ACADEMY OF SCIENCES USSR POPULAR SCIENTIFIC SERIES

• Document Type: CREST
• Collection: General CIA Records
• Document Number (FOIA) /ESDN (CREST): CIA-RDP82-00039R000100240012-2
• Release Decision: RIPPUB
• Original Classification: R
• Document Page Count: 330
• Document Creation Date: December 22, 2016
• Document Release Date: March 20, 2012
• Sequence Number: 12
• Publication Date: March 24, 1952
• Content Type: REPORT

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STAT
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p~,A~T~ A~VD ' XMFi~YS
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~r~de~? the general editc~x~h~.p off' the Co~i~rn~.s~ian c~ the Ac~de~'
cf ~cieneea USSR fcx pub~.~.sha.r~g popular scienti~'ic
1~.~~?a,tuxe
ia.n c~~' the Gorn>~a.asien, ~xeaident 'v~ the ~~adem~r of ~c~.e~cea ~1SS~.
Cha
~.oaderci~ ~. ~. , VAU'l~~
ut Gh~.ix~ C?rrespanc~.rt~ I~mbex of the Acad~er~' a~' Sc~.ences ~~
Dip y
F o F o YCJD~~
~DT'~?R~I~-CHTF~ ',
oxres c~nc~ng r~;mb~x of the Acader~' o~ Sei~ncea U~~R
G p
I,. A. Ivanav
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When ~~Entgen mach '~iiis ~x~eat `di~cavery in 18~~ he could
haxdly have suspected the full. bcdpe of ~.ts ~pplicatic~n. Qn
naw, fin' years ofr the c~.scbv~r~' can ~~ apprec~.ate ~.ts ~~.~~
ni.fcanc~? ~' Wing ar~l~r the b~.~l?~ica1 effect of these ra~sg
`wch pa>~sese atimula~in,~; .:and ~anexatly~ ~la.tiea9 arse ca;
only marvel at the poor and ~~.n~,.f~ce.nce of i th~.s new taal
~rhich hay been p~acad at the d~epasal a~ eex?imental ~ciehce.'
~~ny three ~D yoa~?s a va~~ a~aoux~t of e~~er~~ts :and
c~bee rod t~.ar~s have been pe rf arrr~ d de~nar~s tra ta~n~ the e ~"~'e c t of
~~rays on aural end ~ala~t lif?m During th~.s pex~.ad the i,nte~~
e et which biology; ~ ?~ have Shawn ~.n tx~a ~ee rays h~ s floc tua ~e d
Althau~~a the:eacp~ctatians off' the fire investi~atars in the
Held of ~aiedicine haves in ~e~ra~.,~ been reala.~ed9 after the
f~.~?et b~?a.ll~.ant success a'per~.od af' diailluszan set in b~fare
the successful use of -thee rsys way conclusively established
far diagncsi~ and cure of nun~eraus 'diseases.
Si~rd.~.ar fluetuatid~ls in' attitud+~ can be n~ied ~.~ the
evaluatdon.~c~f bidlo~ical effects of X-rays dep~~adin~;'r~n
the success ar failura of the experiments . performed4
ln~,~ x~~ s t ~.n the effects' of : shart waves a~~eare d shartl~r
after the~.r ~ d3:sco~rery but the peak cf interest was reached. ix~'
1927 when ~~aller made; h3.s 'annnunaero~n~t abqut the multi~`old ink
crease in the number of mu~tions. obtained by the use: of Giese
rays on ;fruit `f~:~es. Cer~a,in of osr bin/?~~strs responded quick-
1y to Mu1~.axt s discovery. It seed that a .method had been
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action. As a result a sceptical attitude devo1oped in relaUon
to the poss?bility of stiniuIatiz-g plant growth by radiation of
seeds or sprouts. Only in the last few years have worke bea
to appear (in the United Statee as well ae in the USSR) which
indicate that with appropriate dosages of these rays it is possible
not only to std~ulat the growth and development of agricultural
crops, but even toincreaae their yeld?
We feel that our sketch should be prefaced by\ a short
explanation of the physics of these rays. But first of all it
is imperative to say a few words about the nomenclature of the
raysm
Roentgen called then X~rays since he didn't understand
their properties clearly. But in due time their name became
so closely associated with his that they quite prpperly received
the na of Roentgen rays. This is particularly' fortunate since
it i,s flloet canvenieft for the coining cif such new terms as
roentgenbiology, roentgennutatjon, etc. However, on certain
occasions we will reevrt to the term X-ray for brevity, as.
n iv contemporary authors do. [sic] [In this English trans.,
lation, however, the `term X.'ray will by used exclusively.)
'E PHYSICS XwI~AYS
Investigators .in recent years have shown that atoms, which
were considered indivisible, are actually complex structure
consisting of a central nucleus: and an outer shell, composed of
the smallost eaclstirg particles ` of ;matter, carryng rne` ative
charges. These particles are called electrons. In 1
order to
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obtain X?raye it is necessary to have, not only electrone from
the outer shell
of the atoms, but also electrons separated from
atoms or "free- electrons". The process of formation of
X-rays
tares place in a vacuum tuba, filled with rar fled gas, where
by application of high voltage across the tube free electrons are
lit off (from individual atoms`) and directed at e. very high
speed toward the anode, thus forming a beam of "cathode rays".
The electrons are propelled along their path at a speed approach-
ing that of light. In order to impart such speed to electrons
it is necessary to expend a large amount of energy which is de-
rived from a special source of current. If a meal- play is
placed in the path of the electrons, they will strike it with
tremendous force, their speed diminiehea abruptly and at that
Lnstant X~rays are formed. This plate has to be made from ?a
metal with a very high melting point since electronic bombard
merit generates a large amount of heat. Such a plate is called
the anode' of the tube whilethe tube itself is called an X-ray
tube. The mo8t prevalent type of electronic tube is the so..
called 0colidge tube which was first made in the United States
in 1913. e X~ray, tube is, essentially, a container 'from whidi
air has been pumped out. Inside the container are two electradee~
one is corn?cted to the ar3ode, the other to the cathode) which
constitutes the source of electrons (fig. 1. X-ray apparatus
b
in action). X-rays are short wave radiations ranging in wave
length from 0`.02 to 1100 angstrom units; a unit of length which
is equivalent to ,1/10,000 ' or 1/10 milli ' The energy
radiated by the tube is not unU'o consisting of both short and
long wave radiation. The qualitative composition of the rad$a'
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ion deper ds on the intensity of the voltage : the `greater the
vo1tago tho greater is the quantity of short wave rays produced.
X~raYa are invisi ile and can only be detected by their:
action. Tree basic runs are used to dethct rays. (1) fluore
,
cences (2) photsgraphY9 and :(3) lonizaU-ono The first method
,
_ ask
is based upon the ability of" rays to cause visible 1umfx~e8cence {
In certain ,3ULstanes8' If a screen is made from such substances
it will light up every thc thaX~rays touch it (fluoroscopic
, ,.
examination is based upon this principle ) The photographic ac
`
~tion of X rays is similar to fluorescence but more pronounced.
It i$ pos ible9 for exa p1e to obtain photographic prints by
means of X~rays. These prints can be retained as documentary
evidence for an indefinite period. At the present time Kostov
(i3) has had great success in photographing ears of wheat by
means` of X~rays; a, method which enables him to obtain not only
he external morphological appearance of the ears, but even the
extent of development o the kernels` inside the ear, due to the
penetrate. action of X~raya. Such photographs, produced by
the action of X-rays, are known as radiograms.
The ionisation method is !based on the ability of X rays
to cause air and other gases to -became . conductors by i xr arts
ing posiU`=e or negative charges to `ions of neutral ga. mole
cules. Tern amount of current passing throug the ionized gas,
with a given difference in potential is proportionai to the ins.
tensity of the X-'rays ? which cause the ionization. Special
ionizing chambers which are used in order to measure he degree
of ionization of the gas, can serve as dosage meters, for quan~
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titatis measure ent of X rays.
In their optical properties Xwrays show all of the
characteristics of light rays. They can be 'refracted and re~
flected. They exhibit the phenozr of po1ari~at3 on and di
fraction. The ability of Xcrays to penetrate deep into the
tissues and to induce vaxious changes there is. of particular
significance in biology. It should be noted that only rays which
are absorbed exert any effect. Rays which penetrath. 'feebly or
Chase which are completely re f1ecd have no action. The 'abil
i.ty to penetrate increases as the wave length is shortened.
Dorm the very first year of discovery of Xrays
people working with them experienced serious inila Cations of
the skin accompanied by iris of hair. It soon became' apparent
that he action of the rays i not limited to their . effect on
the skin, but that all of the internal organs of men and ani~
ma.28 are affected if the amounts of exposure are sufficiently
great, Eadia Lion therapy is based upon' this phenomenon.
19Q3 Heineeeke amazed everyone by his discovery that
niice and gu i ea :pigs can be killed by X-rays. Autopsy of ani-
male ki~.ed in this manner showed that their' spleens were small:'
and darks while a hiw o ogica1 examination revealed ai ectra
ordray' increase in the aariount of pignt, and the disappear
ante of follicles and cellular ele nt5 of the spleen. S ir1,~
lar diiargs took 'place in the lymph nodes, in bone marrow, a to.
x
In `cases of eub-letha1 dosage a regeneration of lymphatic tisanes
con ences aft.er several days. Roweyer,, a diecusaion of ths ae-
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pect of X~ray action is .outside the scope of this book; those
interested in this phenozi non are referred to the book written
by Nenov in 1926.
hE MEASUREMENT OF ENERGY
Before proceeding further with our account some concept
of the units of xneaaureirient `of X .rays should be established.
This is particularly imperative in view of the fact that the use
of various units of meaaurern~nt by the numerous workers in this
field increases the difficulty of `comparing the results obtained
in the radiation of various plants and anim als
The reiosV widespread unit of measurement, up to recent
years, was a dosage bearing the abbreviation fED (H gh~Erythetna?
Dose ) This dose is equivalent to that amount of rays which
results in a reddening and peeling of the skin in roan, ten days
after exposure to radiations But this n~asure, as most biolagi..,
cal unit. j too inaccurate since the indicated signs of actio~
v xIy, dependi on the age and condition of the given patient
and on the pigmentation of the skin. Glark, who a camined
ry
patients in numerous clinics, has `shown that the erythenic dose
varies from 14po r to 1?QO r (roentgen units), depending on in
dividtaa]. diffe
rences and that the average dose equals about
8S0 roentgen units e
i4ary ; Geran and French investigators used the Holtzkxcbt
unit (H), which is equal 4 about of the erythemic dose,
measure' X-rays,`
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 of energy required. a re8ult or cornp.Lete 1on1za1,Qn J.u
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This .latter unit. was abandoned .in favor of a new inter
national unit x (roentgen Wit) which corresponds to the amount
Go D,Gc4~
airs at zero degrees centigrade and normal atmosphere pressures
ch~rgas 'of cne electrostatic unit per cubic centiu i r
of the irradiated object.
Dosage rretern used for quan .tative deterri.natiOnvf
X.rays 'give the u asuremeent in roentgen `urats ? The majority of
investigahrsq however, are not content to indicate the amount
of X~rays in roentgen Units; they also indicate the source of
radiations 'the voltage, the amperage, the distance of the object
from the tubes he duration of ` exposures and the quality (or
absence) of 'filters.
Filters are very thin plates (of aluminum,
and sew to dispex the X'rcys of
This
copper, etc
specific wave lengths.
makes it passible to ach eire aor~ un i.fcrmity (of wave lengths)
in the beam which irradiates the object.
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C ik D
PHYSIOLOGICAL CHANGES PRODUCED BY X-RAYS
As has been stated above, 'cellular reactions to various
irritants depend. on the power of the irritating agent. At first
normal cellular activity is accelerated; exposures of longer
duration to '? stronger doses produce depression; prolonged' ex~
poeures result in death. This thesis has found confirmation in
the woes of the most recent investigators and was most prey
cieely formulated by Arndt, "Weak irrits.nts stimulate activity;
medium 'irritants depress themy strong ones halt them." This
formularization has been 'extended by Schultze to include all
processes of animal and plant cello, of diseased, as weell as
of healthy organisms. This generalization has become accepted
in literature as a basic biological. law.
Since X-rays have an irritating effect on the activity
of organiams, it was necessary to determine to what extent they
conform to the Arndt 'Schultze law.' There was little objection
on the part of the majority of investigators to that part of
the law which states that strong doses of radiation tend to
suppress normal activity. Numerous experi nts testify the
depression of the development of organisms or tissues which
have been subjected to strong or prolonged dosed of radiation.
Concerning the other part of the law, however, opinion is sharpy
1$ divided. While some point out the stimulating effect of
weak doses, others deny thin effect altogether, or else th
y
adkaowlede it s a teaipprary apur of ?;actlvity whlah is
followed by normal, or even depressed, development. For exarapl
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in 1923 Haltzkneoht wrote: "We have no basis for assuming that
,
is harmful that, in Some amount or other,
becausea substance
it may be beneficial. oncequent~'', since X-rays fatigue and
kill cells, we ..cannot conclude that doses exist which stimulate
or accelerate cellular development. Fifteen years of roentgen/
.
ology have established only the depressing effectw of X-rays on'
cellular activity."
It should be noted that the term ," stimulation" is am -
includes negative, as well a~ positive,
biguoussnce stipulation
rcponses to stimu].i. Undoubtedly certain differences of opine
ion stem. directly from the use of this word. In order to
misunderstanding the teas r"stimulation"
avoid this .:type of
more generallY accepted sense, i. e.1 to
will be used. in its
indicate the positive reaction.'
e of X-raYS which stimulates the life prOO?Ssea
The . dos
ommonlY referred to as an "irritating dose". a
of plants is c , 'ram
.
the ;numerous `tiona of various euthors it is evident
; ~.nvestiga
that some of them have succeeded in establishing irritating
.:..
dosages for the subjects of their experiments, while 'others
,
ossibility of the existence of such dosages.
..deny the p
t investigators of the biological ef-
One of the ea.rlie s
fects of X-rays was chobey, who in 1896 set out to study the
B
'nature of X-raysin order to. detelmine whether they were iden-
or not. Schober selected sprouting oats
tical with light rays
as the most light sensitive object' he could find. saving
. ,
sprouted oat seeds in darkness, he selected the most vigorous
specimens ( tars high) and planted them, five to (1-2 centime .
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cup, in four small cups.
He then placed the cups in a darkened
box and subjected them to the action of X-rays for ono hour,
with the source of the rays `being one centimeter
away from the
box. Careful observation of these pants, did not reveal in
them any evidences of (heliotropic) bending. But when the box
with the cups was placed on the window sill and a .diffused light
was permitted to enter through a. narrow slit, after an' hour it
was possible to observe a bending of the aprouis-in the direction
of the light. After 2 and ) hours this bending was even more
pronounced. On the basis of this experiment the author con-
cluded that X-rays were not identical with light rays.
In 169.7, Atkinson set up experiments. in order to doter
mine the effect of prolonged irradiations on plants. Having
established, by preliminary experiments, that radiations of 1 to
10 hours on 1eaves ` of Cam, on flowers of Bed, and on
various sprouts, do not produce any visible signs of injury;
he sett up an experiment: with prolonged irrs.diations (L-hour
exposures). This experiment was conducted in `darkness. The
irradiated plants behaved identically with the. controls, which
had been 'grown together with them in a dark room. When the ir-radiated and control plants were placed a n light both groups ex.-
hibited pale green color. The difference lay in the fact that
the plants which had been exposed to X-rays attained full color
more slowly.
than the plants which had not been subjected to
radiation. This observation forced the author to `asme that
X*rays exert a' certain detrimental effect on plastids. The
author did not succeed in establishing any other di Sferenceffi
in the plants despite the prolonged exposure.
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Lopriare in his experiments an 1897 found indications of
the irritating effect of X~rays? By ? subjecting parts of plants
of Vallisneria s ralis to 'X-rays, the author .discQ:ved an
of the movement of plasmaz in the cells. 1r? subw
acceleation
jesting to the action of X-=rays sprouting pollen tubules of
: s
~ succeeded in slowing
two p/ante en:sta and paringtonia) he
down 'the process of sproutinga.
] diney i~d Thouvenin whO, in 1898, were the
., __
z-ar~v
A.-
w
perform eXPeriments with irradiation of seeds (they used seeds
,
of nvol ulvus arv , idum sativum, and Panum miliaceum) ,'
,. A, ,.- - - -.-..-.
came to the conclusion that X-rays accelerate the germination
addition they observed that X-rays do not affect.
of seed,i, In
the -formation of chlorophyl. since the sprouts come up from the
ground with the characteristic pale, green color?n.g.? Unfortunate
of seeds used (only 3 in each experiment and in
13r the amounts
each control) was not significant. The question of stiti.1ation
:
by exposure to X-rays, consequen'tly' remained unsettled as a re -
,
ult of in~ufficient evidence. however, by comparison with pre-`
vious experiment, this is a step forward since the authors
and the distance from the source of radiant
noted he amperage
tion.
In 19OL ?perthes ;studied the effects of X-rays on the
is faba which have since become the favorite sub-
seeds of
,~i..,..~..., ,~....~..,,
iments with radiation. By measuring. the length of
sect for exper
roots in exposed and control plants this author established the
, .,
retardation of growth in irradiated plants. In thie way his ex.e periments demonstrated the depressing (inhibitive) action of
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rays. }As a counterpoise to thi
in 1906 showed that he irradiated seeds of beans, radishes,
le ttuce, squash, and cucumbers ex1~bited a greater energy in
sprautixi, a more powerful 'development, and earlier bloom than ;
Koernicke used the seeds of
Brassica napes in large numbers. His experiments were dis
those of his predeces orw. In the first place, he investigated`
the effects of radiation on seeds in various states:
ite and ever increasing, dosages expressed in Holtzknecht
units (H), in the following qunntitiess 16, 20, and 26 HoltZ~
knecht units. With small dosages Koernicke observed temporary
arrest of growth; with more powerful doses, total' cessation of
growth. In the following experiments a dosage of 20 Holtzkneaht
units produced a small acceleration in Vim` faba and a signify
leant one in Brassia napus,The difference between the irradiated
plants and the control was striking; from 100 irradiated seeds
of 'Bra sea nape s0 sprouted; from the same number of control
seeds, only one. In due time, however, the differences between
the irradiated and control plants disappeared entirely.
.Iri 1907 q.~ifeminot used for his experiments 'seeds of
the gillyflower (Matte a) in batches of 20. ' Sixteen batches
rere subjected to proportionately increasing dosages
f KMray'
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ceived irarious exposures to radium and, 'four were fsetained for
Contro .
Two, control batches were
ticns, one at nnaximum dryness, and the 1a Mt` batch under en~
forced humidity. From hip experiments he concluder that re-
tardation of growth is characteristic for irradiated plants,
provided the dosage xs sufficiently great. Acceleration of
a
growth was observed when the plants were exposed to dosages
between 00o and 7000 roentgen units, but the results were too
small to be significant.
By
the time Schmidt set up his. experiment in 1910 it
had become `customary to express dosages of radiation in
Fim
~i~-ryttha~ose) ? This investigator exposed peas, which' had
previously' been soaked in water for b hours, to various doses of
X-rays (1/20, jjio, 1/li. 1/2, and 1 High Erythema Doses).. The
plants raieed from the irradiated plants possessed unusual size
to 0 centimeters) their leaves and pods were twice the
size of normal unirrada.atad) p Rants and conta. j.ned from , four
a
to ftre pea.$
r pod, while normal plants contained only two or
three. The experiments of this investigator brought him to the
following conclusions "It appears to me most probable that thie
fact should have practical significance for horticulture and
ruct?culture." Thus his experiments brought him to confirm
f
the existence of irritating doses while the experiments of
wetterer, in 1911, brought the latter to diametrically opposite
wa 5
coneluoions. ,urd seeds of sunflowers divided into five
qual, parts. The ;first part was retained for control, the second
e
xace~ved ~' ~ ,~~
ltz1(? t unit
left under normal condi-
the third received
10 H,'
the fourth ~- `20 , and the fifth ._ " ho H. Then the control and
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1rrad.aced seeds were plar ted in a soil which had been epriahed
with nutritious $ub$tance d Observations revealed that seeds
receiving doses of rr 10 Holt knecht unite c une up simultan,
eou ly with the control seedu; those 1lhi.ch r'edeived 20 units
came up consideribIy later, and those which received LO units
never came up at all. Further observations of. their' development
showed that while the control plants developed very fully the
irradiated plants
pr'o~
portionately to the increase oI' their exposure to radiation. `
The `experiments of Promsy and Drevon in 1912 a.re inter
esting because the effect of temperature on sensitivity tea -rays
was investigated, and also because the rays were used in frac
tional doses, eedK of lcntle, 'rye, bean s,X white lupine and
kidney-beans were usod. These experiments brought them to the
following conclusions Under normal conditions ( 15 degre'es
centigrade) the sprouting of seeds exposed to X-rays was ; omen-
tines accelerated and sometimes retarded. When the temperature
was ;raised to 354.0 degrees (centigrade) the results became more
exact. The do
es of X-rays applied by the authors under the
showed progressively poorer developments
higher temperature invariably exerted a beneficial influence on
the sprouting and development of the seeds.
In .1913, ;Seh~arz used broad beans (Vicia faba) in his
investigations, They were subjected to 30, 60, goy and ]30 seoon&exposures. The 1$0~seoond exposure 'produced the most :.fair
orable results; plants receiving this dose exhibited the tallest`
and most vigorous; development. Irradiations of five minutes
0 .
r he growth of the e
greatly xet retarded tplant, h however this retards.
;
tion did not dhow tap immediately; such plants cam? up at the
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same time as the others, but eventually their 'growth stopped
altogether, although the plants remained green for a long time.
The author did not observe any sports or atypical developnflent.
kind of discrepancies duo to occurrence of
In order to avoid any
unusually vigorous specimens, in his second experiment, Schwarz
selected the most vigorous plants for his control, those next
i.:n vigor of development for the weakest dose, etc. 4oasuremente
weeks after irradiation revealed that the
of the received a dose of 3 floltzknecht unite showed
the finest development' they had a greater number of stems per
bush and their yield exceeded the controls by 1j.0 . joses'of 7
and 10 Holtzknecht units lowered the yield by 2di, and`., which
the author ascribed to injury of the plants by insects and fungi.
;ql~ Nakamura gave
~~7t~~r;of h
imcnte with QO" sawn (tSiruriki"), the eee s of which were
subjected to 10, and IS minute exposures, after a brief so.a =
17?.
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00240012-2
 Declassified
ing in water. He points out that plants which' had been irrad1''?
ated for five minutes showed an increase in yield.
Komuro, in 1919, also established that irradiated r5eeds
of Oryza eativa ("Hekiyema") show accelerated germination and
that seeds in an air-dry state were Iese affected by X-raya than
plants whioh had been soaked for 12 hours. Optimum doses were
S to 10 Holtzknecht units,
During a fve.year period, from 1917 ? to 1921 Komur+~ set
up expeximents with irradiation of seeds and sprouts of rice.
On the basis of these expert mente 'Komuro came to the
following conclusions: 1. X rays exert a harmful effect on
seeds proportionately to the water content of the seeds.
2. The seeds of some pits accezerate their ddeve1opment; ecds
of other plants retard theirs. 3. Moderate dosages act as posi-
tive stimuli on the seeds.
sensitivity to X-rays.
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4. Yung plants exhibit a variable
192 0 Jungling irradiated pr outs of Vicia f aba and
then placed them in special !boxes with Mass sides?' The length
of the roots was noted on the walls every 2L. hours.' The reaction
to X-rays was judged according to the growth (in length) of the
roots and `the appe'irance of side branches. jungling observed
various injuries depending on the size of the `dose. The ceasa
tjon of growth, ~.n
engt
nd osa.on, of
of nee
side roote he,oonsidered as a maximum' injury. The dose capable
of producing this he called a "full dose! (l'Vol).doais") With
a' smaller dose slow ' growth would continue for two more days 'after
18`
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which the
rate of growth would accelerate and side .branohee
would begin to appear. This was the "medium dose" ("Mitte1~
dosis" )`? Reversing the process he judged doses on the basis
of Length of time it took for side branches (roots) to appear.
If the `ideroot appear two days later than the control plants
it means that 35% of
"full dose" was given. A delay of one 'day
indicates 2% of a full dosen. However, if 10% of a "full
dose" was applied, the appearance of the side roots was accel-
erated (see' Chapter h)? These observations indicate that Jung'
ling W9S seeking a biological' dose meter by his: experiments..
In `1922, three other 'authors, Schwarz, Czepa, and
Schindler, were>iore categorical in their denial of the stimulat-'
n? action of Xrays. Having set up experiments with 12 differ
ent species of plants (wheat, barley,. oats, pe, beans, broad'
beans, mustard, lettuce, Onabrych s and throe field grassee).
they did not succeed in establishing irritating doses. They,
believed that the major error of the majority of X-ray biologists
lay in' the fact that they used insufficient numbers of samples
n their experiments, ; which made it impossible for them to
away with individual differences.` However, they were able to
confirm the depressing effect of large doses of radiation on
plant growth. .
In'ag2?~ Sierp and Robberts attempted to introduce a
new factor` into X~ray research by examining the growth of separ-
ate organs and in that manner were able to analyze their action
in detail. They used sprouts of ,,wens, sva,whose coleoptylee
are extremely sensitive. These experiments indicated how correct
19
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ed examinations imperative: for
~ were in cons~.deri detail
th
uts of ram~.na~ turned out to be
the sheathing of spr0
exarnpls' if only the
is` difficult tO detects
extreme~.y serlsit:I.ve, but it
u clea~~' if separated
growth i observed, but it snows, p
overall ,
hrou h? After a cert~a.n
Ct~ges of dev'e~.apment are' followed t g
~
acceleration of growth the authors obserTEd` its ' retardation
1g ngth ~ of ~~cpo pure we ~
why.
ch became more pronounced as the jncreaed.:
laer`obsex'vations ' to the
ln 1922 ' La,llem limited
of
first stakes of development
lo~~.ce~1 effects of Xrraye on the
~. ~ras~iaa ~,pus'a
e ~,ep~.dium satiirum' ~-~
the seeds of P '
ar~:~
cu1ata' phaseoiu$ ' 1gax i and onion
-
- th day after
Qn the )
th of plants w'as measured
bulbsA the
i the basis of the follo~~
n is
rrad~.aticn. This time was.. selected on
urious effec
jn the it pie $s s the in j
~.ng cons~.deration.~ s
iately apparent'' tlieT be~
/ants do not become ii~uned
of x~raYe on p
Domunc e of time after ex~iosure~ and
ed w jth paa
come more p
tad on wader ind?f~.na.tel~' ~-~
and p],nts cannot be cult~.va
sec side ? roots9 leave
f f ectin~ plant developd,. Roos stemss s
out a.
.
bjacted to measurement.
and ` hyp,~~cotY were su
i atic~ns into two parts ti one
ingest ~
llsman' c3jvided her
rad
`
s, a
gr?li.p ed to weak ' doses of X-ray
>~, , were sub~e~ct,
_ of p lant ,
he first gxoup
f
medium
t
t
str?ng canes. 58eds o
in
others,
o?acs3ca, ud~
rs
~,udeda phasecalue i
for
c ..
' he seeds designated
'
Pa~c ns aGulent&? `
the experiment 8 sevd as cis urn
iw~dad into s'l6 sample s~ ~ 3 V
were , d
~ ~
9
.
and B~ c~thers were. irradiated by -Xrays in' 1/12,
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11
5, 10and 20 Ha1tzknecht units which
y with the apparatus she
was, using, corresponded to 5 15, ' and 30 seconds and 1, 3' 5s 1.0
and '20 minutes. The number of sudsy 4Periding ~?ied on the
Species of `plant being irradiated.
After?rxa.diatlon the p1ants
were placed between ..j0 layers of oatton i
n Petri dishes, and
at given intervals the sprouts seeds
~ were eounted Whose ex-
periments showed that the most diverse dome of xMrays failed to
hir. a baleful influence on either thy ox sowed seeds. ' In ore.
der to test the 'effect of Rpra s an r
a,t$ or growth9 experjrnents
were sett up' with seeds of lentils whey,
to and dried kidney berms...
From these experiments the author oanclu
ded that the weakest
doses of X-rsys do not stimulate
plant growth. In re1atian to
the second paw ofthe expera.ment i? e?, the act1an of Medium
and strong doses of X-rays: Lalleman set out to clarify two..
questjonss (1) de very strong; doses of X,-rays. destroy the abi1.
sty of seeds to gerrninate and ,(2) dc, mediumdoses delay the time
of appearance of sprouts. Experiments were
set up with various
types of beans and lentils, In her experiments the author; did
not succeed in establishing the inhibi
tian or retardation of
sprouting. In exposing 1000 germinet
ed lentil seeds to variou~
doses of x-rays and then measuring the lengths of stems, raote':
and side 'roots it turned out that, decrease in
sire is directly
pzxapartional to increase of dosage received by the. seeds..
The axpexd,nients of Altmann Roc
hlin' end Oleichgewj~sht
were yet up with unusual thoroughness fora, their time (1923) In
the first ;place they used seeds of a pure strain of
Vici& faba
from a dingy crop and of ids
nttoal weight (which ws.s deteran,~,ned
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iii
on" torsion ;scales). In addition they triple checked their ex-
per?ments and irradiated seeds in various state's (dry, soaked,
and germinating). The sprouting and `development of young plants were
most carefully and minutely obser'sred:. the appearance of side roots
and first leaflets was noted; the length of roots, stems, and size
of leaves were measured. These observations brought the authors
to the following conclusions, (l) by gradually increasing dosages
it is possible to obtain a transient acceleration off' development,
which carries within' itself the seeds of a' `depression of develop
mont, which depression is inver$ely proport:Ional to the brevity o?'
the acceleration; (2) the 'irritating dose" changes depending on
the stage of development ('or dry beans it was between 6 and 12 .Holtz..
knecht units, for germinating ones, between 1 and 3 Holtzkneeht
units)
:investigating the problem of the accelerating and rew
tarding effect of XWrays on the development of plants, Czepa,
in 192s irrada ated seed, of Vida f, Vicia saw, Phaseo
lus izuris and lettuce with various doses of rays, ranging
from 1/ to 2 Holtzknecht units, Only in one experiment did
he observe acceleration of growth in the plants used by him.
In all the `others he noted neither acceleration nor retardation
of development; consequently, he ass,.nied that the one exception
was due` to chance and didn't attach any significance to it. ? In
further experiments in which Cxepa used stronger dosages (l,?
and 300 Hoitzknecht units) the er~rgy of the sprouting of seeds
didn ' t diminish at all, while plants which had received lO Holtz-
knecht units actually exceeded the controls'. Czepa dial not con
tinue or intensify his investigatiens, although his results show.
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that stimulating' doses reslde ` somewhere between; 25 and 150.
Holtzknecht unit doses of''the rays. On the' contrary., as a
result of his experiments he came to the conclusion that stixa-
ulating does do' not exist for Vica faba, Vicia $ativa, Phased
lus vulgaria, and aettuwe.
Nartius ; in 1921, also failed to find a stimulating dose'
in his experiments in which he was trying to ascertain the deg
pendence of , the action of the rays on their tntensitye Using
filtered rays of high intensity (130 kilovolta,'L milliamperes)
he observed,
distance
veiopmonts
that' radiations of on)y lad seoonds with a focal
wr
.-
,
f 10 oentimetersIj uffiaient in order to halt de-
In 1921, Geller posed the question of the constancy of
the effect of X-rays on young plants, at a definite dosage, and
carne up with negative conclusions. Comparing the data of prey
vious `investigators he formulated the following conclusions:'
(1)
Xprays either depress or accelerate plant development;
(2) their actxgn depends : (a) on the dose, ' (b) ..on the species
~f
sensitivity, (o) on its internal and external
tste. (plants in normal gz?o`th are more sensitive than slowlyy
a~,
growing ones or quiescent Seeds; and the more favorable the ex-
ternal environment the greater will be the effect of the rays),
(d) on the place or 'area which is being irra.diated (3) since
the action, depends on various factors,' absolute doses for either
depression or acceleration of the rates of growth do not exist, `
there are, only relative doses; (Lip) the author believes that
the permanence of effect is very limited; (5) the initial ao-
23
e
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celeration of `de c1opment i sometimes followed by dopress1on '.
(6) with modexate 1ange doses (those which retard develcrnrent'.
but which do not kill the plant) microscopic examination does
not reveal any particuiarl r' obvious in juries of the cells but
only 'retardation of aotion,' a,nd when the doses are small, scce1-
eraton of action
In 1925, Gambarov, who tried to hedge his experiments
with the most ..thorough precautions, also came` to the conclusion
that irritating doses do not exist. From a huge number of
seeds of v'icia fabs equina he selected several hundred( which
were identical in `weight and outward
appearance. After soaking
the `seeds for` 36 houxs in 'water he transferred ham to a special
iy constructed box with sawdust. The bottom and two narrow
sides of this box were made from zinc, while the wick sides were
of glass. The length of the rootu of the irradiated and the conk
trot plants was meawured through the glass with a rnillmeter
rule for 12 days. In addition the appearance of side roots was
noted. Despite the fact that Oambarav used the socal1ed 'tir
ritatingsr doses, i. e? 1, 2, 3 1I., 5, and 10 (Highp
Eryther a 'Dose), not only did he fail to observe acceleration
of growth, but he actually obtained the contrary effect due
to the fact that the development of the plants was retarded
these dosages
by
In 1925 xven' e detailed and reliable book an the action
of X-rays wa published. In it ha presents the results of his
experiments with seeds of vicia f ba in dry and turgid conditions.
The irradiation was conducted with" ever tncreasing dosages at'
,.
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 lei appeared earlier than in controls, while larger doses re?
-:..
2
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)trays s nxprsssQd in D (HxghE-hemaDose) ranging as?
iows 1/2,o, 1I00' 1I109 1., 10, 18, and 22. When doses of
r
~. ,
1/2S0 to 1/2 HD (H1gh-E7` hens-D0e) were used on dry seed,
{
acceleration of germination was noted. On a, graph curve, it
K,
was possible to obsexsro first a gentle and then a sharp rive
r Erythema Doses) . Beginning. with 1/2 High
(from 1/20 to 1/2 ~lh ~
Erythema Dose the graph curve begins to decline. Consequently,
lven s experiments confirm that the irritating effect of X-rays
an the first stages of plant development fo11owu the Arndt
Sc or twrent days' later, however, depending on
~.ut~e : law. Ten ~'
the jntens~.tIy of growth and external conditions, the difference
between the ' rradiated plants anal controls tends to disappear.
~.
those plants which exhibited a retardation
On the other hand, ~
action of larger doses of rays, were unable
of growth, due
in the course of their development to catch up with the controls
the retarding effect of rays is retained.
then ben used turgid oeedsy the maximuXa rise of the
graph curve occurred at 1/100 ;D (3?gh.rythemaD0se)4 By
eriments with dry and turgid seeds lven demon
these parallel exp
strated the significance of the condition of the plant on its
saneitiv~ty to.~-;rays.
As for the action of ?ray s on developmental procssseS,
in i'ven's experiments) after a definite ins.
it become apparent
erencee between the irradiated and control
~erv'a~. of tints. ~if~',
plants ~h in the growth a' stems and in the time of app.
showed up ; ~ ,.
t laaf? with otinnula.txa doses the first
pe;arance of the firs
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tarded the appearance
that the retardatvnwas
whsh
leaveaa What is mogt jnteresting is
prapartional to the dosages.
dry. seedy received do
Final
es of 22 flED -g~,wErythema ?~ase
no lea VesE appeared at all, stere and roots were shorter, and
the latter shawed ' 'gments.t~:Oh? 1n add~.t~.on they lacked
bla,ak pa.
s Deed turgid deed, to ~.~~'a~'s, all of
~:de roots. ~ Jhe~ lv~n exp _
smaller doses than were required
these `phenornefla appeared with
for der ;seeds.
.
ncel ubl~.shed a series of works
Between . ~~ and 1,~~"~ r~ p
clarify the q~zesta.on of the existence
in ~~hich she attempted to a
of irritat? `ng dc~~es of Xera s for certain plants of the bean _
~.~ ~'
farm' especially and one escu~,en`ta In the first
; PhiSu s
i.fferences in the energy of gerna~
exriment (in 1920, the. d
tion
among jndivdual
roc:ds
(bath control and
were sa great that the author refused to
the X~ray~a De~ide Ancel repeated the experiments of Maldiney
and Thauvenin with the seeds that they had used (Gonvalvulus;and
'
In. the next
likewi8e obtained negative result.
~~,p and nt' 1920 she demonstrated the importance of ths expe.me (
role played by temperattire at which the plants develop after
being jradjated?
lOwex temperature 410] degrees ~- cents-`
A_
noreas$ 3 the Injuries due to strong doses of
grade, of course)
xr temperature (2Q-2S ~egreee) lowers them. In
..a,~a higher ~: ~ ..
he h?raied the queet~.an of the aig~her ; third w~ ~~~~b)
Dance of the time interval/ from the "moment of expo sure to the
.~.
a.
moment , of the appearance of in juXiee due to X rYe of the
e erimente performed by her indicate that the intensity of in-'
tiorLs of the time from the moment
juries due to x.raye are fund
26
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irradiated anew)
ascribe any effect to
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of irradiation; the longer the time interv 1, the more pro?
ounoed ie the appearance of injury In her fourth work (1926.)
n
Ansel examined in detail- the effects of irradiation on separate
parts of pants (in this case of lentils and beans) Tn, one
of the numerous (but' identical as far as reult were concerned)
experiments dry lentil seeds were exposed to doges of L.09 70',
lan9 and oltzknecht unit's. In the measurement of individual`
~.,,~ . ~
arts of the plants, on the )i'th day after irradiation' it be-
p
carne apparent that injure due to. X-rays were not identical for
various pas of he plarztso On the basis of this data we have
,
f.the following table in order to facilitate Compariann.
slvt up
Table 1
(based on :Ansel's data, 1927)
Dosages.
per centages' of , Injuries
knecht units Hain root stem Side roots
Li.O 15.79 9.10 23.8
70 31.71 21.9 31.57
3702L 32.81 ?
..:.loo,..
69.1k 564 21 83.69
lea
hjs table indioates1 that side roots are most suecep"
T 1' to injury, then comes the mein root' and' fin~1 y, the
table t ~"~
stem. Sjnil.aD results were obtained in studying the peroents.ges
;.:: . _.
of injures in beans. ina11y, in experiments devoted to deter-
-
of irritating doses of ;-rays for tdormant buds of ten-
urination
the author argues against the experiments of Weber,
who had estab1iehed the stimulation (by irradiation of such
27
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buds in lilacs. She subjected 16 sprouts of lentils (whose
stemE ranged from 8-10 oentimeter,s and whose roots were pro-
tected by plates of leaded rubber) to doses of 8, Eoltzknecht
units. On the fifth day after irradiation two dormant buds
(located on the sides of the main stem) began to develop. On
the tenth day after irradiation the stems had reached the
length of L.6 millimeters , and the buds supplied additional
stems with an average length of 16 millimeters; while on the
control plants the buds showed no development whatsoever. This
experiment provided the basis for.belieying;that stimulating
doses of XMrays exist. However, if the stem is cut off, after
it reaches 100 millimeters in height, the buds will produce
additional shoots. This fact caused Ancel to believe that in
the X-ray experiment it was the injury of the stem by the rays
and not the effect of irradiation, which caused the buds to
grow. In order to find out if. injury of the stem by
x
rays
will produce the Same effect as cuttings she irradiated the
stem with doses of 20 and LO Holtzknecht units; the dormant
buds began to develop as they had after' the "stem was removed.
In other: experiments when the whole plant (except for the roots)
was subjected to radiation, the stem ceased to develop, but the
buds remained dormant: apparently' the dose was so great that.
it injured the buds as well. From her experiments Anoel comes
to the conclusion that the development of buds in her experiments
with -r Y
8 -
28-
from injury to , the stem, and not from the stimulating, action of
from
X-rays on the, buds.
is the to the phenomenon of compensation resulting
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in 1925 A. v? and L? T? Koletrsov set up experiments wth
irradiation of pea and wheat seeds. They exposed dry and sproutw
ed 8eeds of Trite vu~ lgare lutescens to various doses of X'-rays,
but they did not note differeriees in the onset of various `phases.
In individual cases an increase of total weight was observed in
irradiated plants but it was not possible to arrive at any
kind of defnite'conclusions. Experiments with peas gave more
definite` results as far as acceleration of development was con-
cerned. For example, ` the plants bloomed 34. days earlier than
the controls. However, the total weights varied to such an e*-
tent in their cxperlments that the 'authors were unable to estab.,.
li~h any. rule to which irradiated peas conformed.`
In 1926 Bersa attempted to approach the `work of previous
investigators critically. For example, he considered that the
..
work of Schwarz, Czepa, and Schindler (mentioned previously)
suffered from many shortcomings, but that their merit lies in
the fact that they were the first to point out the many sources
of errors due to the inconstancy of vegetative (plant):..objects.
Zy his investigations Bersa was trying to show how careful one
has to be in ? the. interpretation of the results of experiments.
The first condition to be observed is to have a sufficiently
large number of test-objects. In sprouts of vicia faba the auth
or measured the lengths of the stems, from the ring where the
root begins to the apex of growth and the length of the root
from the ring to the tip. With doses of 0.05 Holtzkneoht units
an increase in length:of the, root of 2 percent was observed, but.
the author did not attach any significance to this since only 10
29
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roots were examined. The measurement of control and irradiated
root8 of 81napis albs did not reveal any significant differences
while a measurement of their hypocotyls showed an incontestable
acceleration. ` The following experiments gave less significant
or negative results. For instance, although an acceleration of
the growth of hypocotyls takes place, this growth depende not
on cell division but on, expansion of the cells, 1. e?' the cells
of thehypoaatyl enlarge greatly as an aftertath of taking water.
In general the cells of the hypacotyl are less sensitive than
the cells of the root. If the root :is seriously injured the
stoppage of access of water affects the hypoeotyl. Weak irrad-
atlons, however, do not affect the water supply of the stomp
substances from. the stem reach the abnormally slowly growing
root; this results in the overfeeding of the hypncptyl, which
accelerates its rate of growth at the expense of the `root. Con?'
sequently, the irradiation of sprouts by weak doses does not alp
ways' result in an acceleratign of growth, but it is always
in.
dicative of the disruption of the balance between the shoat
the root. On' the basis of ' the fact' that even in medical and
zoological experiments flawless results do not exist, Bersa
concludes that X--rays are not capable of producing real irri
taring effects.
Beginning in 1928 Johnson began' publishing a eeres of
works dealing with theirradiation of plants, but since her in'.
ve8tigationa are ;being continued up to the present, we feel that
it would be more appropriate to desoribo them later.
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In 1929 ;Doroehenko setup er1mente with three `planta
(Av by ant m 1 eta and winter e) ...The oonditi ors of
irradiation were Identical in all experiments, only the length
f ;exposures were varied (5, 10, and 2.0 minutes)'. The effects , .J
of Irradiation became apparent ink bant1na from the f1rst
stages of development, especially with the 20-minute: exposure
When the plants which had received this dose had already- given
complete shoots, the contra is showed only individual sprouts.
In the course of its development the stimulating effects became
more pronouncedtufts appeared two days earlier than in the
controls, stems elongated four days earlier, oars were formed
two weeks esr1ier, and since the planting took place very' late
in the season, the control plants never blossomed. The rrad-
fated plants, on the' other hand, which had gotten ahead of the
controis~bloomed lustily.
Doroshenko E' experiments demonstrated
the stimulating effect (of radiation) on the development 'of the
reproductive` as well as on the vegetative organs' of the plants.
However, when foroshenko attempted to increase the exposures
(to 20, 30, and 10 minutes), she found that 'exposures of L0 minim
utes fall beyond the optimum, although still within the threshold
of physiological stimulation. Still greater exposures (1' hour,
1 hour and l minutes, and 1 hour and 20 minutes), however, ale
ready had a depressing (retarding)' effect. The application of
a second exposure showed that it had a stronger stimulating ef~
feet on development than single doges. All of these favorable
data a to , ena zarnt
pp~" A~ by ina._ As for Toulouse millet, the
author did not su~eed in finding a stimulating dose for it.
The same was true for flax (of both the curly and the long-fiber
31..
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var? ,eties). On the other "hand, winter xye, as a result of. iz
r d:Ition9 not only developed vigorously but changed its form
aa: well.
Th
on
irradiation of potato tubers
first 'experiments
were performed by
ments
Sprague and Lenz in 1929.
For their experiw
they selected the tubers of two family varieties: "Iris/
Cobbler'T and "Green Mountain" . Half of the tubers were irradiat?
ed; Mtha other half were left for control The tubers were sub-
jeered to radiation just after sprouts began to appear on them.
One part (experiment 1) of them was subjected to doses of " HED
(High"Erythema.Dose); the other part (experiment 2) waa subjeot
ed to 1HED . The first leaves in experiment had an abnormal
appearance; aventuallya` however, only normal leaves developed,
and they 'remained green for several days longer than those` of
the control plints As for the yield of the potatoes in number
of tubers, it was 8J.
j percent of the controls for experiment 1,
and lOi~.7 percent for experiment 2. But in bath irradiated
groups, the weight of the average tuber was above average (of
the control). So we can see that even" heairradiation, " which
injured. the leaves and decreased the number of tubers did not
diminish the total Crap.
Patten and 'Wigoder" jn their ;experimente in 1929 once
more returned t4 the "olassical objects of roentgenology: the
seeds of beansmustard, and barley. In their opinion the o?'a.
facts of X-raye on growth and development may be studied most
easily On plants where 'cell division takee place" eo quickly that
the progress of ,sprouts rn~y be observed daily. Giving" then seed,
32
I
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a dace of 3 }j figh-Erythema-ose) they observed various
react1on which were dependent on the species of plants (since
other factors were constarnt). The roost sans t ve in their
experiments, were the seeds of large beans, especially if they
were, f.rrad1ated from L$ to 72. hours after growth had begun. d
mustard sprouts the srnallest doses (about 1/20 High..Eryythema.. ose
caused the most rapid growth, while. irradiated barley lagged be~
hind the controls..
The value of Chekhov's contribution (l930~i93l) ijes in
the fact that in radiating ` seeds of 'dry and sprouting plants of
various sorts` (barley, rye, lentils and oats) he paid particular
care to the condition of the plants at the moment of irrada Lion'
a condition which varies with different tames of the year, and
even of the month.` He demonstrated that weak doses of rays had
a stimulating, affect an both germination and on further develop.
rnent~ In experiments with dry seeds when strong doses were used,
which have a depressing effect, the 'graph curve of their growth
does' not fall evenly; these doses do not depress germination and
the force of sprouting, but after sprouting a certain percentage
of the developing 'plants dies. As doses are increased, the nu*i
ber of plants which die increases alao and the time. of onset of
death becomes shortened, 'cry strong doses have the effect that
c~~a ar f &J pE4i T
only 1/10; as many irradiated plants come up ~antrnl
l ;1:9 Cattel demonstrated that the growing parts of
wheat , sprouts are relatively sensitive to X.rays. Ris e~perimente
are remarkable "for' the ;fact' that he performed them on 200,000
samples and kept repeating them for a period of three years. f1x~r
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seeds were thoroughly washed in distilled water: then they were
soaked in it fad 3 hours
chambers were then placed
26 degrees (centigrade.
and planed in humid chambers; the
in thermo`stat3 at a temperature of
After 214 . hours the coleoptyle, leaf
and two side roots were 2 millimeters long the primary root
was m llizneterg in length. At this time sprouts were seleot'
ed for irradiation (they had to be of equal size and compietely
healthy). After irradiation the sprouts were once more paced
in humid chambers which were placed in thermostats for L8
hours, after which the sprouts of bath the control and jrradi.
a,ted plants were measured. These measurements
showed that each
of the four growing parts (coleoptyle, leaflets,' sides and pr~?
mary roots) was effected to a different degree. These changes
could be observed after only a few hours. Equal. doses of radia
Lion produced equal changes. Sprouts which had received large
doses showed various irregularities in their development; roots
thickened, thickenings appeared in cells, etc.
In 1933 thud and Mitchel 'set up their experiments with
extreme care both biologically and physioal1yA As far as the
physical aspect of the experiment is concerned, the authors
suhjeeted seeds 'to irradiations which were both quantitatively
and qualitatively identical (100 kilovolts, 5 milliamperee, at
a distance of 30 centimeters). 'they used an aluminuxn filter
1 millimeter thick. As for the biological aspect of the exper~
iment, the authors used for their experiments : seeds of corn, ; oats,`
and surif lowere, peed them in humidifying chambers' an a layer
of cotton, eoaked in distilled water, and kept it at 22 degrees
centigrade). After 2L hoi the seeds of all `three plants began
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to sprouts Twenty seeds (and aometimes ore) were seIteoted
from the rnas ~ whenever possible on an identical level of dev..
eloprnent and i 'e dl .ded into two parts d One part was retained
for: control and the other was placed on fresh damp cotton R.nd
Irradiated. The optimum. irradiation for sunf lowers turned out
to be 3 rn1nute s f i.. e.
11hr ?enen units) for corn' and car
taro varieties of wheat, the optimum d?Be was 2 minuteE. For
other varieties (of wheat)
sores of 34, LS, or 6o seconda
were sufficient. As soon a$ she rradiatior s were concluded:
the p1antr (e,nd controls) were planted in soil, 'hand, or can, cotes
ton .in the humi.difying chamber. Fxperin'iente demonstrated that
a. varioty of hard wheat (s'Minhaxdin') was less sensitive to X-rays
than "Truntai1 which in turn was -lose , en itive than a variety
known as "Marquis" (Marklz). From 'thisthe authors concluded
that just as there exist varieties which are more resistant to
cold, so there are those which` are more resistant to radiation.
The ? most interesting data were obtained with corn, a variety
called "Nadiion Jellow pent"
Yellow Tooth"
?]
ranslator's note r ` tMadi.r~on
Turgid kerns 1 of corn were di~rided into
three parts; the first was irradiated with an aluminum filter,
the second without one,. snd. the third part was used as a cans
trol. Determinations of raw and dry weight of the coleoptyles
showed that Irradiated seeds are. from 5 to 26 percent heavier
in
the `raw and from 3 to 16
percent heavier
in a desiccated con-
di Lion. It is possible to ascribe this ?ncre'ase in the wei,j'it
` ... .,
ly
f the coleaptylea~
a ftstexr ttilizat.on of the endorperm re-~
serves of the'. irradiated cells. Several weeks after. ,irradiation
p1antwhich had received 13 minute `dasea had thicker stems
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control$ or those which hay'. received?minut,e doses.
than the
'the weight of the .green part of the pant re;ater 1e.
irradiated plants.
hen irradiated) grew to above average height. A.
Oats (when
compar o1s and irradiated $eede of sunflowers gave
~.~on of - controls -
a food graph curve for their development. A group of plants
...
which had been irradiated for three minutes was the first to
experiments indicate that when exper~ment$ are
blossor~. ! These carefully het ups and the proper dosageS are used favorable
:.: ,.:.
nfox~tunatelT the authors did not
results can be obtained. U
follow' through they' experiments until the cxopW could be ha.r'r
vested a $ unf"a they d~.d not repeat their e~per~.rr~ente
~?tunately,
even one more time.
riments of Benedict and artefl are ?nteIS3tifl1
The expo
_ e er~i'ie11t~ rays' w~lich were praCti~
bedause they used a. n their. ~
~va11y monochromatic. They Placed :rains of wheat . in 8pecial
T
a. manner that all of the buds poiflt~
dishes Econtainorsi in such
ed. :l.n the d "~Na,) ` tube, at a dietance of 1 centi-
meter of the ~x~'
meter from the a~ure, in each a manner that it was pos~ible
to insert a n (,f3.ltvr) between the ~eed~ and the aper~
lead ~aree
the : seedy were treated with fungicide
ture, After' irradiation
and aUrnrJ~d to spraut$ then their diaetatic activity, their
their respiration, and their water
reduced), sugar content'
contente,were detexdned proute, which had been exposed iO
radiation fnr, ~ secoride, showed an increase in their-di~.statiQ
content. If the dosage waS increased,
s
activity and their ubar
however, it wa~ pasEible to observe a decease in one or the other
3
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e?ernente? The ;amount of substance used for respiration
of these
~ corepani on with the coi troll, but neither
~. _
didnot ?ncrease b,-~
d ?nlsh with large doees of radiation.' These results
~.d = it dimx
the Gpflditions of thie experiment irradiated
:~ndicate that under
seedu cannot convert their reserve starches into sugars a eae~.w
s and also they cannot use their sugar fog
:1y ell,
, ~,~ n~rr~al c
growth a rapidly as the control CCIlse
The effects of 'radiation on wed of citrus fru,t were
obse red by Ha d fS and Moore
n19
They observed a ouriau$
flaraenon of premature flower3:ng of 'two grapefruit p ants grown
phe
from had been expo aed to doses of 300 and 1300?oen y-
seeds which h
an. aib~:rao .:and' ;con$oquen~ly,
r gen un?ta One of the plants was
was the other developed more or lees normally, de.
:.: s~~vd.asw ~ '
pit! .size, and after , a few months exhibited a second
~. its ~r~^
b1oeso n ng.
The expper m i-1ts of Long and Karsten, In 1936, were set
up with soya under r field condit1on5. The irradiated and control
;
seeds
formeds he 'plants were cut off, a:t ground level, and weighed.
were f'
The weight of the green part of the plant was greater in the ir-
r~d irice the experiment `was conducted with large radiated p~.ents .
o
had beet aUowed . to grow under normal conditions,
numbers which
J
arid s.~np$ the average increase of weight of the irradiated plarxts
.
as cornpared. with th controls, exceeded the standard deviations three
tim over, the authors came to the conclusion that this indicates;
es,...;
true jmulatiaction of the rays, although this should not
ng .
S
be,taken as abdc1ute' proof of it.
37
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produced, all told, l2,7 l plants.. When the first Beane
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1935 rolav used for hi8 investigaticna three'
which had been used
and flax
at
,
si whe
, plnts~ na~+e]~y, . ~a
for radiation intents many other authors. The irradi"
. ~~per by atior)of whet grains not only accelerated development, but
re su1ted in, a 60 percent inorease' in iielddf as compared with
COfltrOl. The eriments in 1936 gave & hat lower results.
a stimulating effect on flax only in the first
X-ra,fs have
stages opmeet after' that a depressing: effect sets in.
of c~eve ~
soya seed, produced negative results, in cones
~.rrad~.~.t~,on
The or a
tradiction to the experiments of long $ Karsten.
In l93I ?u5hryak?va and Vasievskiy set up experiments
with sedge (Cyperus e?culentus) , soya, blue lupine, tomatoes,
..-.-.----.---
on;5. The sedge was treated with X-rags for the purpose
and m~1
of shortening q1ten~.ye`'n~ .d. `j4 gi' rowin}y scam and causing it to multiply vegiw
4 7
tatively? The tubers were irradiated in dry and soaked condi
tions for s',S, and 20 minute periods. At first the irradi-.
tubers lagged behind in their development, but eventually
sated
those which had received l5-minute irradiations moved ahead.
;
Two months after exposure the. difference between the controls
,...,.. ;
and the irradiated plants was striking. The authors pointed
out the diff' erence produced by irradiation which were due to
,
the materials and the experiment. In 1933,
the conditiran of
193, and 19.35 experiments were conducted in the field, and al?
difference in the time of development, the
though there. was no
been irradiated for 35' minutes stood out sharp
plants which had.
height and vigor of development. They, had
:Iy because of their -
. ,
a high xoductivity and a; shorter growing season.
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In eperiments with soya a' shortening of the period of
rp in cor~~1f)a,rison with controls) war observed. The s an?
~,n~.~i~r s ~ x
th rved with blue lupine whoa th,e rracli.ted plants
earlier than the contxbls. The expe4re2.t with
ripe~ l2 ;days
m~l.ons wa nOt rt~ncludc~d but it showed that plants which had
o~
received 2aroentgen 0 unit ripened earlier than the controls.
_
an tornatoe ripened more intensively than the
irradiated Mexic
control ;a and the numb:r of fruit and their weight was greater.
The experiments of these authors should be considered as
prel?mdnary, despite the fact that the ex rimente with sedge
~..
were repeated over a eriod of several years. The small number
~'
d in these experiXnents and the lack of concentraM
of samples use
t ion data were scattered, too widely) of the data
~.On ,~.? e? 9 ,
prevent us from considerif : these experiments completely . eon
olw d ve.
attn. After irradiation the Feeds were planted in growing
c
dishes hind their development was minutely observed. The dis~
cxepaney' between the develapahsnt of the vegetative part of
and their productivity Tsuryup ' s experiments
the plane
different the effects of X-ray can be in their
indicates hod
app o different plants. In experiments with wheat it
~.cation t
riteufl' dui variety 'melanopus, and also those of oats, and
W2.a 'o1asdred. ;that; ,X
~uryupa irradiated seeds of laard wheat
than on ;ragetative oils. On
mend of the reproductive organs
the other hand when oats were " used, the effects were ;just the
1thou h X-rays do . not increase the gex'nina'tion of
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wheat, they increase their sprnutimg power. As far as yield
is concerned, poeitivo effects were obtained in some cases,
-
exceeding the controls by 60 percent. In addition, the grow
i season was shortened $0 that the irradiated plants ripened
In the case of flax the author observed the
ten dye earlier.
ization of the initial stagos'of development, which were
activ
followed by depression, while irradiation of soya seeds caused
development to be retarded from the very' beginning.
A whole series of works were perforrxed by Jehnsoh
(1926, 1928, two` in 1931, 1933, and twp in 1936) ? In fact,
her iens are being carried on right up to the Present.
,. ~.n~rost~,gat
thewide variety of objects, the thoroughness of details, and
.
the multiplicity of aspects which this author investigated
.
place her in the first ranks of roentgen-biolcgists? In her
vary first works devoted to research on sunflowers (1926 and
198 he compared the effects of dceas from 5 to 10 D
:)' ~
(ma~se1 ' on dry and soaked seed. She studied
~,gh?~ryt~ie
the relate.onshxp between' mounts of exposure and growth; she
...
investigated the effects produced by radiation on catalysis,
,
oxidation, and an respiration. Not satisfied with these phy'si-
s she turned her attention to phenologi??
aZagcal investigatian,
cal abservati0n$, which gave 'moat significant results with doses
.
ema1*lQee)? Her experiments showed that
of 10 D (dig
germinatian of seeds ie not accelerated by irradiation?while
sprout ight~ retarded. Sawed seeds which were ire
sprouting ; is s l
medium doses came 'to a complete halt in their
radiated with
in the first three weeks; after that, there was a
development
temporary acceleration. ` The depression of growth during the first
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three weeks (a8 a re ult of irradiations js approx ateiy pro-
portional to the dosages.. Plants which` had 'been' soaked p$
vious to irradiation did not attain the size of the 'control
plants, but they `bloomed earlier. ` In her next work (1931) the
author presents the reeulte obtained from irradiation of two
thistles, r everal Eolanaceae and Viaia faba. ln` order to give
stimulation a chance to show its effects, Johnson permitted her
plants to grow for 25 days (since previous expermente demen-
strated that the effects of radiation tend to vanish three weeks
after irradiation unless a aecond dose is administered). Comp
iIfl
partitive examination of fresh and dried weights of the irradiator
od and control plants revealed that only one plant, "eunberry")
had increased its weight due to stimulation `with weak doses of
X'ray. Since one weak dose, in Johnson's experiments, did not
produce stimulation, she gave a second dose after a certain in..
tervaLIt is interesting that in these experiments . Johnson
ueed the same dose for Vicia faba as Patten and Wigoder, who
pointed to the increase in size of the plant as a result of irw
d
:Des its
radiation. p condition '
the fact that identical" s were main..
tamed for irradiation (seeds were used after being allowed to
germinate for Ij0 hours in water) , Johnson failed to observe any
stimulation in. the roots ?tubers. In irradiation of tulip
bulbs it was obserred that the length of leaves was increased,
but the formation of flowers was "unaffected. In this work the
author presents her observations concerning the increase of
growth of lante' b' , irradiation of ` bulbe and tubers of curtain
plants. She pointy out that when these experiments were set up
under, more or less, working' obnditions, they did not ex
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hibt the effects of stimulation. The irradiation of wild pota-
toes from Colorado gave favorable results in the ;preliminary ex-
periments. However, this experiment wad given up when the atth
or dieoovered that upon repetition with a large number of samples,
this reaction could not be obtained.
Finally, Johnson wrote a large article in nuggarea book
("The Hinlogical Effect of Radiation") in which she sunrn rizee
the results obtained by her, and others, and comes to the conclu&
si.on that strong and tedium doses of rays have a harmful effect
. on the development of plants. As for weak doses, she considers `
that in those cases when they produced results, the authors did
riot give sufficient consideration to the individual variability
of plants
In x.937, Zankevich and Brunet were studying the effects.
ofau ur.uai
'J .1
Individual' develo menu of tobacco (icoti na
rustiea) poppies9 f lax, and rhubax?b? Increasing doses of radi-
'ati'on were used (250,` SOO 750, 1500, 3000, 6000, and 10,000
roentgen units). Doses of 500 and 7 0 roentgen units had a
stimulating effect on tobacco and flax. Irn tobacco the increase
In sire .was of transitory nature, but in flax a certain amount
of `stimulation was retained until the end of the growing' season.
Medium doses of (rays resulted in a retardation of growth, which
way (in degree) directly proportional to the dosage.; Large
daces. had a lethal effect. The authors make an interesting ob-
servation to the effect that tie latent period" of plants i.s
i a a . ' to l0 da
off' dif'ferent, duration: in poppies arid flax it l st 7 ye
while in tobacco aril rhubarb it is off' short duration or ntire3y
absent.
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Zaurov in 1927 [sic, ,1937-?] used various doses of
wrays on the seeds of Indian hemp. He observed an insignifi-
cant increase in the length of stems with 'a dose of ~OO oent-
gon urdts in the first measurement (experiment) and a similar
increase with a dose of 2000 roentgen units in a second experi-
rent. Depression set in starting with doses of LOOO roentgen
units. Sn&11 doses of X=rays (12541000 roentgen units) produce
an increase in bushiness; larger doses'. depress. bushiness.
In 1938 Bres1avets and Sinitskaya, who were interested
in changing of forms of decorative plants, set out to study the
effects of X-'rays on these plants. They irradiated dry seeds
despite indications from literature that irradiation of sprout-
in seeds is more effective. This was done for a definite pnr-
I , poses "Tf the effect 0f `some dose of X-rays an dry seeds were
to result in the change of shape or coloring of the leavers
of the flowers, ' or of both, it would then be ' po ei ble to send
the d irradiated seeds to various horticultural state fares
(sov~hoz), a.nd to amateurs, while sprouted seeds could only be
used by us." Summarizing the results of action of X~rays on
five kinds of decorative' plants the authors come to the conclu-
:Mora that note a single case was it possible to obtain the
correct biol.gical graPh-cu for germination and sprouting
power of the plants. But it is interesting to note that th$ ex-
periments brought, out a variable sensitivity to x-rays of these
plants. This result, however, should have been expected since
U"
the fire . plants belong to completely unrelated families. The
rnoet sensitive (to xwrays) turned out to be sweet pea, which
could have been foreseen on the basis of available data, which
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point out that the bean group is particularr sensitive to
X..rays. stars also proved to be quite sensitive in this re-
. A -
arnatiane and gil1yCZowexs (stock) were so in.sensi
post, while c
tive that doses of 8000 and 16,000 roentgen units have (within
limits) a tinxutin effect on them. Tobacco occupied an interN
~.
mediate poation as far as radiasefsitiv.ty is concerned.
If w carefully scrutinize the vo1'uriinoUs data obtained
e
s authors in their experints on the physiological
by the va,r, Ou
a.ct the ray's, we will see that they lead to completely[
~.on of ,
d oncluaions~ Some authors indicate the existence odiverse c ;
doses ? others deny their existence. This would be
stimulating ~
understandable if it were a questiom of differnt varieties,
since ume ous investigators have established that plants are
n
variable in their sensitivity to X~rays. However, aontradiotDry
results are reported even when the same species of plant is be-
ted. As has already been pointed out, the seeds of
ing nnvesta.ga
a a favorite object in X-ray ; research. It l s take
!icia faha
a look at how various authors evaluate. the results of irradiation
f this plant. Perthes found
observed a temporary eessati0n of growth, Schwarz. obtained a
typical biological curve of development,
i.
e. he ;, ound that
weak doses of X..rays stimulates plant 'development., that stronger
ones retard it, . and that still etronger ones stop development al
.~,a~d
tog?ther. In the expe?iments of AUaanfl Rochlin, and leiohge-
:.
acceleration of development Baas `abaerved; this
w~.cht a ?~enapa rah' /i ' ~:~
was'Qllowed b~ an
failed. to
x e
ra
depressing e1'fect, Koernicke
"the ,eo$rais. Gar1arov
dose far viola f ba ec inn*, high
had been `established mn lverc~ s
xperirneuts?. lJnd, '.rial.ly
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jahnson repeating the experifs nts of Patten and Wigader, was
unable to the stimulating effect of X-rays which
confirrf~
Patten and Wigodex h a& ob erve.d,despite the most careful.
repro uuction of their experiment.
Besides, as Kochtin and aleichgewicht pointed out in
:Vicia f: ie na t an appropriate object for the study of
the stimulating effects of ~-raYs because its threshold of
timuiatien I
too low. it is so `radiosensttjVe that the weak
of radiation cause a retardation of growth a thin
est doses
complicate experiments.
he questian comes up of why' experiments
Naturally ~ dealing with the radiation of one / and the same plant pro.
. ~.~
$ultS. This is not easy to anger. But
dues such different re
if we carefu all o the experiments we will gee that
~.~,y' analyze ,
in the rain the disdrepancie6 can be explained by an insufficient-?
of ddsee (this applies particulax'] to
lY aacuratemea,surement ,
that period when do es were measured in ex h$tlas) The sensi
tiv -
-raY varies cousider~abl" m and to equate
~.`~Y o f ; people ~,
an e dose to w 640 roentgen units as some authors do' is
r"y'tnezs~a c
comp1et?i inaccurate. In general, it is only in the last few
years that we have learned to measure XMray dosages with a
~.
great degree of accuraaY Be~idee this chief shotco.ng' we
into consideration. In experiments with
harre~ :t~- take others
i-. f' whieh'id : the subject of inve~tigatione of many' auth-
ord, it main'; , what sane// numbers of sesde were used for ir"
'radiation. a,tian. The reIeon for this' hioE in the large size of the
s a eub8tantial barrier for performing the
seede,whi.ch crinetitute
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exi?rimnt within the restrictions imposed by the Coolidge tube,
which has l.rrijtod area of action. [ The desire fr great accur
acy forces the investigators to irradiate the seeds eimu1taneoue~
ly'with several 'dgsage, removing frc*r the field of action of the
rays Petri dishee (which contain eeeda) at various interval, of
time, in order to sc+ure the most uniform voltage and amperage.
It is small wonder then that some authors limited their exper
ments to 1? seeds of this plant.
Deapite all atteraptE t
match the seeds in size and weight, it was `still, necessary to
take into eonsiderat1on `the
individual variations' which could
only have been overcome by uEing large numbers of plants. Con
Sequent ', to critici n of. those authors (Schwarz?'
et. aL) who
point out that individual variability of the suds completely
overahadows` the'effecte of the rays, e entirely justifiable.
On expori.ents/ Where the numbers of plants used ie sufficient
ly great to permit biometrie analysis' can establ1 h whether
oh~nge.n size and development result from irradiation. but
i.t is most interesting to note that the
same author. (n.
ry
Cloned above) who, categozized the expex~nens of their preda-
ceeor` ae nconclusve on the basis of insufficient numbers of
test objects' used only 20 seedy in their experiments.
A third , source of errors in the experiments, is found
in 'the small va 1.ation of dosages. Noet authors are content to
vary the time of exposure or the distance from the tube' both'
Of 44Pk are : s;l:ected at ;random,. It s v
authors 'of this'
Y~
q of `
f, experiment, who obtain changes from ran-
doi~ , da se s of ~~raye, ' t
dump to conclusions conci~rning thhe e.a~
tense pr 4$nce of stimuating doiee.'
tab
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perirnents which undertake the investigation of the
effects of
XWrays on plants have s14'l1 another shortcoming.
most of them are incomplete The nmjority of rove; tigatar
either nv only the first phasee of development of the irrad1"
-.' `
; pants and corpare them with the c0ntrv.e, or eke cuts.
atedP
them down as soon as the lrs~ fruit appears and judge) the e1
fact of X. a 's cn this or that plant on the basis of the weight
~'
^?' the fresh (green) or dried ma$e s ignoring the fact that
u,~, the
Doroshenko in 1929) and uryapa (in 193h) accurate
pointed
out that 'rag can affect reproductive organa9 as well as vege
tative ones, and that their effect is e1en mere PronPUTiced on
the 'formers
However, the mo st aerious defect of experiments with irp
radiation of plants , as of all experiments with etimulaton' is
the preconceived notion in the mind of thn anvestigatnr?
if
the latter, on the basis of theoretical constderatiOnsg refuses
to acknawed.ge the action of this physical factor an the devoLop
, ~. ,
olanta then upon getting positive ;results he ascribes
ment o
ahem to ehancev That is what Seide doe$ in his experiments (1929
, .
with the ova of Asca .s mo ra] 1 . Observing in one exper~
a?ment an increase of 26 percent (as compared to the control) In
dividing irradiated ova, and failing to obtain
eim?1ar results
upon repetition of the experiment: he ca11fl]3r? proceeded to chaff
off the first experiment as "due to chance" , instead of determin'
the second e per ent didn't succeed. Johnson acted , in
ing why
the ; a111e way. Observing the increase in weight of the fresh
(green) mass of the plant a a result , of 'irradiation of "sung
berryr seeds, she assumed that it was due to chance and deleted
'.7
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it from her count. ?On the other hand, those investigators who
are px'ejud ced in favor of stimulation of X-rays axe even more
guilty, for they sometimes contrive their expedmentq is~ ;uch
Nevertheless, the great bud of the experiments and.
oberv~ations of plants which had been. exposed to the action
. of x-rays constitutes a unique body of biological data, which
t,rs most interesting, both in their theoretical and their prao
tical aspects.
I STIQATIONS' PEPFORNED IN T.
ELECTROBIOT.OGICAL LASTORR
We were guided by the above eonsideratic'ns when we dew
cideo by a series of systematic experiments on the effects of
radiation on plants of various species, including' agriculturally
significant ones, to answer the question posed by our predecea~
sort as to whether irritating, and consequently stimulating,
doses of Xwrays'exist. But we expanded the question of stimu-
lation to include the question of yield, and consequently not
only brought the plants up to the fruit~bearing stage of their
developnt, but we analyzed the yield a well.
The experiments were first conducted in a laboratory for
the biontizatian of seeds which in 1935 became part of the Allp
Union Institute on Fertilizers, Soils, and Agricultural Engineer-
ing, and later of the Al1'Union Institute: for Electrification of
Agriculture, and,upon liquidation of the latter, of the
Timiryazev Agricultural Academy.
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Our experirrente began in 1931,, and our first subject
This was a very hapl3 choice for, S$ we shall see
later, rye turned out to be very sensjt?ve to X.rays.
e
Bye. In order to find out the regulartty of the pr~
cess we ey~etemtically increased dosages for each aucceed:ing
a
expe owing order r 2 O, .X00., 7 O i 1000, 2000,
r~,m~nt in the following
roentgen Units' Not content with varying the
~,aao, and . Joao a.
we also hied qualtitative variations in
?rays quantitativelym
we used hard and soft rays. Seeds which
our experiments, i. e.,
had b een soaked for twelve hours as well as sprouted ones were
irradiated,. We present the findings of these experirnente eepar'
ately.
sprouts (investigatione of Breslavets, Afanaeyeva,
and Medvedy'evs.) For irradiation we used freshly sprouted
but since we hadn't selected the sprouts fOr ir
seeds of 'es
nt care, some of them had attained )J2
radiation -
with sufficient
believe that this difference in size is
centimeter in sire. we
for the uneven manner in which the trradi~
large responsible
ated sprouts camp upmedatey after irradiation the sprouts
..
were planted in flower pots,, several per pat. The plants de"
for 27 dey A otter that they were planted., one
vel.op?d in them
at a tame, in a heavi1y rertilized plot of grourd. then the
,
p1ants were still in the pots, before planting out, they were
piiQtographed. On thi photograph graph differences in their develop"
on the dosges they had received) were clearly
meet ; (dapes'ding
apparent. Doe of 2O roentgen units already have a depreiBJ.ng
affect an the d.evelapmeat of x'ye; doses of. 5C0roentgen unite
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produce approximately the same effect. But when the dose is in.
creased to 7O poentgen unitsg part of the plant i develop poor-
ly while the remainder die.. A doee of. 1000 roentgen un?ts has
such a, depressing effect on the ?plantaa that they barely reach a
height of 2w3 centimeters. With doses of 2000 roentgen un1ta
only a few sprouts survived; with a dose of li.000 only one sur-
vJ.veUj W1l.L..e
al, not a single plant uurviving.
However we did riot limit ourselves to a ingie opti-
cal evaluation. On the 27th day after irradiation, 1. e. 9 just
before planting out into the field, we carefully measured the
plants, the controls and those which had received various dos-
ages of the ray . The results of these measure rents can be seen
in Table 2 a
Table 2
(based. an the data of Preslavets9 Afanasyeva
and edvedyeva,i932)
Dace Average Height
in Centimeters
Average Bushiness
(Number of Stems
per Plant)
Dose Average Height
in Centimeters
Average Bushiness
(Nurxber` of Stems
per Plant)
Control
x , 50
2,00
1000 h. r.
3.36
1.10
250 h. r.
1i. T?
2.0
1000 s. r.
6.90
1.60
(hard rays
250 e. rt
13 50.
2.i.0
2000 h. r.
1.53
1.00
(soft rage
5o0 h. r.
1,70
2000 's. r.
1.2
1.00
500 e, r
8.S0
?.02
hp r.
1.la6
1,00
750 h. r.
14.(50
1.30.
L000 e. r
1.27
1.00
7508 XI.
l4?
1.20.
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dose of 8000 roentgen unite was completely leth~
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tion of _this table will reveal that a
u~. eXarn~.na
f
care
sufficient to depress
2O roentgen unitS is already
~
dose Of
average height of the plants, but the average bush
(umber of stems per plant) is: still somewhat higher than in
r la
n
s, the 'differences between the effec
the controls. Beeide to of
to show up hares the raft rays have
..soft and hardray8 begin
the /amts. 'phis difference also
a leas drart~.c effect can p
. _ .~ ..
in other doses with the excePtt0f of the 7O roe ntw
shawl up
and
en unit dvses where the, left rays produce smaller sire
the hard rays. hut' in generals
leaser bushiness than
of soft and hard rays in our exper
difference of the effects:
that in future presentation of data, we
~.ments was sq slight will deal only' With hard rays' since they were used in the
majority o experiments.
observations conducted by us in the
The phenol Ogical
spring and summer of 1932 have demnnstrated that the average
sear flowered, and ripened at about
irradiated plant` grew it ~
cOS ''
the same time . the controls oaev er, individual variation8
.
These individual differences are
for some plants were 3?L days
irradiated plans, as we shall see
frequently enaountersd among
when we turn our attention to the study of other aspects.
We should note that only during :flowering did some of
attention by their unusual level-
the irradiated plants attract
apment? Such plants coind be encountered only with dosages of
,
nits Start~.ng with doses of 2004 roent~
up to l0(c roentgen u an
to
degenerate fOX with smaller ears beg
gen un~.ts c~vious]~r
e was ripe, it was collected by individual
show.. up? when the. ry'
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plants and the weight and contente were subjected to analydie,
The number of ears was counted, the total weight of seeds was
obtainedg the total number of graine, and also the number of
health and injured seed$? These resulte are pre8ented in
Table 3.
Table 3
(ba$od on the data of Breslavets,, Afanasyeva,
and edvedyeva, 1932)
Dose Number of Number of Seeds Weight of
ears Normal Underdeveloped seeds
in rs
heeds and diseased Total
.111.7 3S J.6.7 .5.36
Control 3.7
250
10.0
2S
85
310
12.
500
7.0
122
9
181
6.911
7o
82
37
119
4.57
1000
3 ? ~
80.
13
93
3.18
All these data refer to a single plant In analyzing
this table it turns out that a dose of 250 roentgen units
more than doublee the number of ears, and the number and
weight of seeds. A dose of x,00 roentgen units has a much
weak r effect, although the number of ears, and the weight
and number of seeds still show a significant gain over the
controls. A dose of 750
roentgen unite already depresses the
development of ears and seeds, while a dose of 1000 roentgen
unite gives a still more pronounced depressing effect. plants
which had received doses of 2000, and'1~000 roentgen units per-'
fished early in the spring.
2.
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If we were to project line graphs on the bads of these
data., we would see first a sharp rise of all quantitative fac-
tors and then their gradual falling offs This points up our
rmi.stake in setting up the experiment, since for irradiation of
sprouts we took too large an initial dose
Apparently we A hould have begun with smaller doses, per..
h~pJs sb sutalL as
50 roentgen units.. it is possible that air11er
doses than the or we used would have also produced increases
in yield and we would have obtained a less lopsided curve (i? e~,
a more normal, curve).
Our experiments with the irradiation of rya sprouts inu
dicate how wrong it would have been for us to limit our obser~
va.tions to the initial stages of development (as the majority
of authors did with various plants) or take them only as fax as
ear formation (as Long and Karsten did) Comparing the two
tables we can see how much the number of stems per plant has
increased with benevolent dosages with the course of developments
With a dose of 250 roentgen units the number of stems per plant
is not much' greater than that of the controls, while the number
of ears at harvest time is almost triple that of the controls,
and this does net include the number of underdeveloped (those
which had failed to catch up) ears of the same stems.
If we had not carried-our` experiments through harvest
time, we would have had to agree with those authors who paint
to the small irritating effect of X~rays, or to the total
ihee~,
absence oirritating effect.
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received
The seeds (investigations of Dreslavete and Afanasyeva
the carne doses of X'"rays as the sprouts/ but
these doses produced a different effect on the development
of the plants. Just as had been done with the irradiated sprouts,
the soaked irradiated seeds were planted in flower
..pots and on
27th day transferred to field plots. When we set
out the flower
pots according to dosages, we saw a very vivid example o
of the.
curve of the Arndt-Schultze law. (Fib, 2. Differences in
development of rye depending on the dosages of ra y e they had.
received.)
In order to check by means of figures our direct obserM.
vation9 we cured all of the plants on the 27th day after
planting. As may be seen from Table , the numerical data
coincide 4th the visual evaluation.
Table 14
(based on data of Dreslavets and Afans eva .i932
Y , )
Doee Average thgh Average Dushinepe. Dose Average Length Average Bushinees
r~ in Centimeters (:Number of Stoma r. in Con .mete
~ z s thrniber of Stoma
per Plant) per Plant)
Control 12.5 2.0 1000 2.0
~ 3.2
250 20.0 2.$ 2000.
17a9 2.0
500. 23 g 3, 14000
177 2.6
750 22..6 3..1 8000 10.8 1.6
w 514 _
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we carefullY exam1fe our irradiated plants and compare
If
thorn o is when both are in the process of growing
with the contr
some interesting details can be brought to
additional stem$g
that attracts attention is the increased
light. The. first thing
their breadth and coloring are directly' re-
sj~a of the 1eav3s;
lated . With doseE of 2S0 roentgen unit6 It wad
to the dosages
not_pousible to find any obvious differences with the controls'
but with doses of COQ roentgen units the width of the leaves and
their darker (more hse) green oolc~r~.; were mediate]
.. ~.nte
parent? posed of 7~0 roentgen u .ta intensify these character-
istics still more. The clim Ie reached with doses of `1,000
width of. the leaves is nearly double
roentgen units when the
that of the controls eginning with 2004 roentgen unite the
. ~ o
leaves first approaches that, the contrals, and then
sire of the
becomes censiderab3-Y narrower. It was likewise obvious at a
glance that plants which had received w-beneva1entn` doses had
controls. we also measured 50 roots from
larger roots than the
an ooularmicrometria ruler and then trans-
each dole bar. means of
ons these measurements were perfod
po emd the results intd mica
by ments were alwayB made from the same spot
~,payev~.) ? Leasure
on the root, nalnel here the root begins to dfferentiate into
~'s w
the p1.erolUe and the periblem (Table ) (figs 3) e
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 Control
Tible 5
The Average Diameter of the Root, Expressed in Microns
(biased on the data of E.reslavets and
Afana,syev9. 1932)
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250 x 5oO 75O r
303.8 362.9 387 s 14 393.2.
By using the average size of diameters of the roots it
wee possible to obtain the name kind of a graph curve that it
was possible to do by ? using the length of the etas. With a
dope of 250 roentgen units the curve rises sharply;. a dose of
500 roentgen units con .nues the rises with 750 roentgen units
the curve reaches its a pex after which a gradual falling off
begins; and at 8000 roentgen units the average diameter of the
root equals the diameter of the control plants.
320
310,
300w-
1000 r 2000 r Looo r 8000 r
358.6 326.1. 316.8 305,23
Diamets'r of\the Root
MI1Y:YtW1W,'NMiAWMWNt+kY'ffW1)M?4HMW~ WwNNtpvMlMlwwu~hW..w4MM+IMa~M'~W wwwM,ewWVUYh?wY+,r
Urexposed
Fig. 3?
20 500 750
Qurve.representing the changes of diameter
of roots depending On their exposure. to various doses
of X 'ray
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During the spring and swtrnar lphenological observations
were carried on, which showed, just as with the irradiation of
sprouts, that there aren't any particular differences in the
o ions phases of ear. formatiaf, flowering, arid of the ~"
the irradiated plants and the controls. _1n~
rf.psniflg between
d.ividual plants, however: ripened from 8M1o days earlier than
the controls. is a point that will bear looking into, in
This the future.
Upon ripening the ears of certain of the irradiated
~
p ark.y one~and'a~ha ' tines a large as the earplants were nearly
of normal plants in our e riment, which, in turn, were con
r than plants which are grown under ordinary
s~.derably lame
oar8 in turn (i? e?, those of irradir
field canditionse these
aced plants) sometimes exceeded nonnal size by one-and-aihalf
timesd(Fig. 4J, comparat ve sues of ears of control and ir-
radiated rye: a control, b Irradiated.) but especially
amazing was the size of the grains which by their size approached
N
the size of Po~, iih wheat. In order to compare the weight of the
p
initial material with the weight of seeds obtained from rradi?
their weight an torsion scales, weigh-
at?d p1ante we determined
ing the basis of these facts, two var-
each
.able graph cures were constructed. By examining the left curve
we can see that in the initial material seeds weighing Za~? ~
milligrams'were most common and that they varied from 9? to
33.S mi~.~.~g~^arne, while in the r:~ght curve (weights' of seeds obi
material) a weight of 39?a~ milligrams was
tamed from irradiated
predominant a weight which does& t even occur in. the jnitjal
~
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material. The weight of the experimental seeds varies from
21? to 51. milligrams'. A comparison of these two curves points
up the large size and uniformity _of the irradiated grain, which
from the point of view of agriculture gives it a great advantage
over the initial material. (Fig. 5. Comparison of two line
graphs based on the weight of normal and irradiated seeds.
Note: dark solid line is control; broken line is for seeds
exposed to 750 roentgen units,)
An analysis of the identical factors in the yield (just
as analysis of the yield of the irradiated shoots) shows that
all graph curves based on individual factors tend to shift to
the right.
From Table 6 we can see that the average number of ears
on a control plant is Li.5e while plants which have been exposed
to 250 roentgen units have on the average nearly 12 ears. Then
as the doses of X-rays are increased, the number of ears tends
to diminish; however' it does' not approach the control until a
dose of 2000 roentgen units is reached.
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 'a'
500
750
1000
2000
14000
8000
)4.?5
87.0
Lb8.0
135.0
5.29
~.J.. f
,cap. Q
O.(
L44O.
. yb
87
292.
52.0
315
l3!3
8.0
297.0
61.0
358.0
1L..08
7.0
230.5
70.0
300.5
10.714,
14.S
131O
20.5
151.5
5.58
3.0
90.0
38.0
328.0
3.66
3.0
57.0
59.0
86.0
3.21
Only doses of 14000 and 8000 roentgen unite, which have to
be considered strong dose, cut dawn the average number of ears
per plant` to three, i. e., they give a number which is smaller
than the control (Fig. 6.).
Number of ears
Weight of seeds
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I Table 6
(based on data of Breslavets, Afanasyeva,
' and Medvedyeva, 1932)
Number of seeds per plant
Bose Number Normal Underdeveloped Weight
F
in r0 of ears seeds and diseased Total of seeds
Control
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If we analyze the co 1unri with the number , of seeds, we
shall see a similar Picturet the number` of Seeds triples with
a dare of 250 roentgen unite, then it begins to go down gradual
as for the number of ears, the number falls below
:Iys and dust
that of the controls with donee of 14000 and 8000 roentgen unite
(Fig. 7.).
Number of seeds from one plant
Unexposed
Fig. 7, The change in the number of seeds depending
on expoeure to various doses of X-rays
The diminution of the number of underdeveloped and
nd seeds in plants :which have been irradiated by bene
diseas
oses of rays, as compared with the controls' should
fic~.al d
noted. In oontrole b8 out of l3eeeds were underde-
also be
or diseased, that le 3+ percent, while in plants
veloped
which had received 2S0 roentgen units only 63.7 out of 16.5
eeeds were diseased, that is 11.3
percent. This indioatee?f an
increase not only in quantity of the yield' but also in quality.
to this what we know concerning the increase of the
If we add
weight of the seeds: than' the full eignifioanoe to agriculture
of'the proper uee of X~raye becomes evident.
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attention to the total weight ?f the
If we flow turn our
seedy ~e shall see that 2S0 roentgen unfits triple the weight,
it on the same.
and 7 50 roentgen units keep
that eXpasuree of X00
be inning with ]000 roentgen unite doss.
level, and that only ~
the w . roentgen units
~
weight begin to regxese (with 000 and $OOO
it is nearly twice as small as that of the eontroTs)? Thug a
dare of 8000 roentgen units has a strongly depressing, effect Ofl
umber of seeds,. ands, in ~dd3.-
the number of ears, the :weight and n
ber of unde~?dsveloPd and diseased
floe, it ~.nareases the num
seeds (Fig. 4).
s cific plant' for each
s,ooking over the data for every ~ ,
of the doses we find great individual variations 5pi for ex~
from 2 to 21~; the number of
ample, the number of ears can vary
0 the fatal height for any one pmts
normal seeds, from 17 to 93 x
his- great variability is character from
i,06 to 36.~b g~ams? ~' . t~
of all other plans which are subjec
istic not only of rye, but
ed to the action of X-days, as can be seen from literature. If
variability of the number of ears,
we burn our attention to tie
kes our attention is the lame number
the first thing that stri e
aced /onto, which did not have tim
of ssconda` .stems an i~^z adl p .
in turn, increase
o maturity. These secondary eteme,
to come t
seeds and `thus sower the total sued
the number of undardeVelapea
(The word "total" ~, s probably ~.nc or
plant.
weight, for a `single bob meant "avera~;e"'~reght? 3 If we are
reef; the author prv ~' fated
at the present moment' w1'j i,rrad
not in e position, to answer,
whia~i do not reach maturity,
secondary stem
p~,~ate-leave so ~ venture a ( oonjecte.
~,ppaxen,yt~' XMray's, by
we can at least
stimulating plants, open up.,~ss~:'~ilit~.es w~~;~Gka
days/?p~nt , in
6'
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in normal plants never reach beyond the potential stake ? How
ever, these ever reaeli tu11 development because
possibilities n
something is m~ i$i~ f IOU the environment, and this result in
tams and ears. What is missing
the incomplete develapment of ~
oil,~r light, or host, or all of
(nouri~hiflg elements in the
We cannot tell at this time.
these elements
From our pr+e1iflLina7 data it is quite obvious that xwray~
vela ant. of e. If we compare all
act as stimulants on the de
development, and productivitY
of the curves by which the growths
~ we shall of that they
adiat$d plants are reflected,
of the ~.rr
co t of the plants, the diameter of
~.nc~.de? the initial height
the number snd weight of the seeds,.
roots, the number of stems,
under irradiation. Radiations of
all act in a ~-1~,` manner r~ent~?
the curves up steep]'; X00 and 700
250 roentgen unite send
denc ~ and with doser~ of lOnO roentgen
gee unite continue th
units the curves begin to drop.
point out once more the
At this pa~.nt it eeer~s proper to ~'
h the experiments with irradiation
~~ecessity' Of carrying throng
or sprautss tc the point of the ripening of the seeds;
of seed s
ather~ise, it is possible to obtain an incorrect impresr~tan of
the effects of X?-rya ? If we had carried our expeZ`imnts only
up to the point where plants were beginning to send secondary
development we might have came
stsms, from the graphs of the
ose$ of 1000 roentgen units were most
to the Conq~.usio~ that d
reases in i lds. But ,by the time the seeds
cOnducive tows,~ls inc
ripen, doses of 250, OO, and 70 5
of the :irr~d1rpentM
ated p,nte .
unit. dome 9 which by this time
sn uits outstrip the 1040 raentge~a
.62
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is exhibiting effects which are to a certain extent depressing,
since it i? '4th thi dose that s;ns of decline begin to show
up. However' even this dose cannot be excluded from detenniw
t1on of effects of X-rays on athez, p1ants9 for there are tunes
(depending on the problem tiie want to salve) when it becorr~s .
portant to obtain greater initial developr-ient.
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The advantages of X~irradiation are not limited by an increase.
of yield of the irradiated plants, the advantages can be transmitted
to the offspring of the plants. These data were obtained by us as
a result of planting, in the fall of 1932
suds produced from
plants which were irradiated in 1931. For planting we selected seeds
of the plants which gave the greatest yield. For control we used
the offspring of the control 'plants of 1932, from which we similarly
selected the best plants, hence the best indicators of the control
plants of' 1932, by comparison with 1931. During the growing season
phenological observations of the irradiated and control plants were
conducted, and no appreciable difference in the length of various
phases [of development] was observed. The ripened plants were sub'
ected to the same detailed analysis as in 1932, i.e., the number of
ears and the number and weight of grains were determined. The ob-
tamed data are compared in Table 7
Table 7
Average data per single plant
(based on data. of Dreslavets and Afnasyeva, 1932 and 1933)
Dose Number of ears TotJ, weight of seeds number of seeds
r 1932 1933 1932 1933 1932 1933
Control 7?3 5.29 7.T 13S.0 222.2
25p 11.7 ? y?~` 1196 L.93 La.6, 0 168.2
8.7 6. 13.L.3 9.17 3L3.5 279.2
r,
75o 800 7.8 1.O3 10.0Li, 358.0 X06.,
bas 7.0 7.2 10.71 8.78 300,E 277.6
2000 L ,. . $?S8 7. .. 215.7
X00? 3.0` 74 3.66 9.)4 128.0 283.8
80003.0 7,2 3121 8.18 86.0 306.6
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1fatiOf of this table indicates an obvious secondary
An exam
act creased yie~.dS nth certain.. dc~saction which shows up in the in
x~rays. Although we sae that the 1933 yield is :lass than that of
~,se is evident in the total weight of the .
1932, neverthele~e the a.ncre
seeds as well as in their number, In ardor to facilitate the compari
son of the effect of direct irradiation and of its secondary action
we have drawn up Table 8, where results are iveh in percentages, uoax~g the controls as index of 100.
Tab/ 8
oslavets and Afnasyeva, 1932 and 1933)
`based on data of 13r
I D$eS Number of ears Total weight o scecte 1>ui-aber of seeds
r 1932 1933 1932 1933 1932 1933
.
Control
100.0
100.0
100.0
100.0
100.0
100.0
7
7~
250
260.0
}IdI
01.7
63?7
330?7
.
500
19393
93.1
23.6
U8a7
2L
12.7
7~0
177?7
146.?8
266.1
129.?7
26o2
137.9
1000
98.6
203.0
113.6
222.6
12L.9
2000
100 .0
7~*3
10~ ?5
9S . 0
112 , 2
97 ?Q
~Q00
66?6
97,2
69.1
121.9
9~?8
127.7
8000
66.6
6
60.7 ?
106e7
63.7
137.9
From ~ this table it is ssab1e to arrive at the
data , of
that a dace of 7a roentgen units not only
vary def~riltG COnelusion
increases the weight of soed b r percent as a result of direct
,~' ,
retains an increase of ' near/ 30 percent
radiatlCa:n,, but that it
nett learl s leld, a a, socondaz7 action.. Doses of 2O roentgen
ua~its which praauce ~ sut~Y~ ~harp ricrea$es (21 percent), by comparison
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with control, during the year of irradiation here give strongly
negative indications. We W?11 return to this question later ($es
Chapter 3). Doses of 1000 and 2040 roentgen units reveal in their
secondary action a gradual dininuton t o? yield] which takes place
more slowly than under the direct effect o X rays. the secanda r
action is distinguished by the fact that strong doses (LiOaO and 8400
roentgen units) which depress the development of the punt in the
first year .a,y a result of. dtrect irradiation) give . positive effect
ventua1 . Probably the explanation of this phenomenon lies in the
fact that those pints upon which the rays had the most depressing
effect had perished.
Pas; (investigations of Breslavets and Atabekov). (This work,,
started jointly was transferred on the following year to Atabekova
alone) This plant was selected as an object for investigation
chief because it is we known lrr literature for its sensitivity
to X''rays. 13es1d0S2?bhis was falt to be a usefu1 object for genetic
i
investigatIons because it is self pol1~na tig.
Taking into account the sensitivity of peas to X?rays we used
7e'n,
A
weaker doses for i.e., we began with a~0 roentgen units. Pea
seeds which were irradiated under air*dry conditions were planted in
vegetatin dishes on the day after frradiat on. When the irradiated
and control plants formed pods they were photographed. 3S0 rcentgen
units should be considered the most benefi ia1 dose since at this
dace the plant shared the most vigorous development and gave the
greatest number.of pods (Figw 8. DS fer-ences in the development of
peas depending Qn application of various dosages of x rays)
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In this exper merit the yield -could be judged only by the numbex' of
pods wb1ch were counted on the controls as well as an aU of the
.jd p1iLiit$ a The average number of pods per plant with its
probable error is shorn in Table 9
Table 9
Peas. Average number of pods per single plant
(data of Atabekcva, 1933)
DQse in r
Number of pods Dose in r
Number of pods
Control
2.33
? 0.32
La
3.57
? 0.33
5o
2.51
? 0.35
55O
3.38
? 0.39
100
2.6
? 0.3,a
6,O
3A25
? a~39
200
2.S7
? 0.39
7O
3.38
? 0.35
250
3.63
? 0.71
100
2.6L
? 0.1
X50
1..63
? 0.38
From this table it can be seen that a very we dose (250
roentgen unite) makes the difference between the controls end the
radiated plants reliably ti.e.,. grater than the probable error),
while a dose of. 350 ro itgen units significantly increases the
number of pods. Begiffg with L$Q roentgen units the curve begins
tQ drop and with 1,000 roentgen snits the number of pods evens out
with the nulnbei of podgy; in the control plants. These facts indicate
the etpulatng
effect off' X'rays which ehowe `up in the increase of
the number of pods per plant. For a mare accurate eval'L ation of the
quality: the ob ted yield of pea3, the irradiated and control
.s
peas were weighed on , a for ion scale ; in batches of 525
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In 193 (this date should probably be 1933 to correspond with
the table below., or vice versa] the irradiated pea seeds were planted
Out on plots of the Timizryazev Academy with a' quadruple repetition
of the experiments. As the action of X rays ndicates,we selected
the average weight of seeds' collected from a single plant. This
data is compared in the following table.
Table 10
Peas. The weight of seeds from one plant
(based on data of Atabekova, 193)
Dose in r Weight of seeds,. gr. Dose in r Weight of seeds, gr.
Control 3.66 ? 0.28 550 3.56 + 0.29
20 ? 1..29 ? 0.32 650 3.69 t 0.33
35O 5.07 ? 0.37 750 ).,22 ? 0.39
)450 1.26 ? 0.30
From this table we can see that as far as the increase of the
weight of the seeds is concerned the dose of 350 roentgen units has
a particularly beneficial effect.. From this we can deduce that for
peas, just as for rye, an irradiation of the seeds before planting
leads to an increase in yield ,(Fig 9)..
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t
til Leading control
r
Y /
Growths irradiated with a
ddse of 2Q roentgen units
F 9 Comparison of the weights of pea saed8 in contrn).s
and in plants which had received 250 roentgen units
Five years of experimentation with the effects of radial.Qf
on t us toe~ress our cnnv?ctiOns concerning the. sign
plants perms
nificance of this factor in stimulation of growth, development and
yields of plants:. Negative results which were obtained by many
jnvestigatars we explain by insufficient work on the dosages, which
are specific for every species and variety of plant. Let us take an
from our own experience. Despite three years of experience
example
with spring wheat we still have been unable to obtain stmulating
doses far. it. We don' t draw pessimistic conclusions from this that
slimu,at doses don't exist for wheat, but we consider that we
have not discovered them a yet. Nevertheless our experiments with
this plant axs rather irtc3rest.
west (nvesttgattons o.f Afanasyeva) . For our experiments we
ussd, a pure 1e of soft wheat whose suds were irradiated in alr"
ons, soakedsand, finally, in the f arm oi' sprouts. The
ca~d~ti
dr~r,
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a _: Seed wa$ discpvered t be not radii en$ tiVO3 it didn~ t
react in ary way on exposure to small and medium doses of X'rays,
wk es to heavy doses produced a depressing effect on the while while e~-poeur ~'
f'rst phases of development of the plant. In order to study the
~.
4.radatian on soaked seeds they were planted in the lab
effects of X
oratory and in field plots This last experiment was performed under
.
n using land planting by row. The seeds which had
quadruple ~?epe~titio
been planted under laboratory conditions all came up at the same
time (regardless of dosage as the controls. The percentage of irra
hinted $eedo which cane up in no way differed from the controls.
:pux'inb further development obvious depressing effects were noted with
a dose of 16, 000 roe~ nt g en units (,j us t a s in the previous experiment).
Under field conditions it was possible to obserI that as the
dosages increased the viability of to plants went down. The dose
of L6,000 roentgen unite, which under laboratory conditions came up
in the same amounts as the other dosages and the controls, under
tions made a S0 percent lower showing . This Is. explained
field cond
by the lOss of a certain percentage of the day ilopa ng plants` during
the entire rowing season of the plant. Not a single one of the
.
dosage ~ wraY v s iscd ehaed a tmulatig effect and only haws oof x
8,00Q and 16f000 roentgen units exerted a depressing effect on the
and develo mont of the plants. This depressing effect lowered
growth
the y`eld by30 percent for a dose of 8,000 roentgen units axed by
63 percent for >a dose of 16,000 roentgen units.
or neither the dry for the soaked seeds, of wheat were we able
to find stimulating doses of X-rays. She many authors point out
that sprouts are morn sensiiva than seeds, either dry or wet, it
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became cessaXy ~or us to 'firJ out the effect o~ the rays on sprouts.
eedS of wheat were al3.aWed to germinate an wet filter paper oi'
the germ ed out those which had Just begun t
~.r~~ing seeds we p~.c~
shag the sprout'. After irradia4 with doses frog 0 to 8000
roentgen un e planted under iabara~~' conditions
,~ts ~ the se~a~ were
in tree groups, 100 seeds per group. SIoots appeared
in all dosages' and in the control. Fux'th0r observations l Uh!
de~re +eri4dic rneasuremente r~hawed that doses
~.~pmerrt of the .. plants and p
01' 100, and 200 roentgen units do not afcct the growth and
`~. to that of , the
delTelopraent of the plants 9 whose size ren ~,,s s ua
controls With .h doses of OQ = p this is a -rnjstal~e s it should reed ~OQ~
~,s . With ~
roentgenun~tS beta observe weak dep~`essive e~'i'ect.
. :~.t is pQ,~s~. ~,
s cialiy 16,000 roentgen' units
With dosages of 1000, 2000q SQOn and epe
jt was possible tO observe a strong ` depression of develaptent
r doses de?ve~.c~p~d ~.dorrt~.ca~y with the cant
plan ~aa~ated ' by other
h~.ch had
phano1ogical observations have shown that plants w
tx~c~.
received a dose ~a units begin sending secc~ndar~' etas
?
of 1000 raentge
b days later than the controls, and that they begin to ripen ten
,
tame fir from, all of the p1ants w.ch had
days later.... By YarVest
been exposed to this dose had riperred; the ears were sma11 and many
of the seede` were underdeveloped. At the time of harvest the height
of the plants was meaSured9 and thQ,number of ears were counted Cor
.: .
e~rery
plants at harveet tame, the number of ears and 'weight of seeds, are
,
p.
nt for' each of the doses. Threshed. groan was weighted
,
separabe. or each p'lanee,. The obtained average data (height of
~r' ~
aoun~~d
r'. sing1~ plant)' aro presented in Table 11.
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'fable U
(Based on data of Afanasyev&.,, i.9314)
Dn se Height of Number Weight o?
In r plants of ears grain
c a gr.
Dose
r
i eight of Number weight of
plants of ears grain
cm, gr.
r
Control
93 .
3
2.Lil.
?1000
93 .
3
2.39
o
Dry..-
93 :
3
2 a 20
2000
9
3
.
2
l2~
92
3
2 `e 2b
1ooo
92
3
2.19
i. T,
8000
88
3
1.78
sUo
93
3
2. Li.
16000
75
?
O91
7O
93
3
2.33
Ana 'z .ng this table, we can see that the height Qf the p1 me
remains unchanged despite the action of rather strong ddse- of X.rays
(only of 8000 and loo roentgen unite decrease he heig(only doses n the stro est of all applied dosages, ..e., 16401
Similarly roentgen nits, brings the number of ears down to 2 from 3 whit h is
blie , number found on 'bhe controls and on all the plants which had rep
'red smaller dosages. The weight of the grain per single p; nt
caa.~
began to become smaller with the applicatiQn of LOO0 roentgen units
and fell to near/' a third when 14004 roentgen units were used;,
In compar lg the action of radiation on ch7m soaked, and
sprouted seeds of wheat it was established that the la are. the
most sensitive, for doses of 1000 roentgen units had Strong depress
sure effects on them. Surrunari2ing the investigations of the effects
z
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11
of Xrays on seeds of wheat we can see' that for this plant,
Manasyeva s experiments, stimulating; dosages have not yet been found.
This dos not mean, however, that we should stop trying to fend them.
It is possible that b' changing
external conditions (for exarnle,,.
temperature)it will be possible to achieve success,
In any case, of the three plants subjected to the action of
X?rays our small laboratory obtained a significant increase of
yield with peas, and 'a very largo 'increase of yield with rye,; 1
sides continuin~g to work with the irradiation of rare we have already
planted with exper iental seeds half a hectare :of rye at one ; of the
ko Lkhoz of the Moscow Oblast, The results of these experi rents will
be described in another article, We shall only point out that on
both he initial 'development and yield X-rays exerted a mo st bene
\ e/ c r
Licial ~-?...
The inves.gatiQns which were conducted in our laboratory on
changes in the development of plants due to irradiation of seod of.
s
t7e wheat, and peas, were set 'up with the idea
mistakes of our predecessorsa
f avoiding the
1. The,seeds irradiated in our experiments received exact
doses of X"rays carefully measured by moans of dosemeters by expert.
phy3iCist8. 2. The amounts of radiant energy` were graduauy in
creased in order to obtain the full curve due to ;irritating doaa
gas
(i. e., according to the Arndt"Schultza lawn) ,
.. 3 A sufficiently
large number of seedy were used in the experitnents so that ., results
wouldn't be affected by individual fluctuations
.1i. When?ver, poa..
sib/ s ilar condibjo of temperature and humidity were maixairainad
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both during irradiation and while the plants were growing. 5. Thox
ough
phenological obeervatiOns were carried on during the course af.
the entire deve1apment` of the plant from the morent it Came up rrom
the round to the complete ripening of the seeds. The crop was
carefully gathered and ana red. These observata-ons showed that
? nation during the first stages of development doee not coincide
with the increase oz" yield,, at least for the most thoroughly nvesti'
ated object, i.e. spring wheat. 6. Measurements were carried on..
There were measurements of the main xoot of the sprouts, measure -menu o:r the pZants on the twety~$eventh day after planting, and
measurements of ears and stems at harvest time. 7. The seeds were
weighed in such a manner that the weights of healthy and 'cf diseased
seeds could be obtained as well as the total weight. In addition
all of the ,seeds of single plant, selected for the = except1on*:y
large size off' seeds, were weighed, `grain by grain.
As far aseparate cultures are concerned, our investigatins
have shown that e is sufuiciently sensitive to X-rays that experp
dents can be set up with it and a whole series of questions which
of interest to radiation bioiogists can be answcred. These ex~
*?
per ents a 1. showed the Iconteatabie appUcation of the Arndt
5e hue law to the action of X-ray on plants, alb least an rye,
since the measurement of young plants an the twenty' eventh day' after
:ant.in` gave a splendid growth curva small daae (2S0 roentgen
units) stimulated development, QO and 750 roentgen units produced
still ?ux ther aeve1opmant in t1 e same direction, until at 1000
a
roentgen unfits m rnum s'bimulation . was achieved, after that the , curve
began to drop, 2, confirmed the variable sensitivity of 'soaked seeds
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and sprouts~ 3$ showed that dijferences do nat' depend an the qu.litY
~+ ~, the effects of radiatien
o~~ rays;
o ~ the rays, i.e., hard , or ~
an the individuality a? Various
~ad'u in dif~erent ways depand~
shay p
r and weight of seeds a the .length of
p1ants~ which affected the iwmbe
the stems and ears`, the time of x?ipefing, etc.
~ee ocal canoLusians we were also able Besides these pure ' t~~~~a~.
p
to establish the effeCt of x~rays on ire.. Plants which
s~~nut~
had dove~.aPed mQr~ quickbad broader
o
received ~ene~~.caal dosea
` deeper ~re~n Gc1cr than the cc~~.trc~a~. plant8.~.eavcs, ...,and e,n ~bited a ~
` coats foflowed the same curve of growth
The average diameter of the
O roentgen hn~.ts
e lants kith a dose ai' 7
-~h~
as the ther pars of
it gradually dixninishg its size as the
' ~.ncr~ased by ~~ percent, ~~
doses incx?eased. Analysis of the yield indicates that the number.,: of
ears: tees, while the number and weight a
increases nearly' three
e thing , the advan'ges of i?rad3.ti0n
suds per punt does the ~aln ~
are not ijted to the increase cf yield of the 'directly irradiated
` plant; they are also transmitted to the dascendants of the irradiated
pianto Seeds collected f'xm' irradiated plants gave an increased
a d?$$ Of 7~U rcentgen units tae
yie1d on the ~olioing year, with
increase was 37.99 percent.
$a,nc a pea suds are krofl to be 'extremely radipsensitve the
~. s wore used for their irradiation.
weakest pa~~~.b~.~ rinses of ~ray
number of pods per single pant doubled
In the : first e;cpeximent the
units was
troll when ..a dose of LSO roentgen
(in campar~$0n with of ~~~ rp entgen units
used. In an experamen'b of anc7theD year a dose d
'
seeds per Single plant from 3.~ grams . (We
raised he weight (weight
t of .
of the control) to .07 grams.
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The Irradiation of the seeds and sprouts of spring ;wheat
produced neither stimulation of development nor an inorease in :yiold.
Only high does of X-rays (8000 and 16000 roentgen unite clear
showed the depressing effect, .
Sums rizing the results of our five years experixnents with
~.
three plants we can draw the conclusion an ecposure of rye and peas
~
X rays before they are planted results in the stimulation of thelx
development arid an increase of yield. For wheat we have so far been
successful in finding doses which produce one or the other of the
.above results0
Rye 'ass (Investigations of Breslau te). Severa3. years have
passed since the end of our experiments with the action of Xra s an
agricultural crop plants, and the question of application of these
ray s to meadow grasses came up. One of these was the estervalt d
rye-grass, picked out as the most rapidly growing one. Doses of
BOO 300x.04 ~ a , 500, 7O, 1040, 2000, L.Ooo 8400 roentgen units
wore selected for this plant as well as a control,' They were planted
on Fobrua3y 2i, 192 in quadruple. Ff of all the time. of app'ear
ante of seeds with the various dosages was noted (Table 12).
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Taber 12..
(Based on data of Bres1avets' 1912)
Dose
28 February
2 March
L. March
6 larch
control
7]
.76
200
39
8
78
97
300
145
j9
31 j
oa
146
6
89
l(
500.
. 32
81
914
97
750
37
75
83
9
ioc
140
. 7t
8
96
2000
9
8
9S
100
1000
18
39
814W
91
8000
2
15
61
7
These data show that doses of 3OO~ )4.00a. Oa and 2000: roentgen
units cause 11 seeds to grow;throu?h, while in the controls only.
76 percent came up. We have carte across this type of phenomenon
more than once in our experiments with Xrays on the seeds of many
other speo e of pants. he a fac?ts .should be noted since they
show that an amount of the vaiuable seeding material can be saved 'by
application of radiation.
Ty the end of MMarch the plants had developed to such a '!Dint
that it`was no longer pos ib i to leave them in boxes. They were
removed from the wound and, the number of roots for each of the
/ants Lox' each dosage was counted:. In addition the lengths of roots
and stems, end the weight oI the stems' were recorded ('able 13)
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Table 13
Dose '
Based on da
Number o
of Bres
vets, 19h2)
Length
,eight of
;r~ r
roots .
ci ste
of root
st rres
Oontro1
10U
100
100
110
2OO
240
1234.
100
113
Qo
23O
1E;
142
112
:ao
230
110
" 102
112
bo
20o
118
6
a
7;0
290
119
125
176
L0O
124
90
98
80
coo
150
107
loo
87
14000
127
.93
10
As can be seen froi this tab1as.a dose of 750 roentgen units
near' tripled the number of roots, ancrea8ed the length of sterns
by 19 percent, increased the length of the roots by 25 percent, and.
increased the weight of the stems by 76 percent: 'rho above dose
pb isi' stitau taa the growth of the
p1ants9 while a dose o? 8000.
rc)entgen units which depresses plant devalapment can be used to
create the impression off' the unevenneae of ti soil due to the uneven
de1opn nt of the plants e
r the numbed ot roots we can see the typical biological`
carve ai the ~~ndtchu.t~,aw a gradual, rise up to and ine1udig
7O roentgen units and then a gradual Ealing off to the dose of
7
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8000 roentgen urJt, The sie may be observed for length of sums
and we ht of stems From this we may draw the conclusion
..and race
that V tervai d1 s ryegrass is a rad osexadti,ve plant.
i~s~4 w grasses (investigation of Afanasyeva). [e seeds of
?oliowthg meadow grassos were subjeeted to X-rays: bottom
the
grasses red fescue, red tap grass (Agro tis) and Kenbuekr blue
~
grass (Poa pratenis) top grasses, fescue timo thy, brorn gra8s,
,
rr
and American couch or q b grass. Doses of 7O, lO09 axzd 3000
,
Its were used for irradiation The irradiated and don'trQl
roentgen un
seed a7re planted in boxes in quadx~pl. The bores were located
? the greenhouse of the botan cal `garden. Sprouts Of the irradiated
seeds appeared at the same time as the sprouts of the controls. Th
dose used had practica]L1Y no effect on the percentage ref plarLts
:..
which came up., since. we didn't attach any si ni iCance to a sraall
rise in the percentage (36 percent). Then the appearance of the
,.
and third leaves were taunted but no differences were observed
second
hare either. When the plants were removed from the ground the number
~,
ax) roots. of each plant were couxfed, measurements of their.
of iGas
length and weight and cireumf$Z'encs of their root system were made.
This was done L or red f ecue~ for meadow f e scue brome grass, and
u~
for Ameriean couch or grass.. The obtained data failed to show
a djfference by ` of 'Ghe above signs, between the irradiated
, ,ants. The ;doses of X-rays which were used ?aid
and unfrradatod p
,
ing effect.
to ; show either a stiirnulatg or , a depre~s
Data far tae ~ec,and accounting, at the time of appearance of
the thud ~sa~ were ab~td:far:.brams~aes and Fescue. 'These data
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show that separate doses increase9 by this moment, the percerntage of
plants which have developed three or more r00t8.. This can be seen
from the fallowing table (Table ~.
Tab'e i;
Percentage of plants which after 35 days
have developed from 15 roots
Name of
kid (of.. Plant
(based on the data of Afahasyeva,
Oontro 7 r i;oo . 3OO r
No of roots No Of roots No of roots No of roots
l&2 3 .&2.3 1&2 3 l&2 3
Id fescue 61.1 3a'9 .93x1 LF9.29,8 70.2
Brome grass )$3. 5 . 32.1 07.9 2.6 7L..h L~l.8 X8.1
From this table we can see that with a dose of 3000 roentgen'
units the percentage of plants of red fescue having three or more
roots is nearly double that of the controls, namely: 70.2 percent
of the plants (as against 3.9 pexaent) have from 3'5 roots;' whi
a dose of 100 ;roentgen units with brome grass increase the nunber
of roots by 18 percent over the controls
These data give us same 'background for selection of doses
which are capable of developing root systems.
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Atabekova, A
Spr9uts of Peas." Biol. Zhurn.' T 5 ado 1: 99116.
1937. ttT"he Action
CIA-R D P82-00039 R000100240012-2
1936. "The Action of X-y- on the Seeds and
of. X4 s cn Peas ,1t ., Gad. Sens e
I _Rak e. , T 10: 3O1?.3O2
193.
'I oncernag the 6tirnulating Action of X--Rays on
Plants "
Afanasyeva, A.
T 78: 5662.
1936. r1The Action of XMftays on the C : ,ular 1o.a.
rents of. Spring 'Wheat e' . i. Zhurn. lp i17.12L
1936. "The R8tentjOn of Action of X*R r on Wheat." _____?
Moak , Ob.Va w `. T L5: 1334L Oe
1939. ttThe Ac Lion cif X'Rays on . AirDzy, Humidified, and
Sprouted Seeds o: Wheat, o~ ibid, T 18 (1): 1928.
resla ts, ,. P. and Atabekovaj A? I. 1935? rtThe Increase of ?Y eld
Under the Influence of X' Rays. Peas." Tr.' Vyp 8; 2L.
253?
Breslavets, L. P Afana yeva, A. ., and edvedyeva G. 'B e 1935.
"The Action
X 7 ay on e
l T, ?O; `253Q1,?
193
Yield Under the Influence of XMRays e. I. 2 ? Irradiation of
?eslavet , ~ " o ~. ar1d Afanas ev :. A. S. 1935.
Seed.. r~ Tree vyp 3 a ; 2)
ves~tr~. Rcr~t ~nol. I Rad~.o~.
53 ?.'
The Action . o f X i y on ye. 1. Irrad Lion ofSeeds."
vestn. Rent eno1.. Radiol. T 0 w
., 2
~iT Increase of
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Byes vets, L. P. and Sinits,kaya, L. A e 1937 r "Exper ents in
Application of X Rays and Ultrashort Wavos in Horticulture. tl
Tr. ' Eotan. Ca& gu. T l !66
Breslavets L, P. 1937. ttContemporary Developments in RO$ntgen~
b ,o19 yell Ob ar. e o, k. db"'v 'iq `r d. p e
T L.6. (6) s 359369
Gambarov, 0. 0. 1925. "On the Question of the So Ca11ed Irritating
Action of. X-Rd's:.r$ Vein. Reza eno:l. I Radiol. T 3 Vyp g
3U~323
Eoroshenko, A. V. i9?9930. "The Influence of XRadiation' on the
Length o the Growing Season of Plants, rl Tr Po Priki, Bot.
fie:. 1. Via 2
Zhukov, N. I. x93!.. 11 The Influence of X?Rays an Sprouting and
Energy of Gemination of Tobacco' Seeds." Sb. Rabot po Gen.
1 Sel. Tabaka. Izd. Vses. InTa Tab. Prom. Xrasnodar~ 17173,
Zenkevich, E. and Brunt
: Individual?
.v? 1937y a e e tg nov .1 h Luohe,
(L, r
c a R,avit. a lkhork.a L e m 1 Rene
(The Influence of X,Rays on the Individual Devlop1nent of
Tobacco., ,Poppies, . Flax, and Rhubar :) Ira-T Eatar~il~j AI'
Xol e tso v, A. :v. and L. I. 1928. "The Influence of the Radiant
Energy o;L' Radioactive Elements and X Rays an the Growth and
Develaprnant o' Plants. fl
Mikylorciy, A. G. and Oc4! dehteyn, L. M. 1933.
XRays, on Yields of 'Agricultural 'Crops
ia-. P 12 Vy'p ht 21226.
Thy Inf I,ucnca of
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Nemenov, I . I~ 1924. t'Advancee :ire Roentgendlogy'in 2 Years,''
"estn. Rent Brio. Rad.. T 1
1933. R .' T 11; Ii.16. og iz .
Frolav, G. 1936. ;'zh~ Action of XMP ys and 'Uitravic et aays on the
Growth of Plants..Il '1?r., wKh~ Arad. Irn. T` aZev. T I
v 2: 189246...
Chekhov, . v. 1931. t~The lnfuence of X*Rays on plants."
Tomsk Un'-Ta.
T 8s 67ai13.
Altrnafln v., Rochl n B.' und'Gieichgewicht E. 1923. ttUber entwick
lungsbeschleungenden urmd entwioklungshermnendsn Eiluss der
Itontgenstrahlen.'t F brtsc~ir. Geb. R$ rrbenstr. V 31: 51-62.
Ancel, S. 1921i.? ttActlorn deis falbles doses de rayons X sur les grainer
oche ? tl C. R. Soc_. Biota 1i35 1J136.
1925. Its rayons X appliques sur des grainess~chee n' ant
aucune influence sur 1? poque dtap r tion du germs. Bull.
SQC. Bptan. `ranc . v 720 196.
r
1925. "Suz un'phsnom ne de pseudo-excitation deterrninee pax
les rayons X sur lea bourgeons dormants et sr lea cotyle
dona?res de 1enti1e It ibid. ? v 72: 1QS1i.
1926, "Rechexhes du meilleur test de la radio-reactor de
grained des LegumirOusee." ibid. V 73s 7173.
"De ' influence accelatDlces des rayons X sur le deve1oppernent
~.
des planta$."-ch.1.. 5s lob,
1927.
Le, retard de d~ve1Opp,rnoUt datermine par 1' application
des rayons X sur 1e8 graines sst'il du a uno action sir los
3? ?" Tid. V 7 m21..
Vyl?
1-2s 329336.
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Atabekov, AQ 1930. 'Die Wirlcu der Rcntgenbostrahiung ruhendor
and keimender S en; rt Prv lasma V V h 2: 23l2 6O.
.,....
Atkinson, F. 1897. "Report upon some preliminary xper ments with
the Roentgen `raja on plants." Nature ,
V of 600
nedjct, A;. and Karsten, 1931. "Effect of soft X rays on germina'
tion." Plans V 9m No 1'2.; 173i78
13ar8a :: L .1926p "Strahlenbaologische Untorsuchungen... `1. Zur 'rage
der `Rc5ntgenstrah1en b i Keimlingen. S~.tzungsber. Akad. Wins.
in Wiens Abt. 1 D 13, No 1: `L21 4 1..
tt or die Wirk1ng de Rontgonstrah1on auf die Kcrnteilung der.
Wur e1spitzen von` Zea Mays o ? Tbid D 36: 383.L.19.
T t ber den inf1uss der Rk5ntgenstrah en auf die Atmung der
Wurzeispitzen von Vic: a faba a rt /bid D 1362 L o3 19 ?
.Bergonie et Tribondeau. 1906. Interpretation de queiques reaultats
de la radiotherapie at essa,is de la fixation diune technique
rationelle." C. R. Acad. Sc. Paris V ] 3: 98f.995.
I3resvetz L. P. and AfanassieWa, A. S. 193e "Die Wrkun der
Ro tgenatra.h en auf Roggen. I. Die Samen." Pratoplas
D 23, H h: S2O533. .
X3,7 ? "The aetian of K,ra~r on the rye. 2. X-radiation oi'
seeds." a0 V 8 No is O 7.
Dord~, r~ Hf i9 3.. " . chemische' Wirkung der Strahien, insbesondore
der Rdntgans'trah en. t' S1 t~~, . H 11; 368
I
12-2
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C spari, W. 192k.a
Iw
ita s zur bioIngi chen .Crw~1 t der 5trah1en ..
wirkung. rah n 2j_ s V i8: 17 36.`.
192. Zum bioiagischen Wirknngemeclaniemus der Rtntgen
Qondcn, E, and Terr 1, I. 1927. "Quantum` phenomen (Sic 3 in
Cat e1, W. 1931 t'The 'ef'fects of X~ra rs on the growth of wheat
to biological ction of .gays
Y U: 3 L.i
Sc?en. V.73 , N1878g 5333
Campton, A and U1is n, s.. 193. ~ inTheo and E ,axthn8nt .
3d edition. Van 1rstrand, New York
Coat'
s od l iri tt
P.
strahlon. atrahlenthorape. V 2Q 195
tt
Jcnixn. Cancer Rest
1931. ttSensibl~.ty to X-rays of coils in .trO." f rch.
B U.
Cran erA H. 11Zur 'bio1ogischan Strahienw r ng." Strahlei herd
V23a $3,
Czepa,9 A. 192 "Das ?rob in 4r wachst rdsrderjaden urxd w chstum
steigerndan Rontgen~-radjur Wixku tga" Strom hors,
.v 21: 913;
.9? ? tt er E n nss deb R ntgenstxah1en auk" den Iceimunga.
prOOGs8, der Pf 1anzens en. ~t Fort.sck r. oob. Rbntgenst4
y j2
75.
t
esauer, ?. 1923. ,.~ Zur EZ- ,., 4,~ d bialagischen. ~tr~hiex~w~:rkgen~t
.
trahienb ' a V 16 2O8 22
0
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joss 1E r, F. 2L "Uber die bio]og sche ,Strahienwir
Gsb. y0$2 ..
122. "Ubex die a11geme Jien Bedingungen fUr R othesen
bilduren in der R~ntgentherapie. U: Strahlcnthera ie .
i9 s J43 ?
EUinger,;. 1936 "co11ds and bio)oga] effects of radiation."
Na ~iaxe V 138 a 1o],Li0
Esdarn,
of Ff.aren.,""+'crteahr GeU. Rr-tertr. v 33: 528,
Eirier, 1906. "tiler die he lende W ricuug der R&ntgen trah1en bei
abgegrenten Biterungen Veroffenti. Geb. des Milit~,r. Sani
t tsweeens. Berlin.' In'Jahrb, d wise.ensch. }3o.tan k
.v 56 U.,
Geller. 192. t?'Die Wirkung der Rntgenetrahen auk` jugenciliche
Ozganismen.q" Klin. Wochenschr. V??3 g 561.566.
Glocker, R. 1932. "Quantenphy ik der baologisehen R ntgenatrahlen
wirkun
I. 1925. "Untersuehun en uber inw rkung der'RcnL ene rah en
chr,w,l. B 77 653
675.
Glncker, R., Hayer, L and `J1 n ling, 0. 1929. t13jer die biolo i the
Wirkur rachiedener i tgenstrah1enq 1itIt bei Dosieren in
inheiten. ~+ Stxahlenthera,. V
Goodpeed,
32: i..38.
and Uber, `F. M . 1939. ""Radiation and Fonts, wIsing; as source material some
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times wide leaves and sometijne narrow, needlelike ones to create
new forris which ara so prized in horticulture, Changes in leaf
Structure can serve the sage purpose it is suffidennt..'or e c
ample, to produce a Wrhinkled appearance with certain parts of
the lea?? blade Mein up in order to achieve the purpose,
these changes can,. only be welcomed by hor?ticiiltura.li:sts since
.
they increase' the variety ofrai.able plants. To this must be
added the var:} egated coloring of the
leaves which also arises.
quite frequently ss a result of irradiation Leaves lase their.
coioring either completely or partiaUyr i i which case the most
curious distributions of white or yellgw spots or stripes can
be observed, What unusual decoratjve forms
can
be created by
combining changes of form and coloring of leaves; &iually rich
prospects open up before horticuituraliots in relation to changes
of form and color in the corolla, calyx, and. fr?uitb
From what has been said, the tremendous si,iicai3ce of
X' rays :t.n the creation of new for~as should be clear. Many of the
changes which have arisen in plants deserve attention
not only from
'the theoretical but also from the practical point of va.. r
ewm, fh..s
does n'o't mean, however that is is easy to obtain .a - '
~ ~es.~t'abie changes
in plants0 A great deal cif labor is required not on
~ ly in order.
to find doses which will change the form of the plants,
but Aar"ti-
cularly in order to see `I to it that these changes a
. changes are desirable
ones, and that is d fficult and 'very time consuni.Ln '
But no
matter what' difficulties beset the `path of the e
xperimentor they
aro well worth the success which is bound to crown the efforts of
the 'persistant investigator, . ,, s.oo. rer wih. some pants a,nd.later
with, others;
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REFERENC
<
Assoyeva, T V. and Blagov dava, M 1935 "Artificial .Mutations
th Potatoes" S?~ _ts~ t n read c- . No 15: X31 "
Afansr yeva, A. 5. 1938 The Action cf X-rays on Air-d7 Soaked,
and Sprouted seedy of Wheat', 1 .1,: o O v ~P i
TL.
Breslavets, L. pw ?939. t1llorphalogica1 Changes in Hemp induced by
rays! V AN U
Breslavets, L
SR ' Otd piol
Afanasyeva, A
?335 ? 3j3:
and Medyedyeva, 0 iJ a 193
uEffects of XRay's on Rya's Vestn Rant
288 301
enal I Radial T 10
GerasirnoVB, Em N. 19IQ "Tranelocation Between the B and D Chromo~
sores, Trison:c Effect of the B Chromosome in Cre'
Tectorumdt' lxv. A . IB3S1
ray voh, C a. 193g.. IfObta nin of Mutations inFisun by
Ecperiment. mean
of
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by Rays"
1936 "Pear rent 5 m ter lit r Induced by X-Rays"
8$
553:
Dole. 'AN USSR' l
55~
1936 III it:ssib1e to Cbain Tsefu1 Mutations
mental Means?" ; Pr~da. No 9 e 62 -
Sapegirl, A.; A , 193. '
ray titan s as a Source' of New
Ari'cu1tura1 P~:sns~
51~a
'rda o9 z 8
by Experi-
Varieties 'of
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Concerning Delayed MQ.diiications
Cl ronosomes induced
 Declassified in Part - Sanitized Copy Approved for Release 2012/03/20 : CIA-RDP82-00039R000100240012-2
Chekhav, N.
V.
x.9.31.
Un Ta T 8
"Effects of
67135
X"Rays on plants."
Tr, Torsk
A. 1936 ,*wirkung des cntgenstrahlen auf die Elemente
Afanas8ie
von Sommeeizen Triticum vulgare var. caesium) ." ____
o
V 25: 77-91
Andersen E. 1936 ' Induced chromosomal alterations in mare
y
a in Duggar' sivla .cal Effects of Radiation
Ch
At~be~owa, A 1936 "Die irkung der R ntgenstDah1ung ruhsnder and
ketmender Samen. Prolaa8fl V
Daux, E. 1932 nDie Au1?iOsufg
2g ?3h 26O
von Faktcrenutatianen dutch chemische
Labor. Genet
and psikalische Ra ze bei Antirrhinum. r, Tr.
No 9
1930
,~ utat3ans Auf/ sung der Antirrhihum majus.
f. Bot V.
23i 767O2
1932 ,Der Ein fuss von chemischen un
p
Zeitschr.
ikaischen Reiungen
dje Muta ionas??,ate. von ntirrh .rium a u
ind Abst. and Verel~un 8 :a ba: I67473
B1ak?elee,.
A. F 1929
itshr. 'f
t1A ,a]ysis of nature Stramonium plants gXQ
Bze81avetz L.. 1939 I'Mnrphologieal changes in hemp induced by
X.itadiono I :croecopy observtiorie" Bull. de liAcad,
. ,
SC o de i' UR : ' 33 '3Li8
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from seeds exposed to X-rays." Anat. Rec. V hh* 281
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n/_i
Belaunay, L. 1934 "Die Chromosomenaberraten in der Nachkommensehaf t
von rontgenisierten Ah$n einr reinen'Line von' Triticum
vureb..su." AI1. Zeitsc:hr. ind. ?Ab$t' u. ererbungs.
lehre 55: 352355
Duggarm B. . (Editor) 1936 Biological effects of raditio~
Gager, S. and Blakeslee, A. 1927 ~~Chromeasome and gene mutations
Goodsell, ;S. 1930 ?The relations between X-ray intensity and the
Goodspeed, T. tL 1929 "Cytological and other features of nriants
plants produced from X-rayed `sex cells of N eotiana tabaeun."
Bat. Gaz. V 87 563 82 .
1929 The effects of Xrays and radium an species of the
genus N`icot ana o ' Jou n. of , Heredit?r 3 20: 2L.329. -*_ .-,,
1939 " nheritarice in Nic'otiana tabacum. 9. Mutation foi~
Calif. Bubi.
, 0 ~r oo rang of tripIcid and' tetrapi4id individual in
~'~ i c tfil ~6 W~ C~~ aeA q~j`J ~.(y~gyI
yY x
" t y n 7
a x
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Goods:peedo Te H. 1936 r'Indu ed chror Qsornal a1terations," Ch
in'Duggar, 1o1agic~i effects of radiatLon
1937 "significance of cytogenetic alterations induced '
high frequency radiation in Nicer species ~~ypl
Goodspeed) T. and Avery, A 1930
!Nature and sjgnif cant of
structural 'chromosome alternations induced by Xrays and
radium." Cytologia V 1: 3O8327
I933 'a lnherjtance in k ico bona tabactun. 13 ? The cytag neti+v
of deformed' X rays derivative .'+ Gen V 13 d h81-521
.93)4 ''The cytogenetics or fourteen types derived from
single X~rayed sex cell of Nicotiara tabadum
V 29: `32-7~33
It Jaurn. Genet.
Qoodspee ~ T., Avery A and Olsen, A. R.1928.: "Progenies' from.
X*rayed sex cells of tobacco.' Science V 67: L6
Goodspeed., T. and Olsen, A 1928 t'The production of variations
N cot na :species Xpray treatment of sex ce.lls." Proc. Nat.
_._..a,.-*-*
Acad. ah- s 6669
Ooodspeed, T. and Uber~ ]". . 1939 "Radiation and plant eytow
genets cs.'t The 33o tfliC! fi9 ew V 5, No 1s 1*14.8
Qustar'sson, A. and Ewert, A. 19h0 "Two extreme Xwray mutations a
mp hd1ogiea1 interest.' iexeditas V 26, No 4: 2 7 261.
Haskins, C and Madre, C ?. N. 193 "Gruwt i modificattons in Citrus
seedlings grown from X~rayed'aeeds.u Plant Pk 'eio1o~ V 10,
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Hert ig, P. 1932 'tDie khns tliche Er eu Ufl von Mutatioxlen und a h
theoreti$chef and praktiaehen AUSwlrkungGno" Zeitschr. m ci.
Abs -..u Vererbun s~l hire B 61, H 1: i6
Horlaoher W. R. arid Killough9 1).
`f
1931 "Radiation induced var
Lion in cotton. Somatic chanaeS induced in COQ ypium h r utum
by xrayin seeds." Ter V. 22, No 8S 2!362
?1933
radiation. " Amex..Nat . V 67 w 532434
29si hirSUt by
1ShiJima, K. 19 rr n artifi ia1ly induced mutations and poliploid [sic
3
pant; of rice occurring in sub equent generations.' ac.
.Acad. yo V 10,, 38439
Jacob,, '. 19 tIXpray studies in jute. 1. PreiJ1.aary observatiofS m'
Sc. and Ou.tu V 9 (U); 42403
Johnson, E. L. 1931 gEffeCt of .. irradiation upon growth and repro's
ductian of tomato." plan ,o1O v 6: 685p 9
:L3::5
t1 Progre $ s eve mutations induced in
The influence
New ~hr -_L$ 32:
,1 '
V'
i
Xrad i a t iOn on Atr iolex hor teniu L.'
2973Q7
1936 g~]Lffeets of Xx'ay$ upon green plants
Biolo ica effects o Rash ova
Ch 29 in Du?gar,
Katayarna, r. 1913 tIploid formatiorn by X?ray treatment in
Tritia'um monOcoccum." 4 i V s 235237
Knapp,
193 tiUntersuchungen tuber die ~ rkung
von Rntenst1ah1en
an dem Lebermoose phaelcocarpus mit Hilfe` der Ttradenanalyse."
A-
eit$clzr. 3.d. Abet. u. Vere~b s'w19hr V 70: 309*31.9
t co
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a
Lmsbauer K. 1926 "R ntgeno1ogische Untersuchungerr an o 'en and
}~axinen. tt Fcrtshr?. Geb. Rdint enstr. V 3Li.r No 1-2; 241.8,
26287
MacArtur, j. 193h rtX.ray mutations in the
V 29 No 2: 757
tomato . t'?urfl:exed
MacKay, I. and Goodspeed, T. 1930 }1The effect' of X radiation on
. cotton.!' ;cnc No 3 V 71, No 1371; 6
Mal, E. de 1933 -1Muta tions sswohi als Modification dutch 1Wntgen
bestrahiung land die ~Teiungshypotese ' - Ce1iu1e ~;t
F 2: 11-162
Mulier~ n U. 1930 Radiation and genetics."
220a251
T
nr.t V 63
'oeth1in, I. and Stubbe~ H. 193 'tuntersuchungen Aber experir n
tefe Aus1tsung von Mutatioiion bed, A;nt1rrhariurn rna jus L."
Zeitschr. end. Abst. VererbLmgs-lehre V 67, H 1s 2-172
Oliver, ? c. 193a-~ ttRadiation genet CS 1 ' ' ae uu r . R~ cri w of
'V ,, NO
: jo L 4Ou
Rosen, G. 1$12 Hpiumovum muta~.ons induced by X-rayst'
HerecUt V
?8:. 313
33 .
Sapegin, A. 1930 ttRontgenMMutationen beirn Weizen."
V 13, 9: 257-22
M
el ter.
De!
Stadlex, L. 19.26 t?Genetic effects of. X, rays in maize." I oc. S at
Aoa? V lh: 62?.7
//
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Stadler, L. 1928 4tMutatjons in barley induced by X'ray$ and radium
, eno4,l, V 68: 186-187
1928 'tT be ra to of induced mutation in relatjorr to dormancy
temperature and do gage. R1 Anat. Rec a x , 97
1929 ttChramosome number and mutat1an rate in .wens arid Tri.
t ruin r' Proc r Nat. Acad. Sc.
15 87688i
Stadler L. 1930 "Some genetic effects of X~rays in plants."
V 21: 3 19
1930 "The frequency of mutatjon of spec:Lfjc genes in maize.
.I ec, V 17a 381
1931 "The experimental modification of heredity in crop
plants. 1. Induced chromo; omal irreulartjes,E' c 4~
V s 557~72
1931 ttThe experimental modification
of heredity in crop
plantsb a. Induced mutation." Sc. A r. V ltd 6i.661
1932 Udn genetic nature of induced mutations in lante+"
p Prot.
~.r
I . Cpngr. Qcnet. V I; 27 28
933 frd~ the genetics nature of induced mutations ib plant,
2. A haploviable dof'iciency in maize " o a Sba,
1936a
aduoed mutat .ins is plants 'I Ch !o in Duggan Bio
la 'ica1 effects of RadiaUon
/5_
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Stubbe, H. 1930
C ntersuchungen tuber expermentelle Aus1sung` van
Mutatlonen> bei Antirrhinum fl jj'us.; 1. Vnrsuche mit Rontgen-
strahlsn, ultravialettem Dichte9 Temperaturschdcks end Zentri'-
fugierung n." zeitschr. a.nd. Abst..Vererb s 1ebre V S6:
I
38.
1930 '1Samen'- and Keimiingsbelmnd1ung mlt R ntgenstrahlen and
Chemikalien.ll /bid V 6: 202-232
32 "Die Erhhung der Qenmutatonsrate each Rntgedbeetrahl
u , Destrahlun mit ultravialettem Licht, Ternperaturschock
nebst cinigar Berkungn tuber die in diesem Versuchcn
i_n du lertern Variationen /bid V 604 b7L 13
tubbe, H 1933 "Uber die Dez?iehung zwischen Dos un Germutation
rate ,ch . Rontgenbestrahiung m1nn1icher raen.11 V 61x:.181u24h
1935 1"Uber den Einfluss artfremden Plasmas auf day Konstanz
der Gene." /bid V 7Q: 161-169
1937 "Der gegenrtie Stand der Strahtierentik. " Dle
Nato 4$ser $chafte No 3O31: 5O0 O6
1938 r=Oethutation (Handbuch der Vererbun swiasenschal't')
Ternovsky, N. 193 f 'gebnisse der Versuche kfxl t1iche Mulra ionen.
bei elnigen Solanaceae zu erhalten. 't Genet V 17 No /6
9, 6
Young,! . A. ? : 19IQ : "white; Ewer character from Xray treatment of
tomato seed.!', Journ.Heredity . V 31, Na 2s 7879
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Chapter. 3
I TRACELLUl AR CHANGES
n 1897 Lopr cro's article: ?'tThe Anions of X-Rays on the
Protoplasm of Living Vegetative Cells' appeared. Lapriore' s ah-
servations indicated that X~rays accelerate the flow of plasma
(cytoplasm) in cells of yallineria r a raliso then the source of
rays Is removed the protoplasm of the cells regains its normal'
s-b to But if the irradiation was of an hours s duration ;harmful
effects of X.'rays manifested themselves The protoplasm still
continued to move, but it became granular, and numerous vacuoles
appeared in it, rid it acquired a yellow tinge m . Several years later
(in' 1902) Sept repeated the experiments of Lopriore on the stem
hairs of Cucu~rbita Pegg, and also on the stamen hairs of Tradescantia
t
virinica and on the leaf hairs of Tradeseantia sello`.
The hairs were ;irradiated in a humid . chamber after being
separated from the organs on which they appeared', Observing the
.q
mnve5rnon~i^:cxrien 8 of the . plasma -i'n ?n the . i?rr_ada4ed hairs, S~ ..~,..,3 hairs ev _~_Lekt ~_ y _exmi......~ ~ nC?d
mo
'.det
that not only is the movement of .the plasma accelerated due to the
action of X?rays, but that the movement is retained for a longer
period of time than in control hairs, Irradiation for periods of
1. hours hover caused dyiaig ;off, the beginning of `plasmolyss
and sometimes deforrat~.an of the entire cell. The cell could be
returned to its normal condition by placing it ink fresh water.
PhasIfla '
1y~.s set , in much more readily if `s c: : the distance from the (X-ray
tube was dinished, This caused Sekt to assume that in these
oases some other rays aIoaffect the cells,, He became convinced
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STAT'"111
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of this when he setup experiments with spirogyra cells which
in general, are characterized by acute
sensitivity to electric
currents. GJhen the distance from the tube is greatjspirogyra
does not react to the rays, but when the distance is diminished
to 10 to 20 centimeters plasmolysis sets in within the cells in
to 10 minutes. After an hour or two the threads of spirogyra
die. Sekt also set up interesting e cperim .pats with irradiation
of M osa . u c and Oxalis 'cornicu1atao A.t a distance of 2S -
30 centimeters frog' the tube after only 10 nnnutes the leaves
were observed to begin fold ng? After L minutes all of the leaf
lets folded and the petioles arched, if the source `of the rays was
removed in 20 to 2 minutes the leaves would _resume their normal
position. Sekt explains this phenorrienon by a lowering of the tur'
gidity which, in turn, depends on the specific effect of X=rays on
the protoplasm of the cells,
After a considerable interval Schmidt' s work:+Experirental
Investigation of the Action of Small and Large Doses of X"~rays
on Young Cells" appeared in1910. However these' investigations
were concerned only with the irradiation of sprouts, and not with
cells, as such.
Williams' experiments (1923) are very interesting. They
paint up the significance of selection of the proper objects for
X-radiation. In order to obtain quantative data for this action
it is essential to have identical materials, to irradiate cells
at the same stage of development, to perform the experiments under
conditions as identical as possible, and to use homogeneous rays.
As material for his expernents Williams used the skin from the
upper surface of petioles of 5axLfra a unibrosa where, as he knew
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from previous experimentS, cells are all in a mature state and
equal in size, I3esides these cells art completely transparent.
and the circulation of the plasma is retained for a longtime
( L1 hours in distilled water). Williams used special doses of
X"rays in hi.s experiments and in order to obtain more hornogene
ous raysg filtered them. through an alwninux plate. in order to
assure identical external conditipns he conducted experiments in
darkness and at identi?ai temperatures. The skin was examined
microscopically at definite i tervals after irradiation. The
.examinations were carried out under high magni f ications of ard1n
ary microscopes and also with the aid of ultraJn-Lcr?oscopee The
first change brought about by X-rays was the increase of speed
of the 'circulation of the plasma, as could be ascertained by the
accelerated movements of reflecting particles, If the time of
exposure was increased the Brownian movement of the particles
increased. Since temperature during the experiment was held corgi
scant and since the size and optical properties of the particles
remained the same, the increase of the spud of movement' of the
latter indicated a change in the visoosity of the protoplasm.
Stall loner periods of irradiation cause a decrease in the speed'
of the movement and finally to its complete stop, if the action
of X-rays is extended beyond the time required` to accelerate raovew
rant, other changes in the plasma . take place. The protoplasm
begins to withdraw from the tells and, in addition, its pink
color becomes less intenso, It was possible to a sstune that the
pigment was being broken >down however irradiation ` of aqueous
solutions of the juice of red beets does not confirm this as-~
su ii , n~ Apparently, the di colorat~.an i s due; to acids which
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are liberated as'a result of the change in viscosity. An addition
alfactor he passage of solutions from the cell thus dIDn.nish-
is t
irn ` the amount of celluiar juices. When the time of irradiation
was extended the speed:;.of the circulation of the plasma was re-
duced, as has been pointed out earlier, The surface of the proto~
plasm became zno+e and more irregular and son cells became strong-
.~ ly va,auvlizede in order to deterfdfle , ieher this process ?s re-
versible 1 llia~ns transferred the cells into distilled water,
~iowaver, the process of contraction (wrinkling) of the protgplasif
'hued in the water, consequently this process was not revers-
conto.
ib1e
Actually the experiments of Lopriore, Sekt, and Willi s
establish the change of visosity as a result of the effect of
X-Sys on various vegetable ob jectoe In 1923 Weber set up special
for the mea$urement of viscosity in protopla m of cells
experiments
which had been exposed to the action of X'rays. The first experi-
rnents were set up with Snirogyra`r The dosage used was `170 iiolt ~
knecht units. After an hour the' ability of the
hlorophyll` band
(rtbon) to settle out under centrifugation was tested. After
five minute period of centrifugation the displacement was identi-
cal with that of the controls. Consequently, Weber did not suc~
d in determining a change in the cytoplasm as a result of the
cee
primary action of X.,.rays.,
However when the five minute
centrifu-
gation was performed 0 hours after irradiation a remarkable di.f-
snce in the displacement of the chromatophores was observed.
for
In the irradiated 'controls of spirogyra the displacement of
chromatophorrs took place in 88 percent of the cells, while the
X-raYed ehron.to hours were not displaced at all. Another experi
I
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a1 irradiati9T of threads of spiro3 Tra was per
merit with jdentic
n. :
form?d 2!.' hours after irrad~.atbti QnJY 1~ percent ofthe Celle
.
hrar~oto rscres shale ~.n controls 8~
showed csplacement of the controls B}.
s lacemsnt. ~$ hours after ~.r
percent of the cells exhibited
percent ; dip
radiated and un?rradisated cells remained: the `cameo From these
~.
hour after Irradiation there is
experi~ents it f ol1aws that one
no change in the viscosity 4f the plasma,
this plant, despite the fact that they are
hours
the ~.sco5~ty Of the irradiated cells increases. Consequently
se~ ?on of T~. ~ay is observed, ~Jith the dose of ra~rs
cnnda~y action ~x
ra were rep
d no r nrphuiagicai changed in spirO
which was use
-rea1ed and further e ` erllT ents were not su.ccessf ?
his second eriment shoots of hasea
Weber selected for
q . of ~~
~.us r~u~.ta.f~.c~s, ~~~ien he irradiated them dth a dose
Hcltzb~cht ana.ts they did not l11aTl,a.f6st any changes in v~.scc~5ity
.
~.anges in the ~iscosity of the prate
1eithex~ did Weber find any c ?
.p ~,asm. i.n the roots of
extremely sensitive to all kinds of radiattof. From this Weber
concludes that changes of va.seoszty do nnt' take place in living
~z~.t of primary action? Weber considers cells as a rC~S
vegetable ,~ ,
1e since the primary action affects not
tl~.s quite u~~dersta~dsb
the cytoplasm but the ka op~.asm whiff Carries a different oleo'
.. ~'
trice/ charge. He concludes that the viscosity of the nucleius
chaxa;ges from the fact that the nucleolus slip ! aut of it under
cc ntrifugat ion.
he` Japanese scientist lCoriru. o published several works de
voted to ix~.ve~tigatione of cellular changes induced by X..rays.
,.
Actually these were to first detailed cytalogiaal studies`
ar ter 2L. to L.$
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In his work in 1922 in his investigations of the effects of this
type of radiant energy on the root tips of Vicia he demans?.`
traced changes which took place in them. Subsequent authors only
carried these studies further and in snore detail. Root ce118
raised from seeds which had received 50 Ho1t kn cht units maaai.?
felt{d a multiplicity of nuclei, increase in size of the cells`
and. nude 9 a vacuoi zat..on of the nucleolus arid of the c rto .
p1asns an increase in .the number of. n~cieo1. and a dinrLnutton off':
chrorrnatin material, in the prb1orn some cells were found to be
in a condition cif Karryo1ysje and others in the stab ofp
rrosise In his following work (i92L) Komurm investigated changes
at various intervals after irradiati on in the same object, These
observations showed him the succe, s on ( i, e. order:) of the pheno
mend which he described in his previous work, One and a half
a W
hours ai'tor az1radatian
:~
~
l,
?~ _
~
... ,,.
acuo
;
za
tolysis, and irregul.ar. mitosis.
observed. Afez& hours degener=a
t on othe nuclear membrane, pynosiss karyolysxs, and appearance
of abnormal two--nucleiec cells was observed in addition to prey
viols symptoms? After 9 hours gigantic nuclei and nuclei w?th
nu2nerous nuCleali were observed in addition to the other symptoms,
In his work Ic ornuro 'calls attention to the similarity of these
changes with those of maUgnant tumors. This siini1arty is brought
out still more strongly in his following work, in J.92 ,' in which
K omura investigated` the effects of 'X~rays of various powers: Seeds'.
of M' , 'I
of the sarnv variQty Roawere soaked in ;water for
?
72 hours n& f ter irradiation by doses of 20 to p and ..
knech `
were planted in sand. Their ThQts were fixed after:.
owev~2" 1 x
~' ~ he did not observe and significant, diiferenoes in
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action of these three doses. )'at he did describe in greater
the
arise in eells as a result of irradia~
detail those changes which
t an es the extreme rarity of mitosis and theitic, nao~.~general ch
'
m,ult dispersion of chromosomes through the eytopla r~.polar~,ty,, or are chax n addition, the withdrawal of the protoplasm
?acteri~~~.c. _ I
from the cell wall , (l.xi rich case. the cytoplasm becomes `eondfnsed
or ra d) was also observeda The author used thesroughly roughly vi1ized) was also observed The author used these
objects to study the formation of cells with two nuclei and 'showed
than they can arise by various means a 1) by c ons tric ti an cif the
nucleou~, 2) by budding of the nucleusi and 3') by as,metrical
~'
mitas for the formation of polynucloar cells ( those with
~.s? AS : t.
more than 3 nuclei) they arise as a romalt of inhibit1an (or
~'
depression) " the development of the cell wall in conjunction
or
wa.tll one of the fa ?n~; processes 1) tripolar mitosis, 2) a
~,la~.
division of one of the cells in
a trinuclear cell, 3) the exp
clusion of some of the chromosomes or parts of the chromosomes
front the divjsian of the nucl(us in the telophaSe and )4) the
improper distribution of chram~sorfes in the anaphase K amuro
w.
the formation of bi - and palynuclear cells by a lower
explains.
In of the viability (i.e.' sen17 ty) of part' of the protop1a`
abilit of the protoplasl'rr to increase suffjcient1 r for
a..er thea.n ~'
normal cdivision.
in
In addition to the above phenomena he observed an increase
the s and nucleolia eccentric placement (position)
, ~.~e ofnucl~~.
of nucleoli in the nuclei with the separation of the chromatin
material from it vacuoliza tian of nucleoli, and also the
p"
ping out of 'n,ncleoU from the nuclei. , Occaso`ionally gigan'ti o
nuclei prase which willed the entire cell, ls they took hemato
pigmentation strongly and heldseveral 'pale nucleoli. All of these
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phcnarnens were aecoman%ci by pycnosis and karyolsis. The increase
arid a `rayl ke s truc.ti*
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7
of the size of the cells n~.cle? and nucleoli, the hyper and Pe
chrorntic condi "tion.of the nucleic according 'to Kolmzro, all indza"
,
de peneratiV0 or senile condition, of the tissues. comuro
cafe a_
thinks se phenomena remind one of the conditions thinks , that all t~~e -
in the cell: of `naligi a t turnOr8. This reseinblalce`. of the change
he confjimed by bus experiments of tho actionw n:. X rays, on the
a
~.a and
one hand, and tar, on the other, on the roots of ~'a:c ,, -
$a~ vf-
finally in has fourth article .(1928)
P~ sure (~.~~7}~ And,
KOn~iro poits o~t the appearanCe of t..~mors on the roots of Vica
w-M-
Ends cells
~,one In. the cells of the tw~or one E
:.:due to lrrad~:at}
showing xaormal cell tvisLon as well as sells which are in . state
of dacompoga.tion. This 'observation shows that unt l their death
the cells of the tumor continue to growe it is remarkable that
side roots develop completely normally without foriiing any node sv
fn 1921 ' 1926 S trange lays and Oakley observed shah
sin
the celiular tissues ir~iiediately after irradiation or else' 80 rnin~
utes latero In the first series of experiments the first abnorm?~
ality to be noted was the apparance of a granular conditt.on of the.
chromosomes in the metaphase and anaphase. As the time of 'action
f the Xrays was increased a greater number of cells showed, this
the granular changes of the chromosomes became more
abnormality, fri/
and finally some of the chromosomes dander fragments.-
d~.stir~ct
zatian. Under a more prolonged period of radiation ( 60 minutes)
. , ,
the ayteaIso becomes 'granular .,`and less dense. At the same
time the outline of the. cells mangos and becomes abnormal, and the
y
oytoplasnt breaks nc into separate parts. In other cells the
nuc1e us and cytoplasm suddenly' ;fall apart
I
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Ii?
This ra`rlike structure arises from sma11 areas of the
appears. ~
the same time some of those areas contain large
protoplasm and at
of n~ac1ear material. Fzna11y the ceU8 become so
or small_ bits
disorganized only shadows of dispersed fragments of the
disorganized that an.~ .
ve aa,nb Sc metines even they vanish and all traces
and , cytoplasm
pus disappears These c xperimelt5 $hOtJ
of the .. cell and. the nuc~.e
th rer~rkabl,e clarity the growth of changes as doses are increased.
these experiments fail to show whether the action of the
However
rays effect ~.rn~~z?,d.~ ' ~ a t~ y' or whither there i~ definite latest
` takes ef.t.
periado In o clarify this questJ..oXl, the authors Iran d o red
order ~o the irradiated objects
of ` the Culture into an incubator for 30
a
minutes Art e%aa.nata. -an of these cultures revealed that a period
is essential in, order to manifest the action
01 iS to 20 ~.nute ~
Q?:
observed the
ocrn cke after irradiating roots of - ~.
appearance of waviness on them which he a crib c; to Internal diifer'
antes in tension, Xae rflacke traded this down to an enlarg eflt of
the transverse diameter of the cells in the parenchT1Ta of the root
wall (bark) 0 'lhe folds, wbich were observed in the external parts
of the bark, changed the direction o1 growth of the cells. The
which were caught i.n the corners of the f old
cells of the epidermis.
observed the presence of binucleand
were crushed. The author
pclynuclear cells in the periblem 'and plerome? But Koernicke did
not find any injury to the c~rtoplasm- So that the earliest X"ray
.
biological experiments established that the destructive power cf
,
X??rays is proportional to, the dosages used.
The majority of workers who investigated the effects of xrays
.
an cells performed 'their experi]flents in such a ?raanner that primary
.
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efi is of X.rays on the cells were not distinct from the indirect'
effects which depend on the reaction the organism. as a whole.
Wi th this consideration in rdnd~ Strangeways and Hopwood used
the embryonic tissues of chicks in a drop of nutrient solution
for their experiments in 1926. These cultures revea end a wide
zone of cells with 20-104 mi..c mares. The division of the
unirradiated cells requires under these conditions from L.4 to
minutes. A
whole series of experirrents performed by these authors,
indicate that the dinxinut on in the number of cells undergoing mi~
tosis which is found 3O minutes after irradiation is not observed
immediately after irradiation, When stronger doses of X~r ys wee.
used the 'authors observed fra maentation of the chrornosanies their
separation ( or delay) in the anaphaseA and, sometimes, fra,gnentation
of the cells, When the dosages are increased the nurnbor of abnor.
mal cells. increases and also the number of bipolar and multipolar
cells, and then the nurtiber of fragmented cells increases, In order
to establish the reason for the ? rinuti.'on of ~dtosis a whole sere
i.es of experiments wa; subjected to detailed exam nat on, ` This
examination revealed th.t when the strongest of the doses u.sod by
these authors was applied';
ceUs which were already in a state
of mitosis before irradiation ('even if they were in the eery earl
prophase) would complete their diTisionConsegUent1y9 these
observations indicate, that the diminution of the number of cells
which undergo mitosis and the absence of cells in mitosis is dues
not to the injury, death, ox decompoeition of the cells undergoing
rriitasio,~ but to the Inhibition of development of 'those calls which
had not yet entered or had just begun mitosis a b the tune of f*
radiation. This explains the apparent contradiction between the
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conditd.ons of the ; cultures observed' immediately after irradiation
and those observed SO minutes later.'
Tn 1927 Bersa traced the action of X'rays `on cellular elements
.th particular care, By means of a comprehensive series of experi--
n?ients he established that the greatest nimiber of pathological mu-
tations can be observed after 36 and t8 'hours.
A careful. ex mina
tion of these ~ tose leaves the irripresslon that the chroraat n has
undergone substantial changes: ,this chromosomes tend to contract,
to stick together', to fall apart, Those s's indicate that the
viscosity of the piasrna has diminished and that the surface tension
has ncreasedb From this 'stems the tendency towards the stretching
4f the threads ( the bridges that form), on 'the one hand, and the
s trivi
towards ronndixl out into drops, on the other,
In 1927 Pekarek approached this question very thoroughly in
his artcle.
He turhed his attention to studyizi the effects of
X-rays an ndtosis, their nwahors, and on chromosome changes Howe
ewers like Bersa, he considers than the primary effects of Xwrays
should be sought In changes of the 'colloidal condition of the
plasma,
Nadson and Rochlin have performed important works in this
area (1926 and 1933) They examined to cells of the irradiated
epidez i3 of onion scales in a living condition nediate y after
irradiation, after a half hour, after an hour, end so on up to
L,,o hours, The first consequence of irradiation, that the authors
were able to `trace, was the increase in the rate of protoplasmic
movement, However,, a deprae,aiosi araan sets in, this is manifested
by the
3lawing down of the movementof fat lobules and chondrioi-
f q
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sales, and in the weakening; of the T3rownian movelent. This, in
turn, indicates an increase n the viscosity of the pxotop1a ni.
Finally, the movement of the protoplasm stops altos ether, after
an hour the threads, of protoplasm disappear and the protopian
collects in the corners of the cel. At the same time the $oto-
plasm, which was originally fluid and bse ceneous ~. becomes murky.
T is indicates a weakening 01' its dispersion. At this tare the
nude us also begins to change, its granular structure becomes'
mare coarse, and the outlines of the nucleus and the nUc1ev1Us
becorfi more distinct. Tnen fat droplets appear iui the proteplasxa,
`'heir appearance indicates the breakdown of the ipoproteide u:er.
the influence of -rays, Lipophaner psis take s place. A,pro d.-
mate?y at the same time'( i.e. 2 ? to 3 hours after irradiation) va
cuoles appear. Twenty-four 'hours after irradiation the plasm
becomes pale, completely transparent, and watery.
At the same: time the vacUalization becomes core intens e, the
cr,
structure , - ~ ys 9 if after irrc:.diat r n off' Turgid bea we cry them c t
and by thia meaXreduo them to `a .lat ent coact tion~ arid th h o
them again n oaurie thei to grows tI n the' action of the rays will
Tta lif t iteeif 1 aver. Th1 definition of a latent condition was not
acoepted by eyeryO e b What i u .~a 1 iy understood by trxe is tens
period to i time rrxich el pee between rracUa ion ; the f1ret
? f n d this interval t shortened n ore and moron In his experi ent
1; eel .f set o t -tIo arw&ly ze in more detail the previously ob erMved
retarc do of the growth of the plant. In thin exper tent eeee
l hhY N\w d
of V w:re eubjcoted to ~rellin in. boiled water for 24
..
4_S - hour~3, after wi`lch they wrc` placed in glzs diohe filled w&th
water 3o +ked ootton:. Alon de the sprouting seed a aea plate
with grooves wac., plaoed in order to observe the direct growth of
the roots by Meiasi er p o. method (which la bayed on to ntorference
of ii ht wavea of equal length). Tht r thud permitted the deter
minatton o leh th) Of 0.29 .' Before rradxau nn (.e rea mre anta
by ea r ! riethod ~ .o ed ) to roots rew vary qul ok y ~ 0.29
,' sec ade~ i1hsn the pr;auts were irre4iatea twz e by doses of
' ` rnitc th effect, of the ra w, did not becoin cpparont,
burin ; the next to days the ,acme c108e wa' ad riiritered four ttnee
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and each time inda
l afE r trr d ia t-ofl the, growth. ~ cauld cea, ,
only to be followed by auddtn aceleratton of growth? `rheas
observat{ioni3 slew that 1irediate] r t'to M i 'rad ation the roots
grow much faster than before rradiaUo ~ If th done wsri
rug e.!t
ifre sed to 11
"u its then the roots insteaa ox lengthen:.,ngq
would shorten. The probable ( Xpl .raat1Of c this hencrnenon,
aocordIn to Koren, lies in the fact th& at the t wie of irrailat1on
the ce i s always shorten, but thi 3 e f fect i. maskec1 by the
a ,ultaneOus1y ongoing grctwhd The contraction of the cells during
i.rr~tdjatjan _t apparently t,ho suit o4 pias oiy ? e ? The eontrac~
ton of the protoplast i followed by this cont'aeton of the cellular
membrane at..d a c.thortening of he cel lc is obeeried b In order to
eli:iriate the expansion of the root celia due to absorption of water,
a paraffin oil wt s? uaad which doei not interfere with the growth of
the roots. Thi experiment permita the differentiatlon of growth
froni ex anotAn due to absor t1on of waters and it BhOWed contraction
due to the influence of irradiation. In. addition Heare is experi-~.
ments lay a found ate on for i vegti gatiotc of the
effects of various;.
agont9 on the action of X.-rare. When a root was narcotized by water
saturated with Garban dioxide, it ceaaed to gray end wa
incapable
of re &ct:ing to X--r :ys. These experiments perrci t us to establish
the following picture of th action of X.-raya s first f all the
protoplact contracts, it wtt:hdaws from the elastic cel hcicl' a~acaflT fJO1 Of ?aJa1E with kar, of pos that lid an
odd ,t1oxm1 ahrrmQfurne or part od 4 chrommior $The a etophyte
:t(u3ohcB a totr.I off' nine va: iou~s types.
~ o!i X : ai.TO1& twca ~e iaaa~h'sa ( ~4Oa and 19140b) whtoh
are o ~a etiC ~signi f caraoa
for th5..9 part
of our
c pot3itlol of tI c affsots o.' X-rays, we t ball take only purely
alg
r>a3 t,taa R1 3t9 OV1C LllS Qf Qr m tc okra were exooeed
to .-,rays (1940) j~hc ~ ferti.i zod by noamal pollen. Frou 20
ovu1o;3 of 3ilgl:pint via:?y , aferttie plant cci gip; Ewen
o them 1:'lf d non~ca1 ohx`omc~ ova, Cf~ W a trafaloc t The
tDans7iocat~ on ocour ad hetweE;n B and D y the bre Vary etie8
of P?ts
Pr7~, No 9* 28 a 31
Sizova, M. k. 1936. ttSti'uctural Transformations of Chromosarri.es ue
to the Influence off' X Uadiation of 'Transformed
A,N VSSR~ T 193 i9
Oelis
DUKL
Chuksanova, I.. 1939. ttKa Qt eS ? ; Pollen Grains in Tr;Lploid
` ,s 4 ' : ,lar ,r .~: Dolly , Na51. om T 2 , No 3, 219 223.
...:
t
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Afanassieva-, A. 1936 Sur 1a persistance de 1' action des raen
rev1 cytol. et 'cytoph . T 2, nc. 2: io3
Xf'
1936 "'Die'W1rl ng der Rtntgsnstrahl n aiif die Zefenelemente
in si4clcy
B 2?S N? 1 377-39i
Alberti and Poiitzer 1923 "Giber den Einfluss der ,ntenstrahien
vdmr SOmmez weizen " (1 r cj yj xe rar 0
of die Ze W lung'1 Arch. f- m ras .~' nat u..._ t ra.cl :.
V 100, No i'2: 53w109
1921. ' p thnentalbipiogisc1 o Vorstudien cur Krebstherapie
~ortscnr. Geb. IWntge .. B 32, No i'2: 566!.
I92Li. "Uber den Einf1uss der R $ntgenstrahlen auf die Ze1i?~.
tei1ung"" Nitt. Arch. f. ma.ltreslap. Anat. u Bntwaclu ~e
mech B 103
AIrLata, Ae 19fl "Uber die wirksung 'der Hbntgenstrahl n a die
e e f n ? Zelien "" eit chr- '. R ntgns- .0
Anderson J
Bauer,
G.
1936 "Induced,chroncs.anal alteration in Maize
Duggar r sc.a Bf'c.ts o Radti.cns
19272310
H 191ifBie Ent~tehung von Chrenosomennutationen church
}Uintgenbeetrahlung. Line Steilungsnahrna zu den Arbeiten Vt3n
Marquardt Zeitschr,Botan4 38 (1): 2641
Beadle., C. 1937 t1Chramoer~me aberration and gene ;mutation
hramoeame plants of Zea;. Mays" ' Conga J1 e ca s
caes3ura O ;111)
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Bergonle
z' uItats do la r$.didth ap et essa de f ixatic~n d' une `
technique ratinnelj; +.
G. o Acad. V 13 963.98
Blakeslee, A. Bergner, A. Bat nay . ,, Avory, . A. Cartle d e Jr.
and Potter,' H. 1929 P'An&iysis of Batura Stranenium plants
grown from seeds exposed to X'rayeocc Anat Rec. d
V 1L 281
3ordier,4. 1913 taBioehennische Wig der Strah1en, :sbesondere
der R nigenstr h en, c' Strah1enthe r ie B U: 368
Breslavetz L. 1936 "Notes
Bu1J. de biol.
Burthain G. 1932 'IAn interchange in maize giving low sterility ` and
on eXper'ifliental cytology, 1. The
actions of centrifuging on pollen grains
acted. ~.? V U, No 6x 391.39
change conf guration, "
Broc. Nat. Acad. Sc?
V 18* 3t1.
Caspar, Z9 nWeiteres zur.olg,schen Crund:aage der Strah1en?
wirkun
t Tribandeau9 L. 1906 "lnterpretation de que1ques
S th1ent1apie 1 18 z 17.36
Gateheside~ B. G. 1935 !fXizay treatmernt of anothera ehrorosomcs a'c
. '71331
I937 "Recessj e mutations induced in ? Oenothera bl1 ndjna by
. X-rays.,cr Gcn ,tiG V 19; i3 M 2
1938 "The hearing upon the frequenc`jes of XwraYs induced
interchanges in maize upon the mechanism of their 1n;duction.n
191X'n. of Oenetics. V 36: 21328
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C usen~ R, and Goodspeed, T ? 1926 ttlnheritance In N cotiana
tabum. 7? The monosoiic character t fluted 1 - tt Uni~r, Calif'
bl. Bot. 1J4 6i82
`
CaIlinss G. and Ma weJJ. L. 1936 uDelayed killing of _rai seed-
with Xrays, tt Sc1entre V 63 375?376
--
Coiwell, H. 193 Thethd of Actier~adiata.4z~ and-za
v'Mn T.,ssuss Oxford Univ. Press, London
Compton, A. and A1lisson9 S. 1935 rays in ~ ory and rixn nt~
See`, edition Van NO.strand New York
Condon, N. and Terrill,
H. 1927 "Quantur phenomena in the living
biologi a], action of X rays.tt Journ, Cancer Res. V 11:
32333
Cox, p 1931 "Sen~ib1lity to X-rays of cells in vitro
.ll'ars~~ H ll
ttr9I:+*
Creigkiton, Hn l93L. t*Three uses of deficiency' in chromosomes 9 a '
lea .vf ~ t
?roc. Nat.' Acad. Sc.
Cro er, J. 19 L. "Some considerations relative to the action of
X rays an tis8u$:cells.tt ?roc Roy. Soc. H? Vs 96: 2O
7 ill
Crowter, J. 1926 "The action of X.rays on Cod, idiom Co .
p lpoda,,t
of Hey. Soc., London S. H. Vi 7Olis 39O.L.
Prat,
Delauna3r L 1930. t1Die Chromosomenaberranten in der Nachkomen.
schaft von x'ontgenieierten hxen ether relner Linie von
Triticum Vulgate albiduwn? tt . Zeit. f.. mnd.,. Abattam, a ?.
.,. Vere
bunk, a , a5~35
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Beseauer, F. 1923 s' Zul' E r k1 rung der bioiogischen StrahienW Tkung"
trahlentherapie B 16, No 2: 208221,
192k
giber die bielcgisehe Gtrahlenwirkung" Fortschr. Geb..
R ntge~~str. B 32, H 12,
19328
Buggar, B. 1936; Bio1o ica1 Effects o Radiati n Mac Hill [Sick
New lurk LOndol
BI1ingerq F.' 1936 t'Co1ioids and biological effects of rad tic&t
Nature V i3$ d 1OTh Oia
Brdrnaann, i.. 1936 "untersuahung ttber die bh~ngigkeit der ,RWntgen
strahien ?rom WassergehaLt des protepiasmas." PrtL
V 26 Ho 14: 57~S76
Giacker B. 1932
wiring.
ItQuantenphy Sik der biclo gischen k ntgenstrah1en
Q;t~chr. Pypik. B 77: 6 3c 67
Gldcker, R. Hayer., . and Jttngli, U 1929 'Wber die biologische
Wirkung verschiedener Rontgenstrah1enqua .Ut bei Bo$ierux
3_fl rEinheiten." Str~enthie V 32: i~38
Goadsei1, s. 193? ctThe re1atbn between ~.rayintensit and the
?requency of 'defieieney t the maize endosperrm," Anat. e+c
v :b7 8i:3$2
Goodspeed, T. 1929 ."'the effects of X-rage and radium on species of
the genus .iioot3.ar.
urn. Heredity V 20: 21-2S9
X929 "Cytological and; o Lhsr features af? Variant plants pro-
.
a ~ t c Bof~ ~~:
n'c
N ,
on
duCed from X-rayed , sex' cells
26>
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Goodspeed, T 1930 "Meiotic phenomena characteristic of first gent
eration progenies from X~rayed tissues of Nicotiana tabacum."
Univ. Calif. Publ. Bat. V U, No
18
3093
1930 ftiteritancs in Nico Liana taba um. 9. Mutations fo11ow
M.+.W f+. ?A'N~ 'nweM'+4 'H'~MNar4nlouCVi .
trig treatment . with X~rays and radium,"
1930 '1Occurarence of triploid and tetraploid indi dua in
X~ray pragrenies of
Nicotiana tabaouxn. it
mxx~anr~n..~,uw..a,?,.~.w,x :.;xv~amn~w7..wvrc*u..
Bat. V 11, No 17: 299'3Q8
Univ Calif. Pub ?.
1931 d1Die Bedeutung von quantitaativen Chromos menver nder
ungen."
Naturwssenschafi ne L76483
1932 "Cytogenetic consequences of treatment of Nicoti~na
tabacunn species with X rays and radium." Svensk Dot. Tidscrift.
B 20,E No .? ; 7 163
1932 "Chromosomes unbalance anal he asynapti.
Genetic Congross, 67.69
1936. "lnducod chromosomal alterations.,' In Duggar s ia-'
4 ieafeos aaadatiori: 1281?1295
1937 "Significance. of:cytogenetic alterations induced by
Fu i jub.s 961.966
si.
induced Ndcotiana by X.radiation.It Proceed. o tb.
.rrw
;high frequency radiation in Nicotiana species.
.,
nc~sa haploid
condition aS`
Cyto~,pgia
`rho occurrence of a Nico1iana g1uti~
Prat... Nit. USSF~ l st32Q1
I93a ?N Lure and signf icano~e :01'. structural chromosome ante
Goodspeed, T. ,and Avery 1929
rations induced by Xwrays and radium.'1 C o1,ogia V 1 No
3O8328.
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Huskies, C. and Hunter 1935 rtfeots of X"radiatjpn
on chromosomes
in .the ml( A 1
ar
f
r
u
r
u
p
i11j
es o
T
e
ect
, " ?roc RO .. Sn9.
a~~w B
117 22
Husted, L. 1936 "An `anasi of cbrorpsomos structure a
behavior
with the aid of X-ray induced rearrangements "
V 21: 537 553
Geri
Ishijt K. 193h "fin the arti~'ic lbw i 'iced mutations and
_: po
ploidy plants of r.ce occurring in subse uent
~, eneratjons
Joseph and Prowa ek 1902 p' Versuche ~Iber die Einw.ir1c g von
R*ntrcn
strahlen a elnige Or anismen besonder, auf
deren Pslna
tatigkeit. rt
a'ayania, Y. J93L. "Haploid foniiation by. ra
Xys in TritiCUxn i11orioa
"Nwa+yw"'h'Iw4ar.MCe'rWR mm;w!FiWM
coccum.t C,
?__. U$W411ArM!lb.M ~M
old v 5,, ` too 2. 235.237
1935 On a chromosomal variant induced by X-ray treatment in
rri~ticu~i ?ro
. Duo. Acad. Tokyo l:'Uo fl
1935
Karyagenetcs studies in Xwrayed sex cells and their
derivatives j Triticiam monOCOCCUm
Univ. .13. 333362
~Journ Co ?l, A r. Imp.
Kfludsan 19314" Giant plastjda ; fer `ram Xwra ed s 't
~' pores . AA~?le~',,~
n~ ?, t V 21; 7 2.'713
h .
19)tU rf
x,. Permanent changes; of chloropsts induced
the a net~ph es of POiyp1oidj f
um aureutn. Pot,,,,? G;. V 101
. No
721.756
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X-rays.n
 Declassified in Part - Sanitized Copy Approved for Release 2012/03/20
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b
jcOrniche, M. 190 tiUber die wirkung von W ntgen- and di m '
deut th
atrblen auk' pflanzlche Gewebe and Zeller t!" Ber, d,
both e. $ 23, No '8:
i9O r~Weitere untereuchungen tuber die Wa DID von R~ntgen-.
Radiumstrah1en auf die Pflan en." Per. d. deutsch. bot. Ges'.
R 23, 0 7 3214333
Komura, H. 1922 ttPrelinary note on the cells of Vicia faba modi~
fled by Rentgen-rays and their resemblance to tumor ce11s."
p!tani! Mag2.xa T Q V 36, a 2L.
icamure, H. 121. E ici "Cyto1o ical and physiological changes in
Vida faba irradiated with Roentgen rays." Bat. G a. V Lth6
-.- -----,.
1925 "The cells of Vic baba modified by Roentgen rays and
their resemblance to malignant tumor cells with the biological
observations of to no r$. tt Ja an. Journ. of Botany V 2 ` Ra
l3 l! 6
1928 9 ann man der tJog. 'Radkern' der Plaa"mazellen genetiseh
alb em Gebilde sui generis betrachten?" Proc. imp. Acad.
u SGG* Q2
1928 "Uber dada Unterschied; i$chen den ausbe wrahlten
Vices
Laba ?Samen hervorgeg&igen s d. t' Prac. Imp. Akad. s had 14O7
28 n Ubar den Ort der ira Wurze:.spitzen von Vici fba'
gebi.deten Rinbgengeschwulst. tGenn- N?' onsche'Zeitschrft
i ebf orsch. V' 22:
khovSky G. 1938 r.' riine de la via. radiation $t les etres
vans Par
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Lea. Haines
and Collins
1936 r~The mechanism of the bacteriaj, action
of radioacti radtions. i?
alpha and beta particle ,?" c y. oc. V 12O L776
Levan, A. 1937 t9Zy"talo sche Studion an Alliuzu Schoenopras m. r!
Heredas V 22 ll28
Lw?tzky, Go 1931 t1Experirnentai induced alteration of the morphoJ
o of chrorosomes :1r Amex. N t Ali t; %i.67
wits G, 19j0 "A c olag.lcal stun of the `progeny o1 Xarayed
ap . capi]1ari .r~ y-~,oa
Theoretical. 2. perimenta
V 11 N le 1a 9
Loriare~ G. 1897 ;A aona j raggi x sul protoplasma dell, ce:
vegetala vivante" La nuova k assc~gna . Refer. in Bow. Ze ;
bi. 1898 V 73 L5l L 2
McClintock B. 1931 ~3Cytolo icai observation$ of deJiciencje M
v~oly. kinwn .genes tranalo'catjon and an inversion Zea'.
a s a rt oe.1 Igo. Agr. Ix .Sta. Rc s. ` u]4. 163: 1 30
1932 ?A correlation of ring?shaped chromosomes with variega-
tbn Zea hays."
rac.t. Aad~. 18:677.68
L933 t ?s association of non4i nalogous parts of chromosome
Zeit$clw. Zell
arc?,,.mikrosec~ .; A V 19: 1914237
19314 "Therelation of a particular chromzmosom~a), element to the
development of the nuc1 o i in Zed, mays. Zeitschr. Zel1forsoh.
mikrasop. Anat. x 29L..328
w...,,M... .- *.* S..&$
1938 ttThe fusion or broken 'ends of sister ha1r chrommmatids fat.
:Lowirg ehroma.tid breakage at 'meiotic anaphases. a 1 ?
pta. Bud..' V 290: X38
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M.Uet, L~ X36 "vexgleichende Bntersuchung llber die Wirkung`.der
R&~tgcn and G axrthaastrahlen auf die lebenda Ze1e ." trahlen? ;
thorn v ~; g tt 1: 48'57
Marquardt, H. 1937 r1f er ` tU I ust usch zwieohen icht honiolq en
Chromosorien in Mitosis . and Meiosis,' Bex. deutsh. bot' Gel.
va 3J39'159
I937. "Neue l ryo to isehe Prdbler and Ergsbni e 3 ~ Die
R ntgenpatho1o ie der. M tole' " ch v .fat. N 12
57292
1$38 1D1e Rontenthoo;ie der` Mitose. 1 und
3at~. B 32$ 4C1482
1i`n
cites
1942 "Zur Anayse rontenirid uierter GhromosomenverMnderunen
d Chromosorenmutationen ;dine iidezi ng." Zeit ohx. Bot.
19L2 cD?e Verteilung rOntgeninduzierter V4r nderungen auf
den Chromo omen von `Bollevalia romans." Ber, deut$eh Bot.
qe: 60 (2). 98,i2L
Marshal, A. 1?35 "The effect of X-rays on chromosomes in different
stage `of mejoei .'
78di98
f C er r, s. V 9~ N i'3`:
1937 "Tlie effects of X rays on cbromosames in mitosis.'"
!4;4? 4~ v 23`, Ne 7 a 362*369
Mars .and Born n 3$ 36 'The ;absorptIon of .rays by spe c and ery
th?ooytes and ibs relation to, the sus eptibiity of tissues, to
X-days. ex.aurz~~ Goer 2 n$h8.
r.aw...w-r.w.-
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Marith 1922 "Die biologic e
Wir
schjedener we11enn~.
de:' ~ontger~strahlen Very
~~ra~lsn~hear~,?ae
B No
lIather, K. 193)4 "The behvjo of meiotic claromosor~es after:
tion? Pt Hereditas' B I9 h d
X93?
558, 6o
rad
"The - VSyjMdTACfnM; ({~s nw~,,w L ~. .. .
!:! doubling
Mather,. Ico and Spans ~ L0 1933 tr
The effeot ok~gadatUpon
ornatio ~h~`omoSo~lea, t: Journ~ c~~ ~sn~~~cs ' 28: 1
Metz,
Mod
core..
Muller,
kV1b,1e relation to the aheno~ enon of ~utatao
, d o V 20: 1590 16
3
C. 19314 'The rode 'c~the o ~hr?mo o
s ~1e sn~a~ ~.n ~r~1tos~,s and
st~ahl
u d die Te,uns~c~thes P~
~?,,, ~~~.1u~,e V 2 2
151?162
. Fi''Qe
da 1933 UMuta.tior so h1 a18 ~e
d~,"'1oa~lo~a dutch
end Hask
kroc. Roy Soc.. i
- 97,I.1t1'
Xwray : dune j nod1fjcat jons of f ~
.
owe
Color ~,n the Petur~ .~~1a ? J ~rn~~ p,~ ~~ret~~,~
V 26 ~ 3L.:355
rats zur. Strahlc gsj8. U
I'y?. 1936 . r~ ,e die 'Bestirrrniung d
_ es
Verhltnsses der`Mutio
theraje.
V 55 N1g 7276
Nadson~. L925 Peer Ph kg ~~kurig der Radjums t~a~lecn
a die
' L
b
e
ende Sbt
usan, n ~ ;ache
Madsen et '.ch14
V 155: 36,386
-r i
3. effet des xaygns our
1
e
t
pro
a~la~
1e na~yau at le ~
chdndrjome; de a `co ,u ,
le v~g'eta~ d 'apres' 1e
ob
ert
0 '
s
j
sm v1' rit;'i
v
V O No
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Navashin M. 1931 ~r A p$l inary report an some chromosore. ante;-
rations by X-rays any Crams.cr Amer. Water 1i~t v '65'; 2325L.
Nebel, B. x.93? ttChronoeorae structure. 12. Further radiation exper-
rents with Tradeecant w rr er. JQ?1ra. 1etany v ` h : :36372
6 H Chromosomes structure.. 10. An X~ray e aer iient A cr
Genetics V 2.' p w,?
isL, ., Sinota, Y. and a , D. "Effect of he fast neutrons
upon plant . U. Abnora1 behavior of mitosis in Vices fabd "
Cyto1og v 10, No 3 L.O421
Far thasarathy, N. 193$ "Cytogenetical studies in Ory ae and
Phalarideae
.~y~yx+kk~+wR'GMwNabf4f~lrlxMYW~
rice (?ry a sativa L.)." Jpurn oaf Gene ic.s v 31 , No 1: 1~hO
f
Pek~rek, J. 1927 "TJber den E'a in fluss der R ntgenstrahien auf die
Kern,und elltei1uu bey. Wurzel pitzen van Vila fa.."
' Mnvn ..... n1MfMfYl u'a
B h 1/2~~33
Perthes 19Q~ '1versuche tier den E fluss der Runt enetraW:en und
Radium` Strahlen auf die eliteilung.s
~? . '3 17 18
Petry,
Deutch mid. Wohen-
1921 uZur Kenntnie der Bedin ungen der biologi$chen
Wirkur der. R ntgenstraW eai," iochem. el .schr'. B 119: 23.L h
1923, "Zur Kenntni der biolagiachen W:kung
der R& tgen-
strah Wig: VOn G d~ttans~jttOn auf d,.lra mpfind
lchk it. t' iocnem. i4tscbr. B 13SS 3 3 3$3
cq
1. Cyto netics of sonie X ray deri tives i
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Plotnikdwa1 T. 3.931 n'Einlues der Rontgenstrahlen auk' die
`k T. 1920 tlExperimsntel etudien uber die Zell- and Rerntei
sonderer Rjck$icht auf Forme Grosse and ;hi der
:Lungmit be
Ui4v.. V 39
Politzer, G. 193 patho o ,ie; der :iiIitvse Berlin
Po lit'zer , u
tionsteilung von Wsizen" PfnZeX1Uchtu B 16, No L:
662m668
It
Pordes., T. 1923..19211. "Der M haniemus der R ntgenstrahlefr"
Ceb. R Mtgen$trahlen D 31, No 2.3: 287.297
Reiss
bloi q
Z knv8kyr
biologic
1e de+s ra on~ s X:
t~ntersuchungen
Cows depi q?
iliey x. 1936
Tr
tIThe effect of X'raya on the chrome;omss of
cantia gigantea." rtologi V 7, N 2122 131?1L.2,
Chr?mosorrien" ~urr~. Cca
Art l1: 99
sa egin, A. 1930 aR antgenl utatianen Beim 1ieizen. tt
No 9s 27'-259
1937 "Effect of variations in mperature in nuclear and
Sax, ic
yell ' di1'isian. iri Tr deecantia
~
No I 21821
x,93
.lkJ iG.
Bhrntrger r 1-238
r6 roro ,A{1 "6957~.wrr4~ !,1 ichen...._
6i1gle "1 f; ~brti ~ 8.
~
1/31
uber die askufg der Buckj and FWntgen strahlen
teilung 9 of 5t~on h B b2
Zell
'ortsehr
192 o1Zum biclogisehenark~~ngrrech~a.smu der P,cntgenetrahlents
Sarah .en hera . ie B 19 , No
Declassified in
Auer.
auf die
Red
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u ,n. of Boten: V 2L
 Declassified
CIA-RDP82-00039R0001 00240012-2
Sax, K. 193$ "Chromosomes `aberrations induced by X,rays.tt Genstics
' 23, No s L 9L ib
191i1 !The behavior of X? ray anduced chromosomal aberrations
,n . u i root tip ce11s ?1 Genetics V ' 26 (h) , L i8 L25 .
Sax, K. and Bnzmann, H. 1939 "The effect of temperature on X-ray
:I,nduced chromosomes a .teat ans.r
Noy 39715
1r~ac. Nat- .Acad. Sc
x, :K. and Mather, K. 1939 ttAn X. ray ana sjs of progressive
chromosome splitting." Jo?nof Genetcs V 37
0 m L834.9O
Schmidt, H. 1O t1Experi~ientei . Untersuchungen ttb r dieWirkumg
k:.einerer and gr serer Rontgens-trahienmengen auf j mge e11en'~
Berl. k1 in
Jochcneghr. No 21: 97297L.
tr,
Schulze, J. 1910 "tlber die Wirk mg `der Lichtstrahien von 280 e11en
nge auf Pfia~z
B 25 r .3080:
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Schwarz, G 1917 'tExperamentell:le Deity ge zu r Theorie der bipla~
gischen Str h en rirkung. Die LecitTinhypothese'. ~t Fortschr.
Seokt H. 1902 ~'Uber den Einf fuss der XStrahlen auf don pf].anz~.
lichen arganismus
Seide, J. 192 t~Zur kennt der bialogs.chen Strahlenwirksng'
eitsalw. t .' .ss. Zoo :a D 121.
Serebovaky, A. 1929 "A 'general sc1ienae,:. for. the origin of gene muta
eras. nor. Naturalist V 8
nzellen
, ex?0:Ldeutseh. bot.Ges. ' D 20, H Dj.s 8793
Dcihef to Zum Bot. zantralbiatt
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Spek J. ` 192b 1Kritisches Ref erat uber die' neusrcn Untegsuchungen
fiber den physikaisdhen Zustand der Zefe thrend der M tase. "
Arch?. Entwickla V 101
Stadler, L. 1929 ''Chxcmeecmes number and the mutation rate in Avsna
and . Sc )$": 8788
1930 ?fRecavsr ng following genetic deficiency an maize."
Pray. Nab. acad. $c. V 16: 7]172O
:931 "The e per:bental modification of heredity in crap plants.
to . Indue 1 chromosomal irregularities .
t
%7572
Sc. Aric?.
Stadier, L. and Spragu+ 1937 "Qantra. is in the genetic effects of
1tra violet radiation and X"~rays a" Scten V 85: 57#58
Stephan, R. 1920 "tJbr Steigsrung der Zellfunktion durch R8htens
verge?'~ Strah1enthera4e B 21
Ston A L. X933 "The effects of X radiation can the meiotic and
mitotic division of eer i n plant." cry V 7 s
N4' 188 s 8126
Strangeways, T. and Oak1 r 192L. "The ? mediate changes observed
in tissue ce11S after expo sure to sot X rays whi Le grog
vitro
rtl
.
prat. Rohr. SOC a B. V 95s 373-38t.
trangeways~ . a a ,rcaad F. 1926 "The effects of X.'rays upon
mitotic cells in tissue , cu Ltures in Vitro." Proc .____ Roy. Soc.
London s B No loos ' 28 293
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Strau a ~.
1923
er1inente1ie tudien ilber gewisse biolo.ische
Strahler1 irkun en# r
Strah.cnther~e B ]1.k: 89
1923 "Ubor don Einfiuss der R ntgena.trahlen auf ondoce 1u re
andifweche1vorg
Taylor, i
U `C]'uI-ranl~lnrt?6~rts~'try~r+ l 16
B B 1~.
sepaa~.u~svssv+w acP
1935 "Thy efects of X rayed medium on living
Al i4 del_
I Conreeo
Vene 2 . ? 1103- 1i9
.i36 " easux rent'
a,
internat. di ?
of X-rays and radium. " Ck14
B a1 Effects C Radiation
Ternov,y1935
bet
. Duggax1 s
'3Brgebnsse der V ereuche knst1icher Mutation
engen Solanaceae u erh,lte;d, " GentioA B 17,f.
149945146.
rimofeeffReseovky,
tone tt1"aC
afeffseav
icr,
and
Zimraer 1935 "weilen1ll enunabh4r igkeit.
von der Mutataons~rar n der nt en$trah1en bet Drosoah
me1anoj- ter. U
yjinN!MM'IThV~*ryA3r~..
1931 "The earinienta1 production of muta`
Biol. Rev. No 9: 111.
~~rah~~rather~, ie
V 5
T8chaeownikow, N 192$ 'tubber die i.rkun~ der B&ntgenetrahlen auf
den feinen Bau der : heberzellen: beim Frosch. " irc;hr~ ~ arch..
B 269:.i6 ??177
Ube F? and Goddard
X'=ra eux lva1
)o 193U II;w1uaz1ce f death criteria of the
aux. th un u
vee a~ the f ~ s Neurpora:i rs Jan? o'
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r~ 17~ 577.590
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Uber,.and Goodspeed, F. 1935 "Micro~jr~eratxon etud1esA 2
calization
f
h
o
as
yio1d ng subt
sances during mel
possible igriif icance in X radiation phenomena. it
V 97: ' i4J.7 1 ..
wail, S and Freel S . '
, ].92S ,der den E nf1uss der Rtntgenstra
auk' das ZeUp;
923 "Tdntgen strah 1enwjri g unc Pre o a .alms. r .sl c ft
f No 1188 6Li
Weaver, E. i9L2 '~A ?cornparrjso i of ray mutations and mutations occur?
ring spontaneous
Missouri.
3.937
Webor, F ? 1922 o'MetIioden der V .scosjttbsbe tmmung dss 1ebend ri
Pro'topiasmas, c' Abderha Lden+ Handb. Bio).ogisaho Arbeits.
-
methade e
Wi1laris a M. 1923 'rO bseratiors on
ceil.
the action; of Mraya on plants
~, cap Ba lLF6o l72.25
Z:n!er, K.. 3 % " Beitrag zur age reach der Pe ebu wt ohe
IWflbgonstrahlendosis und ;dadurch ausge1~ster Nutat
jonsrato,t
trahlenthr B 1: 17928!
Declassified in
in ring chromosomes in maize.1t ~n ~1? ~a '
V i ?C ow ! s Aroh v 257; 850
u~~denieru der biv,ogi8chn
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trthJ.unn
Leipz :
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Chapter
TIC EF CT OF TIC Q NTFI AND QUALITY OF TIlE RAYS
ON `TIS rFANSFORMATION ' OF PLANTS.
( 1~a4j4.tL the e
In this chapter we are not going to spend `much time. in cosy
sidering the effect of the qu. tity of X'rays on pS Q1)giC. ,
changes since, actually, all of the experiments of this kind in
dicate a close rela Lonsh p between the amounts 01' ray used and
the ciwfges that Caro produced, I1 the Arndt ehultze law is not
4
kI confirmed In all cases It can be ascribed to the fact that the ex~-
~
1, periments were improperly et up rather than to the nature of the
hi
action itself,
}zero we shah emine those e ~perimnts wbioh point to a
dirsct relationsb p bctwecn the dosagos and the morphologic...
change5A both heredi .ry and non-hereditary. Although the re n1LLs
obtained ~ath plats are not as clear but as those obtatned, with
the fruit is iJro$ophala)a nevertheles there can be no `doub
=t
: .
? at the present tine, Ghat a direct, linear, relatiofl8hip exists
4
di1
Ir
` i between they dotage and the frequency of mutations0 How the fre-
quency of rntttations varies, ri i.ri ac .ated corn, as dosages are in
creased can be seen from table 33 ;
e Table 33 , on i"oiowing page
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STAT
iii
 Declassified in Part - Sanitized Copy Approved for Release 2012/03/20: CIA-RDP82-00039R000100240012-2
TM3LE 33
The Relationship Between Doses: and Frequency of Genew''.uta
ti o? in coin suds W (baed Qn he data of Stadtler 1931
~
Duration of
Nuirnb r of
Muta~ Percent of
Duration Number Mutate Percent of
Irradiation
cultures
dons
ntatatioe
of irradia` of
Lions mutaticm.s
in rttinutes
era ,ned
uion in
minutes
u t zx^
o mined
6
2
i
0
b.
11i.&
o
" .
20
37
10
2.66 ? 0,83
6
152
0
. .
?
13
9
6.66 ? 2.iL
6
2
1,27?0?96
21j.
167
2.99 ? 1.31
10
8
2
()52?O36
6
113
9
7 90 ? .5
12
i6
2
1,28?0,90
28
17
6.17 ? 1,80
Q, 2?2,iO
30
177
9
5,08 1,6S
16 161 1,22
In Stubb& S (1933) articie concerning the relationship
between doses and f cuenCy of gene?r utatinns, after i ?.radiatl.on of
male gonads (which is pr sent ed in detail in the second ob pter
this book), the
of
results of irradiation, which were 6btained under
three &.ffcrent set-ups., are pxosented. These results shc~r a regular
r2&Y2C f f
rise in the frequency omutation up to ~ ` xxLts alter which
~,
.'~ i t r~t~~r
the fsquenc, f'aa ls'. l
its and then' ri8bs again,.
2W
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In another portant experiment, in which only- the uGatiofis
in sprouts were courted, a sp,.nd r
tween doses(Qj 3OOO act 6100 ;._ .~_ .
T'abie 3I~)
J ABA 3)
Increased frequencies of mutati+ orn in Antirrhin'wai
- .bus '~r, of tr rra da atai c' '~ ma.a gonads
Short-ware rays, independent o.f
length of
exposure
o:c quality of thEa rays
(based on the data or 3tul)be 193.
Dose in Irradiated Mutations Percent of mutations resulting
-'-C rule gonads
30 X3.7
boo 3616
direct rata was obtained bee.
i ts) and the number x rutations
rpn irradiation
:i38 ? 0.136
2;78 0.266
Our experirtzents an the change of sec ix~..hemp ebtained i
y
irradiation w1th razzous dosas of X rays (,see. chapter two) , and
Lutkov' s e'periments with pea and barley (in the same ohaptex
can serve as goad illustrations of this henamenon
p (i. e. the direct
re~,ationship) ,
ity have indicated that the f're u~ncy of mutations " ~ '
is d.xractly pxo.?
Up to the present time experiments for the
pu:rpas~ Q.C irrvehtj..
Ling the relationship be Gween the 'giant
~ ::. i by of the rays and ~ntx~ta
z
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 Declassified
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ports{:)nal to the dosage,
ID)" Quality of the Ra
This question, as others, has its own history, Way back in
1898 only a year after Roentgen rnade his wonderful discovery Naldw
ney and To'rvenih broached the taestion of th quality
~t''ar
1 vensis and Lepidum with and without filters In other
of the., rays
in their experiments, in that they irradiated seeds of Q,onvQ irulus
words they were trying to det irmine d $'ferences in the actions of
hard rays (using. the fiber) and mixed rays,
Perthes (19OL) on the basis of his ecperiment~ Game to the
conclusion that intensity of the rays is more significant than the
length of exposure to radiation,
11
In .1919 K orni: cke.
in
his ex~eraments with Vica
did not
succeed in. establishing any difference in the action of hard and
soft rays or .in filtered and `unfiltered ones,
Then 'experiments of Burp and Robberts (1922) Tdth the
seedlings of oats, whose coJ,ptyles are most scn`sirM tive to rays,
confirmed this (iae. x riLckes findings),
In the same year (1922) Martius set out to determine what
the biological effects of rays of various wave lengths have, when
the energy Is kept constants. Can the basis of his experiments he
came to the canolusioxi that when the amount of enemy is kept
con-
stant asad theaaveenth haness) of the rays is iraried, that
soft rays have, greater biological effect than hard rays, lven
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S
of the same opt pion. kneel (1928) s the first to experiment
. ~ ona~ ayr ~n p1ax~t$s in this'
with the effects of ~cactx.~~ doses of X_~, ?...
x'P t~ p~c~d to
case.. o~~ lentils, In all 18 e ~ erimezlts 'tfrac~tiona]: doses
than dose g3,ven, a11 i atiance. t~ Next she (Anc4)
be 1esS, harrrifu1
set up a very interesting expeririientn 18 hours' after sprouting, the
seed/ ngs Of lentils which had roots '6 nii11irretere long; were c -
y,14 rpdf f
millimeters,
showed that the length of the roots in Group I was
while that of Group I1 was 1O millimeters. In the authors s op:m-
ion this result indicates that the initial exposure to a weak doe
'ii its rotected the roots from the action of the l.ar
dose and that this is a case of protebtion from. X-rays by X rays,
i.e. that .it i$ a rase of X??rayph~riaxis. Th:1 experiment was re -
prated L8 tiitie and each time the sans results were obtained.
vied into two groups, Group `s received a dose
/
of 3o iurats
and 3 hours later a dose of LO ,'unite' Group S1 received
L5 O n .ts directly. Neasurernents, performed s days later,
The` experiments of Glovkor, Hayer, :and Jung ling were ex~
tremelY interesting for their time (1929) Frortl a theoret;Loal pain
of viewa write : the authors, by analogy with the physical action of
X-ray
it could be expected since soft and hard rays` have identi-
cal effect on ion1Zatiari of air that the biological effect would'
likewise be inddpen.dent
f the quaiity o
the rays. The
rant was performed with seeds of the "notorious' Viola ecjuina.
During the year 1714. groups of 8 to 10 beans each were subjected to
the action of hard and soft .:X~rays., The qualitative data for the en-
ixe series were identical : With ` sraU doses, (up to 2e0
'nit '
soft rays turned out t be more injurious, with lar ~r 'dose's bhe
~r
d.ffe?encp tended t
diminish,. so than with doses of 380
9-,
its
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. the effect of hard and soft rays was identical, So that these ex~
perinents, in general, , tended to confirm
of soft' and hard rays,
the 'biological dentityr'
ke (1929 19O) writes that `when X?ray$ are sad
Dar?sen ,
...
weak currents used in prolonged duration have the
on seedlings,
greatest stimulating effect. FIB. xh voltage currants, even when
have depressing (inhibiting) effects, TO
used. for a short period,
have; a greater. stirnula'ting effect than a single large
small dosea
dose (which is equal 'tQ the two small ones)
Long and Karsten also point out that a greater st.mulation;
rte .
a.s obtained when filters are used, than without them. The filters
used the experirrent decrease the intensity of rats of. all .
in wave lengths, but especially that of the soft rays, `so that if a`
stx~ ? ~nf.ating effect is obtained it must be ascribed to the action
of the ` longwave part of the spectrum, while the injurious effects,
to ? shit wages or the stimulating effect carp be ascribed to the
lesser intensity and injury, to the greater, regardless of the wave
length. Since the authors felt that experiments using soft rays on
large amounts of dry . seeds were wanting, they set up an experiment
.:.,
n which soya seeds were `irradiated only by the soft part of the
~._ dry ,
pectrufli. Since soft rays are absorbed by air it was necessary
s
,
the seeds dose to the (X"ray) tube. For this reason the
La place
authors used the kind of apparatus which held the soya seeds under
he X.-ray machine for only seconds after which they were ejected
into vessels laced below and a second batch of seeds took their
lace and so on. Larger doses were ;obtained by increasing the ex?
osure time from. to 2 seconds. When the voltage increased, the
.24S
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intensity of the wavelen;nth (tnd hence of the whole beam) increased'
proportionately to the square of the voltage ? used. The results of
the experiments were rather i nconalusive,o
{
y Stubbe (1937 and 1939) considers that the question of the
relationship between the quantity of the rays and the frequency of
mutations should be resolved into two questionst l} what qualities
of the rays can cause mutations;, and 2) do equal amounts of rays of
different qualities produce equal increases in the frequencY of
mutati ons
Stadler' s numerous experiments with the seeds of agricultural
plants indicated that many rmutatlons pan be obtained by rays of modem
i um hardness, as well as by hard rays of radium. His and S tubbe' s
experiments with soft rays (the sOMcallod borderline rays) using
8 to 10 kilovolts, were equally suecess ,g
Very lord. rays ('Spa kilovolt) were successfully used on
snapdragons by Noetling and Stubbe in their experiments in x936
All of these experiments indicate that rays with voltages from
8 to OO kilovolts (i. 'e, the entire range of the X-ray spectrum
induce numerous muta.ons
The results of experiments whivh have attempted to establish
the threshold. of action on the ultraviolet end (of the
spectrum) have
been less clear cut. The ;negative results which were obtained in
experiments with snapdragons and Da~osca h~:la have incLcated
,~. that
ultraviolet rays can be effective only ' when they a'
are net absorbed
by the surrounding ;tissues.
The results: 'obtained by N?etling and stubbe indicate that
pollen gxains of snapdragons' are `p,artiaularl uitab
Y , 1o for the
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stud' off' the of feet, . of ultraviolet. light (table 3
L
The 'relationship between dosages and frequency of gene
mutations after irradiation of the male gonads of J ntir-
r.~zir urn maws I. by rays of various qualities.
('based on. the data of Noetling and Stubbs, 1934)
dose
in thLlture mutation frequency of
m~ gene
mutations
loo
2 00 92
1.67 ? 1p3t.
Borderline rays 0,015 and
0,019 inxa A ( filter) )04 . 93 8 34' :o ? 2.83
, 800 9S 14 3.71 ? 1x9)4
1600 44 2 L. a)4 ? 2.97
320 3 6.31 3?
100 118 0.93 ? O,68
200 101 2 I.L8 ? 1.20
Soft X-gays 34-70 kvf o0S,
0.06, 0.08 mm 'Lu (ili~ r) 100 1).Q 3.07 * 1 Y 46
800 127 1. 2.6 ? 1.L2
1604 86 1 o.66 0.87,.
1.70? 13
Table 3S aontinued on. f aUori,ng pag
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C
:.oo 93 1.6S ? 132
200 97 2 1,6 ? 1.26
Hard Xrays 1217 Kv! 0.3
and o?8S rnn ou (filter)IOOO 97 i4 s ? 2olb
800 97 3,0 1,90
l6oo 139 1 ? la 63 ? I. 7S
320Q. o 3 5.00 322
In their experiment' the authors dig ded 'the spec tram into
the smallest parts possible. Pollen grains transmit a significant
percentage of both visible and ultraviolet light and it can be
stated with certainty that they also comsune a definite percentage
of the rays. In comparing the effects of unfiltered rays with
monochromatic ultraviolet rays it was discovered that the frequency
of mutations rises sharply when light from the 280 to. 285 milii~
nnicran ( spactral) line is`used whale irradiation of pollen by rays
[ of the 366 mlllinn,cron lire has no effect in increasing the fre~
qu,ency of mutations. From this we can conclude that the threshold
of action (which 'induces imitation in the pollen grains cif snag
dragons) in the ultraviolet area, :lies in the vicinity
of the 313
m llirnicron line,. although it is not possible to establish any of
feet for the 265 millindcron lined In this fashion these experi'.
rnents established the thresholds for action of ultraviolet rays
(in both directions, that is for longer as well as for shorter i5ays
on the pollen . grains of snapdragons.
In 1936, Stadler snd $pra ue showed that when they irradiated.
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the pollen grains of corn by unfiltered ultraviolet rays, thy
frequency of mutations rose sharply ?.
At the sang time we get an anwer to the first `question posed
by S tubbe (see above) " concernii the threshold af' wavelength at which
mutations can be inducedtt ? The answer consists of the fact that the
threshold is determined by the ability of the rays to penetrate (the Y
tissues); and that in order to induce rmitations any source of rag s
maybe used, begin'riing with ultraviolet rays.
S.tubbe `considers that the second question concerning rays of
different qualities is also answered, since in the region of X rays, ..
from softest to the hardest rays, when identical doses were used no
statistically significant differences an results were obtained.
plants this was confirmed by experiments with snapdragons, in which
equal doses of aof and hard X-rays (as well as mma?rays of ra~
diem) induced identical frequencies of mutations In athar.words9
we can consider as established that the inductive action of X~rays
does not depend on 'wave l.erlgth, This conclusion is splendidly con
f.rmed by the table taken from Stubbe (Table 36).
TABLE 36
ind 9f? rays
The action of equal, doses of rays of various
qualities which induce mutations
based on the data of Stubbs, i93!)
Dose
in
Irradiated Mutations Percent of.
Male Gonads mutatiOns' re
suiting from
- irradiation.
3
Table 36 continuer on folioing page)
'G
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(1), (2) (i)
Radium .? 3000:
X-Rays 506ICv 3000
)1ays J.84 v 3000
Radium
600o
X-Rays X06; icy 6000
X":ys
180 Kv 6oop
O
103 31
1937 6a
1536 14?
15pi 56
1171 60
110 60
From this table we can see that when equal doses
used (3000 ".
liits in one
~
case and 6000' in is in
other)
hard ~Days x,06 kilovolts) and soft ones (180 kilovolts) produce
equal increases in the number of mutations (as compared with the
control) It should be noted that the number of mutations induced
1by the 6000 'unjt dose is nearly 3 banes as great as that
produced by 3000
units.
low let us turn to the question raised in 1923 by as to
whether administration' of doses' in fractions' affects the appearance
of mutations. The most clear out experiment that was set up for
the clarification of this point`, was also that oi~ Stubbe who tested
the difference between. the action of rays
of dif fex'ent qua1iti s
:Ln, concentrated doses and in doses which had been divided into `tree
(fractions) and administered,at 2J hour intervals (table 37)
2-70
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.
1;c3,. 0,30
.1.`31 ?O.2
1,22 ?` 0.28
1,89?0
35
3.29 ? 0.52
3?
0
53
of iys are
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TABS 37
The action ofconcentrated and fractional (three
fractions at 2L hour intervals) doses of rays of
various qualities on the frequency of rnu ations in
An?t~arl~zaum rya us hd
(based on the data of Stubbe, 193L.)
Kind of irradiation
Dose in
1rradi.td
mutations
Percent of mutations reg.
and `qua1 ty of the
r
suiting froi~ fradiat1on
gays
180 Kv 3 fractions
3000
1209
2,22.? O,L2
o6 ICv' 3 fraction;
3000
1973
62
1.33w k 0,2
180 Kv concentrated
3000
1036
14.7
122 0,28
06 acv concentrated 3000
1068
t8 .
1+23 ? 0,27
180 ICv 3 factions
6000
114a
6o
3mL ? o,a3
506 Kv 3 fractions
6000
1170
60
3?29 ? 0,52
18o Kv' 'Coicentrated
600o
1187.
)
3,Jj ? 04 5
506 Ky Concentrated `
6000
1337
79
1107 ? C
All of these experiments indicate (despite the indi
cations of earlIer e gerixente to the contrary) that neither the
cIualjty of the , i'ays9 ;nor the fractionirig of `doses have arty e:L'fect
an the number of mutations but that the only signifjcant factor is
the size of the dose (i,e., the number of Xwray units),
This was confxuad by our experiments with rye, These
experiments ;established that
there is no differeno~ between haze
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rclatton to $ta ulation or depreSSlon (Bresi,avets,
and soft rays , in
Nedvedeva, and Afang,s' YOva
i93!)r
One more intereStin~ queS Lion 'remains. This Is the quostion
of the qua., of ene mu.tat:9.ons obtained by exper?~.anta1 nieansd
~..~.ty
Muller nota.eed the identical quaia.ty ref"
:fin h.~.$ very' ~irst works
porLtanc~oUS mutations and those obtained by
exper meat l rneari$S.
ITned kfor all of tcie objects examined. The n erou
This is confi
the action of X?-rayS in snapdDagons
~;e~xe~r~ut~,t~.c~ns obtained through ctJr71 and .~~
n ,ro~u?.1a had all. been encountered previously among; spon
A h
orp
tanaous rriutat? ns, They pnsSons the Same properties, affect all
:~.o
` s and
of the ph Sia10 i0al propez ties of the ox M
ane and all ~
are 1.i1ewiSe usually reces$ivs (few few being daridnant) r
~.
The first indication of dif ferelces in the quality of sponw
obtained mutations crap up in Stubbe's
taneaus arid exporxmontally'
In the first experirnonts it was f ound'
experiments ( 1932 - 1..933).
tho buds the percentage of gene. mu`tatiOns
that after irradiation of
n
in the ve cells rose sharply, in comparison with thc eontrols,
~;et~t~vc G
hash deteX'WLin~ changes in
while the percentage of hoso r atatiofS w
did not rise, In another experiment, wider
the shape of the flowers
irradiation and semi-hard rays, it was found that as the
by soft a~ -
doses were increased the percentage of previaualy encountered mum
tat ?r of new (not preVa.r~us~,y' en.c~ountea^ed)
~:ohs d1m~.nit~h~d ~il~ the ~u7nb
mutatisns . increased. $tubbe, explains tahi s by the fact that the ace'
ton of 1argc~ . doses shake up the structure of he genes to an ex
tent wh?oh cannot aac under normal cv~ ens or with weak do ea
ur
`
(of x. As a oonsequen s after' strong doses,; even genes
ra~).?
f wh iriguiShad by great stabi~,ity, under~;o
~.~~~ ,. In general, ; are d~.st
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mutation,
The majority ?f experiments in this area, however, ixidiaate
that experzzzental].y obtained matationa do not, in any r, differ
from sparttan~Qus ones, if mutations induced by irradiation, which
waxe riot encoun~Lexed previou ly, (turn up it is, in all probability,
due to the fact that none 01 the objects, with spontaneousJ,y arwic
an tinutati.cfrs, iere Su$fcient1y observed to uncover all of the
poEtsible mutations0 in addition, we know that in every more or
loss thoroughly investi ated'object spontaneous mutations, which
had not been encoimtered in X?ray experiments, crop up from time `to
time,
'the regularity of distribution of oxperin~entaUy obtained chromo.a
sozuia:1 aberrations, confirrs the assumption concerning the lack of
speeifioity in the 'effeo s of short wave radiation which should have
been G~:pected on the basis of the physical prcpert is of the rays,
The foUowin facts indicate that various sectors of. she
wave sp& ctrum do not producespecific effectsz 1} Stubb& s Exper
xrients which were set up with a? i o1ated se ent of ultraviolet
rays 2) the fact that as a consequence of irradiation the same.
kinds of mutations arise:. as are observed under' natural cand itiar
` t
(Contemporary findings indicate that the aq on of ultraviolet rays
does diffez frog the action of X~rsyc) 3) the appearance of rep
verse mutations do ta, action at short waves,
The s
lcn fa?:~ the `narese of. mutations is the
axtio?u t of ene~ aced (the number offr'tmit`s) and not the qualit
1 Y
of the rays (voltage).. The frequency of nautattons rises in direct
proportion to the dose.
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REFERENCES
L. P. 1939.
pp 335.318.
BRESLAVETs, L.P., AFANAS' YeVA, A. S., and MEDVED? VA, G. B. i935.
"Increases a. )the Yield of Rye due to Action of X-rays.t'
-TR UA. v!P. 8 x 245-253 ?
DOROSHENKO, A. V. 19291930 "The Influence of X-rays! on.`the Length
of the Growing Sea,on in Plants," TR. PEIKL o BOT. GEN., I
ANGEL S. 1928. 'PDe l'effect du fractionnement des doses de rayons X
l
sur des graines germees. C. R. Soc. B o1e9 V. 98 223w22L.
ANCEL P. et Lallemand` S. 1928. Sur la' protection c ontre 1'acti on
GLOCKER R, HATER E. Und Jf~NGLING 0 1929. fiber die bi olog sche
1919. Die !tlirkung der R ntgenstrah1en auf die Pflanzeno
Fortschr. Geb. R8ntgenstrahlen. V 27, No` 1 661
MAT INE" ~ `et, THOUV1%NIN. 1898. De 1'.influence des rayons X sur la
germination, Rev. gen. Bot, V, X, No 1' ; 81.?86,
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Wirkung verschedener R8ntgenstrah1enqua1i teen bei Dosier-
ung in rwEinheiten. Strahientherapie. V.; 32 s 138.
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NOELTING W. und. STUBBE F7. 1931. Untersuchungen fiber experimentelle
Auslosung von Mutationen bei And Zeitschr. indQ
Abst.: Vererbung-Lehre L. 67 ~I. 1 s 152-172.
VARIABLE SENSITIVITY OF ORGANISMS TO X-RAYS
Ti the first chapter we have. already pointed out that
X-rays have a different effect, not only on different species and
finer subdivisions of the classification system, 'but also, frets
uentl on organisms 'of a single species. Then we only broached
the question, in this chapter we shall attempt to clarify it. The
first premise Dressed on this subject was that of BERGONIE and
TRIBANDE I1 in 1906. In their article they point out that accord
ing to the data of all the doctors, irradiation, which induces
the death, of neoplasm' cells, loaves the immediately adjacent cells
of the normal tissues, including those of the gelling, uninjured.
In addition, experiments with living tissues have indicated that
the rays have a selective action on healthy tissues. These authors
ha\'e shown that while X-rays destroyed the seminal cells of rat ova
they did not affect the interstitial glands. On the basis of
this, the authors assumed that it was possible to formulate the
following law (which is known as the B .GONIE TR rBAN1T~ ~U law)
"X-rays act; with greater intensity on those cells which have a
greater: power` of reproduction, on those which have a longer karyo
kinetic future, on those whose morphology and function are loss
27S
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definitely' fixed." On the basis of this, they find it quite u>,.-
derstandable why irradiation destroys tumors without offecti.ng
the healthy tissues
In H +, TVrlCi t s experiments (5n 1911) with the action of
radium on unfertilized frogs eggs the particular sensitivity of
reproductive cells (embryonic tissue,' testicles, ovaries, skin)
is likewise pointed out. The greater the demands placed by life
on the c oil (and e specially on its nucleus), the greater are the
changes in the nucleus and cytoplasm; and life places its greatest'.
demand on embryonic tissues, whose cells grow, multiply, and df-
ferentiate into other tissues.
SCI*!ARZ (1913) in a similar fashion, raised the question
ccncerzing the stage of development at which irradiation must be
given in order, to have the most beneficial effects His experiments
have indicated that acceleration of growth is observed only when.
seeds (before sprouting), or veryyoung shoots are irradiated,
wheroas in old t:.ssues he was not able to induce this acceleration.
A survey of KiINICIE ? s experiments (191 ) indicates that
various species of plants display a variable sensitivity to Xuras.
In a1,&itian to this, the more intense are the life processes of
a plant, the more sensitive it is to irradiation. In order to
clarify the reasons for the variable sensitivity of various species
to 1C-rays, K inted, the hypothesis that it is related to
~1~dICKE Moor
the 'number" and sire of the individual "cells, which, in turn, " re-
lated" to the `amount of energy absorbed by : them. His experiments
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indicate that Vida, f ba are injured by da8es of 1OO roentgen w1 ts,
do$ e5?.
by of 20 roentgen units, but that even when donee' of
l
ea.
00 roentgen units are used no traces of injury are observed in
~ .
P aver sarnniferum. In addition, the extreme resistance of bac-
teria to X-rays (where the cells, amounts of protoplasm, and water
are to and the extreme sensitivity of large cells (which con
taro large amounts of protoplasm and water, as compared: with vege-
tative cells ) such 'as sexual forming cells, should .tdlcate the
direction in which the reasons for selective radio sensiti \d tY
should be sought.
NADSON (1920) also points out that radium does not have the
same effect (in relation to both quantity and quality) on various
Species of yeasts, and even on individuals or cells of the same
species. "The more Intense and rapid are the life processes and
development of yeasts, 'the more sensta.ve they are to the action
of radium' for this reason young cultures ` are affected by radiiii more
quickly, and to a greater extent
P:ETRY (1921) considers that the experiments of KoRNICKE and
Izr
SCHWARZ that resting (dry) seeds` are not sensitive to X-rays, even
if they are affected by doses 30 ta.mes greater than those which
affect sprouts of the same species. This proves that the sense.-
tivit; of various parts of the cells to X-rays is determined by
the physiological condition of the cell,, which is based, according
to PEf RY, on eheznical differences.. Mowever, the sprouting of seeds
is not a simple chemical transformation, it consists of many
pa? e 4P'
var1ous (processes: which determine changes of composition
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and function of the seed. Conclusions concerning the action of
the rays can be }eased on the insensitivity of rest uig seeds only
on the condition that we know which of these processes is reopon-.,
Bible for sensi..tivity to X-rays. The simple swelling of sproutang
seeds mayR;.make them more sensitive to irradiation, just as the
accompanying actiization of proferments (Pro enzymes) may, or
the enzymatic breakdown of the h gh molecular reserve substances;
by increasing the number and ? kinds of molecules, or the diminution
of the size` of.'molecus which increases the ability. of substances
to ionize. The presence of oxygen as 'a result of soaking the seeds.
in wager may have an even greater. significance
P ETRY also `studied the action of other factors in conjunc-
tion with i.rradiati on2as for example a temperatures cyanide pois-
'ing, absence of molecular oxygen, etc. Experiments showed that
sensitivity to X-rays is not necessarily related to the energy of
respiration and growth,and may be separated from them. 3y its
low temperature coefficient the reaction of X'rays i.s not only
considered to be an independent process,. independent from the.
metabolic
processes but on the basis of this low coefficient
PETRY classifies it as a reacts on of light. Experiments involving
cyanide have demonstrated that sensitivity to X~rays is not related
to the depression (inhibition) of growth. From this it is possible
to conclude that only the composition of the cells is responsible for
their' behavior towards . the ?harmful influence of `X-rays,' and not the
transformation of substances which , takes place within the cell.
From these experiments PFTRY comes to the conclusion that 'the
"278
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process of injury to'the cells is a purely.{photoahemical.' reaction.
The absence of molecular oxygen does not inhibit sensitivity to
X-.rays` The final conclusion from PETRY's experiments is as fol-
lows: the reason for the sen51tivit3r o X--rays during the time
of sprouting, lies not in the changes in respiration,
but in the
commencing constitutional, transforrnatons The question
wheter tie process of swelling of the seeds, as such, exerts an
influence, or whetber this swelling is a phenomenon with farreaohing
consequences (which accompany sprouting) which formsP.a light-sensitive
system. So, the X-ray reactions form an independent process, al-
lied to the photochemical one, in all probability consisting of
,,
photochemical changes of the important components of the cell.
Although PEL THESE (1922) article was written, in order to
clarify other questions, it is directly related to the question
of variable sensitivity to X-rays. His first question is: "do
X-r. aYs' have a selective action rr , All of the facts point to the
variable sensitivity of various types of cells, but P3 ?THES ae
sines that all cells are sensitive to X-rays in ;some degree.
Nevertheless, the selective action of these rays is the essential
PERTHES' attaches ;a
basis for the effect they have on tumors
special significance to the latent action of irradiation, both for
practical purposes and for theoretical investigation. The time
after which the influence of irradiation begins to manifest ~ tself
is extremely variable a the k4gher the dose and the' sensitivity of
the irradiated ` cell, the shorter is the time intera11 .rERT11ES
xp fact that irradiation of the coin
lal ns the latent action by ` the i
ponent parts of the cell, chiefly, the `nucleus, induces chemical
. 2-7
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changes which proceed slowly, and gradually increase the amount of
harmful substances, (probably insoluble products) which increases
first their stimulating and later their harmful action FERTHES
considers that his hypothesis completely corresponds to the avail-
able data. In the first p]ace it shows` that not only latency, as
such but its duration is, to a certain extent, in invefse ratio
to .the intensity of irradiation In the second place this hypo-
thesis offers a good explanation for the fact that during the
latent period L.e o before the appearance of the main' effects,
the foremost of which are the abnormal processes of the cells)
cellular. `reactions slow downq etc. It also oxplains why in some
objects the stimulating effect becomes harmful and injurious to
the c&.Lis and tissues. Observations indicate that the substance
which a `feets-the cell is, at first, found in a minimum amount
which ; radually increases because of the influence of irradiations
ALBERTI and POLITZER (i921.) in their first article, devoted
o the action of X-'rays on 'organisms, write that even the very.
first biological X-z?ay.
xperiments showed that resting c ells were
more stable in relation .to the action of X-.rays than cells in
mitosis, or any of the stages of mitosis. If. the various stages
of mitosis' are projected on the X~axis (abscissa) and the sens-
ti v,
ity to X-rays (expressing, for exanple, the nurnbe of perishing
ova of the horse ascaris) is projected on the YwaxIs (ordinate),
we will then (get a curve at whose peak will be the metaphase. Con-
sequently a small ;1rrad!ation will disrupt only those mitoses
which are in the metaphase, higher doses Will destroy :those stages
which are dearest to the metaphase, both bafo~ a and after. Finally
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there exist stages which will `lead' to the necrosis of all stages of
;mitosis. In order to kill the resting ce113 greater doses than
any of the previous ones are required. .
CELERY s observations (19210 on irradiated beans, peas,
wheat, and lilac buds permitted him to conclride that, the action
of rays i.s determined first by the pOCi OS of the plant and secondly
by their condition. Unsprouted seeds are more sensitive than
sprouted ones swollen seeds are more sensitive than -:pry seeds,
The age. of the seeds does not affect their sensitivity. Changes
in external conditions (temperature, humidity of the air) show'
up under irradiation.
JUNGLING't s observation concerning the 9 m-
portance (signir'icanc) of the 'point of application of irradiation
is very interesting. when the tip is irraitiated growth in the tip
slows down when the whole root .s irradiated the side roots
cease their development.
:n sEanE's (l920 article we find a sumTrL ri ation of ex-
periments with zoological objects irhich indicates that strong
injuris occur .in early `stages of developmnt vith moderate dosed`
of X-rays and radium, while older stages are completely unaffected
by these doses. SEa DE's own experiments indicate an extreme sen-
sitivity of the stages when chroroosonles began to form (prophase)_0
Stages when the chromosomes are completely formed (metaphase) are
considerably lass sensitive. IE.OCHLIN and. Or1 EICHCxEWICHT `(1925) compared
the action of Xwr'ayd~on sprouted seeds of .Phaseolus vulgaxis and
vicla faba. By irradiating them .wa.th identical dosages they estab
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lisheds (1) a variable sensitiv?ty of these two plants` and (2
variable sensitivty among individuals . What is most interesti
g
that the authors d3.scovered that the more sensitive. the object
the more quickly is stimulation followed by retardation. In
Vi.cia eq4na retardation ja induced by very small doses, for
which reason' this
object is not s 4table for investigation of he
stimulattg effects of Xwrays s So to study this effect the authors
used Phaseolus vul,ar instead. hen tree seedlings of the two
plants were irradiated by identical doses (1 High Erytheme Doses)
two weeks later sprouts of Pha 1s only slightly retar ted
their growth while sprouts .of Via stopped growing altogether.
NEM1:LNOV (192 and 1926) consid?rs' that old cells are most
sensitive to X'rays. The age of the cel ., according to him, is
determined by the ralation of the already lived part of its life to
the total life span=. of. the given cell. The essence of the. biolo-
gical process in the X 4rradiated cell, which 1oad to the aging
of the cell, apparently consists of the disruption of metaboli8m
and of the retenj; Lon of the products of mctabolism within the cell.
1VEN (192) also takes note of the varied sensitivity of
X~rays which varies chiefly with the species of the plant.Com-
paring the data of numerous experimeritS he canes to the conclusion
that small seeds have. a low sensitity to X-rays and. seedsc.bf,
:1m e size, high 5ensi1ivity. The most sensitive obiect5#are the
seeds of Vida faba, which are the large`st ?r1 size (of those ob
jects which have been subjected to irradiation). He also notes;
the ;importance of the physiological condition` of the , seeds for
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successful irradiation, pointing out that turgid seeds read, t
irradiation ? :o a considerably
Cytological examinations have revealed that nuclei of vegetative
eels are more stable than those of sexual cells'. The vegetative
cells, in turn, manifest a difference of behavior; cells of the
points of growth of'.the root and stem are more sensitive than the
cells of the ;permanent tissues'.
ANCEL (1927 ) finds shortcomings in the works of many authors,
who point out the differences i.n the sensitivity of C?y and sprout
ing seeds, in the fact,Uhat these authors failed to follow up
thei?? stud?es after the appearance of shoots. She .find it neces-
sary to 1111 out their experiments in this direction by the study
of chan es In sensitivity of lentil seedlings from the moment
greater iEfree than dry seeds.
that srrouting begins to the time the roots reach a length of
2L centimeters. Her observations showed that the sensitivity of
lentil sees continued to rase from -the moment of the appearence
of sprouts, that it reached its riaximur wren the roots were 10
millimeters ion
fter 'which it went down gradually to the time
the roots ere 21~ millimeters. C'Either this should be centimeters
or else the previous measurement n this same paragraph, see above,.
should be "2!. r Ili meters. i These results safeguard experimenters
from mistakes, if they set up experiments with seedlings whose
roots have different lengths..
STADLER (1928) was the first to s~i?w that with equal doses,
timas as many mutations appear 1n sproutTg seeds, as in resting
seeds, Mutability
jn generative' cells at the time of formation of
R Y.
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Sexual cells or gonads is stud1ed ''estar, detail,
Gaon5pEm (1929) pointed out that certaan stares of reduC-
tlon~d vis~.on are Parta.cularly favorablefor obt,aiing chromosomal
aberrants, bu?t whether the sam r
e can be said in relation to gone
mutations has not yet been determine
STtTBI3E (1932) showed that when
srapdragon buds are irradi-
ated by equal doses that more m
uts.ta.ans occur If the irrada~t1.on`.
takes place after roduc t~on~d~:~risian than if it takes place dur--
i n
x
g o
before ite
.tn the first chapter the result's of r~a~aents of B' ESLAvTs
and her co~waz'kore conecrni.n, the
~ variable sensitivity of turgid
seeds and sprout s
of rye were presented (BREsLA ~?'' .,
~ X7.5, AFANAS ' YeVA,
and OULDEV,, 1933) arZd also of
dry and turgid (soaked) seeds of
peas (ATABEKOVA, 193f)?
JOHNSON performed many e eri
merits for the purpose of
clar~fy~ng phY8io1og1ca.l changes of ~
,.~ ~ ].ant,s dui to the Influence
of X-rays, Two of h ra ~'
x works have a d.:.r, oc t bearing on
In ~~he first :of ?Lhe ~' ..our subject,
se wo:rks (193) the author a.nvosti atoll
'effect of irradiation ~ .the .
on three varlpta.es of French spy-mach
(At?~1lei h_nrtens1s) a (1) variety. with pale, almost yellota
..leaves, the; spinach which Is most fr
equen?tly used us food,
(2) variety with dark
red leaves which is cultivated ast a ds
t~.ve plant, and ' corm
(3 ) green variety Which ealhibits the most luxuriant
growth. Expariment5 with different varieties of the sane spec io
were set up in arr~er to clarify whether or not the smech, B. ion, N, 1/2 r 83-J,09p
192~. 1r3menta1b.i. ,
ologi sche Vorstud1en 'zur
sthcrapxp.
d :R~nt~c~r~~tr. Kreb
rortsehr P 32, H. 1/2 S66L
ANCEL, . 1927 Sur 1.as vsrMxat ions cue r
sens:fbilite aux rayons X d
ra~.n~ , germ~es suivant le s ., ~ e
t :4rle .otu dc~rel.oppemente G. ~~.
Soc. Bj.ol.~e 96 No 13 r 986989
EioNi J. 9 et TR1BQ JDI, U~ L. 1906, Interp1 >'
8tatiOn dQ qUeZqUGB
resu.ta F s de l "rack ptha r'a
ple et essay, cue fixation c~ 'u
ne
technique rit1one11e, G, R. AC. S, Paris, V. 3 d
; 983 , 985.'
C~us'rjA. 1936
To di..f'ferer~~ stabui.to
chromo~cmes and the
na turn p;f' ? ..
ma.Uos:.$, eX'edtas. v,. XXII, 28L...33
~
1936, er rersc.
hj.edene Sensmiiit and }1; , ;;
Ghroma
$Omona Bots~j.ska Notj zer; .
4g8.512
GENTC}{p, 0. and UPS TA FSSGN, A. 1 939? The double chromosome rc- ;
pzryoducta.on in ~~a~.z~~,cea and s. causes iZ An X..ra ,
Neredjta.se Y sxpcr~.me
V. No1 :371-386.
2332380.
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NE~N4V, M. I. 192g. ?Cxiti ca sm of Teat;ha.ng ~LLout the Bo14gica1
.
?
Action of X raY S e
tr (ftosntgenologY? Chapter x~V
19~~i? rf~,~NTG~NOLO~xYa
Concern ~ical :A~;ti.fln Of X:r-aye? )
-~.n the - ~ ~.o~.c~~~
t er Bnflus~ der I~dnt~;en?
iLBERT, ~~.: and BO7~T`f Z1.,I~,B (~. 1923
en auf di e Zeiltaiiung. Arch. f r
S trahl
h. B. 100, H. 1/2 @3-109.
~'xtih~~~a:{~l~lun~s~nec
Vor tudien zux Krebstherapie.
.192k? Lx~aea~~.m.cn~~a~b:~.oLo~a. sahe
F'or Lschr o ; RbntgefS ur B. 32, H 1/2
( do
ANGEL, S. 1927 . Sur e vrinUors de sensiblaux rayane X -~.de ,,?lu d~ eloppem~nt. C. ~?
~;rai.n~~e germes ,u,~.vant le ~~..
. Soc. B' o1. V. 96 No 13 A 98&?989'
U L. 1906. nt~;rp~'eta~,~.an de cuelque~
BBh'CCNI~ g . ~ . et 7: .~3CD~~.A. ,
et ~eda~. d~ ~'~.xe,~~_can cl ~l~ne
_ r+~s~z:1..t,a1. e de la rach.otherap
~1ecY~.3.c^;ue` raUOnelle. C. R. Ac. Sc.
M
,, :tl.it ~ of chromosomes and the
,SSON' ~,. 1936. The c~i;~f~aa enr stJab~
rtrS'T~~~'
(.
. nature of mitoea.s. Herec3a.tas. V. XXII: 281~33~
" ~.~.t~.~ and Chr orao
~.936. ~ber ~rerec~.a~edene Sens:~b~
comm.
Bo1.ika Not er: L 88 12
N',A. x.939? The double chromoSame a~G
G~~I~`CI~B~'~ C . and C~US.,A~'S~ ~C
~~ and :~ts'cauees IT. An X~ray eXper~.mcnt.
in ~ ~~~na~c
pr.oc~uct~.nn -
Hereditas. V. x:xV, No L. s 371u346.
..
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HEP.TWIG' C. 1911. imbefruchteter and ire Ent-
r?' umb?strahlun
~ra`cklurig Hach g mit normalen Samen. 'Arch. ~.cr.
~e :Cruc ~~.
Anat. V. 77 ? Na 2 s 16-209?
~umkrankhea.t t9.eri:echex KE mzF,llen.
~T~IG 9 ' G . . 1.911 . Die . .~ ~~ ~.d~.
Arch . micr Anat V r 17 9 No 1: 1- 9~ 9 97 X164.
. . ,
durch Radium Und. R8nt,gen5trahlen.
HT.SR`I`~1.G , P. 1927 ,? Kea-mE~~SCh~di~~~en
der VE erbungS r SS' . 13. ITL
Handb.
IVY, Ha 192g. Neue tersuchUngen `fiber die \ ix'kun; der
~'e n
strahlen auf .E'flanzen~ Strablenther Pie B. XIX, H.
3 413-461.
Rontcn
~eatibility of Seventy spec 7.e~ of fi].owering:
JoHNSON , E . 1936 ? Sus~ ~
diat 0n. Plant Physiology. V. 11. No 2
plants to X ra
319-342e
K~RNICKE, M. 191~ . er die l~irk~ng vershiedenen starker R8ntgen
str. ahlen auf Koimung and ~Jachscum bci ricn h8heren p flanzen.
V. 6 s %16 430
Jabxb YZSS? Bot.
~ arches Sur 1es T'act~etxre de 1a radio-
T.,A~3ER`~a J. 1933. ~sah _
~h?rphalo-
enc~~enes
?7?
SenSibill.te ;tassulairo en dehox s des ~
iOteS baolc lqu s dos tissUs latc3nts.
giques. Ices propr
44,9 ~ e 4 6~1?" 14~
e Bipl a T.
Arch
LONG, ~.936. Stimulation of growth of soy
T. and K~RS~~N, H.
begin seeds by soft X-ra $. Plant Phy$i ol?gy. V. 11g No 3
~' .
61621.
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PERTHI{a~S? 1922. Die jotogischen ~irkungan der Rontgenstrahlenp,
-
b
ie, ' U, lL , H. 3 s 738~.760.
Strahlenth?rap
?s der Ber~ingng~n der b~.ologiSchen
PE'~1~, E. 1921 . Zur ~{enntn~.
enstrahlen. Bi'ochem. Zeitshr. ? V. 119
Vir1uflg der R8ntg
23wLL
193k. Pathologic der` N~.tose ~ 1~238?
I~CIL~'1uER9 ~?
tumsre~.~ der rtdntgenstr. ahlen a,a.f
SC't~iAF.2~ ~ E o 1913.. Der W ac . s
p:i'lanzliches und teriScl~?s Ge~iebe? Munch. mid. ocherisehr.
.
(a.
No 39 s 216g.
en ber e~erimcntielle Ausl~sun~;
SBEE, H. 1933 ? Un~~e~ suchung
,h~.num ma `us IV. er die Be2eih~g
van ~ .oven bea_ Ant ----- ....,~ ..
azionsr~te Hach ~~ntgenbestr~hlung
zwischen Dozi.e and Genrnut
nannlicher Gone.
Zeit schr. and Abs. Ueraxbungslehre.
E. L, F? 3/h ; 1.81~2QL.
e Stand. der Strahlengenet k. Die
gegenwtrta.g
1937. Der
Naturwi ssorisdh.
No 30-31 : L83-L90; OO Q6 ?
193$4 Genmutation.
Chungen xur etperimentellen Erzeugung van
19~D~ Neue Fors.,
alb/. V a 60, No 3/L : 113~126.
Nutata:on?n. viol?g: , Zentr
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THE BIQLOGICAL DOSIMETER
As soon as the biological effects of X?ravs were established
the neect for expressing their action in some kind of una.ts 1mmedi?ately arose. Since the action of the rays on
human skin soon
became apparent (the skin. of people who worked
with X-ray. appar,ati
became reddened and. peeled) the so-called er
ythema dose was pro.?
posed as a dosimeter, This dose had an international
d.esa.gna-tion
IlED, a contraction of the words Haut..Er thema~D
Y osis the dose which
~ ~.ch
caused the irritation of the skin). Howev3r this
reddening
varies very greaiy dap cnd'. ng on the pi.mentata on of the
skin -
(blonds ore more `sens. tive to X-rays than brunettes ) , and the
age and" health of the patient. Consequen-tl a r' al
Y need arose for
the precise dete:rani nation of the dosage, This
.,~ need arose chiefly
due ?;o the medical practice of applying X-rays for tr'eatment of
tumors and for d:'_agnoses of ailments. 11e cannot
linger on the.
history of this question' we can Only point out that at the present
time exact d simeters exist based on the io
n~.zati.on of air by
X*.rays. The number of roentgen units r) u
~ _ used in experiments i:s
determined by mexxns of dosimeters. These da
simoters have deter-
mined /the extent of inaccuracy of measur. ev en
t of roentgen una.ts
:in HED's (High }erythema Doses). As Clarck'
s c'xper, i.ments 9 which
Were conducted on numerous patients in seve:rai c ti '
nxcs,.have
shown that, depending on the indivi.dualit ofi
Y of the. patiant and on
his condition,' the li.ED varies between Li.OO and 1200 roentgen units
with the average lying around 8l9 roentgen
nit. If we Will recall
frown our data the fact that a plant which 'a
s s v,nsitive to X~r. aYs,
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such as !'i urn Ssvitum~ is stimulated by a does of 39 roentgen units
and depressed by a dose of 1000 roentgen unite we shall underStan.
that such :a dorneter (as the H ED) is worthless.
:n this manner an accuracy was attained in determination
of the dose but that d d not do away with the need for a biological
dosimeter. In -investigations of..the effects of X-rays on plants
it becomes necessary to 'feel out doses, which are capable of inducing
desired changes, by a strictly trial and error method. Only by
analogy can we judge concerning the effect that the rays w..ll have
on a certain plant, ? and hen only approximately. The reason for
this lies in the variable sensitivity of plants to X?rays and in
their physiological conditions AS we have pointed out previously
(chapter. Li., section B) sensiti ty to X-rays varies greatly depend?
,
ing on the genus, species, and even variety of the plant. This
clearly expressed property of plants'of reacting variously to
irradiation has forced (and is forcing) investigators to seek a
biological; dosimeter which will enable them to determine quickly
to sensitivity (to X-rays) of a' given plant.
In 1931 Packard proposed the use> ?f a4la eggs as a
biological dosimeter. This proposal opens the possibiitieS of
a new biological method.
Johnson. (1936) points out~.hat two SPCCI(?S of plants:
Nemophia and Zip are also sensitive to X-rays. Although She
does not point' out that the plants can be used as dosimeters,
nevertheless the extreme sens v~.ty of these species to iy'radiatl. art
, ~.t
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(oi
oreUcal interest because it permits the observer, who is
i~ of the
fects of radiation on plants, to judge the
oxper~.enced ~.nthee_f
approxirnate dosage on the basis of the extent of the injury. The
out that two considerations ~njtigate against
author herself points
the use of the plants as dosimeters; The changeability of the
that the effects of irradiation do not show
plants and the :fact
Even of the plants are given lethal doses, they
~..atel
up a.mmed y
can continua to live for several weekS, while leaves (tho most
ens arts in respect to rays) can reveal: abnormal
s~:tive of the erg
developments a long time after the irradiation.
Preliminary experiments have indicated that for 11 day old
sprouts of Nema h 1a insigniS a dose of `2 OO roentgen units was
.~,....~'
lcthal? In a second experiment 28 day old sprouts were irradiated
by doses of 1 OO, and 2000 roentgen units, and this experi~
0
100 , ~
L
menu showed that 'a direct relationship exists between dosage and
the pc, arcentage of survival among the plants. On 110th day after
.
sprouting 70 percent of the control plants survived, hO percent
of the 1000 roentgen unit :group, 27 porcent of the 100 roentgen
una.?t gz aup, and E percent of the 2000 roentgen unit group. So we
,
rr ority of plants which deceived a 2000 roentgen
See that the j
unit dose perisheS while those which received a 1000 roentgen
,
units show retardation and cfiminut,lon of growth. A dose of
1 oo roentgen unite e Chib is an intermediate developrncrnt. Cone'
it is asstble to estimate the dose of X-rays by the
ver seJ.y,
number of surviving plants.
Co
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Prel9.rr nary experiments with ; Zinn' a, revealed its sensitivlty
to X.rays. The effec t of the rays on the growth, the weight, the
iL ormation of abnormal leaves, and the diminution of branches all
shotir a direct rein tionship between the in juries end the dosage.
The size o:: trte dose can be estimated by the extent 0f the injur .
ies, just as in Ne~moph~la.
But as the author herself points out, the results require
a 4d?
several weeks to how up (110 days for Nerno.h a) for that reason
it is doubtful whether this method can be used as a biological
dosimeter.
tJe feel that this question should be broached on the
cytological level. Pr'o;nsy and ?lrevon (1912), Kornicke (1920), and
others point out the increase in the diameter of the cells due to
X'radiation, the irregular course of,1Karyokinesis etc a This Is
also well demonstrated in our wore with rye (Breslavets and A.fanas t Ye a )
with peas (Atabekova), and with wheat (Afanas'yeva),
c/5
seta take
by way of an example thentracellular changes obtained from Xratated
seeds of rye (for amore detailed description and diagrams see
Chapter 3). By irrathati.ng seeds o1 rye with doses of 2O, 500,
750$ 1000 2000, 1,000, and 8000` roertgen units we can observe the
following changes With a dose of 250 roentgen units In 30 roots
examined only one nuclear plate was found with a triploid number
of chromosomes and one th greatly shortened and thickened chromo-
somes. In general t;he' cells and the nuclei were cornpl, tely normal
in division as well as' in tho resting stage. l'ith a dose of 500
roentgen 'units only in two roo~s nuclei which migrated' from one
cell to another grid fusion of dells ;observed,' One tetraploid plate
4.
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To these criteria another one 'may be added, which was proposed by
Johnson, namely the speed of reaction, On the third or fourth
day after the irrada.a Gaon of seeds and on the second day after
irradiation of sprouts we can already have our answer on how a
certain plant reacts to a given dose of X-rays by j.ts 'intracellular
changes, In order to establish these changes by a worked out
methodology, of crushing the rots and staining them with aceto-
carmine, does not present. any difficulties,
REEEEENC +'
Bf ESLA.VETS, L1 P. 1937 ? ?Contemporary Developments in X ray
Biology (Tlants).H YuLL. MOSK. OB-VA ISP'YT, fLiRa T.Li.6
(6) . 359369.
?
RT,)BINSl:;YN, D. A. 1933. 'tTowards a Methodology of a Biological
Dosimeter of Xwrays " VESTN. k~4EJ\TG1~NOL. I RAJJIOL. J; . 12
V'. 6 ; 389395.
BRE SLAVETZ L. and A.FANASSIEV A n 1937. The ac tjon of X-x?a rs on
the rye 2. X-radiatj.on, of seeds. CytoIogia'a
10427.
No 1
C1LARCK. 19314. The effects of Xradation on cell structure and
growth Sy tposium on qu~amtitative Bi?logy. 2 2L 9x263.
JOHNSON, E? 1936. The relation of X?ray dosage to degree of injury
in Neniop?y;L and Zinnia. Amer. Jaurn. of Botany, V. 23,
No 6 ; '1 lJ~W 'l8.
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93d. Susceptibility severf y sped es of flowering plants
X~radiation. Plant ? Physiology V. 11 0 319-3Ij.2
KGRNICE, M. 1920. Die Wirkung der R~ntgenstrahlen auf die
Pflan~en. Fortschr. Gcb R8ntgonstr. V. 27 c 661
PAGKARD, 1931. The biological effects of short racUations.
Quart. Rev. Ilio1. Vp 6 259?280
hE RETENTION OF THE E t FLC TS OF X RAYS
n the 'great majority of ;experiments with :irradiation of
seeds, the latter are planted immed.atel3r after a:i-radiation.
Naturally.the question of haw long the action can be retained comes
up. Very frequently for theoretical purposes and particularly
for practical ones it is iraportant to retain the action for a long
?timee 0u ? laboratory has frequently received requests for irradl..
ation of seeds from outlying places where good X-ray setups and
dosimeters do not exist. If the action of X.rays is retained for
only =a short time it is obvious that it as impossible for us to
accept such
'equests. Since the literature on this .question .is
very contradictory it became necessary to set up our 'own experiments
to clarify ito
Guilleminot (1910) writes: "The injurious action of irradi.
ation is retained for two years ;in
original force."?
Schwarz (1913) considers that the act: on of X-rays is rei..
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44
14
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vorsible, ire. that irradiated seeds rotain the :irritation induced
by X-rays for only a short h,lne.
This opinion is 'seconded by Geller who concludes, on the
basis of hio experiments that the irritating action of X?rayrs is
replaced in a few days by normal` development, except vrhon irri ta-
tion i such that it leads to gradual death of the plant
:GIN (192) ' tries to prove on the basa.s of the experiments
b
r
of Guilleminot andpr, that even after being stored for several
months, irradiated seeds retain the full farce of the primary
effects of irradiatioi:
The data of Muller (1928) and Hargis (1929) on. ' hi- sub-
ject are very valuableb On the basis of their experiments they have
pointed owt that the frequency of lethal` mutations was .dantical
among sporrfl taken imrnedi.ately aftCT irradi ate on and in those taken
several .ys afterwards.
A fang s' yeva
(1936 and 19313) used a plant pith which she bad'.
done a groat deal.` of 'work, namely: spring to at Tr cum vulgare
var caea:~.um Cll1, (pure .strain)
the seeds of which were irradiated
by a do;3e of 16000 rOent;en units. The ;author was aware from pre-
vious experimentsthat t+h s
changes -n. 'the cells. These
doss invariably produces the same
chanties include': inc1u3.oneifl the
cytoplasm, formation of micronuclei occ irr face of ;two and more
pe
number
~;rotoplaets within single ee11s, changes in the and she
of ahe chroasoma
arreguar nuclear plates, and, finally, chromatin
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bridges. In the first e aera rnent the sec ds were planted at five
a:Ctt?r irradiation, after l month, after
different periods. one, day
3 ~tonthspaftor b months, and, finra.llY, after ayar. From l able
1~p e can see that allof the above described changes appeared
ta
after a day and also after all of the other periods
year
including one'
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`LAEIE !s0
Frequency of cytological.. i~^re an . ,, .:tics in the somatic yells of wheat
-
Date of 2`ime of P3 er er
Average
root
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Number of IInc .,luslons Metaphase bath
t x
= '~.
s ,.. A olynu,. i Abner mat Cb~: in Formation
~lear in the Clanged Changed t y
r r n.. ape I plates. eridges~ Wo Or ,.
c;e1l ,-
c ? la ,number r shape
w ,_ t r ~~top~
more
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weakening or iianlshlng of the effects of X-rays as not
observed. .n the contrary, a certain increase can be, detected In
the changes, For example, the number of mJ cronuclej a n j `
t~
nuclear cells Increases somewhat, Three months after irradiation
the number of micronuclei was 2S per root instead of 12, six months
afterwards it was 31, a year after rds:tthe number went down again
2105e to the original one a It seemed interesting and important to
determine whether the action of irrar i bjon can be retained f Or
even longer periods, say for 2 or 3 years, Simuitaneously with the
planting of the seeds which were irradiated in 193L, two controls
were planted. The first control consistsd of the nnirrad1ated seeds
of wheat of the 1933 crop which were kept in the same c onditior
as as
the irradiated one 1 Seeds of he same pure strain of the 193S
crop served as. the second control. In the second table (Table Lii
data of the cytological examination of 1937 wore compared with those
of.193v
:fa
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:f ter Irradiation iicronucle ~` . ; Poly-nuclear Chromosomes _ Metaphase
_ Percent of roots.
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..fig can be seen frbm Table 11, the act?on of X-rays ors calis'
is not lost for' a long tirne. All intrcei1uiar changes induced by.
X-rays, and observed in roots fixed seVeral days after irrad?ation
are alsc 'observed in roots which are fixed throe years after irra-
diation. The number of observed changes, the number of micronuclei
in pc rt1cu1a.r, actually increased, as -did he number of polyprotoplast
cells Examination of both control groups dI.d'not reveal any changes
in trhe nuc1ei~ Consequently the experiments.nc1icate that the changes
induced b . X-rays are st b1 e and do not disappear or weaken even '
when `?rraci:.ated suds are stored x or? three yoars.
Afanas'yeva, A S. 1936, "Retention' of Action of Xrrayson 1'heat."
SYuL ,I QS , OE VA TSPYT> ` e 1. L (6) : ).3 341J,o o
1936. , 'Stabi1ity cf Action of 'X=rays on the. Seeds bf Spring
Jheatm" SB. PAWTaT1 V, N 0 LYuI3IFENKO - AN UK. SSR : 1 1.15 ,
.... .
Afanas'sieva ft 1936 SUr la prrsistance de:1'action des rayons X.
Rev..de ;cyt.. et de cytchys. veg. T'
Geller F. '9 ; ilia lrk~tn 1der 'Rntanstrahler: auf jugendliche
rganismenl.n~acinrichr No 11
Gu4le ,oat f~~,r :~ Q-'a~r r~e s s e .i M~ on des rayons X et des
~~~
i r d r ~! rJrfr~j rt r ~y rn ~r a r'}t'~ S.~^4 i ~' F r a i e~
t Fm . t ~ ~ ~ y Y ~,, ~ i
~ ll ah~`i t ' i lp ~~ }~>,~ ' ~ M r d'C r4 ~ Iy ~~i 1 1 S i i -
} ~N' ) n 4, a i t G' a h 4
J: r Y i ~r4p}x~y t~wr+ F~1~1ry95 a`I~~~ke ~ ar~~ht~A~9 ~!r ~n i ~i
;~ a,d1um~~ 1ade'latente. C. R. Soc.
~~ ari Z 1 rid (fib ~~My hVW ~~1 i~ni{I ~f`eS~'dyA.~' t~9 i'~4 ~i j'~lry^Id i ~Ik`'l~ h~';V , dw~ , 1 t >
;~
is A f ~1~7 i r r i
r l f } a { 4 .~ r~r` ~{ fly ~r +a~ ! '~ w Syr a 61fd 4r yn~H~~{~ ~~ ~~ u 1f i;~ k r H
,~ a ~ I v~~5x r - r 1~1 h ~~lCM:1~" e y~ 4' ~ I rl ~p4 Ada f s y
~CT~~fri 4
s ea .1, . tiL68 y, ok
Cry
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lven, H. 192g. ~euere Untersuchungen fiber die irkur der Rtintgen-
strahlen auf P1'lanzen. Strahlentherapie. i'. 19, H. 3 Li.13'161
Jever, 1'. 1923
Latenperiode rontgenhestrahlter ruhender Samen
Wiener K1i n, V1AQhenschr. 8.
NATURAL RADIATION ANT) COSMIC RAYS
In 1928, Muller in his First works dealing with the increase
of frequency c1' ~nutatrions induced by X-ras, expressed the
premise that spontaneously arising mutations are possibly due to
natural shortwave irradiation. This assumption was all the more
probable eiince all of the expert.mentally obtained mutations [di.d
not differ in any way from natural mutations
Olson and Lewis (192B) in their articles "Natural Radio-
activity and the Origin of ' Speciee," writethat In nature all
living creatures are constantly bombarded by gamma rays of low
intensity which are due to a' wide distribution of radatoacta.ve
substances. On the basis of materials