.5
I I I I I I� I I I I I I' I
(3) 03F+n+aA nweiu+un
T+~� JibTecucf~c 0329 37 - 20 18 2 61 - 38 32 13 100 - lOQ 59 67
I~tOCK00CKOq O24II 39 1~ ~ - ~ 3 2 - 3~ 12 0
3{IAIOq pON~b
BAr?ca 85 61 65 50 18 78 61 14 15 - -
Key :
1. Number of days of vernalization of seeds
2. P1anCing Cimes
3. Winrer wheat
4. Lyutestsens 0329
5. Moskovskaya 02411
- 6. Winter rye
- 7. Vyatka
There is a lower mortality rate in planCs from countries where winter crops -
are frequently sub~ected to damage under deep snow covers than in plants
which rarely spend the winter under an abundant snow cover. Thus the
fewesC specimens died in the group obtained from Sweden is frequently
combined with warm winters; strains that are resistant to damage under the
- snow have also been formed in the mountainous regions of Austria. -
On the b asis of experiments with an excessive snow cover which were
conducted in the All-Union InsCitute of Crop Growing (City of Pushkin near
Leningrad) in a year typical for damage to crops under the snow cover,
N. N. Yakovlev [44] gives the information which we are printin~ in table -
11 .
- From these data it is clear that Che strains of wheat from regions where
~ there are rarely conditions which lead to damage to the plants under the -
snow (Central Asian, Transcaucasian, Southern Steppe and Southern Forest
5teppe ecological groups) are less resistant to damage under the snow than
strains of wheat from the Northern Russian and Forest Steppe Volga _
ecological groups.
- Interesting data concerning the content of sugars in winter and s~ring
crops and their destruction after wintering in the Estonian SSR (Kuuziku
Agrometeorological Station) in the winter of 1957/58 are given by K. M.
- Pyyklik (Table 12).
- 19
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~OR Ol~i~'ICIAL US~ ONLY
- I~'rom r~ble 12 iC is clear ChaC Sangas re ~nd Priyeku:Li wineer ryes ~ccumul~~Ced
approximaCely the same qu~nCitiy of sugars in Che ti],~.ering nodes, but
SangasCe hnd 1~1 percent mora sugara in the leaves. During rhe Cime of
wintering Priyekuli expended relatively more sugars and approximately
twice as much of ir was desCroyed. F'ive sCrains of wineer wheat were
rested. Puuk and Kuuzilcu accumulated more sug~rs Chan rhe others--gn
average of 20 percent more than the st~rains of Ci~e second group--Luun'ya _
Uluchshennnya and Gibrid 599. The results of the wintering showed rhaC
an average of 30-40 percenC more plantia of the second group died.
Table 11. Resistance of Strains of Winter Wheat Co Perishin,g Under Snow
(poinCs) (Pushkin, 1965)
- 9-10 5-6 _
Lyutestsens 116 Ukrainka -
Moskovskaya 2411 Odesskaya 3 ~
Kolkhoznitsa Stavropol'ka 328
Priyekul'skaya Voroshilovskaya
Borovichskaya
Kuuziku
Luun'ya
8-9 4-5
U1'yanovka Novokrymlca 204
Sandomirka Zemka 4
LyutesCsens 329 KooperaCorka
7-8 3-~+
- Lyutestsens 1060/10 Grekum 433
Eritrospermum 15 Dolis-Puri
6-7 1-3
Lyutestsens 17 Shark
Lesostepka 74 Surkhak _
Ferrugineum 1239 Irody
Eritrospermum 917
- In addition to the strains mentioned above, Mandorf winter barley and
Kauka and Diamant spring wheat were also tested. The barley contained
approximately the same amount of sugar both in the leaves and in the
tillering nodes as Kauka spring wheat but 60 percent of the barley
plants died and all of the plants of both strains of spring wheat died.
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' rOR Oi'P'ICIAL USC UNLY -
: .
I ~ a
~ ~ s ~ ~ N N ~Q7' M ~I N t0 0~0 O
M K
~ _ ~
~
d ~ ~ ~ ~ ~ ~ 8
~ � Q ~ R ~ -
~ ' '
~ - -
Q) p
A ~ a C o~ r~ o 0o t.
~ ~ R ~ ~ ~ ~ M ~ I I -
~ a
~t.. a ~ t. eo 00 ~c o
" ~ ~ ~ ~c ~ ~ ~ i I I -
~y v Y
~ ~Y a
u m ~ s 0 O ~ N 00 C~I o0 u~ M
' a = ~ c R ~ c~ ~ .Qi c�~ ~ ~
O ' x
~ 6
C,) u ~ ,^r ~ a
~ U A~' R= G~ O N h ~O O~ ~Y' a0 h
_ � v ~ c") M e'~ N CV N ~ ~ ~
~ o
- ~ v b� , F N Ci C~. C`I t0 ~ C': t~
~ C~~ ? _ o oi ai o rn o 0o i: ~c u~
_ d Y u n~ r"{
'PI Y O Y ^ ~C
~ ~0 A V 'j t~ A
O
N E ~ n" 3 s : t c`~ ~ ~O Cf Q1
a
o~_ x~
o~~ o I~ ~T ..c.~ ~ a0 O t~ G~ C ~C ~C >C ~C X >C
U u
_ w ~ ~O h c0 t~ h t~ v~ eM h
O CO ^
- ~
L ~r{
~ ~
41 QI
_ ~J 1J
C~. o ~ ~7 ~ eO ~0
~.r.{ ~ ~ S 7 7 7 ~ 7
r ~ ~ ~ T ~ y ~
~ p"~ _ G!"' C G. ~ G~- a A~ l~ ? ' 21 7 F -
~ v n �t ~ = _ t = ~ ~ G '~J~ C 7
N G L 0~ Y ~ G ^ = - - -
s r. ~ - i,= - r~3 a Q :
~c:~TFCnrr~i y,n~'~_^,$7'.J~=
~ O V O~= O C O~=_:d L, a,=~ i at~i
~ O O F O G
~ ~ ~ ~ ~ ~ ~ ~ ~ ~
N M ~Y C
H ~--I r-I r-1 ~-~-I r~-1 ~--1 r~-~ r~-1 N
v v v v v ~ v v v
21
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7
roH o~~tctn~ ct~~ oNLY
K~y:
l. Crop gnd ~tr~in 10~ Le~v~eg
2. pl~nring Cim~ 11. p1gnC~ -
3. SCnge of developmenC ~t ~nd 12. ~~ngtt~Ce wintpr ry~
_ of growing periodl 1~. p~uk winter whe~e
- 4. Contenr of dry gub~tnnc~ 14, pri'yekuli wint~r rye
in c~11 ~~p 15. tuun'y~ Ulu~hghenn~y~ wint~r
5. Contene o~ gugarg (9~ df wh~ae
dry gubgC~nce) 16. Kuuziku wine~r whegC
6. Deneh by the end of 1'l. Gibrfd 59~ winter wheat
wineering (9~) ' 18. Kauka gpring whe~t
At beginning o~ winC~r 19. M~nford winter barley
8. At end of winter 20. biatt~nt gpring whe~t
9. Tillering nodeg
lIn the tillering sCage only the number nf ghnz~tg i~ indic~e~d.
Erom the results of the determinaCinn of sug~rs it b~~gme cle~r that
at the time c?f intensive growth of the plnntg in the gutumn they Contgined
3-9 percent of the suggrs in Che leaveg end 7-12 percent in the till~ring
nodes. From the beginning of the periad of tempering Ch~ overall quantiCy
of sugars gradually incre~se~s both in the lexves and in the tillering
nodes.
When a deep snow cover l~sts for a very lor.g time the content of hydrocer-
bons significantly equalizes by sprin~. This happens under tt~e conditions
of the northeast ~uropean part of the USSit and frequenCly in th~ uegtern
regions of tl-,e U5SR. The differences in the destruction of various strains
are evened out considerably (Table 13).
One of the essential reasons for the considerably smaller differences in
- the resistances of strains of winter crops to damage under heavy snow
is that under a deep snow cover, especially when the snow begins to melt
and the humidity in the plants' environment reaches 90-100 percent, fungi
develop on them.
M. I. Rybakovn [31j suggested an indirect method of determining the degree
of wlnter hardiness of strains of winter wheat and rye--observing the
- dynamics of oligosaccharides. In addition to the plants' resistance to
Ereezing, she also studied the reaction of strains to fall and winter
thawing and the destruction of plants fram remaining under a heavy ar,,,w
cover. As standards for various resistances of strains to unfavorable
factors ~f wintering she used Zhitkinskaya winCer rye and U1'yanovka winter
wheat--winCer-hardy strains; wheat-wheat grass hybrid 186--a medium winter
t~ardy strain; and Motsinave winter wheat--a weakly Winter hardy strain.
During seven years a total of mare than 50 strains and selection specimens
- were tes[ed.
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~
~OK d~rICIAL U5~ hNLY
_ 'T~bl~ 1~~ ~ffectg d� Length o~ WinCer perind nn Wineering n~ Winter
Whpnt (3~ n~ ~urviving p1.~neg)
~ 1 ~ I1p0~0AMCNflAbHOtTb ~NNN (~HN)
G~pPf
t ITO I I10 I I6tl te0 I `!00 I 750
~
~ KonxoaHNU~ ge,1 ~~,g 4A,G 43,4 ~G,~ 30,4
G
~lap~~b 97,4 g1,A A7,4 4;2,6 35.7 28,3
5 1lbtecueNC 329 94,~ 90,6 40,1 ~8,~ 30~) 15,7 -
(6 Kdoncp~topKa g4,0 81,1 38,5 32,~ 29,3 20~7
Key :
1. burntion of winter (dgys) 4. Uyurabl'
2. 5trnin 5. LyueegCeens 329
3. Kolkhoznitg~ 6. Koopergtorkg
In the tillering nodes and legves nf winCer pla~yts under the conditione of
the Moscow area there are alcohol soluble hydrocarbons of oligosaccharides,
sucrose, glucose attd frucCose. In the gueuron there is usually a predominance
in the winter plants of sncrose--from 30 Co 50 percent (depending on the
str~in and autumn conditions); fructose--from 10 to 45 percent; oligo- ~
saccharidea--from 10 to 30 percent (in individual periods, up ro 40 percent
in rye); and Chere wns leagt glucose--from 10 to 20 percent (Tnble 14).
~ Table 14. Content of Sugars in Tillering Nodes of Wheat and Rye Plante
(9~ of total sugars)
~9x ~xi
eopr ~~ro- I e,x ' I I~D~~ M_~ I I~Pr~.
~ 1~ l~f~pN1Y po� raaKO~i to~a tlf~pN1Y pw~ r~A~aoi saoa
BAtKa a~ocKOectiaa 77,6 52,6 4,A 15 28,2 30 6,6 35,1
S'.16Ati0Bli~ , I~,O 50 9.~ 2~1 17 35 9.5 38.6
IlflC�18G Y0,6 48 14,2 17.1 9,1 3~,5 18,7 37.5
9 Kooneparop~a 13,`l 50,2 12,1 24,2 10,3 33,5 11,3 44.7 .
Key :
1. Strain 6. Vyatka Moskovskaya
2. Oligosaccharides 7. U1'yanovka
3. Sucrose 8. PPG-186
4. Clucose 9. Kooperatorka
5. Fructose
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After ~dmpleting the winter th~ md~e r~gul~r cdnnectinn with the wittter ~
hr~rdine~~ of t:he ~pecimen~ df winC~r Crdpg ig found in the cnnC~nC of
alignsnccl~nride~. 'I'tiere ig t~ 1C~~ r?~y:cut, buC predi~tab1.e Connpctic~n -
h~~twr~~~ tl~r truct~~~~~ r.nntr.ne r~nd wine~r hnrdin~~a~ ('Cnb1r 1S),
'Tgble 15. ConCenC of Nydroc~rbong in Tillpring Ndd~g df Str~in~ ~f Wineer
Whpge and Ity~ (9~ of bry Subgt~ttce) 14 April 1965
I O,~Nro�
~OPt K2~~xipNed ~~~~x~po� ( ~4f.~nRwi ( ~~gcKro~~
J
~ %KNTKIINCK~q a,00 11,6fi 0,04 5,9~
~ y1bp110~K1 ~,~4 g,5~ ~,~2 5,5~
Mnpo~io~r?caa 808 2,00 9,b6 1,08 5,64
hfll'�t86 I,10 5,96 1,12 4,~4
(1q Aiouwia~c 0,86 ~,16 1,1~1 7,58
Key:
1. Strain 6. Zhitinsk~y~
2. Oligosaccharides 7. U1'yattnvka -
3. 5ucrose 8. Mironovgkaya 808
4. Glucose 9. PPG-186
5. Fructose 10. MoCsinave
_ M. I. (tybnkovn (31], nn the basis of roany years of analyses of the dynamica
of the content of oligosaccharides in plants, recommends uCilizing them ~s
an indicator for evaluating the winter hardiness of various strains (Table
- 16). The change in the content of oligosaccharides and Cheir significant
_ expenditure is manifested especially in winter with slightly frozen soil
and minimal temperatures at the level of the tillering nodes that are close
to 0�C. 5uch were the winters of 1961/62 and 1965/66 (Tab1e 17).
Since the most significant changes in the contenC of oligosaccharides boCh
in winter rye ar.d in winter wheat Cake place during warm winters, one
3 must presume that this indicator can be utilized for characterizing the
resistance of strains to damage under the snow. In a comparative evaluation
of resistance to damage under tt.e s?~ow, the content of oligosaccharides is
a considerably more reliable indicator than the content of sucrose and
f ructose. But when determining them it is necessary to take into account
the origin of the strains and their adaptability to one t,ype of winter or
another. Thus analyses of oligosaccharides have confirr.ed already known
information to the effect that the most frost resist~~it strains of steppe
origin, where the snow cover usually does not lasr long, are far from always
the most resistant to damage under snow as Well.
.
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~ox n~~~~ic.'tAL U5L dNLY
`C~ble 16. Cdne~nt dt Hydrd~~rbdn~ in ~illering Nod~~ of 5tr~in nf Wint~r
Whe~t nnd Itye of nr~? 5ub~e~nc~) 15 Aprf1 1g62
~1~ Ca~~ O.~Nro~ ( C~~~ ~i I
2 ,a~PN,~e~ ~ D~ ~.,p~o� S o~~ra~~
6)K?+rk~u~cKaa 2,58 13,~7 1,10 6,g4
7 Ynbp~~oeK~ 1,12 11,06 1,1~ tU,76
E3 Aaa6acckan 1,17 9,d2 1,38 >i,bg
9~~ti~poNO~c?taa 808 l,il8 17,9 0,10 11,56
~Lp KyHitl'U~keq 4~ 1,06 9,8A 0,0~! g,2p
11 Mout+tt~~c 0 ~,I6 0 ~,g.~
Key '
.1. 5train U1'y~novk~
2. Oligosncchnrid~g g, Algb~sgkaya
3. Sucra~e 9. Mironnvsk~ya 808
4. G1ur.ose 10. Kuntg~v~kaya 45
5. ~ructose 11. Motsinav~
6. zhiCinekny~~
Table 17. ConCent of Oligo~gccharideg in 'Tillering Nodes of SCrains of
Winter Wheat and Itye in Varioug Years (9; of Ury 5ubst~nce)
tv.t~r~ ~~s~ca i~,c~ �c~;w ~~ss,cc
C�pr I y x I y y ~ I y ie
(1> ~ N 3 ~ o ~ ~ ~ ~
�
2 BNtKB IUOCKOBC~:IA 5,9 2,7 10,0 6,3 12,8 8,3 9,4 3,8 9,4 2,8
J'ii~anne~a 3,0 1,1 ~1,0 1,9 8,7 5,5 6.9 2,2 3,Y 0,0
~ n n r�186 2,1 0,6 4,0 3,0 14,8 2.7 9.4 1,1 5,8 0~6
j K~neparopKa 7,0 Cae- 3,n 1,0 10,9 3,0 5,1 1,2 4,4 0,5
A61
Key :
1. Strain 4. PF'G-186
2. Vyatka Moskovskaya 5. Kooperatorka
3. U1'yanovka 6. Traces
G. Salcheva [48) in Bulgaria studied the dynamics of free amino acids.
As a result of applying the method of unidimensional chromatography on
paper and visual determination of the quantity of individual amino acids,
it Was shoan that in the process of tempering (with a reduction of the
temperature in the autumn and the beginning of the winter), the quantity of
free amino acids increases in the leaves and especially in the til?ering
nodes. There was an especially marked increas~ in the content of aspar~agine,
25
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1~UEt O{~ f~ tC tAl, 11;~1. c)NLY
gerine, gluCnmi~c gcid, nl~nine ~nd proline. There w~~ glgo ~ po~itiv~e
corr~ldtion b~ew~~n the ncCUmulatinn of prdlitte in the eillerin$ nodeg
nnd the ~rn~t re~igCdnC~ of thp ~trgin~ d~ wt~p~e.
G. 5~leh~v~ ~~nclu~e~ecl experiment:~ with rpduCed (45 p~rcent) , in~re~s~d
(g5 per~.~enC) ~nd opeim~l (60 p~r~~nr) mdiseure enne~nt itt the gail. 'Tt~e
r~~e~rGh ghow~d th~e i.n ~he proc~gg df rempering wieh inaregged moigture
~~ntent in th~ gdil eh~ qu~nCiCy u~ fr~~ ~minn ~cidg deCreggeg itt ehe le~vpg
~nd ~illering n~deg. M~or~dver in p1nnC~ grown with reduced mdi~ture
rnntent df ehe snil, moge of the proline is gccumulgeed in ehe Cill~ring
nodeg during the period o~ segge~ I~nd I~ of eempering.
, WiCh tt lnw~rin~ ~f th~ eemper~eurp, in wineer whe~e ehere is g~hift of
the enzym~ r~~cCiong in Che direcrinn of ~ predominance of hydralysis
dv~r ~ynthegig. Ac~drding to ehe regegrch df N. N. Sis~kynn, wiCh ~ cprt~in '
d~pth of th~ ghift of enzyme reactions in Che dfreGtton of hydrolygis,
whiCh he c~ll~ the "bnrder df denth," there ig n decompengaeion crf
phygioldgiC~1 processe~ which ~auses the pl~nts Co die.
L. N. Itdm~nov~ [30j conducted rege~rCh on the coefficier~Cs of the shif.t of
enzyme reactions in struins of winter wheat with varying degrees of winter
hardiness. The coefficients of the shifC were established�by dividing the
indicatorg synthesig/hydroly.sis ~C each date of analysis wiCh respect eo
ehe ~ame indicator in the 5eptember sample (Table 18).
Tabl~ 18. Coefficientg of Shift of ~nzyme Reactions wiCh a Change from Che
Wgrm Autumn to the Cold Winter Period itt SCrains of Winter WheaC.
IfItNM! ~NMtt~ ~~{~O~~RqMtqT t1fltiltNNw
C2~ fM~QOAN) 1
Copf
~1~ JO IX 30 X JO IX I .0 X
llatccueuc 230 0,96 1,18 1,0 1,22
~ MocKOec?;aa 2a53 0,91 1,05 1,0 1,15
f1Rf �IH6 0,92 1,00 1,0 1,08
7)Kooncpatap?ca 0,90 0,6~ I,0 0,93
Key: -
L. Strain 4. Lyutestsens 230
2' Ra[io g~'nthesis 5. Moskovskaya 2453 �
hydrolysis 6. PPG-186
3. Coefficient of shift 7. Kooperatorka
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i~'C)It ~ll~ i~ 1 c; l At. I I51: C)N1,Y
Th~ dgeg dbt~ined by L. N. Itomgnovn r~in~id~ with ehe regulCg di the regegrch
df A. V. Slagov~gh~h~nekiy. Whi1~ fdr e~tglyeig in wine~r hardy whe~C
he obeained Q~~ m~..35 ~nd medium winCer hnrdy wh~~C--Q10 ~ L.78, in ehe
~xperim~nC~ n L. N. Itomanova wiCh Che winter hgrdy ~h~ae LyuCe~Caeng
23d ~h~ obCained Q10 ~ 1.59 and in Che weakly winter hnrdy grr~in
Kdop~rgCork~--Q~~ ~ 2.02.
'rhe resQarch of L. N. Itomanov~ algo ~greeg wiCh the ideng oE H. A. Itubin
whn gh~wed,~h~C Ch~ quantity Q10 ig nae ~ congCanC, bue dependg boeh on
rhe ~dndiC~on of th~ develnpmene of tihe pl~nes and dn th~ in~lu~nce of
gurrounding cnnditiong. -
Along with thp acGUmulgtion of suggrg and amino ~cids ~nd elle dehydr~eion,
pra~egs~g thne are essential for the eempering of plgntg t~ke p1nCe in st~ge
II. They ~re related to tih~ ch~nge in Che hydrophily, Ch~ vi~~n~iCy ~nd
the perme~bility di the prdtoplnsm. Wieh a reduceion in e~mper~eu~e eh~
_ visco~ity of Che protoplasm incxeaseg gCCnmpanied by u reductidn in Che -
_ ~peed of bintogic~l proce~ses in the Cells, thug conCributing Co their
Changeover to forced dorm~ncy. The permeabiliey af protopl~sm and ies
adsorptive C~[~~C~CY also ch~nge.
A.~ one cun see from a brief survey of the work physiologists and phytn-
pathologists conducted itt the northeastern and northwestern regions of [he
_ 5oviet Unidn, Che main reason for the deaCh of winter crops is the -
exce~sive loss of hydrocarbons, exhaustion and ehen sCarvation of ehe
plants which leads to a decomposition of proteins when Che pl~nts remain
fnr a long time under a deep snow cover und the temperatures are close to
0�C ut the depth of the tillering node. InfecCion wiCh fungnl diseases
- is a secondury c~use of the death of plants when they are damaged under the
snow, since the development of fungi is possible only on plants that are
already essentially weakened and physiologically ill.
In the northeastern regions of Che country's European territory during
years when there is a deep snow cover and deep freezing of the soil, winter
plants rarely suffer from damage caused b,y being u~sder the snow and in the
southern steppe regions where there is almost never an excessive snow
cever the winCer crops most frequently die from freezing and ice crusCs;
but in the cenCral belt of Che country's Europet?n territory winter crops
- freq uently sub~ected both to the effects of strong frosts with lengthy
periods when there is no anow and to damage under the snow when the snow
falls on soil that is thawed or only slightly frozen.
The majority of research on Che process of tempering winter crops involves
a study of the plants' resistance to freezing. Only a very small amount
oE work has been devoted to clarifying the correlations between tempering
and resistance to perishing under the snow.
27
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rn~ n~rYCrn~, usi, nNLY
M~ny physi,ologistn hgve been ~nd s~i.11 gre eng~ged in re~enrch on gugttrg
in connecrion wieh Che r~eigCgnce of wine~r cr~ps Co low ~emper~Curee.
'Chig re~earch bpggn with N. A. MAkgimnv~ A, A. ItikhC~r and I, i. Tumanov.
CerCain wriCers ar~ fully cnnvin~ed by ~heir duta Chat there ia a direct
link be~ween Ch~ cdnCenC of ~ugars and the re~i~egnce of winrer planes
(7J while othere hnve mad~ g numb~x of ad~u~~menCs in connection wieh the
- gtrgin nnd nCh~r peculiariti~s of win~er cropg and glgo ggro-meeeorologicgl
condieidng of vnrioua g~ographicgl r~giong [44~~ -
N. N. Y~kovlev [44~, wiChouC denying Che role of eempering and ~ccumulaCion
of hydroc~rb~ng in incren~ing ehe frost resigeance of wheat, citea ehe
data of a number of rese~rchers who think ehat ehe cnrrelgCion between
ehe content of hydrocarbons be�ore beginning Chp wineer and ehe winCering
of wint~r wheaC ~re noe ~lwnyg clenr enough. Like many oCh~r wrieere,,
h~ Chinks ehaC eh~ mnst gignificgnti tihing for the wintering of p1anC~ ig
not sa much Che conC~nC of guggrg in the ~ueumtt period gs ehe complex
- o~ f~ctors in Che winCer gnd early spring period.
- One cnnndC fully ~gr~e with ehe conclusions of N. N. Yakovlev since,
- ~lthough in a number of cases no direct connection is observed between the
- resistance of unfavorable conditions and the content of sugars, the protec-
Cive rnle of hydrocarbons is quiCe significant as is shown in Che works
of physiologista and selection workers who deal with the change o.f spring
forms into winter forma [29J in Che process of selection of such winter-
- hardy strains as Mtronovskaya $08.
~ The development of fungal diseaees plays a decisive role in Che subsequent
period of winCering of plants under an excessive snow cover. The period
of development of fungal diseases and the damage and death of plants
because of them, as was already pointed out, is the third and final phase -
of the perishing of winter crops under snow.
The damage to winter crops by snow mold and collar roC has been investigated
in greatesC detail by A. V. Pukhal'skiy [26J and S. M. Tupinevich [36-38].
Sclerotia are solid formations which are black on the outside and white
or yellow on the inside. The size and shape of sclerotia are extremely _
varied; usually sclerotia are 1.5-10 x 1-3.5 millimeters. They are usually
circular or oval in shape but one does encounCer sclerotia thaC are long
and irregular in shape. In the majority of cases sc?erotia are flat and
their thickness does not exceed 1-1.5 millimeters.
In certain years sclerotia are formed mostly in the axillary part of the
leaf and less on the surface of the blade of the leaf, where they are
Ioc~ted under khe epidermis. In other years scleroria are formed only on
the surface of the blade of the leaf and there are none at all in the
axiltary part. This is explained by the conditions of the year (depth of
snow cover, temperature and so forth). The number of sclerotia on one
plant can be from 1 to 25 and more.
28
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_ I~0[t d~'rtCIAL USi: tlNLY
SC1erdCia nt fungi Ch~1C fnrm in ehe spring r~quire ~ Gertnin p~rind df
rip~nittg in th~ light under n~rur~], condiCinny in order eo germin~te. -
'I'li~y do ndt germinnte in r1~c ~pring df Che year et~ey are �ormed. Cnnver~ely,
iC ~urly in Che gpring the young ~clexoCig ot fungi ~re plnc~d in the ~oil
nt U depCh dL 3-6 cenCimeterg, wiCtiin 75-60 dgys ehey are fully dpcnmposed
undcr el~e cffect~ oE xAprophyric fungi, bacCeria nnd nemoeod~s. SC1eroCin
germin~rte nnly itt Che auCumn afCer Che compleCinn of ripening on Che
gUtEF1Cf.' nf th~ ~di1. In orcter Co germinate acleroCin ne~d~ lighC, ~ high
content of moigeure in the gir and goil ~nd ~ f~irly low eemperaeure--
(x-12�C),~' Und~r ehese condiCiong ehe germittgCing gcl~rnCia form ~n
gfCOtilyCP.CnUy fruit be~rer for ehe fungus--~poehecia wieh sack ~pores. In -
- Leningrgd on Che field s~cCinn of tihe A11-Union InsCiCuCe of I'lgne
proCection ehe germingeion of sclerotia was observed during h~1f of October
in 1937 and 1938. Under naturgl condiCione at the ~alenskaya selection ~
sCation Che germination of sclerorig and the formation of fungal npothecia
were observed ~~C Che beginning of OcCober in 1936 and 1937 [44J. -
In the autumn wiCl~ the alternaCing rainy and sunny days the mature futtgal
sgck spores are broadcase into ehe air. ~alling on winter plants ehe s~ck
spores germinare easily and infecC them. The infected planCs enter Che
winCer tt~i~ way. In the spring uttder the snow wiCh n low temperaCure above
zero fungnl mycolin develop on them, which gt Che end of eheir development
on t}ie infected plants form new scleroCi~a. ConsequenCly, the form~tion of
sclerotin is the final stage in the cycle of the development of the fungus.
In addiCion to wheat and rye, sclerotia infect many feed grasses (CimoChy,
rye grass, sescue, wheat grass, meadow grass, cock's-fooC) and grass
weeds on which the fungus can maintain itself on the fields. Fungal
srleroCia can be reCained in the soil for 1-2 years and do not lose their
capacity for germination and the forniation of apothecia with sack spores.
A. B. Pukhal'skiy [26] did a large amount of work to study the resistance -
of strains of winter wheat to collar rot. He devoted special attention
to searching for forms of winter wheat that are resistant to collar rot
and characterizing various ecological groups of wheat with respect to
this indicator. During three ye~ars he managed to evaluate Che resistance
to collar roC of more than 2,000 specimens of winter wheat from the world
r.ollection of the All-Union Scientific Research Institute of Plant Growing.
_ During epipl~ytotic years sclerotia of fungi were found on all specimens
of tiie collecrion included in various ecological groups, which showed
their infection with Sclerotinia graminearum Elen, but the growth of the
specimens differed. Certain specimens developed satisfactorily despite the ~
infection with the fungus, formed young leaves and subsequently entered ~
the stem extension stage. bored spikes and produced a harvest. But the
maJority of specimens of the collection eiCher completely died off or only
part of them survived.
29
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roK orrtr.tnt, us~s dNLY
A. V. Pukltnl'ski.y noted ~ fairLy Clpar-cue gruuping in eermg df ehe degr~e
_ oE gpring ~fCergrowth of wh~at gmnng specim~ng from varioug ecological
E;rnup. 'The hi~;he~e d~gr~~ af winter h~rdine~g gnd Che bese afeergrowth
durin~ the gpring period characCerized represenr~eives of Che Scandingvian,
W~sCern ~urdpe~n hybrid, Ale~y and Ndrth American hybrid ecologicaL group.
Sdmewhnt behind ehem wieh r~gpecC Co these indicnenr~ w~re specimens from
rhe northern Ituggian ~cnlogical group. 'The wor~C ~.ndicgeorg (degCh nnd
poor ~fCergroweh) were obs~rv~d in gp~cimens from Chp norChertt ~uropeatt,
U~nube, Balcnn, easCern Asign, Ka:~hir and Kashg~r ecoingic~l groups. He
included serg3ns of winter wheat from ehe Northeaseern 5~leceion =
Center (now Falenskaya selection aCgtion) among the specimens Chat were
most resistant to coll~r rot.
A~ n result of tiis study of Ghe infecCion of winCer wh~ars with nollar
rot, the auChor came Co the conclusion ehae Che hardiness (registgnce) of
specimer.s is deCermined by their capability Co change over Co inCensive
growth in Che spring period, to form new roots and leaves and thus to
survive Che dam~g~ cgused under the snow. In this respect the role of _
- agroCechnic~l factors !s of exceptionally great imporCance. In Che works
of S. M. Tupinevich (36, 37] he points ouC the significance of good agro-
technology in the fight againsr the destruction of winCer crops by collar
rot.
Snow mold, develaping strongly on starving plants and accelerating the
expenditure of proteins, leads to Che destruction first of Che leaves that
touch the soil, then the bases of the shortened stalks and subsequently _
the tillering nodes as we11. The leaves of infected plants bend down to
Che ground and lose their color. An accumulation of mycelia appears on
them, which are white at first (hence the name "snow" mold) and then the
fruit bearers cause a rose tinge. The mycelia of Che fungi are spread
from one leaf to anoCher and it is as if they are glued together. The
disease rapidly moves from one plant to another, covering ever larger
sections of the ~lanted area.
There are various degrees of damage to winter plants. The f irst degree is
wlien the leaves are partial].y green, the tillering nodes are normally
turgid and with a rapid disappearance of Che snow in the spring and the
beginning of sunny and warm weather nPw leaves grow. The damage to the
leaves and part of the severely damaged plants die. The plant stand on the
section becomes thinner. The second degree is when the leaves are parCially
infected with snow mold and the tillering node is not very turgid. When
- the temp~erature is about 2-3�C for a prolonged time the mycelia proliferate
and the winter plants are frequently severely thinned out. The third
degree is complete destruction; the leaves of the plants are dark brown in
the spring, sometimes with a white or silver film on them and with
sclerotia of fungi of collar rot. The bases ~f the plant and the tillering
nodes are yellow and not turgid. A typical sign is a slight separation in
places where the tillering node is attached to the rooC sysCem which is
usually not infected.
_ 30
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~ ro~ ~~r~c tnc. USL (1NLY
ru
.
Andther ~ung~g Ch~C causes gimilnr d~m~ge en winCer crops, ~'ueArium nivn~.e
(~r.) C~g, ig no longer widegpre~d. ,
Snow mnld is c~uaed by a number of orher parhogenic fungi gs well. M~ny
of them ~re polyrhgges which int�ecr noe only win~er crops, but also many
~p~~ies nf wild grasses. Depending on soil condiCion, Che apecies nnd
the straing of plants, winCer crops c~re more frequently infectied wieh
coll~r roe in some regiong and ehere are varioug �orms of Fusgrial wilt
in nther regions. Agroeechttical measures direceed tnward proreceing planred
are~s from perishing under ehe snow reduce the harm cauaed by fung~l
diseases in Che ma~~rity of cageg.
ItC5~AltCH ON THE PROC~SS~S OF GROWTEI AND DEVELOPMENT OF PLANT5 IN Tli~
AUTUMN-WINT~It-5I'ItINC P~RIOD AS FACTORS IN THE FOItMATSON 0~ WINT~It-I~lAltbY
WINT~K CROI'S
te h~s long been knowtt ehaC increased growth of winter crops during a warm
- and moisC auCumn worsens t}~c wintering of the planted areas. In order to
_ protect rhe planCs from freezing and perishing under the snow, in agricultur~l
pracCice the crops have been mowed in the fall and it has been recommended
that potassium top-dressing be applied in ehe fall which somewhgt impedes
the growth of parts of the plant ChaC are above ground.
The connection between early curtailmenC of the growth processes in Che
auCumn and the winCer hnrdiness of plants has been repeatedly noCed by
I. ht. Vasil'yev (3] and I. I. Tumanov (35]. It is known that the most
winter hardy strains grow more slowly in the autumn, form short, narrow
leuves and are characterized by a split bush which grows closer to rhe
ground than those of less winter-hardy plants [7, 12].
For a long time the idea has prevailed among physiologists, crop growers
and agrometeorologists that with the beginning of the winter period and
the curtailment of apparent growth of winter plants, all thefr organs and
tissues pass inCo a state of deep dormancy, ~ust as they do in woody
plants. "
Indeed, as the research of P. A. Cenkel' and T. M. Zhiwkhina [7] has
shown, in winter wheat there are not very prolonged, but clear processes
of separation of protoplasm from the cell walls which characterize a
condition of deep dormancy. But in the ce11s of leaves that are less
tempered and especially in "overgrown" plants of less winter-hardy strains,
_ the condition of dormancy does not last long. In the plants of less
winter-hardy strains under the conditions of the Moscow area, the number
of cells in which there is a separation of the protoplasm does not reach
a maximum until November and in the beginning of March Che planCs are
already completely out of the dormant condition.
31 -
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Ih?ring yenr~ with n],ong, warm aueumn when the gnow fg11g on god gni1,
- no c}inngeover of winCer pl~nts to a deep dormanC condiCion is observed.
In eheg~ yegrg, ag I. I. 7umanov has nlready noeed [35], even un~~er a deep
snow cover ehe eeiolaCed leavea of winter whenC grnw.
Mnny researchera hnve observed a sllghe growth of veg~tative organg under
_ the snnw~ noting eh~e the balance of the overall ~xpenditure of gug~re
frequenCly doee ttoe corregponcl Co the c~lculated figurea for Che loes nf
sugars. Cnnsequenely, some of Che sugara expended by th~ planCs under deep
- snow apparenely go fc~r growth procesaes, although insignificanC ones. -
- The weakening of the growCh processes, as many researchers have noCed,
is one o~ the necessary condiCions �or the p1.anCs' changeover from a
grnwing condition which is not resietanC to unfavorable conditions of
wintering to a period of dormancy and ehe second phase of tempering for
, low winter temperaCures [35, 37~ and this condiCion is also necessary
for the plnnts' resistance to winCering under a deep snow cover.
How are Che growCh regulaCors and seimulators of growCh processes firsC
- inhibited in the proceas of tempering the plants in the auCumn period?
What is the interaction and relationship among growth regulators--auxins,
gibberellins, kinins and various growth stimulators and inhibitors of
winter plants during Che au~umn, summer and early spring periods?
Many scientists attach a grpat deal of imporCance Co the role of auxins
in the process of tempering winter wheat. Until recently in ehe literaCure
little atCention was paid to the question of the dynamics and relation-
ships of grow th sCimulators and inhibitors during the process of tempering
winter plants [4]. In the works of V. G. Konarev (1959, 1964) it was
suggested that the influence of all f~ctors nf the external environment
is exercised through the functional activity of the nucleic acids.
More interesting experiments to investigaCe the growth processes when the
plants are changing over from a growing condition to forced dormancy have .
been conducted in recenC years by V. V. Vinogradova [5]. She investi~ated
the frost resistance of winter wheat in connecCion with the inCensiveness
- of growth and the content of endogenic stimulators and inhibitors in the
autumn, winter and spring periods; she studied the influence of treatment
with data-indoleacetic acid on Che growth and level of native auxins and
inhibitors under various temperature conditions for tempering; and she
also investigated the changes in the condition of desoxyribonucleic acid
in the chromatin of the cell nuclei.
The intensiveness of growth in the experiments of V. V. Vinogradova was
detet-mined by the sizes of the leaves of 60 plants and Che length of the
vegetative cones of 15 plants were measured under a microscope. The
biological activity of the growth substances was determined through
stimulation or inhibition of the growth of cuttings of the coleoptiles
of winter wheat.
32
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I
1~'dk Ol~ P' CG 1 AL USI's UNLY
A comp~ri~ott of ehe darn on ehe inC~nsivenes~ of growth of planes ~nd
Che 1eve1 0� their frose resisCance ~howed (F'ig, 2) Chat Che differences
. in the leve~, of frose xesiseance o� the a~raina inves~igated by the
- gurl~~ur--U1'yanovkg, Borovichskaya and BezoaCaya 1--depended on the
ictr~etgivene~e of eh~ g~oweh c~f the plgnes' leavee during ehe gutumn p~rind,
Iteduced intpnsivenee~ o� growth in ehe ~utumns of 1968 gnd 1969 con-
Cributed to Che development of more highly resistant planes. V. V. -
Vinogradova gccompanied the research on the dynamics o� froat resigeance -
of plants witih measuremenes of the vegeCative conee (Table 19).
A reductiion in resisCance in the experinentis af V. V. Vinogradova (5]
coincided wiCh the perioa of growth of the vegeCative ~ones or~ as Che
author formulaCes iC, with a period of "concealed growCh" of the plants.
' 'I'he regulrs of the seudy of the dynamics of growth substances in the same _
experimenta showed ChgC begintting in AuCumn~ durtng the course of
tempering, ehe sCimulaCors in the tillering nodea of wineer whear are
almosC completely inactivaCed. Moreover, it is very interesCing that
- U1'yanovka, a highly frosC resigtane gtrain~ is diaCinguished by a lower
level of stimulaCore, high speed of their inactivation and earlier
attainment of the autumn physiological maximum in the contenC of
inhibitors (Figures 3, 4). For the weakly frost resistant strain
Bezostaya 1, iC was characteristic to have a alow inactivaCion of growth
stimulaCors, later aCCainmenC of the autumn physiological maximum in the
content of inhibiCors and the prevalence of Cheir acCivity over that of
growth stimulatora.
From the data obtained by V. V. Vinogradova it is also clear that
during the winter period there is a gradual reduction of the growth
inhibitors and almost a complete disappearance of them from the tissues
of the tillering nodes during the period thaC precedes visible growth.
On the basis of these and figures from literature, the author arrives at
the conclusion that during the second half of the win~~er, even with
negative temperatures, there is a change Ln the intensivenesa of certain
metabolic processes in the tillering nodes of winter wheat which lead to
a loss of the tempered condition.
_ This is also confirmed by an analysis of the condition of Che DNA in the
vegetative cones. In the weakly winter hardy strain Bezostaya 1, in the
differentiated tissues of the vegetative cone during freezing and
especially after thawing there is a marked loss of some of the nucleic
acxds as a result of their degredation. Unfortunately the amount of
physiological research that uncovers the mechanism of growth processes
is not very great so far. _
33 -
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roit c~rt~~r.r,rnr, usi~: ntv~,v
_ ~eC
16 a)
f? ~
B
I
4 iJ
\
~ _j
-a r r
_a 1
f0 ~.1J !0 10 31 !0 ?9 JO 10 ?J 3f f0 ?ID 3! JO TO?B f~ 110 31 f0 10 3D
/.f 1( J(1 ,tll l ll lll lV
- B~
6) _3 A
c~r ~
~ A 6
13 -10 ~
_ \
~ -fS
~ /
S , , ,a,~
~-e...e~
-S 6 f
~ f 7 J 456 1? J 456 ~3 ~
~ / ~ , -!0 ~
-~s
-s
` ~ ' _
-~o ~ 3
. ~ l~!
,
-~s / j
~
/
-~o ~ -
u~ x? ~a r u ia n~
Figure 2. Frost ResisCance of Winter Wheat, Depending on
= Temperature Cdnditions of the Soil and Intensiveness
of the P1anCs' Growth in the Autumn
_ Key:
Temperature conditions of the soil at the depth of the plants'
tillering nodes
G. Length of leave~ (from first to sixth) at end of autumn growing
periods of 1967 (I) and 1968 (IZj
34
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ro~ nt~CrC1AL USL ONLY -
Key (co*~tinued) :
v. Itesist~nce of strgins of wheaC in variou.q years cri.Cicn1 C�C
(death o� 50% of p],anCs) .
A. U1'yanovka
~ezosCay~ 1
B. Dorovichsknya
1. 1967/68 -
2. 1968/69
3. 19fi9/70 '
Table 19. Sizes of VegetaCive Cones (mcm) of WinCer Whear in AuCumn-
Winter-Spring Period (1968/69)
91 X I ~ Xfl I 48 111 I $1 111 I t2 IV I 15 IV I 5 V
. ~1~ yJIbpII00K~
.~36 f 8 ~ 736 ~ 12 ~ 742 � 15 ~ 253 � 18 ~ 284 8 ~ 338 � 14 ~ G49 71 _
(2 ) 6e~ocraa 1
':GO � 12 ~ 270 � 13 ~ 276 � 15 ( 300 ~ 20 ~ 312 � 21 ~ 390 f; 17 ~ 690 � 33
Key:
1. U1'yanovka
2. Ber.ostaya 1
There is considerably more work and more observations of the growth of
plants in which changes in the morphology, size and form of the vegetative
cone are investiga ted by the microscopic metho d and in recent years the
condition of the stages of organogenesis in the autumn, winter and spring
periods has been studied.
Beginning in 1936 F. M. Kuperman reputedly neted that under the conditions
of positive temperatures in the soil during the winter, especially during
the thawing period, there is a certain growth of the vegetative cones,
although insignificanr [7]. Moreover, if the process proceeds normally,
when the plants are transferred to a greenhouse the vegetative cones
rapidly begin differentiation of the rudimentary spike and enter stages
III-IV of organogenesis. Among the methodological aids, in 1936 F. M.
Kuperman recommended diagnosing the condition of areas planted in winter
crops by observing the vegetative cones by the so-called method of "bringing -
it to the rudimentary spike." To do this some of the plants in the
pore samples fox~ 20-25 days (from the time they are removed from the field)
- were placed under conditions of 24-hour light and then the rudimentary
35 =
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~dK OI~~ICtAi. U5h tlNLY
% nf conCrol
f30 a~ f1 ~ ~ D
f?t~ ) t.3:!
1f0
29 /X fQ0 - ~
~
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~ f10 f~2,5
fS X u f01~ ' - - _ -
a - -
~ ~
0 617 -
u
1f0 1~7? ~:s
3f X ~ f00 _ _ -
~ ~ - - - -
~ ff0 1:3 f~?
u.
74 Xl~ 1A0 - - ~ ~
~
!f0 1~1,3 f~3
?4 Xll 10a -
90
1f0 f3~ f
?0 /l ~OU -
~ro ~ s. ~
30/ll ;00 - -
~ ~ ? ~ i i ~ 1 i
~0 0,? 0,4 . 0,6 0,8 f,0 0 0,? 0,4 ~ 0,6' O,B Rr
Figure 3. The Activity of Growth Substances and the Relationship
Between Stimulators and Inhibitora (figures over histo-
grams) in Tillering Nodes of Winter Wheat (1968-69)
Key :
1. % of conerol
2. Lncrease in coleontiles
36
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1~ Ok cli~ t' CC t AL tI51, hNLY
~nik~ wns nnglyzed. Thi~ method wn~ 1~tpr d~gcribed in more detail
in 1953~sncl 1956.
Aft~r ehe publicgtion of eh~ work which nnnounced ehe main p~teern~ of the
gtggeg of orggnngenegi~ [7~ nnd Che schemes wE are pregeneing in figures
5 nnd 6, Che peculiarieiee of organogene~is of vegeCgtive cnneg during ehe
aueumn gnd winter periods were investigaCed ~.n deesil by many researchere _
C10, 12, 14, 18, 19, 22, 25, 45, 47J.
x .of~ control
f?0 0~ 6~
f10 35~ f
?6 /X 1Q0
~
a~
~ ff0
2B X oMaO ~
v
~
u1f0 2`~ f:5
~sxi ~~c~
~ s~
~
~ �o ~:s ~.6
3oxu,~~oo - -
~ -
n~
24// f0I0 -
_ ~ `
1f0 23: ~ -
?6/l/ f00
~ -r ~ ~ ~ ~ -
0 D,Z 0,4 0,6 O,B f,0 0 0,? 0,4 0,6 O,B R~
Figure 4. The Activity of Growth Substances in Extracts From
Tillering Nodes of Winter Wheat in the Process of
Tempering, Wintering and Renewal of Growth in the
Spring (1969-70)
Key :
1. % of control
2. Increase in coleoptiles
- 37
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Thc mnJoriey nC ~Crnin~ of winCcr wheue~ u~tder thc condirionq oC Chc m~in
rcgionH wh~r~ eh~y nrr cultiv~e~d. nr~ in st~g~ II of organog~n~~is U~for~
_ th~y enter Ch~ wineer. Ag F. M~ Kuperm~n [7, 10], V. A. Moigeychik [22j
~nd mnny aCher ~uthorg note, at th~ beginning of ~uCumn the vegetative coneg
in variou~ strains ar~ characterized by relatively eimilar siz~s, egpe~it~~1y
if Che e~mperature drope rgpidly (below -5-8�C).
'I'hus gt the beginning of November 1966 in Moacow Ob1asC in such eCrains
at U1'yannvka and PPG-559 ehe length of rhe veget~tive cone was 0.19
mm, in the less winter hardy atrains Moakovskaya 2453 and Odegakaya 16
th~ lengtt~ amounted to 0.2 mm, and in the relatively weakly winCer h~rdy
strain Bezoataya 1 ie was glea 0.23 mm. I~'rom Table 20 ie ig cl~~r rhgt
with varicws plgneing Cimes gnd in years wieh various meCenrologicgl
- condition3, in pYants of all atrains (with exceptions, as will be shown
subsequently, of anomalous years and exCra early planting times) th2
vegeCaCive cones are in stage II of organogenesis before entering Che ~
winter.
Table 20. Growth of Vegetative Cone During Winter Period in Various
Typea of WinCer Wheat, bepending on Meteorological Conditions
and Planting Times [12]
I
- MH(WHJetK~ll 808 K>pucuKU 43 Yae~HO~K�
a~ra noceri aar~ ea~rN~ ~41 ~S)
(1) ~~~~oau
npupoer npNpoer npnpoet
(6) I ~~1j (6~~ (~9' ~T~S ~f~~
25 I i I 1966 r. 21 X 1~J66 r. t 1 100 1 I 100 I I 100
12 1 V 1967 r. I I 147 t( 1 a 5 I I 130
7 X 196fi r. .~9 X 19
~6 r. I I 100 I( 100 - - -
12 I V l~J67 r. I I 1 Y~1 1 I 119 - -
10 X 19fi5 r. G t I I 1 X'i5 r. I I 100 1 I 100 I I iM
_ Y2 ( I I 1 SK'iG r. i I 1~N I 1 138 I 1 120
1 G I V 19Gf, r. I 1 115 I I 141 I I 266 -
d,N.a ~9~a~,~,
;loTi sa~tM4
A~T~ naeu ~p�by .
~1~ ~2~ fi rm ( ~pl' ~ ~6~m I np~Qar (Xl
~ ~ ~ )
25 I 11 196G r. 21 X 19fi6 r. 1 I ] 00 I I 100
1? I V i S?G7 r. i I 136 I I 167
7 X 1 QC,fi r. 'lf) X 1~J66 r. 11 100 f f 100
12 t V 1967 r. I I 119 I 1 109
- 10 X 19fi5 r. fi X I I 1~JGS r. I 1 100 I I 100
i I I 1~~GG r. 11 132 I I 130
1G I~' 1y66 r. II 135 11 144 -
38
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~0[t 0~~~'LGtAL USL nNLY
Key :
1. t'lnntin~ dnC~ 6. 5eag~
7~ pnC~ ~nmple teken 7, Incr~~~~ (9;)
Mironovpkny~ ~OS 8~ ~nnnl ~
4. KunC~evc~k~y~ 45 9, K~khl~nd
5. U1'y~novkn
In rhe gutumn Chere w~g v~ry lieel~ differen~e itt the size~ of the
vegetative con~n nf such ~traitt~ ~s Che wineer hgrdy U1'yanovekgy~ nnd
Mironovk~ 808 ~nd ehe weakly winter hardy Kokhl~nd gnd ~gnal. Under Che
Gnnditidns ~f 1964/65 when th~re wgg an pXCE88~Vp enow cnver in Mnecow
dblgse, in nlmo~e all of the sCraing Ch~C wer~ ee~ted ehere wge ~tt
~xtenginn of ehe lengeh df Che veg~eaeive coneg nnd in ~'~brugry the
incregge in Chpir lengthe ~ignifi~~ntly ~xceeded ehe norm. But before the
disgppegranCe of the snow all sCraine were itt sCgge II ~nd r~rely iti
gtnge III of organogenesig. The atraitt di~ferences itt terms of the
differentiaeion in eize of the vegetative cone in straing with varying
wint~r hardin~gs were mnse eagily reve~led from the gftergrowth of the
plants in ~ greenhouse during the winter or under field condiCions in the
e~rly spring (Tnble 21).
5imilar datn concerning the change in the length of the vegetaCive cone
were obtained by V. I. Ponamarev. As one can see from table 22 the
intensiveness of Che gruwth of the vegetative cone during the winrer
period under a snow cover depends to ~ eignificant degree on the gge nf Che
pl~ntg, the strain peculiaritieg gnd the weather conditions. Thus more
- intensive growth of the vegetative cone in the same sCage of organogenesis
was observeci in "older" plants with an earlier planting time. Plants
planted in 5eptember were distinguished by the slowest rates of growth of
th~z vegetative cone under the conditions of the Moscow area. The length
ef [hc vegetative cone changed equally predicCably depending on the degree
of winter hardiness of the strain--the least increase in the length of the
- veg~tative cune was observed in the most winter hardy strain, U1'yanovka.
~ Winter growth of the vegeCative cone also ch~nged depending nn the
_ temperature and waCer conditions of the autumn and winter. Thus in the
dry autumn of 1966 the plants of all straina and planted at all times
formed a shorter vegetative cone than in 1465.
The connection between the intensiveness of growth processes (and especially
the intensiveness of growth of the vegetative cone) in the early spring
and the frost resistance of the strain was excellently investigated in
experiments conducted in the winter of 1966/67. The more intensive
the growth of the vegetative cone under the conditions of an excessive
snow cover in 1966/67, the greater the percentage of deaeh ~f the
plants (Table 23).
39
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y
~ ~y ~ ~ N~ g N _
~ ~
,~W > o o g, I I 1 I
y
~ b � -
a? ~ ~ ~ g o~ o ~ 8 e
~ o
ro ~
I I I~'" I= g I I I I _
a
roro
a,
~ ~3 > ~ o ~ o ? c~i � > c`~
~ a~i 1�
o�1oa ~ I~ ~3 I ~ i ~ ~ I
= c~ "
~ M r
~ y
~ ~ ~ ^
~ O ~ ~ 8 ~ - 8 ~ N : g -
N o 0 0 o
w p -
O M p er 1.y S F9 ,y o0 CNi~
~ ~ O ~ :
Cl u p^ O` 0 O
DO t/~ ~
N ~ x
o ^ I I I~ N I I I i~ R I
,G M ~
7 N ~G
N~�~ ~ ~ 8 R ~ 8 8 0` 8 c~
w ~o � � ~
~ o o~ -
~ a h ~ ~ A ea
~ ~ o a ~ " ~ a ti A
tA O U t0 r'' eda r' r' 6 v r' n~ ~
V) V ~/I ~ ~ S~ ` a ` V~ ~ f~ = a'�
r. ~
a;v cQ i. o� oQ =
q N ~ _ a - ~
0~~ C~ p,- Y s Gp'= x j ~ O~_= Y~ Ov~" K -
v ? 6 O r
VI J-1 ~ O F~ ~ Oc C!- e7 R C F, ~ a F0 a
- ua~o ' _
ooc~ =0~~_ h
c~o > 3 m aQ ri cr ae ~ m a� d m ae ~i
r oo v~ ~ oo c~ u, o0 0+ ao c~
4+ C v v v v ~
~ N vv v~v v e
n
~ Y r ~
h r1 y~ V Y OC
o V m O m ~
Cl a
~ ~ O G~ ~ ~
d G ~ T V
~ ~
M ~ ~
H ~ ~ v
40
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~o~ orrtr.inL usL nNLY
Key: "
1. 5ernin 6. Bezngeayn 1
IndicaCor 7. SC~g~ c~~ nxgandg~n~sis
tll''y~novkn 8. X o~ over~il growth of plants
4. Mironovgkaya 808 9. LengCh o~ vegeraCive cone~ mm
5. Kun~~evgk~ya 45
~ Noee: A--Ch~ condieiott of Che plante on Che day Ch~ g~mpl~ w~g e~ken
from ehe field; ~--the snme, g~Cer Cheir growrh for 3U dnys itt
~ greenhouge.
Table 22. Change in I,engCh of VegeCaCive Cones of Winter Whe~t Plants
of Various Ages During Wineer Period
' CpoK tes~
Cept 18 VIII I 48 Vt~l-29 Vtll I 10 IX w
~2~ A 6 I A I 6 I A I 6
_L_
1965~66 r.
Ynbn~ionKa ( 339 I 153 I 262 I id6 I 177 I 128
M~tpoeoecxax 808 409 165 324 157 240 137
1966I67 r.
3 YnbaNOSKa 252 122 193 109 150 10~f
43 M~ipoHOecxaa 808 ( 314 I 140 I 252 I 129 I 189 I 111
~ Key :
1. Planting Time 3. U1'yanovka
2. Strain 4. Mironovskaya 808
Note: A--Length of cone at beginning of spring growing period;
B--in % of Length of cone before entering winter.
Tl~e influence of growth processes on the change in winter hardiness of
plants is noted in the works of A. I. Mitropolenko (1975). Table 24
gives his data concerning the frost resistance of planCs of various ages
under :he conditions of Krasnograd (Khar'kovskaya Oblast). Table 25
shows that during the course of the winter period a change in the length
of the vegetative cones of the plants was noted and the influence of the
age of the plants, the strain peculiarities and weather conditions was
clearly manifested here.
Plants of the Kuntsevskaya 45 and Nemchinovskaya 495 strain had a longer and
more differentiated cone both at the end of the autumn growing period and
at the beginning of spring aftergrowth than did plants of the U1'yanovka
strain.
41
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~nit 0~'~'YCYAL US~ ONLY
~ ~
l~
~
~
~
, ; d ~r 8 ,
B soa~? Tpemu~ Kywe~ue Ba~rod e p
~g) ~b~ (c) mPy~Ky
(d) -
~
f ? 3 ~ ~ ~ '
~ I'4 1 '
, ~ ,
~ ~ ~ ~
~ ~
~ '
g
,
� !
,
' ~ ri q, k
; -
~ ; ~ ~
:
Pam rnxk~~ YI ~ B~N~vuia- L(~
r~
ov.~oA ~c~o~mo
~e) ~~f~~ ~8) cn~hjme
Figure 5. Phases of Development and Stages of Organogenesis of
Winter Wheat
Stage I--undifferentiated vegetative cone; stage Il--differentiation of
~ rudimentary stalk into nodes and internodes (beginning of the formation
of sheathes of stalk leaves); stage II--segmentation of lower part of
vegetative cone and formation of rudimentary covering leaves (bracts);
stage IV--beginning of formation of spike tubereles; stage V--formation
of blossoms on spikes; stage VI--formation of anthers and pistils;
= sCage VII--formation of sex cells (gametophyres), growth of length of
segments of spike shank; stage VIII--formation of spikes; stage IX--
blossoming. fertilization, formaL�ion of zygoes; stage X--formation of
caryopsis; stage XI--milky ripeness (accumulation of nutritive sub-
- . . stances); staye XII--waxy ripeness (Cransfer of nutritive substances
into reserve) and ripening of seeds;
42
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Key: 1, 2, 3-~ subgequenC fnrmntion of pollen
5hnoeg d. 5Cem extensinn g. Illosgoming
b. 'I'hird l~g~ e. Spike formnrion h. Milky, w~xy
c. T111~ring f. 5pike formaeinn ripenesg
(1) mnj w (2) arinw (3) naouv~ii1NnNOCrN _
fl Op~tt~Nll! !lMIIN. I1N101~lplH4N~{INN N pOtf S~� y nd�1lS~11 Stk071IttT~~
I r4~ Bt1t0Ay Iwl PoAUwe~u� opr~HO~ ~ I I/~2~~~TOtt ltOMNN�
~ \ paereHHA
1 ~1
~11~~lplMIINBUNM OtNOSaHN~ C~~NTyt pOtTlMM~
7petMR JIMlT, kOH)'t0 N~ ~14oT0~MY! )'lJIY~ ,y (ShcOT1. ~Nl10
IG\ MyWlMN! MlJKAby~1NR M GTlOJIt~N! AN� ~~~~T?t~), NOimQIN� `
tT?II ~ 4NlHT NyWtNNf,
~ ~NMOt10AN0lT?
1 -
1~~. AN~~tDlNYN~4N11 fAalMOI) OtN I~1,4~4Nt10 t.llMNkO~ (
~lM~tO~MOfO t04slTNII M Gp~K� koroeoeoro etep~en�
teA
- i !
H wieo ~MxoA? ~I IV. 06D~~o~aNNe roMyeoe napaera� I~ 15 )4eero �oroero� I
I~6~ r tpyGKy I NNII sro oro nop~A~s (roao� 'Y konote, ~acyxa _
lN0~41t ~yfOpMO/) )'tt0A4N~OGT\
~ 1 i
~ buxoll ~ Tpy6Ry - V~ 3aKa~A~a IIOkp0~MY1I oprano~ 1 4Neao u~et~o~
M~M~110 litatlNMM11 I I UbtTK~. tM4MHOK M flttTNNO~ I~a16, ~ ROJIOtK~R I ~
~
. I VI. ~OpMN~O~~MNt tOY~lTM11 M -
LL!liMS ~MNNpO�M~Mpanopore� I I
Nt~)
1 mlQTN 71MOCT~
VII. faMetomMtoreHa, par noRpo~� ,,,,I u~etKO~, n~oteoet?
I MY7I opnHOe. yAANMtMM! rae- I ro~oa, ~apamA�
NMMU~ K070C0~0~0 [TlpIRNY ~~7\ kOtt?
l
V111. t'aMetoreNe~� ~~~epweMNe npa I
I~8~ KoaoweMMe I-' ueeco~ ~OpMMpO~OMN~ tltx 0p- I
raMOe eou~erM~ N q~etRa
i !
I(9, ~(~tTtNN! I-.I IX. OIIJ16AOf~OptNN! M OQQa70~~� I,,,~ O~tQNtMMOtT? I
{ NM! 1NMiM 1 O kOAOt~
~ 10
~
I X. POlT N ~OpMMpO~~NN! !t~ I!I � I
MO~RN 7~f1~tAN~NN> >tpMO~~M
\ ~
f +
~1~~M/ CfMtNM. xI. N~~011.7CIIM! flNTiTlt?IIYt
I M0710YM~� CIIt70C1~ (~I t4lCT~ �~tpNO/11! (CtMlMMI y ~~OatC ]tpMO~RN:
j + )'C10A~N~Ott~
x~l. ~l t~ ~ R ~ytO~t~M
BYCLLO~I� N 110.7N~11 ~ I D D WlNN! 11NTaTlA?MYx
K~~\ !11laMt? I~~ Wt[t/ � Jif1~CMYt ~llqtliN "
J � .~t~1NO11R! I~lMlMN~
Figure 6. Formation of Productive Elements of Wheats in Various
Phases of Dc:velopment and Stages of Organogenesis
~+3
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Key :
- 1. 1'hasea
2. Seages -
3. Produceive elem~nts
4. SprouCing of seeds,~ahooCa
5. Third leaf, Cillerittg
6. Beginning of stem exCenaion
7. Stem extension--beginning of stem growth
8. Heading
- 9. Blosaoming
10. Filling in of aceds, milky ripenesa
11. Waxy and complete ripenesa `
1Z. Field germination, density of plane atand
13. HabiCus of planea (heighC, number of leaves), tillering coefficiene,
winter-hardiness
14. Number of segments on apike shank
~ 15. Number of spikelets in spike, drought-resiseance
16. Number of blossoms in spike
17. ~ertility oE blossoms, densi..y of spike, heat-resistance
18. Crain content of spike
19. Size of caryopsis
20. WeighC of caryopsis; resistance to dry winds
I. Differentiation and growth of embryonic organs
II. DifferentiaCion of cone base into rudimentary nodes, internodes
and stalk leaves
III. Differentiation of main axis of rudimentary flower cluster and
bracts
IV. Formation of vegetative cones of second order (spike tubereles)
V. Establishment of integumentary organs of blassom, stamens and
pistils
VI. Formation of flower clu~ter and blossom (micro-macrosparogenesis)
VII. Gametophytogenesis, growCh of inCegume~;tary organs, lengthening
of aegments os pike shank
VIII. Gametogenesis, completion of processes of formation of all "
organs of flower cluster and blossom
IX. Fertilization and ~ygote formation
X. Growth and c~~;opsis formation
XI. Accumulation of nuCritive substances -
XII. Transformation of nutritive substances into reserve substances
in caryopsis (seeds)
When there is a long auCumn and the snow falls on thawed or slightly
frozen soii at a temperature close to 0�C, even at the beginning of
Novembe~ the vernalization processes are completed in the plants, the
tempering decreases and there is a sharp decline in the resistance both to
critically low temperatures and to conditions that cause perishing under
the snow.
44
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~OEt d~~ICIAL U5~ dNLY
_ Tnbl~ ~3. InC~ngiven~~~ df Groweh dE VegeC~tive Conee buxing Winter
P~riod in Winrpr Whe~C WiCh Vurinu~ Uegre~g df WinCer H~rdiness
Under Condition~ af Mog~ow Obl~~t
~I X t~65 r. t4 ~V 19G~
CoP* ~2~n ~l w~~ ?co� ~z~n ~.3i~~s ko� 4~, ktNON1t
pacreH~~n
Opl'~HO� NyC~ HO� opreuo� NytO H~� fl0~.tr JNNtl9�
~1~ ~l11!]~ plCt~NNp P!Hl~I p8C10HNq Kii npN bet�
CHpMt6@
S YJIb11NOBKA ~ ~ ~,Q~ ~~3~ ~0,~
6 biN oHOectia~ 808 II 0,~0 II b,.~.~ .~g,0
7 nr~r�186 II 0,28 Il 0,46 ~6,0
8 Kyi~ueecKaa d5 II 0,26 II O,aS d4,0
9 6eaocraA I I 1 0, 33 1 t-(1 I 0, 59 20,
~~Q IC98M0TeC 0~30 0~57 23,0
_ 1~Dai~a~ 11 0,3G (I 0,59 10,0
- Key :
1. Strain 6. Mironovskaya 80$
2. 5Cage of organogenesis 7. PPG-186
3. I.c~ngth of vegetative cone 8. Kuntsevskaya 45
- 4. X of live plants after 9. Bezostaya 1
wintering without snow 10. Kvametas
S. U1'yanovka 11. Fanal
Tnble 24. Frost resistance of Mironovskaya 808 Winter.Wheat With Various
Planting Times (February 1972)
~1~ ~2~ ~3~ ~43ewnep~rqp~ npowop+~nuxHw (�Cl (2/
~~T~ Caro~nMe ~oMre~ ba npowo- 16 I 19 ~ 41 I 23
naeaa H~p~tT/NIHI Nl II QlMtN~~MN~
~rine opr~aorena~ (koxrpa~)
/5~ tOx(~~NNAOC~ Q~tTtMMA
\
~ 7 V I I I CN 16it0 B6iTAH}rT�
aurroede Bam~ 81,9 55,5 30~0 0 0
23 V 1 I I B~rAHyr, .i~ cTOBtie
ean?iKn ~7) 90,0 61,6 a3,3 ?,7 0
l IX N ava~o edrArNea�
uuA noAeneNNe
m~rrq~
x eanN-
xoe l~ 1 100,0 100,0 91.6 81,9 16,7 -
? 1 X Haaano estrArnea-
HNA (9 ) IOO~O IOO~O IOO~O TT,7 34~5
15 IX CorroaHee nepexo�
A8 K fit88�
- NN10 ~1 IOO,O IOO,O IOO,O IOO,O IOO,O
[Key on following pageJ
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K~y:
1. rl~nting dgC~
- 2. C~ndiCion ~f Vcg~C~CiVe eon~ in ~t~g~ II df arg~ndgen~~i~
3. WithouC freexing (cnnerol)
4. T~mp~rnCure of f r~ezing (�C) (24 hours)
5. Surviving plgnes
- 6. ~xCr~mely ~longaeed lpnf v~lliculg
7. ~longaCed l~af vallicula
8. Beginning of elongation ~nd gpp~grance of legf vglliculg
9. Beginning of elong~Cion
10. Condition of chgngenver Co elnng~tion
Tnb1~ 25. Change itt Length of Vegee~eive Coneg of Wint~r WhegC pl~tttg nf
Vgrious Ages During Winter Period (main shoot), 1964/65
/2~ CQOKN
l
~o~ 15 VI11 IZS VtU ( S tX I tS IX I IS Vill I2S Vfll l 5 IX I 15 IX
~3J~t NH A t~tf~UHN (MKN~� I ~4~ �.r1t0~ M~~ MMrpf~
(5 Kyi+ueecxaA 45 I 338 284 212 185 12T 132 119 109
(6~ HCM4NIIODCKaA 49S 328 297 224 191 138 123 112 108
~~I YAbAilO8K8 I 288 260 165 169 125 122 105 101
Key :
1. SCrain
2. Planting ti~s
3. Length of cone at beginning of spring growing period (mcm)
4. In X of length before enCering winter
5. Nemchinovska;~a i.~5
7. U1'yanovka
It l~as been establiahed (7) that when vernalized seeds are planted Che
plants' resistance to low temperatures decreases sharply and when they are
planted too early they suffer severely from freezing.
It has been explained that wi[h early planting times the plants from
vErnalized seeds enter stage IIZ of organogenesis in the autumn and strains
oE souther selection (Odesskaya Selection StaCion) complete vernalization
more rapidly than do strains of Mironovskaya and Khar'kovskaya selection
and therefore are more severely damaged.
I. I. Tumanov (34J also noted that the capability of tempering in
vernalized plants decreased apparently because of the fact Chat their growth
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wng more intensive. xt w~~ ~lmngti ehe rul~ rh~e plgnCg from vernnlized
~~~dg grew m~r~ r~pidly ~nd eh~ lpgve~ wer~ langer; I. I. T~mgndv did not
Check Ch~ eondieion oE th~ veg~e~eiv~ ann~. -
[n Che w~nC~~ o~ 196ji/65 we ~tudied tlte wintering of five gtrains o~ winter
wtieut: 'U1'ynnovkn~ Mironnv~knya ~64, wheaC-when~ gr~gs hybridg 1~6 ~nd 55~
nnd Be~d~Cny~ 1, I'lanein~ wag don~ with vernaliz~d ~nd dry ~~edg nC
fnur tim~~: early for Mogcow Obl~gt--5 gnd 15 Augugr; opeimnl--2S Augu~t;
nnd l~t~--10 S~pC~mb~r. W~ al~~ p1gnC�d Ch~ Vy~ek~ ~tr~in n~ wineer
ry~. `1'h~ winCer nf 1964/65 hnd a lot of ~now ~nd ehe winrer cropg--nnC
nnly Choge plane~d nt the e~rliesC Cime, 5 August, but ~1so thoge pl~nred
t~n 15 Augugt--wer~ significgnely dgmgged even by th~ end bf ~ebruury.
tn the gpring Che plunts plgnted a~ ~tl, eimes with v~rn~liz~d ~~edg h~d
u11 di~d nnd eh~re w~ ine~ngiv~ developm~ne of gnow mdld nn Chem while
70-9U percent c~f the contrdl pl~neg, plgnt~d on 25 August ~ttd lU Sept~mber,
gurvived Che wittter.
The resultg of this experiment convincingly prove thae after the completion
of vernalizarion procESSes there ig n gh~rp reduceion in the plants'
resistance Co dam~ge under Che enow.
With inten~ive growth of the vegeeative cone even during 1-2 months, there
is nlsc~ a sharp reduction in Che plantg' resigtance Co perighing under a
snow ~nd ice cover, which is cle~r from Che data of 1970/71 (Fignre 7)
~10~� '
uX ~
, v
ef
_ Bo ~ ?
x 3
o y
~ S
60 ~
0
~ x
" o
.
�o.
?0 .
.
~i i
0 Qf Gl? t xM
Figu!-e 7. Dependency Between Thinning (7.) of Winter Wheat of the
Mironovskaya 808 Strain as a Result of Freezing and the
Ice Crust and the Length (mm) of the Vegetative Cone
Thickness of ice crust: 1) mm, 2) 10-20 mm, 3) 21-40 mm, 4) 41-60 mm,
5) minimum soil temperature at deptli of tillering nodes -15 --17�C:
lg u= 1.5277 lg 1+ 0.3694 lg m+ 2.0360. _
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;
In Che winCer o~ ~.~70/71 Che wi,nCer haxdine~~ o~ ~ighe gCraing of winCer -
whe~t (U1'y~novka, I~PG-18G, Mixnnovgkayg 808, Yubi~,eyn~y~ S0, 8ezdgC~yg 1,
Avror~, Itnnny~y~ 12, San-p~~Cnr~, '~aezium 39 gnd ehe Vy~tk~ str~in df
winCer rye) w~tg investigaCed on Che experimenC~l plot of Moscow SCaee
Univ~rsiry. Th~ plnnCing wes done nC eight Cimes, b~ginning extr~ enrlv
(f'rom 29 Ju1y ehrougki ~ Oceober 1970). Three conditinng of Che snow cover
wer~ cre~eed: 1) nnt~r~l--10-2b ceneimeters; 2) excegsive--up eo 60-70
ceneimeters; ~nd 3) wirhnut sndw from 0 eo 2 ceneimetiers. ~ach dgy Che
minimum eemperaCure ne ehe deprh of Che Cillering node w~g tnken w ith AM-17
remoee cc~nCrol ehermomee~r~ and ehe depth of the ~now cover ~nd the depth
of freezing of the goil were meagured. In the wineer the condition of the
vegeC~CiV~ cone wgs deCermin~d and the plantg were ehen grown in a
gre~nhouse with gub~~quenC cheCkin$ of the vegeCgCive cone. At Che ~ame
~ime g~mple~ nf whege nnd rye �rom various regione of ehe US5R were
invesCigaC~d: in December, January, February and Che firsC of March.
Inform~tinn abost th~ condiCion of planCed areas wgs processed and
romp~red with agromeeeorologicnl conditians of the wineer and the resulCs
of ~feergrowth and ~pring investig~tion of the planted areas.
. Th~ winCer of 1970/71 was unusual in the European parC of the USSR. The
snow cover in the northern belt of the CerriCory (northern Vil'yus, Bryansk, -
5aratov, Ural'sk) was estublished very early--in the Chird ten days of
_ October. The soil under it was thawed or slightly frozer.. In the second
ten days of November, ns a result of very warm weather, the snow cover
dis~ppeared and the soil thawed everywhere except for the northeastern '
nbl~sts and the Ural area. A slight snow cover (2-5 centimeters) remained _
on the fields in the ma3ority of western and central oblasCs Chroughout
the winter. The minimum temperature at the depCh of the tillering node
ranged f rom 0 to -10�C.
Even in the auCumn the inCensive growth of the vegeCative cone in OcCober
led to a reduction in frost resistance. A comparison of the sizes of the
vegetative cones of Che main shoots of the plants wit.h the agrometeorological
condiCions of wintering of winter crops showed that they have a great
- influence on the change in the sizes of the vegetaCive cones during the -
winter [18].
tn the plants planCed at optimal times, as one can see from figure 8, the
length of the vegetative cone on 20 February had a sufficiently well
expressed direct linear dependency on the minimum Cemperature of the soil
~~t the depth of the tillering node t3 and the depth of freezing of the
soil H. Tl~e coefficient of the correlaCion between the sizes of the
vegetative cones and the freezing of the soil was 0.74 � 0.03.
Analytically the dependency between the length of the vegetative cone 1
oE the main shoot of plants that had bushed normally in the autumn, the
minimum temperat~lre of the soil at the depth of the tillering node t3 `
and the depth of the freezing of the soil H on 20 February, according to
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daea frnm the winrer o~ 1970/71, is expressed by the following mulriple
regreesinn equnCions: for Vyatkn winCer xye -
1 = 0.0'15t3 - 0.006H + 1.377~ (2)
I M~
~~5 -1 b=-O,OOOSH-Q,037J C3�0,4.939
� I! 1=-0,0060H-0,01Jt3+1,317 -
- ~ �ao� ~
~,o ~ 0 2
x3
0 0 ~ ~ �4
8 ~ �i �
o ~
0 O p � �
O �
~~5 ~ 00 C �
~ ~ o o ~
v~~ ~ x o o i �
v3'yyx~~~~xx ~ �
. \W y X �
v ~ xv~~;� vxx x Y.x~1? �ll
L'~ r ~ l
a~'~.l
0
SO f00 ISO HcM
_ ~ ~ ~ , _
- p - 5 -10 -/S EJ C
Figure 8. Dependency of Length of Vegetative Cone 1 on Minimum
Soil Temperature at Depth of Tillering Node t3 and
Depth of Freezing of Soil H on 20 February
= 1, 3--winter wheat; 2, 4--winter rye
_ for winter wheat ~f the Mironovskaya 808 strain -
1 = 0.0370t3 - 0.0005H + 0.4839. (3)
The coefficients of the multiple correlation are.0.94 � 0.01 and 0.91 +
- 0.02, respectively; the standard deviations of the equations m are -
f 0.013 and 0.028 mm. The equations are valid with minimum soil temperatures
at the depth of the tillering node of 0--10�C and with a depth of
Ereezing of the soil of 0- 150 centimeters.
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I~UIZ ~11~ i~ I C; I 111~ tl;;l: t1NI~Y
AK onc cnn ge~ from rhe equnCions ~nd Gurveg in ~igure g, the higher ehe
minimum temper~ture n[ the ~nii gnd th~ l~~ger the d~peh of fre~zing of
th~ ~oil, Chp longer eh~ v~g~t~tive cune nf pl~ne~ Ch~t h~ve develaped
ndrmall.y in the nutumn.
An unttlygig of Che ~prin~ inveatiggeion of tihe plnnCed areag, from which
p1~nt ~pecimeng w~re t~ken. ~nn~irm~d eh~; Che intett~ive ~rowtih of rhe
veg~tntive rone in eh~ ~ueumn ~nd winrer. reduces Chp fro~t reei~e~nce uf
the pl~nts. The Chinning of winCer wh~~C ~nd winCer rye wgs congiderably
grenter in pln~:es wher~ Che gizes of the vegeCative cones were significantly
~re~ter than usuaL. ~
'The fro~t regigtnnce o� whe~C, ~cedrding Co daCa from obeervaeione in ehe
winC~r nE 1970/71 frnm m~tporologicgl stationg locgted in the cenrral and
e~gtcrn oblaets of the ~uropenn parC of Che U55It, cnrreldtes wiCh ehe
lengths of the vegeeative cone. The coef�icient of correlgCion between
thinning and Che sizes of the cone on 20 ~ebruary was 0.71 [18).
l~:~sed nn datn conCerning Che reverse dependency between the inCensiveness
of growth proces~ea and the degree o~ hardiness of the planCs, in the
~xperiment wiCh Mironovskaya 808 winter wheat epecial attention was
devoted to an analysie of daea concerning the length of the vegeCative cone
and the connection between Chis indicator and the degree of thinning of the
plants. From Table 26 one can see that here too Chis dependency was
Clearly confirmed.
'Cttble 26. Thinning of Mironovskaya 808 Plants, Depending on LengCh of
Vegetative Cone with Various Planting T'imes and I3epths of Snow
Cover =
~ ~1~ ~2~ G~e~cnwfl no~po~
_ Cp01fN C'3> t~TttT~tNMYR ~4~ N]blff'O~MYR
tt~l
A MOM)'Gl ?'e N~pe� /1 kOXrtl X r~pe�
5 (MN) :lilMIIOGTN 5 4N) 71ttMMOCTN
~
29 VII 0,9 62 1,1
11 VIIt 0,6 60 1,0 84
25 V111 0,5 50 0,T &4 '
3 IX U,4 30 U,5 45
18 IX 0,3 0 0,3 10
~
Key:
1. Planting times 4. Excessive
2. Snow cover 5. h of cone (mm)
N~tural 6. % of thinning
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,
Simil~r d~e~ were abr~ined wieh oeher strains a~ w~1J,. Moreover in the
e~r1y rip~ning, 1~gg winter h~rdy sCr~ina (nezngrgya 1, Avror~, Itgnny~yn 12,
5nn-pn~Cnre), especinlly wieh ~arly planting Cimes, th~re was more ineendive
- growth of Che veg~CnCiv~ cones and~ correspnndingly, considerably greater
~ desCruerion of the planCg.
T71ug the datg obtiained in ehe experiment and Che resulCg of the analysig
of n?~g~ obgervaeions of the aondiCion of winrer wheae and winCer rye ~r
mpCeorological geaeion~ ghowed th~C eh~ winter hardine~g of wineer whegeg,
boCh under the condieion~ of ~ norn~ai gnd an excege snow cover, glong wiCh
other phygiological-biochemical processes, depends on the condition of Ch~
veget~tive cone. The quantiCaCive d~pendenciea of the Chinning of planted
aregs on rhe harmful effecta of unfevurable winter conditiong confirmed ehe
grear significance of growCh processes during rhe period of the gutumn and
winCer and the gize of the v~getaCive cone whtch, in turn, is determined
by the plnnting eime and the agrometeorologic~l conditions of ehe aueumn-
winCer period.
Ttie quesCion arises: does increased growth of the vegetative cone when Chere
� is a prolonged, wgrm ~ueumn ae a Cime of prolonged thaws always and in all
years lead to the denth of ehe planes?
- As our research showed, in these cases one should diseinguiah, on the one _
hand, types oE growth of the vegetative cone and agrometeorological
conditions in which pr.ocesses of the growth of the vegetative cone takes
place and, on the other hand, temperaCure conditions in the second half
of the winter and in the early sprint~. An anulysis of the vegetative
cones showed that iC is possible to distinguish four typee of sutumn-winter -
growCh of the vegetative cones of winter crops.
'I'he first Cype of growth of the vegetative cone in the autumn and winter
(figure 9) is more widespread in all regions where winCer crops are cul-
tivated. In all years before the temperatures at the depth of the tillering
node stabilize below -5�C, the growth of the vegetative cone in the auCumn
is of the first type. The longer Che period of positive temperaCures, the
greater the size of the vegetative cone. Moreover in the ma~ority of years
the growth of the cone takes place in stage II of organogenesis and is
measured initially in millimeters and then parCS of a millimeter. In
the second half of the winter when the temperatures at the depths of the
tillering node are close to -5 --10�C, Che growCh of the vegetative
cone continues almost until spring, although it is insignificant and
retarded. If during the winter or early spring period the plants have not
been sub,~ected to the effects of temperatures that are critical for the
strain, usually in the spring when the average daily temperature rises
to +3 -+S�C the vegetative cone very rapidly moves into stages III-IV of
organogenesis, the rudimentary spikelets are differentiated and the plants
develop normally in keeping with the agrotechnology of planted areas and
the course of spring-summer meteorological conditions. -
J
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8'
~
>.i
` t,y Cj~!{
~C's~,.'~"+ I '
i.,~~ � ar~+~fA~�
. T.V'~ , ~i~IS1~~S_.;
- Figure 9. F'iret Type of GrowCh of VegeCative Cone of Winter Wheat
Key:
a) at beginning of autumn growing period--stage I of organogenegis;
b) before enCering winter--stage II of organogenesis;
c) at beginning of spring growing period--planCS' transiCion to~stage
III of organogenesis.
Insignif{cant growCh of Che vegetative cones according Co Che firat typea
within stage II of organogene:ais is a widespread phenomenon, especially
in the second half of the winCer. As the research of V. V. Vinogradova
- ahowed, during this period there ia a sharp reduction in the content of
growth inhibitors in the tillering nodea. It is almost a rule Chat the
firat type of growth is not a factor which causes thinning or death of
winter crops.
The second type of growth of the vegetative cone (figure 10) is typical
for the conditions of a long warm autumn. It uaually takes place with
very early planting times on well fertilized fallow fields, especially
with strains whose growing plants quickly pass through the vernalization
processea with comparatively high temperatures of the air and the soil
at the depths of the tillering node. In these cases Chere is not only
increased growth of the vegetative cone in stage II of organogenesis,
but in plants in whici~ vernalization processes are being completed, there
ia a transition to stage III and in individual years, to the beginning of
stage IV of organogenesis. We have repeaCedly observed cases like these
at strain teating atations of Krasnodarskiy Kray, Rostovskaya, Odesskaya,
Khersonakaya.and Krymskaya Oblasts, in the Dagestan and Kabardino-Balkar
ASSR's, and in individual years in Vinnitskaya, Poltavskaya and Khar'kovskaya
= Oblasts. The same cases of the plants' transition to stage IV of organo-
genesis in the autumn of 1974 under the conditions of Voronezhskaya Oblast -
were reported by the agrometeorologist N. G. Kolesnikov and the physiologists
of the Mironov Scientific Research Institute of Selection and Seed Growing
of Wheat, I. V. Moroz and N. A. Kryahevich. A transition to stages III-IV
of organogenesis was observerl in experiments of the laboratory of the
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binlogy of p1anC dev~],npmenC in Ch~ auCumn on gxegg planeed in Bezd~e~yg
1~nd ddeggk~y~ S1 winCer whenC at early Cim~g in n~es qu~ntitieg,
.~t`
0 1.
, r
- ,
i~
'.a. -
fY:4bR ~
~ ~ ~F
1
~ ~7".
i 7y~~
~igt~re 10. Second Type of Growth of VegeCaCive Cone of
Winter WheaC
Key :
a) at beginning of autumn growing period--sCage I of organogenesis;
b) in February under deep snow cover; at beginning of spring growing
period--plants' transition to stage III of organogenesis.
If the Cemperatures in the winter period and the early spring do not reach
critical levels and there is no freezing of winter crops, the plants that
have pass~ed through the winter in the condition of stages of III-IV
form a large spike wirh an increased number of multiblossom spikelets in
the early spring. In such years areas planted early in winter crops
under the conditions of good agrotechnology realized to a considerable
degree the potential productivity of the strain. The yields of winter
wheat and eapecially of winter rye are record high.
But if after a long warm autumn before the snow falls there is a sharp
reduction in the air Cemperature, the soil fro:ezes rapidly and deeply .
and the tempernture a~ the depth of the t{11_zring nodes and in the zone
where the vegetative cones are located (above the tillering node) drops
below crieical, the main shoo~s and even whole plants die. Plants in
stages III-IV of organogenesis are usually poo~ly tempered and for them _
the critical temperatures are significantly higher (2-3�C and more) than
for well tempered plants whi~h are in stage II of organogenesis. There-
fore if in the autumn plant~ appear whose growth takes place according _
to the so-called sec~nd type, the fields with these plants must be kept
under conatant attention, especially if minimum temperatures in the soil
at the depth of the tillering node approach critical level.
. 53
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7'h~ flrst type d~ growCh o,� Che veg~eative cone (figure 11) is mor~
fr~qu~nCly abg~rved ~nd mainly in the winCer period in thd~e ye~rg when
Che ~now fa11g early on ehgwed or slighely frozen soil. '1'he eemperatuYe
remgine aC a lev~l of 0-~ 2�C under g deep snow cover for a long time.
- Under ehese condirions in th~ darkn~sg of vegetaeive con~ grows rapidly
in lengtt~ but does not move into atage IIT of organogenesis. There is
u coneiderable elongatiott of the vegetative cone~ a eo-called undifferen-
tinted growCh of the cone in stiage II of organog~nesie, a kind of procees
- nf prolification of the vegetaeive cone. The lengeh of r.hese cones
_ frequenCly increases 2-3-fold over normal (figure 12).
a e
, , ,
1'~'
~
_ J
.*Y~ ~
y . ~a
' 1., r ; , . i,
r. ; ;t `t'~'' '
�1 ' 3t~
,.f `a~. ~ t- ~ i
L;, k, '
+'~.k^~FR ~ =qr .
, . ' �r
� ~ ;i',a~ ~~~:y
~?,~,~�i:`..c`-`?l,~'y~'
:~;~~%i~,,t}~~ ,}'a~
. :"~ti~.~'l:.a ~ .
Figure 11. Third Type of Growth of Vegetative Cone of Winter
Wheat
Key :
a) at beginning of winter after snow has fallen on thawed soil in sCage -
II of organogenesis; b) in the middle of winter--under a deep snow cover
in stage II of organogenesis; c) before the disappearance of the snow--
at the end of winter "prolification: of undifferentiated vegetative
_ cone.
This kind of growth of the cones is observed when the plants are perishing
under the snow cover. They frequently die during the second half of the
winter from starvation when there is a snow cover for an extended time, but
_ more frequently in the spring when, being in a condiCion to enter stage
III of organogenesis, they are first sub~ected to an attack of fungi that
are changing from saprophytes into a parasitic form. Frequently these -
shoots weaken the entire plant and sharply reduce thei: resistance to
frosts. As a reault the plants die even with the return of relatively
froat in the early spring.
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~d(t d~~ICTAL U5~ dNLY
A
M , '
a;
t�~
i
i+ 1
, ~ 5
{1
!
~Y d 1
~ k: "
~ i~/iit
.
~
S
J ~ ~T ' , ~
~
~ �~Y' lv:~ f,~
!h.,' "s'~,~~:
, , . ~r;~ ~ ~`h,"
a. t . . S~ ~,~p
Figure 12. Third Type of Growth of Vegetative Cone of Winter
Rye '
Key:
a, b) at end of long auCumn growing period--ins:ense formation of leaf
vallicula, stage II of organogenesis; c) in February under deep snow
- cover; d, e) at beginning of spring growing period--stage II of organo-
~ genesis--"prolification" of undifferentiated vegetaCive cone.
The undifferentiated growth of winter plants, first noted by F. M. Kuperman
as early as 1951-1954 and then repeatedly observed by F. M. Kuperman,
V. A. Moiseychik and M. S. Bykova [10], is one of the leading causes
(in addition to hydrocarbon exhaustion and disturbance of protein exchange)
of the death of winter crops under a snow cover. Perishing under a snow
cover is observed especially frequently on areas planted at early tim~s
and most frequently in winter rye whose vegetative cone can grow at a lower
+ temperature than that of winter wheat can.
We discovered the fourth type of vegetative cone (figure 13) in Mironovskaya
808 winter wheat. It is clearly distinguished from all of the aforemen-
tioned types. It is known that in the ma~ority of strains of winter
wheat which we investigated, like Kooperatorka, Bezostaya 1, Kavkaz,
U1'yanovka, Odesskaya 51, Priboy, the wheat-wheat grass hybrid 186 and
others, the shoots coming out of the tillering nodes each form one vegeta-
tive cone which is subsequently forn~ed into a spike. The noints of
growth of the shoots of the second order in the axils of ruc~imentary leaves
(leaf vallicula) in the ma~ority of strains of winter wheat die at the
. beginning of stage II of organogenesis and thus a shoot is formed with a
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gtnik ~erucCure ehat ig epecific fdr gr~~gee--~txgw. In Ch~ m~~oriCy df
wine~r crnpe br~nching eakem place dnly in the lnw~r nodes under ehe ground
and in th~ eo-c~11ed Cillering zone~ When Chexe is dgmage (~or var3,nug
reason) of Che veget~eiv~ cone, Che ahooe, d~prived of g normal vegeeaeive
cone, diee in eCages IV-V. Yn strains of Mironovgkaya 808 ~nd also
cerCain'gtraine obCained ehrdugh hybridizaCion wiCh Mironovskgya 808,
in the axile of the leaf vallicula when the ahoota are moving inCo at~ge
Ii of organogeneaig, growth pointe of the second order gre ~stabli~hed~
~y the e~4d of the auCumn growing period ehey form cle~rly expreseed
vegetative cones in st~ge I of organngeneais, or, gs M. Kuperm~n c~llg
them, "reserve buds" (figure 13). The eignificane contene nf growth
inhibirora in the vegetative cones of Mironovskaya 808 leads to a
situgtion where growth ie rnCarded farly in the vegetative coneg and
frequently when the temperaturee at eh~ depCh of the tillering node are
-8 --10�C it practically s~ope. The retarded growth of eh~ vegetaCiv~
c:ones is compared ta BezosCayg 1, Avrora and Odesskaya 51 is gleo typical
of Mironovekaya 808 ge Cemperatures close Co 0 or t 2�C. This explaing
- the compreheneive froet resistance of thia strain which is registant
both to low temperaCures and to an excesaive gnow cover.
~ `.l
y~
r�
~
Figure 13. Fourth Type of Growth of Vegetative Cone of
Mironovakaya 808 Winter Wheat
Key:
a) vegetative cone of main shoot (stage IV) and "reserve" vegetative
cone (stage I) at beginning of spring growing period; b) base of
vegetative cone and "reserve" buds.
56
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Th~ retgrdaridn ~nd ehen glmoee ~ompl~ee h~lting o~ rh~ growth df ehe
t~rminnl vegeeativ~ cone l~adg eo aceivgCion af ehe grawth df ~id~
reserv~ budb which come out nf e eCae~ of dnrman~y in th~ ~uCumn ~nd gp~nd
eh~ winCpr itt ~Cege I of organog~npgis. Ag on~ can ~ee from th~
phoedgr~ph (figure 13) Cnken by A. S. Yar~~hev~l~~ya t~ti.th a acanning
~1pcCron microecope. they ar~ locaepd gt the bggeg of vegeCgtiv~ con~~ df
autumn ghonCs. In th~ nuCumn ~nd wineer Ch~y grow ~xtremely glowly ~u~e
ue ehey db in plgnCe with very 1a~~ planeing Cimeg ~nd ae the ~ame Cimei
gg di~tinc:t from the dnrmgnt budg of Che eill~ring nodps, vernalizgCidn
processes take pl~ce in Chem. ~
Thus vegetgtive coneg with diff~rent d~gree~ of fxogt r~ais~gnc~ gre
combined on one planr and on ehe game ehoota. During yearg when rhp
vegeCgtive conpe of gutumn ehooCg wineer norm~lly they chgnge ov~r ~arly
to differentiatinn of Ch~ rudimenCary epike and form apike~ wiCh high
_ producCivity. During thege y~grg the vegetative coneg of Che "regerve
buds" which are more Chan ewo etggeg behind, die withouC forming regroweh
when Ch~ upper veget~eivp cone moveg into gCage V of organogenesig. BuC
in years when Che upper vegetative cones of gutumn shoote are damaged by
Erogtg or "prolificnte" when there ia an excegaive snow cov~r for a long
time~ the vegetative cones of the "xeserve" buds d~velop r~pidly intd
~ productive shoota and regCnre Che plant stand that was r.hinned during the
winter. 7'hese ehoots from theae vegetaCive cones, like late plantinga, -
` produce a somewhat smaller yield, but because of Che peculiar properCies
of the growth of the vegetative conea, reatore the producCive plant stand
without replanting or under sowing and suppreas spring shooCs of weedy -
plants.
The discovery of four types of growth of vegeCative cones during the
autumn, winter and early spring periods made it possible to refine the
essential differences in the nature of Che damage and death of plants
from freezing and from harmful conditions under the enow.
When plants perish under the anow, as V. A. Moiseychik notes [22J, the
temperature of the soil at the depth of the tillering node is high, a
deep snow cover is established early and the soil is noC frozen to a
greaC depth. All these conditions contribute~~ to the appearance of the
third type of growth of the vegetative cone ~rhich is characterized by
undifferentiated elongation and "protification" in st~ge II of organogenesis.
Perishir~e under the snow is a lengthy process; intense growth of the
vegetative cone, like the death of winter crops, takes place if favorable
conditions remain for no less than 80-100 days.
Conversely, freezing of the plants is observed when the temperature of
the soil is low and when there is no snow cover or when the snow cover
is shallow and the soil is frozen to a great depth. The growth of the
vegetative cones is very retarded. Under these conditions the first type
of growth is encountered most frequently. Damage to the cells by the
~ low temperatures and ~ne rupture of the tissues lead to a rapid dying off
57
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,
~dit O~~ICtAL U5~ ONLY
o~ th~ v~g~C~eive ~one~~ Af~er the ~v~~~g~ e~mp~~~eur~~ ri~~ e~ ~�C
the d~~d ti~~ue~ of the v~g~t~tiv~ con~g mg~ex~~~ (figur~ 14) gnd th~
plantg whieh are d~mgg~d by th~ fruer~ ~r~ frequently ~ub~~et eo gte~~k~
oE fungi--gnow mold, cnll~r rot and ~o ~orth.
Figure 14. Necrosie of VegeCative Cones of Winter Wheat Damaged
in Winter and Maceration of Tissuea
Key :
a, b) vegetative cones of different sizes; c) top of vegetative cone
(micro-photo-scanning electron microscope)
Freezing, as distinct from periahing under the anow, is mogt frequently
a short-term phenomenon and the plants die within 1-3 days. The effects
of low temperatures are especially dang:.rous for planted areas on which
the second type of growth has already t~een noted in the fall (figure 15).
Tf~e freezing of these plantQd areas takes place at comparaCively high
degrees of freezing. Por such planted areas the alternation of long
thaas with freezing ~seather is especially dangerous. This frequently
takes placc in the soutern ateppe regions of the Ukraine and not nnfrequently
in the central nonchernozem regions. The death of winter plants ia
obaerved in the second half of February and the beginning of March.
In the central regions of the nonchernozem zone damage and destruction of -
areas planted in Winter wheat are caused by a complicated cotnplex of
unEavorable weather factors (figure 16). Low temperatures in December
and Jan~ary when there is alaast no anow or a light snow cover and be
the cause of freezing of the planta. And, conversely, the early snow
in Uctober and an excesaive anow cover for a long period of time and
almost the same kind of snow (which ia eapecially frequent in the no~th-
enetern regione country's European territory) can lead to periahing o~
the plants under the anow.
58
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~d[t n!~'t~'I~IAL US~ dNLY
t,ti,ww
~ ~~.r~~~s.~~.~..a~+e~.~~~+~~'~~~'e"""
~
~ �r
~
,f'~'lb~,-~;r,,,;,r".."~i,~�:~''
~
0,4 t~'~,~,~a_--~t~,._
~
_ ~ ~ c�4
_ ?0 + 2
0 0 ~
~ `~.i i~j -2
-w- f
y~ -vw 2 ~ '4
~ ~ / -a
~ ~ y ~ . . -
e . , a
ao ~ s 9 '
~a~ ~ s m a
Mtw /p
X ~l Xll l !I lll
Figure 15. Growth Dynamics of Vegetative Cones of Winter Wheat ~
Planted at Optimal Time Under Conditions of Natural 5now
Cover, 1972/73, Moscow State University
Key:
Soil temperature at depCh of tillering node under snow cover: 1--
"natural", 2--"excesaive"; depth of snow cover: 3--"natural",
4--"excesaive"; de~th of freezing of soil under snow cover: 5--
"natural", 6--excessive".
Strains: 7--U1'yanovka, 8--i~ironovskaya 808, 9--Odesskaya S1, 10--
Bezostaya 2. lkn--length of cone (mm); h--depth of snow cover (em);
11--depth of freezing of soil (cm).
iMder the conditions of Ryazanskaya, Kalininskaya, Novgorodskaya,
Moscow and adjacent oblasts there are frequent Winter thaws With precipita-
tion in the form of rain and wet snow; vhen the freezing weather arrives
- they lead to the formation of an ice crust. In the nonchernozem zone,
especially on i[s southern border, although it is rare there have been
c~ses of early disappearance of the snow (at the beginning of March) with
a subsequ~nt return of ehort-lived but severe frosts.
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~dtt d~rtCtAL USL cINLY
t~~,ww
,r'
0~8
r'
Q6 :+%-...~.,_,.v~_
,~w~..~.~
~
~y~ :ft,..e.r...arr~:.~~ttiil~..s~.~'~'~'"'~
1',r+~y~
,.!".rL'-~._r--.a~_`:::..':-::~..=..`
.:.aaa.~~.
/y~ ~
h "l~~~rwrYiT:L.:..L'=.`.�`�~ ~,...~~~r....~~~~~.+~~i~ t ~
0,? ~0 . +r..r*��.u1-ny.~-~u,u!n..ui '~.....�-,t 4
~I i ,
~ ~ i
1 , I 1 .s, ;
~ ~
0 0 ..~...s ~...,,~i. ~
i .~r~~ ~ ~ �-...-.ti�.,
zo ~ 2 , ~iI"~..~.;~,;
~ ~ ' , % , ; -4
~ ~ , - ~ ~%i;.
< ra , -s
- ~ S . . ' .
ao s r~ -s
~~o ~ : ~
NG? ~ fy ~
.C JU ,Sli / l/ Il!
~igure 16. Grawth Dynaroi~g of Vegetative Cone of Mironovskaya
808 Wintpr Wheat with First-~ourth.Plgrtting Times and
Varioug Depths of SnoW Cover (WinCer 1972/7~).
Moscnw 5tate University.
Kpy:
Conv~ntional gymbols lk , h and H, and also 1-6 see fiigure 15.
."Natural" snow cover: ~}--first planring time, 10 Angust; 8--second
planeing time, 25 AugusC; 9--third planting time, S 5~ptember; 10--
fourth planting time, 20 September. "~accessive" snow cover: 11--
first planting time, 10 August; 12--second planting time~ 25 AugusC;
13--third planting time, S 5eptemb~r; 14--fourth planting time, 20
5eptember.
In individual years (such, for example, as the Winters of 1973/74 and _
1914/75) there is very little snow on the fields in the central chernozem -
oblasts nnd the winter crops spend the winter in temperatures nf 0 f 3�C.
~i'he short~ge of aarmth limits photosynthesis and the differentiation of
the vegetative cone. At the same time the cones become elongated and
"prolificate." Plants with these cones are infected with root fungi in
che early spring.
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tn Ch~ cenCrn]. reginn~ of Ch~ nonCh~rnnr.em ~on~ in individu~l peridds of
th~ winCering of p1anC~ dne ~~n dbs~rve eh~ ~ff~~t~ nnt only of individu~l
C~rtnr~, bt~r of an ~n~ira cnmpl~x nf them. The diversiey nf fnCeorg ChnC
c~u~~ dnm~g~ ~nd d~nCh to ar~~~ plane~d in wine~r Grdpg m~d~ it n~cegynry
in this zone to aearch for atraina which would hbv~ ~ compr~hensive
r~~i~tnnce en low C~mp~raCuxeg, close to criCtc~l, (~nd in whiah eh~
grdweh o~ eh~ v~g~e~eiv~ can~ i~ b~gic~lly of eh~ ~irge Cyp~) gnd ~1~0
td t~mp~r~ eur~g of gbout 0~ 1�C.
In nrder to egegbligh m~xim~m winCer hgrdin~s~ in rhig znne, it ie
impdrtant to hgve gtrains with th~ first Cype of growth with a wide r~nge
af t~rnp~raeur~~ from 0 to -$�C. But the mg3ority of etrnins previougly
rpginn~liz~d in Chig xdn~ havp b~~n ch~rg~C~riz~d by eh~ ehird ~nd ge~and -
typ~g nf growth. Ie w~~ importanC Co find duC whi~h w~re chnr~Cterized by
one type df grdwCh or ~noth~r ~nd th~ n~C~re nf ehe grnwth dynamicg nf rhp
veget~tiv~ cone,und~r vnrious winterittg adndieiong.
Tnking into ~cCOUnC the diversiey of Che complexeg of un~avorgble wintering
C~nditinns, Che l~bor~tnry of biology o~ p1~nC developm~nt in conjunction
with the U55it Hydrometeorologicttl Center during I970-1975 (5-10, 16-22]
conducted morphophygiological research of the winter hardiness of winter
crop~ s'multnnenusly egginsC vgrious backgrounds, the mg~or ones of which
were: n) a natural ~now cover whose depth usually rangtd fra~m 10 to 25
centimeters; b) gn excessive snow rover wirh a depth of 50-60 centimeters
nnd more; and c) so-called ennwlegsne~s or more precisely liCtle anow, with
~t snow covpr of no mc~r~ than 1-3 centin~eerg. In all years the planCing
_ wag done at four times; the firsC, early--5-10 AugusC; the second, opCimal
for Moscaw Oblast--20-25 Auguse; the Chird, medium lgte--5 ~eptember;
- and the fourth, very 1gCe for these regions--20-25 5eptember. AgainsC these
backgrounds they tested strains that were regionalized in the USSR and
also a number of promising new atrains of winter wheat: U1'yanovka,
Mironovskaya 808, Bezostaya 1, Bezostaya 2, Mironovskaya yubileynaya,
Odesskgya 51, Priboy, Avrora, PPG-186 and Vyatka and Khar'kovakaya 60
winter rye (lOJ.
In ~11 the variants of the experiments AM-17 thermometers were used to
tuke the minimum, maximum and periodic temperaCures of the soil and the
depth of the tillering nodes; frost meters installed on platforms at
various depths of the snow cover were used to measure the depth of
freezing of the soil and snow measuring rods were used to determine the
depth and uniforn?ity of the snow cover on the plots. The rest of the
necessary meteorological data (air temperature, precipiCation. number of
hours of sunshine, moisture content of the soil and others) were obtained
in the observatory of the geograph-~ department of Moscow State University
Which is Iocated 50 meters from the experimental plot of the laboratory
of bioloty of plant development.
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I~n[t OI~i~'ICIAL iJSL nNLY ~
In rhe nuCumn nnd spri,ng the number n� pl~nrs and ehoots were cdunted on n11.
n[ Che plntg.
'I'hroughour Che enCir~ nueumn-winCer-spxing pexiod every 10-15 days, n
pneumnCic dxill wieh u sp~de wus used to Cnke samples of 25-50 planes
frnm enCh varianC of Cti~ experimette. They were sub~ected to a deCailed
morphophysio].ogical analyais 3n which the condition and eizes of ehe
vegeCaCive cones, Che Cillering nodea and the lenves were eaken into account,
The vi~biliCy o� the pLants wgs Cegeed itt n greenhouse. To do this the
veg~el~'~qf, Che plgnts (5-10 from each apecimen) were forced to the phase '
- nf hegding and b lossoming. Then in the qreenhouse observaCions of rheir -
p~sging ehrough sCages II-TX of organogentnis were conducted regularly.
In recenC yeara mn~or atCention has been devoted to an analyais of groweh
proce~ges of Che vegetative cone uttder Che condiCions of various temperatures
in the ~utumn-winter-spring period and Co rhe discovery (wiCh the help
of n micrngcdpe photographic aCtachment on an MSS-1 binocular magnifying
glass and nlso g scanning electron microscope of the Khitachi 11-B type
with various magniCudes--from X70 to X1600) of differences in the types
of damttge from f reezing and from perishing under the snow.
Data from observations of the dynamics of Che gt'OWrh of Che vegetative
cone were of greaCest interest in research on the phenomena of winter
hardiness. Regulur observations were conducted for the first time with
various depths of Che snow cover and various minimum and maximum tempera- -
tures at the depth of ehe tillering nodes and various depths of freezing
of the soil which were created in connection wiCh this. In the first
place, these were observaCions of plants of Che same age, planted at the
snme time, on the same aection and under identical soil conditions; in the
second place these were one-time parallel observations of plants of
various ages with intervals in planting Cime of 15-20 days, beginning with
very early times and ending with extra late times (20 SepCember) for
condiCions of the nonchernozem zone. And finally, which is also signi- -
fican[, we inves tigated for the first time not only regionalized and
promising strains for the nonchernozem zone--U1'yanovka, PP~-186, PPG-599, '
Mironovskaya 808, Mironovskaya yubileynaya--buC also strains used as
indicators of ~!es~ winter hardiness which had been regionalized in the -
southern oblf.sts of the USSR--Bezostaya 1, Bezostaya 2, Avrora and
_ others--and also the semi-winter strains San-Pastore.
Througho~,C the entire autumn-winter-spring period the winter plants were
sub~ected to detailed morphogeological analysis. The determined the
coefff.cient of tillering. the number o~ shoots in various stages of
org:mogenesis, the number of leaves on each shoot, the length and breadth
c~c the leaves and their condition; with a point evaluation they determined
the condition of the vegetative cones, their length and breadth, the number -
of leaf vallicula in stage II of organogenesis and the number of segments
and~ in cases where they had entered stage III, also the number of
spike tubercles in stage IV of organogenesis.
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For mass samples the accounting was done on Table 27 [9]. Throughout
_ Che winter period w1Ch unfavorable conditions~ as a rule, Che leaves were
dr~maged first. The degree of damage large~,y deCermines rhe condition
of rii~ planta in Che spring-summer perioci. Tn rhose cases where the leaves
were not damaged (green or parCially yellow-green) a.~Cer the disappearance
of the snow cover and the rise in Che air remperature Co above 0~
_ phoCosynChesis begins in the plants. The development of these planCs
takes place normally in the spring. The condition of the leaves of Chese
planCs is evaluated by points 5 and 4.
If in the winter and spring a11 leaves or a large part of Chem are yellow
and the vegetaCive cone.s are alive, a certain period of time is necessary
for the appearance of new assimilating leaves after the resumption of the
- ~rowing period, as a result of which the growth and development of the
planCs are retarded and Che yield decreases somewhat. Then the condition
of the l.eaves of winter crops is evaluated with point 3. _
Plants whose leaves are brown in the winter and spring are evaluated by
point 2 and dark brown--by po3.nt 1. In the majority of these cases
there is also damage to the vegetative cones and tillering nodes and
thinning or complete destruction of the plants.
Table 27 gives an evaYuaCion of the condiCion of leaves of plants with
- various degrees of damage by unfavorable wintering conditions~ IC takes
into account not only the color of the leaves, but also the number of
_ plants in percenCages with various degrees of damage to the leaves (Table
27, columns 3 and 4). _
The vegetarive cones have a greater resistance than the leaves do and, _
moreover, in the majority of years in winter crops they are located in the
soil at a depth of 1-3 centimeters. The viability of the shoot, the `
growth and development of the insipient spike and, consequently, its
productivity depend on the condition of the vegetative cone. Therefore
' the condition of the vegetative cone, as was noted above, determines
the condition of the plants as a whole during the wintering period.
- Depending on the degree af their damage in the winter, the conditions of
the vegetative cones can be determined in the following way: five points--
- the cone is transparent, alive, turgid and slightly opalescent; three
points--the cone is a~ive, white, turgid, dull and not opalescent; one
_ y point--the cone is dead, brown, wrinkled and macerated. The condition
of the plants is evaluated taking into account the number of shoots (in ~
percentages of the overall number in the plants that are analyzed) which ~
have varying degrees of damage to the vegetative cone.
The indicators for evaluating the condition of the plants relate-'mainly
to cases of damage of winter crops by low temperatures, the ice crust and
drenching. When winter crops perish under the snow one takes into account
~ 63
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'I'nble 27. I:vnluation oZ the CondiCion of Leave~ (A) ~nd VegeCative Coneg
((3) ~F I'1~n~H in Autumn-Winter-~pring Period.
(1~ .hNCtb�
n p0UlNf p~tTt�
OfIN~~11Ng 011lHK~ ~~~~a~ NMtpWllt 06wNR 6uA
\ 2 ~ ~O~A 1) 1~MN_
_ _ N11 CL1.1 ~ J ~ -
( 4 )
i z ~ `
A
_ Ji?~cTbp aeaeF~de ~6) 5 100 b~100 50p
l00 ~-~-b,0
5 75 (5~75) (4~25) 475
)Ke,iro�ae.~e~~~e 4 25 ~pp ~ 100 ~ 4,8
3eaeF~we (8) 5 50 (5~b0) (4~50) 450
iKe.tiro�ae~e?~me (9) 4 50 ~pp ! 100 `4,b -
4 100 4�100 400
100 100 ~ 4~0
H(earde ~10) 3 ~ (4 ~~100`3�~, � lpp ! 3,5
3 l00 3.100 ~0 3,0
100 ~ 100 ~
- s)'Pae (11) 2 75 ~3'25) (2~75) ~ 225_ 2 3
]00 lpp ~
2 100 2�]00 200
100 ! 100 ~ 1' ~
4epno�6ypde (12) 1 100 1� 100 100 ~
- 100 - I00 ' 1~0
~13~ hON)'G M1pftTlHN~
I1QO~tMT II06!-
_ ~0~ C kOMy~
~~NN R~MM0~0 OqlMK1
tOGTOlIINt KOMrt~ OLLtNK! (S~AA) b~~~~
_ (14) (3) 15 (16)
S s ~ e
6
(17 ) Konyc Hcneoti, T~�proptidA, 5 100 5� 100 500
cncrha onanecut~pya� ~ 1~ - 5,0
utm7
(18 ) To ~ne 5 75
N~nooii. 6eiW~1. TpproP� 3 25 (5�75) (3�25) _ 450
(19 ) nwii. a+yTUafi, ncona~c� 100 ` 100 ~ 4 5
cn?~pyawnti '
(Table 27 continued on following page]
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Koxye H~poct~uHa
Rpouenr node�
fd~ C KOHy OUtHKI
COCTMHM! kONrt~ 04~NIf~ (OIAA) ClMN A~IIN0~0
_ (3) iaao (16)
" 6 8 7 ( 8
s
(20)H(NeoA, Typrnpn~il, ona� b b0 (5,5p) ~3,5p) ~Op
necuHpyauti+fi 100 ~ 100 ~ 4,0
N(eeoil, Gen~rti, Typrop� 3 50
~21) udk, Myrtta(1, uconnne�
cuNpywu~ufi 3�100 300
~(HeoA~ TypropHdA, 6e� 3 100 3,0
~22~ ~dA, MyTHWIf, Heona� 100 100
~18)To *eNPYbwt+A 3 75
(23fylepredA~ 6ypmA, c~iop� 1 25 ~3�75) (1 �25) ~ 250 ~ 2~5
weHNwA, MautepHpoeat~� 1~
H61A
� ~(aeoA, Typropuwb, 6e� 3 50
~22~ cw+~ autNp~,� Neonane� ~3�50) -~-(1�50) 200 2 0
Meprst~A. 6ypd~, cn~op� 1 50 100 100
~23~ nieeNmA, ?tauepupoeaH�
HWA
~22~~(NBO~~ TypropHdA, 6e� 3 25
ndA, M}~THWR, Heonane� (3~25) (1~75) a I50 ~~+5
cuHpyauti~A 100 100
(23~MepTedA, 6ypwA, c~top� 1 7b .
WeHeda, MauepNpoaatt-
NdA 1 � 100 100
(18)To ~ce 1 1~ 100 ^ ]00 �
Key: ~
1. Leaves 16. Evaluation
2. Description 17. Cone live, turgid, slightly
3. Evaluation (points) opalescent
4. Percentage of plants with 18. Cone live, turgid, slightly
- given points opalescent
5. Overall points 19. Live, white, turgid, dull, not ~
6. Green leaves opalescent
7. Yellow-green 20. Live, turgid, opalescent
8. Creen 21. Live, white, turgid, dull, not
9. Yel~ow-green opalescent
_ 10. Yellow 22. Live, trugid, white, dull not
11. Brown �opalescent ~
12. Black-brown 23. Dead, brown, wrinkled. macerated
13. Vegetative cone
14. Condition of cone
15. Percentage of shoot~ with
cones with given points
65
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the numbpr of plants wieh elongaeed vegetative cones ttiae have undergone
"prolific;atton." Here one determineg the overn~.l length nf Che vpg~t~Cive
cones ~nd~ s~paraCel.y. ehe 1~ngCh of ehp elongnted, undifferettti~ted pare
of the vegeeaeive cone. The vegerative cone ig dxawn (if pogsible, with
n drgwing device). 'These d~Ca are glso included in the Cable. An account
is made of euch shoot on eypical planCs. 'I'he degree of deviaCions from
- the norm ~nd Che percentage of markedly elongated vegetative cones ~re
also determined by points. _
One can ~udge Che wny varioua depChs of the snow cover are reflected in
planCS of'bne and the same strain planCed at ehe same opCimal Cime
(25 Augu~r for Moscow Ob1asC) from datn conc:erning Che changes in Che length
of th~: vegeCative cone of Che U1'yanovka sCrain (Cable 28). Thus under
a naturnl snow cover or when there is no snow, by the end of November
1972 the vegetative cones had reached 0.40-0.46 mm and under an excessive
snow cover, when the snow remained on the p1oCs in the beginning of -
November, Che growth of Che vegetaCive cones continued and by the
- beginning of December reached 0.50-0.55 mm. Then in a11 variants with
various snow covers the vegetative cone remained un~changed, which was
typical for tl~e first type of growel~. It should be noted thaC in the
winter of 1972/73 there were no great ~liff~rences beCween the natural
and excessive snaw covers.
- The efEects oE the planting times are very clearly traced in the same
experiment. Thus with the very early planting time of 10 August and
a natural snow cover, the length of the vetegative cone by 25 December
had reached 0.50 mm; with planting on 25 August--0.46 mm; 5 September--
0.35 mm; and with a very late planting time--0.20 mm. Whil~ there were
considerable differences in the length of the vegetative cones deter~nined
by the depth of the cover and the planting times, throughout thp winter
period their sizes increased quiCe insignificantly. As one can see
from the figures, the first type of growth of the vegeCstive cone during
the winter perlod is typical for the U1'yanovka strain which largely
determines the great winter hardiness of this strain which has be~n _
regionalized in a number of oblasts that rarely have conditions which
cause perishing of winter wheat under snow (figure 15).
There was a similar situation during this same winter of 1972/73 on areas
planted in Mironovskaya 808 winter wheat. In variants with various
snow covers the vegetative cones differed little and in variants with
various planting times they differed quite distinctly (see figure 16).
During this same winter of 1972/73. with compazatively optimal. wintering
conditions for winter plants (maximum temperatures at the depth of the
tillering node under a natural snow cover were mainly -2 --5�C and only _
Eor a shorC period of time did they drop to -11 - 12�C), even with the
Eirst type o� growth of winter crops it was still possible to discover
differences in the length of the vegetative cones of winter-hardy strains
which for rhe Moscow area are Ul'yanovka and Bezostaya 808 and relatively
66
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- legs wineer hardy on~s--Od~ssk~ya 5L und eapecially Bezostaya 2(aee
figure 15).
Considerably greater strain differences were noted in the winter of 1971/72
(figure 17). ~ven under conditions of a natural snow cover there was
- a continuation of insignificant, buC steady growth of the vegetative cones.
This took place least intensively in the highly winter hardy strain
Tsezium 39 ~nd the winCer hardy s~rains U1'yanovka and Mironovakaya 808,
and less intenaely in such sCrains as Mironovskaya yubileynaya, Bezostaya
2 and Odesskaya 51 and the mosC marked growCh was noCed in the Avrora
strain. It is interesting that the curves representing the growth of
srrains with varying degrees of winCer hardiness fully corresponded with
their winter hardineas, which shows the existence of cerCain correlations
between the intensiveness of growth processes and the resistance to low
temperatures (the winter of 1971/72 was colder and there was less snow
than in the winter of 1972/73).
l,M My
0.6
Q4 e�~�-�~.:~.�.-�
h c,w .,i
- ~ ~ . _ ,~...~~.?,i�.
~
~�~:e-o:=�-ar��
. ' " ~
- ~ ~~~~~~!i;l l~l: t~C
; . ~ ' 2
0 0
?p ' ` ~ %'i;' ;%i ~ . 0
?
40 p 7 ~
3 ` ~
- ~ - 9 ~ /j~ , ,j j/ 4
60 '
4 J~ ~ \ 6
Bp 11 ~ ~ ~ ' \
~ S ~ \
~ ~ 6 3 ~ ~ ~
~
- x xi x~i i u ui
Figure 17. Growth I?ynamics of Vegetative Cone of U1'yanovka
Winter Wheat and New Strains of Winter Wheat in 1971-72
With Optimal Planting Time, Under Conditions of a
"Natural" Snow Cover. Moscow State University.
Conventional symbols: lkn 1 h, H and 1-6, see fig. 15. 7--Avrora.
8--Odesskaya 51, 9--Bezostaya 2, 10--Mironovskaya yubileyanay, 11--
Mironovskaya 808. 12--U1'yanovka, 13--Tsezium 39
67 ~
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1 -
In the winter of 1970/7L the snow cover in the "excessive" variane was
considerably than in the "natural" variant. The minimum eemperature~
at the depth o� the tillering node wieh ehe excess snow cover did no~
drop below -2�C. During this winter there was a very clear manifestation
of the third Cype of growth o� the vegetaeive cones, especially in the
less winter hardy strains such as San Pastore, Avrora, Rannyaya 12 and
Bezostaya 2 (~igure 18).
One can ~udge Che way early planting times affect the growCh of vegetative
cones of weakly winter hardy strains of winter wheat from the curves of the
_ growCh of Che cone in the Bezostaya 2 strain. With an early planting
time, even under conditions of a natural snow cover, Che growth of the
vegetative cones conrinues until December and is renewed as early as the
second half of February, long before the snow disappears from the plot
(figure 19). Moreover, even under the conditions of a naCural snow cover
the growth of the vegetaCive cone was considerably greaCer when the planCing
= time was early than it was with Che third and fourth planting times.
7'he winter of 1973/74 was relatively warm and the snow cover was established
- early. Under these conditions, even with Mironovskaya 808 planCed ar an
early time, there was almost continuous growth of the vegetative cone,
especially in the second half of the winter (figure 20).
It was possible to trace the effects of the winCer conditions of 1972/73
even more clearly in the Odesskaya 51 and Bezostaya 1 strains, which were
tested as indicator strains. As one can see from figures 21 and 22, these
strains which are adapted to the steppe regions are unsuitable for the
central belt of the country's European territory since the relatively
rnpid growth of the vegetative cones in these regions and their inadequate -
winter hardiness lead to considerably more severe damage than Mironovskaya
808 sustained under the same experimental condiCions. _
The growth of the vegetative cone of Bezostaya 1 proceeded approximately
in the same way as that of Odesskaya 51 throughout the entire winter
(figure 23) both under the conditions of a natural snow cover and an
excessive one. These data once again confirm that selecting even the
most productive new and promising strains in other zones, it is necessary
_ to examine them in detail under the condiCions of various winters which
are typical of the nonchernozem zone.
The high growth rate of strains like Odesskaya S1 and Bezostaya 1, which
are comparatively resistant for the countzy's European tezritory, even
in snow winters under the conditions of the chernozem zone can lead to
"prolification" of the vegetative cones under the snow throughout the
- winter and to severe damage with the return of frosts in March during years
with an early disappearance of the snow cover. In these cases planting
times that are too early are especially inadmissable.
68
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FOR OFF'ICTAL US~ ONLY ~
Tnble 28. Uyanimics o� bevelopmenC and CrowCh of Ul'yanovkg 5train of
WinCer Wheae in Autumn, Wineer and Spring periodg of 1972-73
Wieh Various Planting Times gnd Various Varianes of the Snow
Cover and TemperaCure Conditions of the Soi~, at ~he Depth of the
Tillering Node (3 cm).
~R~�~~RR
~ ~RR~RR
~~.(H011 ~NNYV p00000 OOOOCOO
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~ ~ Utli
v
r. r. r... r..~
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~G (RII) _
- t~M071 ~NNYY 000000 0000000
u ~ ~ _
~ nrrrrrr r.n..~rrr
~ ~ Wli rrr.rr~. rrw..o..+rr
I -
c
e
- g ~ u Y0~ McOMC+~e+~M lOehMC9MMM
~ ~ �iQ00 OY'~Mh
Y
C V
a ~ -
x ~p~p~p G
y ~ v (1111) M ~i' d' V' ~1' d' et' ef' ~d" at' ~a' ~fi ~
6 ~~1~N071 ~NNrt QO0000 OOOOOOC
a
. _
_
4
~ I\ w~M Irl r~ 1~
u 7 1l1 ro'~r.~�rr~rr~ ..~.n.....~rrar
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�
q n or~en ~ u~~n ~ N u~
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-
0 ~ L~ll rr~..rr ...r r
~ ~
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~ a oa v~ ,o co eo ~o ~o ~o ~o ~ ~ ~o ti t~
-s~ou ov~wn ,
i
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c.' r~ ~ ......::.~"9~
~ N >C~CYCXX>C M
g�s ~ r C�1 ~ ~j v~ M a0 M N
_ , ~O a ox ~ ~ ~ ~ ..t~. ~ CV CV N ,
r p
~z
4=
_ [Table 28 continued on following page]
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~OK OrF~ICTAL U5L ONLY
~ ~,oooQ 0 o0o o~~~n~n
^~1 (IIN) M77T~1'~1' ~i'V~~3'~'tl'tl~lT
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Y ' ~ V~u r~rrr~r~ rnrrrrrnr
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~J~OU Or~Nh ~~~NN1~ ?ryu~1A~~N'!~i
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N ~
V~ 1111~ rrarrrr r~rr~.nrrr
~ V
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~OU Or~Mh
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- v
~ N
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- " u aa Mc"~MMMM MMehe"JMCOM
~ ~'pou oraxn
a :
~ : ~ (NNl ~'~:e v~~"~
~ ~ ~o~fio~~eMrv o00000 0000000
Y V
~ N v
'ti ~ ~ n~js rl nn r~., r -
~
N QOU01CJwA ~~1~~~~ ~~~N~~~
(n~) 53~~~~`~ ~�u'~b~~ia'v~�
^ t~Xi+oxtNrrt o00000 0000000 -
v
~ - ~ ~
" ~ ~ ..y..~"..r..:...~. r...r..... ~ ~
o v nsa~
_
~ Y~OJ IA t0 t0 e0 t0 tp t0 t0 t0 t0 ~D t~ 1~
�iQ01! OfDNA ~ _
~ ~
Y -
~ ~ ~ � ~ � _
~a~~,-� ~.~i ~exx~x~e e~
~'O~ V b~ tO~MI~..N Q~f ..NMoC...M^~
~w~~ .~CV..N .+N~CV.+
qi
_ [Key on following page] -
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Key:
1. NaCur~1 snow cover, 10-25 cm 4. Numbex of ehootie u
2, r1.~~nting eime~ 5. SCgge
ll~te binlogical exgmination 6. LengCh of con~ (mm)
~onducCed 7. Snowlessnese (0-3 cm)
_ ~~N MN /
~,e , ~
/
_ ',s I
~,4 ~
~
,,o j ~ ~ -
:
i -
~ ~.~i~/
. ~i~ / /
~
~ .~i~~~--~_'/'/ � _
~~i~..~..~..~ ~'r/,.~,._.....~
a4 =z
J~~
~
~ ~ t~s
- ~I
~
! ~~i~~l ~ ` I~ ,
_ O~ ~p ~,III~ '~,I (I1II~~'IIII''I(I 4
i ~ . 2
0 0 l .y~..~ I , . ~ 0
ZO l. ~~'.%~%;N~~''~ -2
i ~ ~
~i0 ~ 5 ~ 4 -
2 9
60 ~ 3 ~ 6 !0 \ 6
7 t! \
- ~ , ~ 4 e n
x x~ x~r i n ~n ro
- Figure 18. Growth Dynamics of Negative Cones of Winter Crops
= in Winter and Early Spring Periods (Winter Wheat
and Rye with Various Degrees of Winter Hardiness) in
1970/71 with Optimal Planting Time Under Conditions of
"Excessive" Snow Cover. Moscow State University.
[Key on following page] _
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rak ~~rr ~ c i ni, us~: c1~vt.v
Key:
= Conveneinn~l symt~nlH 1kr~~ h, t1. .~nd 1-6, ~ee fig. 15. 7--Vy~tk~,
8--5.yn ~n~Cdxe~ 9--Avra~{~, 10--R~nny~~y~ 12, 11--I~ezngtay~ 17--
Mironov~kc~yn yubilpyn~~y~i.
ry
~
_ ~~N MN
1
~A .
Q6
~r,..._.._~:...'=:`..'r.i,:~::~
_ as ~
r.,..-
- ^y %~r.:~�~
~..~~f}it�-~~..'~~.� ~ ! J
~4
Ss~~ u~r~~~
~t tM t~~/ ~'_:i' �-=~"";~~"~~�:~..~~:u
~~~a.G....~~....~� l~~
0.? 40 u,".--�s~s~ 4
, ~ ~
?0 ~ ~ ~ i
2
r,. . -
O D ~ ~ MM~ ~N
0
. ,,i;� ,
n1 ~ j~j~ / _1
~ p g ~ ! /
~ 3 9 4
60 [1.~~ 4 1~ . ~j /
_ ~ S � \ /
eo Q ~ ~ -a
s _w_ ~ ~
- 7 l4 10
N tw
X X/ XI! /
Figure 19. Growth Ilynamics of Vegetaeive Cone of Bezostaya 2
Winter Wheat Under Conditions of Various Snow Cdvers
With First-Fourth Planting Times (1972/73). Moscow
State University
Key :
_ Conventional symbols lkn~ h, H, and 1-6, see fig. 15. "Natural"
snow cover: 7--first planting ticne, 8--second planting time, 9--
_ tlilyd planting time, 19--fourth planting time. "Excessive" snow cover:
1.1--�irst planting time, 12--second planting time, 13--third planting
time. 14--f.ourth planting time.
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tR~ MM
- Q6 ~
0.4 ~~,,,,,,,--,~~.i
-~--,--~-~.~.~/'~''~""~.d~~"'I~
A CM 6
Q2 f0 �~_..~cL~ ,~~._..~~~~~~.-~~~,r"" 4
,,;i ~ _
~
~ ' . ~ r
O O .l~ .~r~..~+~~r ti /,'r,/ ! , / O .
~ r~~.~~~+,.. \ ' ~~j ' 2
.--w~. ~ i i
~ ? E ----J S . \ . 4
v~
~ ~ 3 ; 6 0 -6
~Ip
e
~ ~ E Y
A A1 ~ ~V ~Q
Figure 20. GrowCh Dynamics of Mironovskaya 808 Winter Wheat
With FirsC and Third Planting Timea and "Natural"
and "Excessive" Snow Covers in Winter af 1973/74.
Key:
Conventional symbols lkn, h, H and 1-6, see fig. 15. First planting
time: 7--"excessive" snow cover, 8--"natural" snow cover. Third planting
time: 9--"~excee!sive" snow cover, 10--"natural" snow cover.
Of especially great interest are data from a morphophysiological analysis
of Mironovskaya 808 winter wheat which in the last few yeara has been
regioanlized in the central regions of the nonchernozem zone. In plants
of this strain with various planting times the vegetative cone even in
stage II of organogenesis is somewhat greater than that of U1'yanovka
in the autumn and early spring, but they come out of the state of dormancy
at almost the same time as U1'yanovka does. Consequently, in terms of
Erost resistance (taking into account their coming out of the condition of -
- dormancy in the spring), the Mironovskaya 808 strain comes close to
the Ul'yanovka. In Mironovskaya 808 plants under conditions of an
excessive snow cover (with the first planting time), even at the end of
February and the beginning of March one observes a transition to stage
III and even in the second ten c~~ys of April in 1974 the plants remained
_ at stage III in their development and there was continued metamere growth
of the segments that subsequently conditioned the formation of a large
- rudimentary spike. The t:ansition to stage IV of organogenesis began
later, after there was no return to cold weather in the spring. -
73
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~ `~~~i~V~ A
a ` .,a . / ~ji'" +.w 1'yj'!r.+ . ,
P;~~ ..~.Y~ /.~~~-,t� � Ww.' ~i1
1~~y
~.f' ~ ~~`~~~~a
~ ~v +1,~'
~ ~
`i ~
~
' -
~
r.
Y
Figure 21. Condition of Vegetative Cones With Inadequate Snow
Cover
Key: _
~ Vegetative cone of Bezostaya 1 strain: a) x234, 6) x468; vegetative
cone of Odesskaya 51 strain: b) x200~ r) x400, 16 April 1973
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Figure 22~ Effects~of Planting Times on Condition of Vegetative
Cones of Mironovskaya 808 Winter Wheat Under
Conditions of an Exceas of 5now (depth--about 60-70 cm)
Key: .
a--necrosis of central part and base of cone; 6--deep longitudinal
ruptures and necrosis of tissues.
Thus in terms of the condition of ti~e vegetative cone (level of develop-
ment and growth rates), Mironovskaya 808 is very close to the U1'yanovka
atrain. But among the strains that are included in group 1 in terms of
winter hardineas, Mironovskaya 808 plants, as we know, have one peculiarity
that is very important for the strain's productivity. Namely, as distinct
from U1'yanovka and other highly frost resistant strains like Lyutestsens
_ 329 And Lyutestsens 1060, Mironovskaya 808 plants are capable not only of
maintaining a live tillering node at critical temperatures when the
- shoots above the ground die, but also of developing new productive shoots
in the spring. As was already noCed, in the autumn one observes the fourth
types of growth of the vegetative cone in Mironovskaya 808. It is
precisely this propeYty of the plant that makes it possible when wintering
_ conditions are unfavozable to obtain yields that are close to average
while areas planted in other strains of wineer wheat during these years,
because of the severe thinning of the plant stand, must be undersown with
barley in the apring or repZowed and replanted with late spring crops.
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r
r
_ ~j:
~
, -
~ ~l~;
' ~f
.
.
w,
w
Figut'e 23. Types of Damage Co Winter Wheat in Winter of
1972/73 (planted on 25 August). Bezostaya 1 sCrain.
Key :
Transverse and longitudinal fissures in vegetative cone with "natural"
snow cover: a) x200, 6) x400; b) longitudinal rupture of vegetative
_ cone in absence of snow (x400). 16 April 1973 -
As was already pointed out, one of the peculiarities of this strain is the
formation in the autumn of shooCs that are in various stages of organo-
genesis. Even as certain tillering shoots are developing at the same time
in the autumn and providing for a uniform plant stand of productive shoots
in the early spring, another group of potential shoots (third and fourth
- order of tillering) remain in stage I. Th ,e rudimentary shoo~s in the
form of so-called "reserve" buds pass through the winter well. When
the winter wheat plants have spent the winter well and their main shoots
t~ave rapidly moved into stages IV-V of organogenesis, because of the
law of reduction the rudimentary "reserve" buds do not begin to grow and
completely die off. But if because of extremely unfavorable conditions in
the winter et~e main autumn shoots which are in stage IT of organogenesis
are severely damaged and are behind in their development or have completely
- died, then shoots from the "reserve" cones develop, which are better
protected since, in the first place, in stage I they are more frost
resistant than shoots in stage II and, in the second place, since they .
are in a dormant condition, as is the case with U1'yanovka, vernalization
processes have taken place in them auring the winter, although slowly,
w}~ich has enabled them to move into stages IV-V and subsequent stages of -
organogenesis in the early spring. Because of this they develop into normal
productive shoots. ~
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An ac:count oF the number of planCs thaC have survived rhe wiitrer in
relation Co the planting times shows that the main shoors of the plants -
which have been planted early in the majority o� years, because of the
completion o~ the first sCage of developmenC in Ch e autumn, are more
sinsiCive tu unfavorable conditions (].ow temperatures and excessive _
snow cover). This was established in the comparatively seve~re winter of
1971/72. As one can ~ee fram the figures in Cab1e 29, in Ch~ early
: spring of 1972, under the conditions of Che variant with a natural
snow cover, in Mironovskaya 808 winter wheat 65~percent of tlle planCs and
- 51 percent of the shoots remained (when planted on 10 August) and with an
optimal planting Cime--98 percenC of the live plants s,nd 95 percent of the
shoots; under conditions of an excessive snow cove r with planting on 10
August S2 percenr of the plants and 62 percent of the shoots remained alive
- and with planting on 25 August--77 percent of the plants and 70 percent
of the shoots.
Table 29. Number of Plants and ShooCs of Winter Wheat that Survived the
WinCer. Moscow. 1971/72
EcrecieeHUe+il cne~uuA ( Nsbwtovr~dA cxe~;i~wp
~ 3 ~ naKpoo ~4 ~ noKpoo
_ ' Copr CpoK caBa ~5 ~ *Hemx (yf,)
~ 1~ (I~CTCHN~ ~ noaeraa 6 pacrexxti ] noeeraa
~8~ MFIPONOHCKBA SOB 10 V111 65 51 52 62
_ 25 VIII 98 95 ?7 ?0
5 1 X. 97 97 62 58 -
20 [X 100 97 100 99
~9~ MiipoHOecKaR ro8i~� ]0 VI1I 62 58 50 62 -
~~e~tta~ 50 25 V I I I 96 94 76 78
_ 5 I?c 61 57 75 77
20 I X ?8 84 76 69 _
~lO~ YdbAHO8K8 10 VI1[ 82 49 46 47
20 V 1 l I 96 94 b~ 61
5 1 X 96 96 69 68
20 1:C 100 98 76 86
- ~1Z) 6e3orraA 1 25 VIII 83 92 46 44
5 I:C 95 96 63 T
(12) Aepopa 25 X`~ 95 95 62 69
Key:
1. Strain 7. Shoots
2. Planting time 8. Mironovskaya 808
3. Natural snow cover 9. Mironovskaya yubileynaya SO
4. F.xcessive snow cover 10. U1'yan~vka _
5. Live 11. Bezostaya 1
6. Plants 12. Avrora
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t~c~~i t~ tc rnt, usi, otvt,v
!
1n nrder to get ~ berter idea of the signific~nce of pl~ntin~ timeg ~nd
winterinF; conditiony~ one ~}iould discu~~ the nnalysia of figures cnncerning
the reduction ("di~cerdin};") oE aueumn tillerittg ahoots in various strfling
witti vurious plnnting times, A morphophysioldgical ana],ysi~ of the
tillerin~ shoots reveals, in tlie first pl~ce, ~rrain difterenceg in terms
of this indicator nnd, in the second place, the ~ffects of Che planting
time. All thie is undoubeedly related ro ehe quesCiott of ehe c~nsequence~ -
of winter d~mngeg.
'Cable 30 gives figures from the autumn~ spring and summer ~~ccounCing for
the number of rillering slioots itt two sCrains of whent--Mironovskaya 808
~nd Iiezostaya 1--with various plnnting Cimes. As one can see from t~ble
30, in the first place, Mironovskaya 808 pLnnCs bush coneiderably more
inCen~ely itt Che autumn and more shooCs xemain until harvest than wiCh
I3ezoseayn 1 planrs. In the second place, ehe significant differences in
th~ number. of nutumn ~hoots with various F~lanCing Cimes disappear
througtiout the spring ~nd summer; the plan~`s lose from 50 to 90 percenC
of the til.lering shoots as early as the end of July and the most autumn
stioors--both in abaolute figures and in percEntages--are retained when the
planting time is opCimal (50-60 percent). When ChE planting ti.me is too
early a large number of Che ~hooCs die as early as the end of May and by
}~arvestin~ time onty 25 percent of them remain. With a late planCing "
- rime, Mironovskaya 808 plants formed no more Chan three shoots in Che
autumn ancl Cillering does noC begin in Bezostaya 1, and in Che summer
up to 50 percenC of ttie plants die.
Tiius in 1972/73 rhe plantings that were thinnest by harvest time were those
on plots that had been planted very early and very late. It is inCeresting
to note thaC even at the end of May, especially with early planting times,
it was possibl.e to observe a significant retardation in the development -
of vegetative cones at the ends of the bush: while some shoots had already
passed into stages VI-VII of organogenesis, others remained in stages II-
III and gradually died off.
- Ti~e dying off of shoots in the spring is determined not only by rhe
unEavorable effects of winter conditions. When evaluating both the
wint~r tiardiness of strains and the wintering conditions, one should take
Into nccount the dynamics of "discarding" of shoots: the more favorable
tt~e conditions for the development o� plants in the autumn-winter period.
the tess "discarding" of winter shoots and the greater the productivity _
of the plants.
?
tn cases where winter wheat externally seems to have survived the winter
normally. in plants that have been planted at earl~� times and have been
under an excesstve snow cover there is frequent~, deformation of the
spikes even in stage V of organogenesis (figure 24). Additionally, the
_ more snow resistant the strain, the lesser the consequences of winter
- damage. This is quite clear from a comparison of the rudimentary spikelets
of Mironovskaya 808 and Bezostaya 1 winter wheats.
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7'able ~0. Ilynamics oE IteducCion of Autumn TiJ,lering Shoots nf Various
Str~ins of WinCer Wheat, Depending on P1~nCing ~imes
- C'2> C{IOKN t!/~
A~r~ y~icr~ 10 VI~~ 43 ~'ll~ 16 IX 43 1X
~1~ ~e A o I G a I a ~
r
~a~ MHpOHOBCKeA 8O8
1972 r.
16 XI 10 100 5 )00 3 100 3 100
1973 r. '
21 V 10 100 5 100 3 100 2 66
26 V 4 40 3 60 2 66 2 66
20 V 11 1 10 3 60 1 33 1 33
- ~2~ CpoKe tee~
11~r~ yver~ 10 V111 ZS Vlll S IX I 23 IX
~1~ c I 6 a I 6 a I 6 ~ c I 6
~4~ bt30CT8A 1
1972 r.
I 6 X 1 6 100 4 100 2 100 1 100
1973 r.
21 V 4 67 3 75 2 100 1 100
26 V 2 33 1 50 1 SO 0,5 50
20 VII 1 17 2 50 I I 50 0,3 33 -
Key:
_ 1. Date of report 3. Mironovskaya 808
2. Planting times 4. Bezostaya 1
_ Note: a--overall number of shoots before beginning of wintering~
G--number of reduced shoots -
At the same time, plants that are planted late, which have smaller
vegetativc c:ones. are significantly behind in terms of development and
tf~c productivity of the spike in the spring. For this reason, despite
ti~c better win.tering, they cannot be recommended for production. The
effect of the ;:onsequences of da~mage under a deep snow cover in the winter
ciepenJs considerably on strain resistance of the wheat (table 31). As
one can see from table 31, darnage to winter crops under snow in the
79
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~dEt d~F'ICIAL U5~ dNLY
- winCer of 1965/66 had a n~gaCive effece on eha lle~ghC of the plgneg, the
pge~ing rhrough Che segges o� oxganogeneais, Che coefftcient of bushiness
nnd the number of gpikeleea in the spikea.
- ManifesCationg of damage to crops under the gnow are observed considernbly
more frequ~ntly on areas plgnCed in winCer rye in rh~ northe~stern, eastern
~~nd northwegCern regions of the ~uropean territory of Che 5ovi~C Union.
When the ~now cover falls early and remains far a long eime tihe hydro-
carbon exhaustion of Che plants led to their sCarvaCion and Chen these
plants were sub~ecC to attack by �ungi when there was a high moisture
conCenC under ehe melting snow cover. And only in very recent years has
ir become clear that undifferentiated growth of the vegetative cones is
one of tt~e facCora that lead to the death of ahooCs, primarily main shoots,
to laxears, and to a considerable reducCion in the produceivity of the rye -
spikz.
- As one cnn see from table 32 and figure 25, the vegeCative cone of winter
- rye can reach large sizes even in Che autumn when the conditions are
favorable for Chis but then under the snow with a temperaCure of 0-+2�C
it continues to grow even in the winter months, as was noted in the winters
of 1972/73 (figure 26) and 1973/74. By apring the shoots of these plants
have died off (figure 27).
As was noted in the experiments wiCh various strains of winter wheat, in ~
winter rye the growrh of the vegetative cone eook place more intensively
under Che conditions of an excessive snow cover and with early planting
times (figure 28). Thus, for example, in the winter of 1972/73 under
a natural snow cover the length of the vegetative cone of plants from the
� first planting time had reached 1.5 mm by 28 February and under an excessive
snow cover--1.9 mm; and with a late planting time, although the length of
the vegetative cones was also unusual (0.9-0.93 mm) the differences
between their sizes under natural and excessive snow covers were not great. _
The undifferentiated growth, elongation of the vegetative cone and its
necrosis in winter rye can be illustrated with photographs (figure 29).
Thus even at the end of March in stage IV of organogenesis there were
clearcut differences in the condition of the vegetative cones of plants
tl~at had spent the winter under natural and exc~ssive snow covers (figure
30). These differences were manifested even m~~rE~ clearly in April (figure
31) when the plants entered stage V of organogeri~sis. In the photographs,
even with plants taken from the variant under an excessive snow cover, one
can observe a loss of turgid3ty by the tissues of the undifferentiated
top of the vegetative cone and the appearance of dark spots, which is
especially cJ.ear on the upper parts of the vegetative cones in plants that
have come out from under an exeasive snow cover. In figure 31 a and 6
the tops of vegetative cones from an early planting time (10 August) in
stage V of organogenesis under different snow covers (natural--a and -
excessive--6) are shown for comparison. In the plants that have come
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~ ~OE~ O~~ICIAL USL ONLY -
�y
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Figure 24. The Effects of an Early PJ.anting Time (10 August)
and "Exces9ive" Snow Cover (50-65 cm) on the Formation _
of Spikelets. Stage IV of Organogenesis.
[Key on following page] _
8Y
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t~ott c~rric;rnr, tr5t: aNt,Y
Key:
13ezoseaya 1 winter wheat (x333): a--the upper spikelets are behind in
developmenC, have turned dark and are beginning to die off, 6-~-
spikelets in the central pnrt of the spike; one can see prolificaCion
oE the spike and spikelet axes; Mironovskaya 808 winter wheaC; b--
~pper spikelets axe behind in development. their reducCion is r~vealed
in seage V; r--normal spikelees. 27 April 1973. -
Table 31. Effects of Winter Damage Under Snow on Growth, Development ttnd
Organogenesis of Winrer WheaC Plants in Moscow Area [12~
~ ~ 1 fp~nni Ddeora
nocteneiw puTa ~~~~cno 3ron AnNN~ ~IHC.~o 4HC.~o
Copr noepe~~teH� uNfl ~~�~r~~ax opraao� Konocn KoaocHOa OON08617t
uaTH ~~M) aecrbeo reaeaa (MM) e Ko.~nce ndseroa
f~1 l'~~ (5.~ ~~a ~Z~____~$~.~
~9~M11p01108CN8A 1 ~5 3,3 VI1I 94 18 1,?
- 808 2 S? 3,3 VI1 60 14 0,2
' 3 37 ?,8 VII 25 12 0,2
(lo)nnr�186 1 g,5 3,4 VII( lOQ 19 1,0
2 56 3,~4 VII 7l 17 0,6
. ' 3 a0 3,~ VII 28 13 0,6
(11) KyuueecnaA 45 1 76 3,3 VI I t 109 18,8 1,3
2 50 3,0 VII 58 15,2 0,0
3 4g 3,A VII 49 16,5 O,U
4 3~ 3,? Vll 16 14,4 O,U
~12)~a~ian 1 58 ~,2 VII 5~ 19,9 1,1
2 39 4,0 Vll 23 17,6 Q,3
3 38 3,8 VI-Vti 2�! 15,5 I o,n
4 25 3,3 VI-VII 8 1-4,'? U,0
(13)3poc 1 66 3,5 VI I 53 19,-! 1,5
2 43 3,9 VIl 20 19,0 I 0,6
3 43 3,5 VII 13 18,0 0,3
4 27 3,3 VI-Vli 5 13,3 ; 0,0
- (14~{oxaaHn 1 5? 3,~ VII 4? 19,9 ( 2,1
2 40 4,1 VII 20 17,? 1,6
3 31 3,9 V[-V1I 8 15,2 1,2
4 25 3,2 I VI-VI! 5 12,8 Q,0 _
Key:
1. 5train 8. Number o~ side shoots
2. Croup in terms of degree 9. Mizociovskaya 808 '
of damage 10. PPG-186 _
3. Height of plants (cm) 11. Kuntsevskaya-45
4. Number of green leaves 12. Fanal
5. Stage of organogenesis 13. Eros
- 6. I.ength of spike (mm)~ , 14. Kokhland
7. Number of spikelets in spike
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'l'ablr )2. 1lynumics of DevelopmenC ~hnd Growth o� VyaCka Winter Rye in -
AuCumn~ Winrer and Spring Peziods of 1972-1973 With Various
I'lanting Tiraes and Various VarianCs of Snow Covers
~~s~~o~ Q~ vv~ ~~~g~��
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a
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Fpa~ N O~ ^p~t~...u~ p~ ..~e`~oo.~cOGV
r s v M CV CV N.~ CV CV
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[Table 32 continued ton following pageJ
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a=
[ I:ey on Eollowing page ] -
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- Key:
- l. NArurnl snow cover, 20-25 cm
2. Date of biological examinaCion
3. Number of shoots n
4. Stage
5. Length of cone (mm)
6. Excesaive snow cover, 50-60 cm -
7. Snowlessnesa (2-3 cm)
_ out from under a snow cover and begun to grow later, a sma11 part of the
undifferentiated top of the vegetative cone remained until Chat date
(27 April--figure 31 G) but somewhaC later (by 5 May) iC dries up.
Thus, as a result of many years of experiment, it was established that _
- under a deep snow cover, in addiCion Co processes of re~piration on which
- supplies of hydrocarbons accumulated in the autumn during the course of
tempering are expended, in plants where the temperature is 0 f 1�C and
t~igher in the zone where the tillering nodes are located there are also
growth processes. They are insignificant wnen the planting times are laCe
and extremely marked in well bushed plants with 7-8 shoots and more.
The increase in the length of the leaf blades and their etiolation do not
' ai~uys lead to Cheir rapid death alChough in a number of years one
observes that they die off prematurely, which somewhat impedes the plants'
de~~~lopment, especially in years with a cold and dry spring. Frequently
these leaves turn green in the light ~.nd for some time participate in the
photosynthetic activity of winter wheat and winter rye. In the ma3ority =
~at years abnormally elongated, etiolized leaves 2-4 (counting from the
bottom up) completely die ~~ff.
Considerably more destructive for the plants is "prolification," undif-
" ferentiated growth of the vegetative cone. Necrosis of even the uppermost
parC of the vegetative cone leads to a sharp reduction in the productivity
of the plants and complete destruction of the vegetative cones leads to ~
a ~ansiderable redu~tion in the yield. There has been repeated observa- '
tion~: of. the so-called "white spike" where some of the spike glumes are
deprxved of chloraptiyll; in these spike;s the blossoms are sterile. "Idhite
spike" can be caused by various factor:s--damage by concealed stalk pests,
a:~i~oxtage of a number of necessary trf~ce elements, and so forth. But
Ln tlie m.3,jority of cases, especially in winter rye, "white spike" is a
result of winter damage, including perishing under the snow.
IC unc is to generalize the data obtained from physio7..ogists c~ncerit~nb
- the processes that take place in winter plants that remain for a long time
u~zd^r- a deep snow cover at temperatures of 0 f 2�C, one can classify them
as t~yclrocarbon exhaustion because of the plants' expenditures on respiration
and ~;rowti~ processes, starvation and decomposition of protein components
a~ wc~ll. as undiff.erentiated growth of vegetative cones. Under conditions
that are favorable for the development of fungal flora, weakened, anomalously
85 -
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LNNMM
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x x~ xu ~ i~ u~
I~igure 25. Growth Dynamics of Vegetative Cone o.F Vyatka Winter
Rye With Various Planting Times Under Conditions of -
Snow Cover of Various Depths t1972/1973)
(Key on Collowing page]
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ro~ nr~rr,rnL usc nNLY
Key :
Temperature of soil aC depth o~ tillerin~ node under snow cover:
1--"natural", 2--"excessive"; depth of snow cover: 3--"natural",
4--"exces$ive"; depth of freezing of soil under snow cover: 5--
"natural", 6--"excessive." "NaCurAl" snow cover, planCing time:
7--first, 8--second, 9--Chird, 10--fourth. "Exceasive" snow cover,
planCing time: 11--first, 12--aecond, 13--rhird, 14--fourth. 1972/
1973. lk --leng~h of vegetative con~ (mm), h--depth of snow cover '
(cm), H--~epth of freezing of 3oi1 (cm).
"prolificated" organs of winter plants are infecCed with snow mold and
collar rot which destroy the entire above-ground mass of Che plants and
attack and fully destroy winter crops.
MAIN ACROMETEOROLOGTCAL FACTORS CAU~ING PERISHING OF WINTER CROPS UNDER
SNOW AND PATTERNS OP' THEIR SEASONAL CHANGE
A study of the ~results of the research on physiological processes on
perishing of winter plants under snow made it possible Co come to the -
_ conclusion that the destruction of winter crops when they perish under the
snow is determined mainly under the condition of the plants in the autumn,
the soil temperature at Che depth of the tillering node and on the surface
- of th~ soil under the snow, the depths of freezing of the soil and the -
length of time a deep snow cover remains on the fields.
Investigation of the temperature conditions of the soil at the depth of
the tillering nodes of winter crops showed that wiCh certain combinations
of the depth of the snow cover and the de~pth of freezing of the sojl, the
temperature remains for a long period of time within the range in which
perishing of the plants takes place (0 -5�C). From the results of
observaCions from 1965 through 1976 at the Nemchinovka Agrometeorological
Station in Moscow Oblast with slight freezing of the soil during the period
of the formation of a deep snow cover (depth 30 cm and more), the minimum
temperature of the soil throughout the entire subsequent period of the
winter was within the range of 0--3�C (Table 33, 1965/66, 1967/68,
1969/70 and 1970/71). -
The soil temperature at the depth of the tillering nodes of winter
crops (zn average depth of 3 cm) when there is no snow cover on the fields -
depends mainly on the air temperature and the depth of freezing of the soil
[18, 22J. 1'he daily and periodic changes in it are similar to the changes -
in the air temperature, but the absolute amounts are higher than the
air temperature. With a slight freezing of the soil, the difference between
the soil temperature at a depth of 3 cm and the air temperature is greater
than when there is deep freezing of the soil.
~
_ 8~ -
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~K MM
l,6 ~
.
.i' f~,~
~a
. l'' =
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0,6
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~ ~ ' � � i~/;;~i ~�'~.�i.ii;.:�i;;,. 0
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. ~ j~. ' _
t
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60 ~ ~ ~ 6 -6
~ -K�- g r _
.HtN ~ 4 g -~~--f0
i i i i i ~ -
X XI Xll I /l lll lV
Figure 26. Growth Dynamics of Vegetative Cone of Vyatka Winter
Rye of the First and :iecond Planting Times Under -
_ Conditions of "Natural" and "Excessive" Snow Covers.
Key :
Conventional symbols l~n, h, H and 1-6, see fig. 25. First planting _
time, snow cover: 7-- excessive", 8--"natural"; second planting time,
snow cover: 9--"excessive," 10--"natural". 1973/74
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t'
1 R)
j.
fJ
k
I ~
~
;
; r~
I ~ ' t.
1yt r
1 l ' j 1
I . , . . ' t 1 5 _
. ` ~ ~1 ` ' 1
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. ~ r.. .
_ ~ ~ ,
.
- ~ ' ,y, .f. '
1.. .
Figure 21. Perishing of Vyatka Winter Rye Under Snow in Winter
~ of 1965/66
- Key:
~xternal appearance of plants from specimens taking from agrometeorolo-
gical stations: a--Charozero (Kalininskaya Oblast). 6--Igra (Udmurt
_ ASSR), b--Cherdyn' (Permskaya Oblast)
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; p ~ 6 -
, ;
~ ~v '
_ ~
; Figure 28. Condition of Vegetative Cone of Vyatka Winter Rye
After Wintering on 3 April 1973. (Stage IV)
Key :
a--with "natural" snow cover (normal co;~~itions), 6--with "excessive"
snow cover
Wt~en the snow cover is established on the fields of winter crops the ~
, effects of the air temperature on the soil decrease considerably since the
snow has great heat insulation properties (Table 34). Its heat
conductability is 10 percent of the heat conductability of the mineral
= part of the soil.
; The heat insulating properties of the snow cover becomt greater as its
; depth and reflective capacity increase and its density decreases.
, During the course of the winter the density of the snow cov~r increases;
in January-February it amounts to an average 0.18-0.22 g/cm ; and by
the end of the winter it increases to 0.25-0.35 g/cm3. The heat insulating
role of each centimeter of the snow cover turrts out to be significantly
less by the end of the winteL than in the first half. This is clearly
confirmed by the data pres:ented in table 34 (temperature gradient in the
grass stand of wintet crops). As the depth of the snow~ cover increases
~1t the end of the winter and the air temperature increases. the soil
_ temperature at the depth of the ti,llering node in the second half of the -
winter period in regions where the snow cover persists always turns out
to be greater than at the beginning of the winter [5, 16, 22J, as a result
= of which the processes of perishing of winter crops intensify during this
_ ; period. But when the snow cover is packed beginning at the time it is
' 90
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formed on the fields, the soil temperature at the dept:h of the tillering
node ia lower than when the snow cover is natural, no~; only in rhe wintier, -
- but also in the spring during the period of thawi.ng ok the snow, which is
very important for protecCing the plants~ The result~ o� our field -
experiments for studying the role of the density of Che snow cover with -
the wintering of winter crops at the Belogorka Agrometeorological SCation
_ are preaented in Table 35.
~ r;
~
~ FiEure ~9. Condition of Vegetative Cones of Vyatka Winter Rye
After W.ltering, 21 March 1973 (Beginning of Stage IV)
Key:
a--normal condition with "natural" snow cover, 6--elongation of
vegetative cone with "excessive" snow ~aver (X 140)
The soil temperature after the formation of th~~ snow cover is almost =
always higher than the air temperature. On1y during significant warming
periods (up to thaws) with a deep snow cover a^.d deep freezing of the soil
can it turn out to be lower than the air temperature [22].
A change in the depth of the snow cover has an especially strong effect -
on the temperature of the soil at a depth of 3 c~n when the snow cover is
initially not very deep (up to 5-10 cm) but when the snow cover is as ~
deep as 30 cm and more it does not have a great significance since the
gradiant of the temperature with the depth of Che sn~w cover decreases
- sharply. _
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a ~ ' '
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F~igu~~e 30. Condition of Vegetative Cone of Vyatka Winter Rye in
5prin~ With F:arly Planting Time (10 August 1972)
~ and 'Vatural Snow Cover of 20-25 cm. 15 April 1973.
Key :
a--rudimenrary spike, beginning of stage IV of organogenesis (X60),
~i--top nf spike is undifferentiated ~A~t all spikelets are one stage
behind those of the middle part of the spike (X222)
According [o thP resulta of our research, it turned out that tor each
centim~ter of depth of the snow cover the soil temperature at the depth
of 3 cm under the snow increased in comparison to the air temperature -
by 0.5�C when the snow cover was S cm deep, 0.3�C when the snow cover was
- 30 cm deep and 0.2�C when the snow cover was 60 cm d~ep.
Witl~ an increase in the depths of the snow cover ~to 30 cm and more the
_ soil temperature at the depth of 3 cm and on the s~arface of the soil under
the snow changed very little. regardless of the fl.uctuations in the ~
, air. temperature (17, 22J.
The daily fluctuation in the temperature and the snow cover rspidly -
disappears. According to the data of B. P. Karol', in Jan~.ary 1946 near
Leningrad the change in the daily fluctuation of the tetn~erature dependin~ -
on the depth of the snow cover was as follows: on the surface of the snow--
30�C, at a depth of 5 cm--15.8 �C, at a depth of 10 cm--11.1�C, at a depth
~f 25 cm-~2.1�C and at a depth of 40 cm--0.2�C.
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I~igure 31. Conditfon ol the Ve~;etative Cone nf Vyatka Winrer
Rye After Wintering. 2~ Arril 1973. Cnd of Stage IV.
Key:
' Under conditions of snow cover: a--"natural" (normal conclition),
6--"excessive" (top of cone is not differentiated).
The results of observations at meteorological stations for four years showed
that during periods of lowering of air temperatures in the winter, the
soil temperature at a depth of 3 cm when the snow cover is 30 cm deep and
the soil is slightly frozen decreases insignificantly, almost by the same -
amount as when the snow cover is 50-60 cm deep (Table 36). With deep
Ereez[ng of the soil and a 30-cro snow cover there was a greater reduction
~ in the soil temperature. When the snow cover was this deep there was almost
no change at a depth of 3 cm. The maximum amount of the daily fluctuation
of the soil temperature at a depth of 3 cm when the air temperature ranged
from 2 to 27�C amounted r.o 0.8�C. In 83 percent of 190 cases the daily
amount of chaage in the soil Cemperature ranged within 0.0-0.3�C. And
when the snow cover was from 11 to 20 cm deep the amount of change in the
soil temperature was 20-40 percent of the amount of change in the air
tempernture.
E
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- 1~Uk (11~1~ iCIAL (?51: c~N1.,Y ~
T~bl~ 33. Seaeonal Chnnge in Minimum Air Tempernture C(�C), UepChs of
- Snow CovE~r h(cm), Depth of ~reezing of Soil H(cm) and
Minimum Soil Temperature at Depth of Tillering Node~ of Winter
_ Cropa t3 (�C) gt Nemchinovka Agrometeorologicg]. SCaCion ~rom
1965/66 Through 1974/75. _
x~ xtt t it trt
Data~rea~ -
- ^~Y~ 1 ~ a 1 I 9 I ~ t I 2 3 1( 4 3 I I 4 I 8
1965/66 C.
t --4 -22 -20 -7 -10 --9 -14 -27 -25 -3! -18 -11 -6;-18 -8
h 4I 11 4 5 4 25 30 33 41 41 62 51 44 43 21
H ~1 12 15 15 18 20 18 16 18 22 28 27 23 19 15
UI -5 -3 -1 -4 -2 -2 -3 -2 -2 -2 -3~ -2 -2 -1 '
1968/67 r.
, t -17 -4 -4 -24 -2? -18 -70 -28 -33 -32~-24 -17 -5 -12 -9
h 9 13 U -0 25 25 39 35 42 d5 45 44 45 41 28
H 6 0 9- 39 53 58 58 62 68 76I ?8 79 75 68 61
-4 -4 -1 -5 -4 -3 -2 -3 -4 4 -3 -3 _2 _1 ,
1967/68 r.
~ ~ 0 -2 -1 S ~ 22 -2a -28 -30 -32 -30 -25 -25 -25 -19 -:S -8
h� 0 1 6 20 26 28 39 48 50 47 57 67 63 65 0
tf 0 0 2U 20 18 25 23 31 40 37 42 44 42 40 32
1~ 0 0 -2 -2 -3 -2 -2 -2 -1 -1 -2 -1 -2 -2 -2
1968/69 r.
_ t -7 -18 -19 -28 -?8 -16 -27 -32 -31 -31 -27 -18 -23 -19 -17
ti ~ 5 9 14 1~ 18 20 2I 24 31 23 25 26 26 10
N 12 22 26 19 21 20 34 49 66 74 82 89 98 104 107
ta 0 -3 -4 -3 -3 -2 -5 -6 -6 -4 -5 -5 -6 -4 -4
1969/70 r.
t -13 -2 -3 -8 -19 -28I-25 -37 -:i2 -20 -24 -20 -16 -15 -12
h 16 0 0 0 9 10 29 a:J 35 44 65 59 57 64 61
N 0 0 0 2 14 61I 73 71 70 74 79 0 0 79 79
t3 -3 0 0 -4 -6 -10 -10 -6 -5 -6 -5 -6 -6 -4 -4
1970/71 r. -
~ -17 -4 -9 -14 -14 -20 -20 -20 -3 -18 -24 -22 -21 -18~-11
18 U 5 8 IQ 11 16 S II 6 18 12 10 l0 0
K (1 0 15 1T l5 15 27 34 34 35 43 3 93 1071 109
~3 -1 --1 -3 -2 -2 -2 -2 -4 -2 -4 -6 -4 -5 -4 -5
(Table 33 continued on following page]
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~
xi xii i ii iu _
t1oM~~~t~~? _ -
~1~ 4 I~ I I 2 I 0 1 4 I~ 1 I 4 ~ I I 1 I ~
1971~72 r. ~
_ t -8 -14 -9 -10 --72 -24 -24 --31I--26 --26 --1 --16~-20 ~--20 -8 -
h 0 2 3 10 y 16 17 18 41 18 18 13 13
~1 n ~J H 5 1 S 23 31 49I 6U S 1 86 88I 93I ( U2 100
13 -2 ~--3 -1 -2 -2 -3 -4 -6 -7 -8 -S -5 -8 -10 -3
~974~15 r. -
~ -G --6 -8 -11 -12 -8 -19 -2-1-13 -15 -?6 -18 -20 -7 -7
h 2 U 0 9 9 10 13 I1 '?1 26 40 36 ?3 0 0
H o 0 11 l0 12 li 16 19 20 20 21 22 2I t9 0
r~ -2 - -2 --3 -5 -3 -4 -4 --2 --2 -4 -~3 -3 -4 0
Key:
l. tndicator _
'11ie ab~olute soil temperature at a depth of 3 cm throughout the entire
pertod when a snow cover of 30 cm and more fell on slightly frozen soil
(lesH than 50 cm) remained within the range of 0--3�C, thaC is, wiChin
- the r~nge ehat is mosl� favorable for Che processes of perishing of
plants under Ctie snow. When a snow cover of 30 cm and more was formeri
after e~ie soil was frozen to a depth of 50-60 cm, the soil temperaCure -
near the tillering nodes of winter crops is considerably lower (down to
-6 - -8�C) . .
~ With very severe cooling of the soil (when it freezes down to 120-100 cm
and deeper) before the establishmenr of a deep snow cover on the fields,
the minimum temperature of the soil during the winter can be even lower
than -20�C but when analyzing the materials of observaCions from the past
25-30 years we did not discover any cases like this in regions where
plants perish under the snow. Therefore when a 30-cro snow cover is formed
on deeply frozen soil the plants are in a dormant condition throughout
the entire subsequent period while it remains, respiration processes are
very slight, supplies of sugars are expended sl.owly and there is no perishing
of wtnter crops even when the snow cover remains on the fie].ds for a long
tim~.
't'he de~tl~ of freezing oF the soil in the nonchernozem zone changes from
year to year within a very wide rarzge (from 0 to 150 cm and in the
northeastern regions, even more). But most frequently at the end of
the winter it is less than 100 cm. A rapid increase in the depth of
freezing of the soil takes place when there is no snow cover on the fields
or it is not very deep and the air tE:mperature is abnormally low (Table
37, Torzhok Meteorological Station, Saransk, 1952/63).
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- Key:
1. IndicaCor 7. Aeprh of snow cover (cm)
2. Minimum temperaCure (�C) S. DepCh q~ freezing of aoi~,
3. Air in cabin 9. Height af grass stand
4. At heiBhr of grass atand tT 10. Difference nf eemperatu~~,>.s
5. On surface of soil t~ 11. Gradient of minimum temperature
6. At deprh of tillering node in grass stand (�C/cm)
Table 35. Influence of Density o� Snow on DepCh of Freezing and Minimum
Temperature ef Soil at Depth of Tillering Node. Belogarka
Agrometeorological Seation~ Leningrad Oblast. -
~
1964163 r.
n01f~7~TlAb YV~CTOK
~ 1~ ~ 2~ He~ps I me~p~a~ w~pr I~?pG S?
B~rcora cFie~~~oro noKpooa ?ia l 23 30 30 6
xoHeq Mecaua (cM) II 2U 34 35 16 -
~ Ilnoriiocrb cncra Ha KoFicu Me� I OJ8 0.25 0.30 0.29
(8) cAUa (r/cM~) 11 0,34 0,34 0.36 0,39 =
~9~I'ny6ii?in npoaiepsauHa noved 1 75 88 98 100
Ha Ko~tetl MecA4a (c~+) 11 80 93 100 !05
(10)~~~~NU`ianb?+ae Ten~nepaTypa I -5,8 -2,7 -2,7 -2,7
novnW Ha rny6Nee 3 cM iI -7,3 -6,5 -3,9 -2,7
sa ~+eceu (�C)
_ (1~)CpeltNaA MCCA4H8A TOMflCQBTY� I -2,4 -1,9 -2,1 -1,3
pa noyad (�C) 11 -4,0 -3,2 -2,5 -0,8
~12~MMNlIIIAfA.9bH:lA TE~II1PpeTYPB ~ -.~~3 -30~8 -a~~3 -~~5
803AyX8 3A MCCHU ~ C~
~ 1963/64 r.
1~+. ~~TtJI? y~ 2~K ( / I 4y I PP I !C I
411~b iH\,Jtr ~tl'i~Ab IJ~T ~~t.1?
)
(7~ Bacora cHe~cFioro noKpoea xa 1 l2 9 25 14 7
KoHeu MecAua (cM) II 12 l3 29 16 12
- (g) flnoteocTb cNera Ha Koeeu we� I 0.22 0.25 0.22 0.28 0,28
c~u~ (r/cM~) I t 0.30 0.38 0.38 0.40 0,39
1'nyGnua ~POMCP38NNA noyew I 25 55 72 81 79
~9~ Ha xoueu Mecnua (cM) II 57 67 96 99 97 _
M~~iiN~~a~bu~A reMnepaTypa I -1,2 -1,3 -2,t1 -2,4 0
- (lU) novnw ua r.~yGnne 3 cM II -7,1 -8,0 -5,0 -4.R -0,5
- 3a ~iecAU (`C)
(11) Cpcn~~s~A Mecav?ian reHnep~ry 1 -0,6 -1,4 -1,5 -1~6 -}-0,8
pa nu4aw ( C) f I -2,7 -2,2 -2,9 -2.8 -0,3
A1i~i~i~~+an~uaA TeMnepaTypa I -28.7 -20,7 -28,? -29.2 -6,?
~12~ 003A)'r~ 3~ AICCAL( (�C)
[ Kc~y on following page J `
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Key :
1. Indicneor
2. SecCion
3. January
4. FQbruury
5. March -
- 6. hpril
1. Dipth of snow cover at end of month (cm)
8. DensiCy of snow aC end of mont~ g/cm3)
9. Depth of freezing of soi~. at end of month (cm) _
10. Minimum soil temperature aC depth of 3 cm during month (�C)
11. Average monthLy soil temperaCure (�C)
12. Minimum sir temperature during month (�C)
- 13. December
Notc,: L--section with natural snow cover; II--wirh packed snow. ~
'I'~7ble 36. Limits oE Drop in Temperature of Soil at Depth of 3 cm ~dith
Various Drops in Air Temperature, Depthos of Sn~w Cover and -
bepths of Freezing of Soil.
` , Bwcor~ cHeMnoro noKpoea (cw)
~1~
RptilAM ~ I 20-22 I 30-55 I 60 I 60
nonM~ene~
terneairyp~ ~ 3~ ~ayaHH. npoHep�eNa naved (ew)
w~ays~ (�C)
~o-ao I ~o-eo I ~o-eo I~o-~o I ~o-eo I so-so I so-so
-5 ( -0,8 -2,0 -0,4 -0,1 -0,1 0,0 0,0
-10 -3,3 -4,5 -1,T -0,3 -0,1 0,0 0,0
--15 -5,8 -7,2 -2,9 -0,T -2,1 0,0 0~0
-20 -8,3 -10,0 -4,0 -1,0 -3,3 -0,2 -0,1
-25 -10,T -12,6 -5,4 -1,2 -4,5 -0,5 -0,3
-30 -13,3 -15,3 -6,5 -1,5 -5,8 -0,7 -G~6
Key:
l. nepth of snow cover (cm)
2. I,imits of drop in air temperature (�C)
3. Deptt~ of freezing of soil (cm~
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x I ~ I
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. ~ ~ ,
A ~
~ u~NO'd' NOM N.+OCV ~OCdt~e'~ N~tOC'~ `
~ g N ~'I I`''I I'"I I'"I; ~ I ~
s ~ ' ~
I . 00
r~ a ~ ~ a -
v .roo.� oc~o... ooo.� ooo.-. oo~n.-�
o U v - -I- I -4- ~ ^ -I- -4- ' ~ ~
a ~ . 'r'
u ~ , 3
C1. GJ M ~r~a~te�o~ ~ v ~C v ~ ~C ~ v ~l" v Z ..~~C ' ~ v ~C w ' e-1
A u ~ v r-I
~ O ~
~ N cd t+~ e~ tp tp W
~ N ~ N ~ ` = ` . ~ . ~ _
L v ~ ~ ~ T ~ ~
- b~o 7 ~ .b
~ oo Ev ~
x ~
~�r~l H a Q Y m M
r-1 41 u = Y a ~ ' ~
c0 rl ~ _ ~ a a ~ ' O
~ rl �
O rl td 7 ~ x 4 = x
u~ H A a ~ p, o a ' ~ r~
c0 1 r v Y., a U a.. . xa.-. e'~
tn O~ ~ ~~a OU Q~ ;
o,~ x y ~ V~ a
~ o~ ~ee x~ r-1
s � ~ ~ ~ yc~'< A> a0 ew Ka ~p~aHe� Ne~auaN r
(3) (Q) ~S~r 6
- ~ Xopowue(7) 1-5 y~0,99x~}-t,n2 (7) 0,5~ 0,92
3 Ynoeneroop
te~~,uwc l~~) 6-ltl y.. O,~x 0,40 (8) 0,35 0,9f'i
7 MoxNC~g~ 11-12 y�.0,52x 1,02 (9) 0,32 0,88
~~~~~4 V a ~~n7.C 1' ~~0
t Ovei~b nnoxE~e~ ~`15 y--U,dTx 3,6T (il~ 0,21 -U,91
Kcy : -
1. Evaluation (points)
2. Wintering conditions
Number of 10-day periods with h 30 cm or more
4. Equation ,
5. Mean square error of equation Ey
fi. Coefficient of correlation r
7. Cood
8. Satisfactory
9. Poor
10. Very poor
'19ic rclation between the productivity of the Vyatka and Vyatka 2 strains
of winter rye and the Mironovskaya 808 and PPG-186 strains of winter
wiie:i~~ c~n the one t~~nd, and a number of surviving stalks in the spring
,~s compared to the autumn, which was considered in the works [17, 22],
turned out to be sufEiciently close (r = 0.80 i 0.86): The number of
stalks on the ].Oth day after the renewal of spring growth characterizes
not only the condiCion of the planted axeas in the spring~ but also the
winterin~ conditions for the plants. IC takes into account the degree
of. winter damage and possible consequences for the remaining plants in the
gubsequent period of their life. The number of stalks in the spring as
compared to the autumn can increase under good conditions (as a result
~f sprin~ bushiness), according to our data, by 30-50 percent caith
plants tl~ut were well developed in the autumn (3-4 shoots) and by 100-200 _
125
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t~Uk c)l~~'tCtAL U:~i: ONLY
percent in weakly develaped pl~nts (1-Z ghoots). WiCh pnor winCering
Conditions, e~peCiglly with dnm~ge to win~er cropy ~g ~ re$ult ~f peri~hing
und~r the t~now, a numb~r nf gCa],k~ in rhe gpring on ~n ~vergge for the field
eurn~ out eo bp considergbly les~ th~~n ie wgg in eh~ ~ueumn [2x~.
_ Cn Hever~ly Chinned plnnred ureng as ~ resule of perighing of Che pianeg
under the snow during thp wineer the cnngequ~nce~ ~r~ m~ni~~gted eo a
gre~eer degree. The reduceion of eh~ produceiviey of winter crops in ehe
- nnnchernnzem zone during warm winr~rg wieh 1~CC1~ gnow i~ explgined
primarily by ehe d~aCh from p~righing und~r the snow of n l~rge quanCity
of ~ealkg which were �ormed in th~ gutumn gnd are several times mnre
producCive than g~alk~ gppear in the spring.
On the f.ields of the Scientific Regegrch Institute of Agriculture nf
Centr~l Reginn~ of the Nonchernor.em Zone, wiCh n high level of ~grntechnology,
the produceivity nE PI'G-186 gnd pPG-529 winter wheat ran~ed from 26 to 33
_ ryuintal~ per hectare when the number of live sCulks in the spring amounCed
to 65-93 percene of the ttumber in the auCumn and from 40 to 45 quintals
per hectare whett in the spring the number of stalks was 139 and 154 percene
of tt~c number in the auCumn.
M.yny years of observaetons ~?t the Royka Agrometeorulogical 5tation (Gor'kiy)
Erom 193~ through 1963 showed that the productivity of Vyatka winter rye
und U1'yanovka winCer wheat, with idenCicnl bushiness in the autumn, has
a directly linear relation to the quanCity of sCalks that survive after
wintering. The coefficient of the correlation beCween Chese amounts wiCh
weak development of the plants in the autumn (in the phases of shooCs and
the third leaf) turned out to be 0.88 � 0.02 and with well developed
plants (3-4 shoots), 0.80 f 0.03. This made it possible for us, wiCh daCa
from observations of the mass network of ineteorological stations (1962-1970)
to calculate on an electronic c~mputer the depandency hetween the produc-
tivity of winter grain crops and the number of stalks an the lOth day
aEter the renewal of grow[h in the spring. The supplies of moisture in the
soil in the spring in the nonchernozem zone, as a rule, are good and
_ thereEore they have not been introduced into the equations.
_ An~lytic:~lty, this dependency is reflected by the equations:
1) witli winter ry~ plants oE thc Vyt~tka and Vyatka 2 strains that are
w~~.~kly developed in the autumn
~ y = 0.108P - 1.3, (12)
wherr y--productivity (quintals per hectare), P--relation between quantity
~~f stalks in th~ spring and quantity of them in Che fall on the average
for the fic~id multiplied by 100; r= 0.80 f 0.03; n--the number of
tnst~lnc�ea. exactly 55. The mean square error of the equation F~ 1.58
c~utnt.ltti per hectare; the amount of. the coefficient at P--quintals per
hcctare;
126
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~o~ nH~~~tctnL us~ dNL~r
2) with'~1~u~h~d pl~neing~ a~ wine~r ry~ -
y ~ d.U~~kp, ~1~~ -
, _
- k--~verng~ CO~~E~C~.~riC of bushineg~ df p1~nCg in the ~utumn; tt e 150,
_ r~ C.~4 ~ 0.04; 2,3 quinCalg per hecCare~
~or winter whent of eh~ Mir~nov~k~ya gOg gCr~in ehi~ rel~~i.on ig
~xpregg~d by the equaeiong:
1) for pl~nted nre~~ ehaC hnv~ noe bushed oue in Che nutumn
y ~ 0.12P + 2.00; (14)
n= 65, r+~ 0.89 � 0.02, ~Y ti f 2.1 quintals p~r heCtnre;
- 2) with plnnCed ~rens th~C have buehed ouC in Che guCumn
~
y = 0.0866kP + 1.02; (15)
_ n~ 180, r~ 0.86 ~ b.01, = 22.5 quintals per hece~re.
_ Tt~e conventional symbols are the same. The equaCions ~re valid when k is
equgl ta or less Chan 6~nd P---from 30 Co 200 percenC.
'fhe average number of stalks in the spring ie deCermined by using these
equarions from dgt~ of aut~mn and sprittg examinatione of plgnting nreas
~nd [he equntion
kg uSg
P = k (100 - 100~~ (16)
0
where kB--the coefficient o~ bushing of winter crops in the spring,
ko--ttie same in tl~e autumn, u--the average thinning of winter crops from
- four repetitions (percentage) from data of a spring examination of the
planted arens, S~--the area of the field (percentage) with this amount
oE thinning. Zfiese amounts can also be calculated from the dependencies
= presentecl~~above (Ei~ures 36, 37, 38) .
rrom tf~e resulta of nnalagous calculations made for various values of
Iiu~hines5 of plants in the autumn and wintering conditions, quantitative
indir.at~rs were obtained for evaluating the condition of winCer crops in
tlir gpring on specific fields (table 45). Here bne assumed, as was
.~lready indicated. a certain r$tio between the point evaluation of the
condition oE the planted areas and the possible productivity of the winter -
crops with optimal supplies of moisture in the 1-meter layer of the soil
in tlie spring and the modern level of agrotechnology (table 46).
127
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1~'OIZ Ul~i~ It;1AL l1;il~ ONt~Y
'1'gbl~ 4S. QunntiCative ~ndicaCox~ di th~ ~vnluaeion df Che Condieiong
~ (pointd) nf Areas p1~nCed in WinC~r Itye of Che Vy~tka Srrgin
(I) and Wineer Wh~gC nf Che Mironov~k~yg 80g SCr~in (~Y) in
Che Spring on 5p~ciEiC ~ield~
CaAMt! ~ KOf~~NUMIMT MyltNtilfCtN plCtlNNA OtlH~10 (4MCa0 110dlfM)
n~omy~ no noia
s (~1 c kawveer~o
ao~.eWMNM ~etN
A p ~ ~ ~ ' g �
naeuwM IX or Mx
OtlMMl~O
~1~ ( Z~~t�p t il t I 11 t It I t~ It 1 I ~t 1 it
~p ~'S0 1 1 2 1 2 2 ~3 ~3 ~3 ~3 E3