COLLECTED WORKS ON RADIOBIOLOGY

• Document Type: CREST
• Collection: General CIA Records
• Document Number (FOIA) /ESDN (CREST): CIA-RDP81-01043R000200180025-9
• Release Decision: RIPPUB
• Original Classification: K
• Document Page Count: 272
• Document Creation Date: December 22, 2016
• Document Release Date: April 2, 2010
• Sequence Number: 25
• Publication Date: October 11, 1956
• Content Type: REPORT

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Sbornik Rabnt po R&liobiologii N. I. Nuzhdin,
[Collected d rWorks on Radiobi to ogyJ Editor
1955, Moscow, Pages -160
TABLE OF CONTENTS
Page
Foreword 1
Shapiro, N. I., Nuzhdin, N. I., The effect of different
dosages of Xray irradiations on the survival of
mice 4
Shapiro, N. I . , Nuzhdin, N. I . , Kuzin, A. M., The action of
estrogens on the radiation reaction in mice 24
Shapiro, N. I . , Kuzin, A. M., Nuzhdiri, N. I., The role of
the physiological state of the organism on utiliza-
tion of protective reinedLes against he damaging
action of penetrating radiations
71
Shapiro, N. I., Nuzhdin, N. I., Volkovich, M. A., Concern-
ing the role of damage to hematopoietic organs in
the course of radiation reaction
89
Nuzhdin, N. I., Shapiro, N. I., Petrova, 0. N., Sterilizing
action of ionizing radiation on mammals
137
Communication I; Effect of Xray irradiation on the
fertility of male mice
137
Nuzhdin, N. I., Shapiro, N. I., Petrova, 0. N., Kitayeva,
0. N., Sterilizing action of ionizing radiation on
mammals
189
Communication II; Effect of Xray and gamma irradia-
tions on the oestrous cycle of female mice
189
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Nuzhdin, N. I., Shapiro, N. I., Petreva, 0. N., Nechaycv, I. A.,
Sterilizing action of ionizing radiation on mammals 252
Communication IIi; Hereditary nature of sterility
induced by the action of Xray irradiation 2:2
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COLLECTED WORKS ON RADIOBIOLOGY
FOREWORD
The adaptation to practice of the production of atomic energy
has opened to mankind the possibility of utilizing for peaceful pur-
poses its boundless resources. The fulfillment of this possibility
has been brilliantly effected in our country where there has been
established the first electric power station in the world which
operates by means of atomic energy.
In addition to this most important performance, nuclear radia-
tions have already found extensive utilization in a number of other
domains of human activity. Thus, in medicine there are methods based
on the use of penetrating rays for the diagnosis of diseases. In
addition, for the purpose of the therapy of a number of disorders, more
extensive use is steadily being made of external irradiation with dif-
ferent forms of radiations. Likewise, the introduction into the organisii
of natural and artificially produced radioactive substances has found
application.
15
Ionizing radiations are extensively utilized in the most divers-
ified fields of scientific research and, in particular, in biology,
medicine, agrology, and agronomy. The method of tagged atoms makes
it possible to solve rapidly and accurately many scientific problems,
and in a number of instances it is only as a result of the use of this
new method that their experimental solution has become possible.
Utilization of various kinds of penetrating radiations has found
no less application in engineering ?- in checking the quality of production
(flaw detection) in particular.
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^ fora -oin; . t ? apparent that nt tae ynresel1t time
~r~~,,~,. .~
man encounters with ;steadily increasing frequency various types of radio-
active radiations. It is beyond doubt that on further progress in the
production and utilization of atomic energy an increase will also be
effectod in the contingents of persons who, due to the nature of
their work, are forced to come in contact with nuclear radiatiot;s.
It is well known that one of the characteristic features of
ionizing radiations is their high biological activity. In the prescnce
of relatively high degrees a exposure of the animai and hu;:ian orb snis~n,
these radiation: affect all the organic fuir:tions inducing the so-called
radiation damage. Therefore, it is entirely natural that a steadily
increasing interest is being;hown tine study ai the biological action
of ionizing radiations, the evolvement ofothods and mean; for
biological protection of the crganism, and the therapy of radiation
induced disorders. Moreover, cognizance of thv regularities of
biological action o? radiat .o L~ .iu also neceary for the development
of the most effective uethod of radiation exposure in the treatment
of various diseases (neopla, et al).
Although the study) of the effects of ionizing radiations upon
the organism was imitiated a long time ago -? practically speaking,
since the discovery of ;frays W- it must be admitted that at the present
time science still lacks a satisfactory general theory of the biological
action of penetrating radiations. Furthermore, the factual data
accumulated are insufficient for the solution of a number of important
problems which have arisen in connection with the use o nuclear energy.
Hence, the necessity of further studies of the action upon the organism
of different types of radioactive radiations is fully evident.
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rncc symposium being nereby ub:nitted to the reader includes
a series of experimental papers concerned with two problems; (1)
study of the mechanism of the action of certain substances which
safeguard the organism fron} the harmful efiecta of ioni~inf; radia-
tions; (2) analysis of tcie action o1' the penctaating radlatioix O:
the fertility of maranals.
Among the substances tested as protective agents, speciai
attention has been given to elucidation of the action of estrogcn^.
The correlation between. their protectivc action and the co~c and
time of administration hac bccn ascertain, ua woil ,aG~ the physio-
logical condition of she :has-protected animal. The conditions
whereby the protective action of estrogons i prolonged have been
elucidated.
In addition to studio of the protective action of estrogens,
a special investigation habeen carried out oli the effects of im-
plantations of sp1n and injection of homologous bone marrow, in
the cage oirradiatedaimaI, In both cases the benefici:xl effec?
of these procodw'es upo;i the; ,.ourse of radiation reaction and the
survival of th.: ax;ina1. haw boon ascertained.
In three papers the results of studies of the influence of
ionizing radiations (single exposure to Xrays and chronic gamma-ray
irradiation) on mice fertility arc presented. Analysis of fertility
was carried out by the method of breeding the irradiated animals to
nonirradiated, as well as by means of histological studies of the
sex glands of animals expoaed to radiations. Embryological studies
were al$o made of the offspring of irradiated parents. Finally, an
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analysis of the fertility of irradiatod females during their oestrous
cycle was studied. In these papers new facts are presented which
embody the characteristiva of the sterilizing action of ionizing radia-
Lions in the exposcd aninia1s and their offspring.
The above-enumerated papers are preceded by a contribution
in which data are presented on the correlation between the irradiation
reaction of rnicc and the ovormall degroe of exposure to Xray irradiation.
This contribution constituted the indispensabie prcr.quisit; for all
the subsoquent researchc:s whici? we conduct d on jaico.
The papers being paeblished i. the p~?esent symposium couotitutc
only a portion of the investigation On the effects of ionizing radiat 4 oi~
carried out by a team of assoeia'to5 o1 the cytoloical laboratory of
the Institute of Genetics of the Academy of Science USSR (N. I. Nuzhdin,
O. N. Petrova, d. N. ~itayov, a, M. V. Volkovich, and I. A. Nechayer),
and of the radiobiological laboratory of the ifstitute of Biophysics
of the Academy of Scio, ces USSR (N, I. Shapiro, A. M. Xuzin and Ye, N.
Eolodiy).
THE EFFECT OF DIFFERENT DOSAGES OF X-RAY IRRADIATIONS ON
THE SURVIVAL OF MICE
N. I. Shapiro and N. I. Nuzhdin
in recent years the attention of biologists has been attracted
to a steadily incx3asing extent by questions concerning the action of
ionizing radiations ~ i animal and plar;t organisms. This interest is
due riot only to the significance of the radiation method as concerns
t!~e study of the s L ucturand properties of l?vtrjg mutter, not only
to the fact that ionizing radiations are utilized in medicine on a wider
scale from one year to the next, but primarily and mostly to the prospects
which often open in connection with the possibility of utilizing atomic energy.
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illtnau; h w~tidy of JLuiogic~ a~L .oL vi I1Jt~ L1 t:~, c~t~autivl,u
il,itlatod u long tine ado al.d H vast aiuoul't of fac~uai data has
accumulated in this #i id, t just nc'ver't:cless be admitted that at tie
present tame we Lw v .cat ~1.iy 1Iu t is2 story 6ei. ral t leorr of
Luoloeal actio j X rWdicttio, it a ccnsiderablc po11t o1i c hu
data is deficient ant:i tlcre ox'c.' car.Ylot G41ways be utilized.
The unsatisi'actury riatuof a teeter portion of r'adic+!~io1 ;11ca!
researches, especiaiiy thu crIy ?s due prirnar~ly to in,dequatc:,
or eves: totally
a tL as concerns ap~;zt cxpui ~3.
addition tO dosirctri v dci: c :r,Cios ir,lot `o the rvaearcho i, r;V
determinationz were math 1' tic indE:.pensa ~l. utitativ~: poets
thc biological action. of 1iatior.. oreovc,r, J' w~ consider that rte~ry
of the problems whic i am; o::' immediate concer,i at the pi' :sent time w
"sit studied heretofore, i econ~cs evident w.y- .:~itrnsLChv use ca,i ..~;
rde of the 1 earlih n 7t .~ .a wh sy it ,~ 7. ~, ' 4' r? .~ . .y t ?!-o carry out
..Y' ga.. 7: which it was
reported that administration to mice of estradiol benzoate prior to
irradiation with Xrays increases the resistance of the mice to radiation
exposure (Treadwell, Gardner, and Laurence, 1943).
In 1949 two other contributions on this subjeot were published
(Patt, Strande, Tyree, Swift, and Smith, 1949; Graham and Graham, 1949).
- 25
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in addition to the confirmation of tiia results of the first corninuniea-
tion, in one of these contributions (Graham and Graham, 1949) factual
data are presented, and the conclusion is derived to the effect that
another estrogen, viz., diethylstilbestrol, is completely devoid of
protective properties. As a result, the impression is being conveyed
that the protective action constitutes a specific property of estradiol
benzoate and not of estrogens in general.
Such a conclusion appeared entirely unsubstantiated in view of
general biological concepts. Therefore, it was decided to carry out a
special investigation of estrogens as concerns their protective properties.
For this purpose, selection was made of diethylstilbestrol and synestrol,
which are synthetic estrogens widely utilized in medicine.
Determination of the Existence of the Protective Action of Synestrol
and Diethylstilbestrol
In testing the protective action of Synest.rol and diethylstilbes-
trol,,the objects of the investigation were male white mice of strain A.
Ten days prior to the Xray irradiation, the animals of the
experimental series were subcutaneously administered 0.2 mg of the estrogenic
substance dissolved in 0,2 ml of olive oil. The control animals were given only
olive oil. The single, general Xray irradiation was carried out under
the following conditions: voltage: 160 kv; current intensity 5 ma; filters:
0.75 mm Al + 0.5 mm Cu; focal distance 40 cm; dosage rate 12,9 r/min.
+otal dosage of exposure was 500r. Irradiation was carried out in wood
containers which held 12 mice of which six were of the everimental
series (having received the estrogenic substance) and six were of the
control series. Since our task was a study of the effects of estrogenic
-26-
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substances upon the vital stability of the organism in the presence
of damage inflicted thereto by large lethal doaages of Xrays, we
selected as the baoic index the one that was of greatest interest to us,
i.e., the survival rate. Observations of the experimental mice were
continued for one month after irradiation. Once every 4 days the animals
were weighed. Data on the effects of estrogenic substances on the
survival rate of the irradiated mice are shown in Table 1.
As is apparent from the tabulated data, both the preparations
investigated sharply increase the resistance of mice to the damaging
effects of radiation. Synestrol increases the survival rate of the
animals about one and a half times. Diethyistilbestrol exhibits a
considerably greater protective action, and yielded in all four series
of experiments good, well-defined results,
In Table 1 of the addendum (see addendum at the end of the
present paper) are shown the summative data on the time of death of the
animals following irradiation, for the experimental as well as for the
control series. These data reveal that the mean duration of life of the
mice which were given estrogen is somewhat less than that of the control
animals. Figure 1 shows the curves which represent the time of death
o:f the animals following irradiation.
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TAffiE 1
THE NUMBER OF MICE THAT SURVIVED AiQD T}IDSE THAT DEED IN THE EXPERIMENTAL (ESTROGENTREATKD)
AND TUE CON TROT, GROUPS
of
No of series and Total number
animals that survived
j,nimsls that died
Mature
treatment
designatIon of animals
Number
percent
Nuhnber
Percent
t
lt#
9
50 .0
s
50.0
7
Adminjst1at1on
1
Saperimen
3
16.3
15
83.
of $ynestrol
Control
1S
2
xperiment
:i3
ZZ
67.9
9
2g
32.1
69.4
Control
3#.,
21
~0.~
g riment
4,9
14
X3.2
1~
26
51.6
0
50
3
control
30
15
50.0
.
h4
7
1
42.8?_S.E4
^t i'1
44
+,..
5
.
I
For
Txperiment
ad
0
29
34.5?5.13
c: c
5:
65.515.28
all
Control
13
series
5
20.8
II
24
29
X9'2
istr$tion
1
~cperinent
35
0
1~
65.0
,n
of diethyl ..
Control
20
7
.
St3lbestrol
2
Experiment
24
23
g
95.5
32.6
1
15
4.2
67.4
Control
24
g
Experiment
24
17
70.8
7
a r.,
29.2
50.0
15
30.0
Control
41.1
gperiment
17
10
55.9
C
?0.2
Control
iS
5
2
b
89
69
77,5?4.4
2e
22.514.4
for
Experiment
z;
60.9?5.1
all
Control
92
36
39.1?5.1
series
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The results of observations on the course of radiation injury
in mice treated with estroenic substances and in those not treated
are indicative. The radiation injury had a considerably less acute
cours, in the experimental animals than in those of the control group.
Th18 was especially apparent in the series wherein diethylstilbestrol was
administered. The animals which had been given the estrogen had a
better appearance following irradiation than the controls; the behavior
of many of them differed but little from that of animals which had not
been irradiated. A milder course of the radiation Injury in animals
which had been given the estrogen is also confirmed by the data relating
to their weight, Table 11 of the addendum shows numerical data, and
Figure 2 shows the curves which characterize changes in the weight of
the animals during 30 days following the administration of estrogen.
To characterize the changes in weight of the irradiated animals we
used not the absolute values but an index (ratio of mean weight of
mice on a given day of observation to the mean weight on the day of
irradiation, which is taken as being equal to 100). A compa?ison of
the curves shows that decrease in weight of the animals of the
experimental group is somewhat less than that of the animals of the
control group.
Still more important is the fact that the animals which had
been given the estrogenic substances begin to recover appreciably sooner
than the controls. When the mice of the experimental groups begin to
gain in weight, those of the control groups still show a decrease in
weight. Thus, animals which had been given diethylstilheatrol reach a
day
minimum in weight on the second tc seventh/(on the sixth day in the
case of Synestrol) after which recovery sets in; among the controls
w 29 ?
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loss in weight continues to the twelfth to thirteenth day after which
a gain in weight begina. The dynamics of changes in weight of the
mice following irradiation indicates conclusively that the recuperative
processes take place more intensively in ariimals which had received the
estrogen.
For a more complete characteristic of the protective action
of diethylstilbestrol, analyses of the peripheral blood of the
irradiated animals were carried out. Four groups of animals were used
for the blood studies. Ten days prior to irradiation, the mice of tlac;
first group (36 animals) were given a subcutaneous injection of 0..2 mg
diethylstilbestrol in 0.2 ml of olive oil.. The seconl group (33 animals)
were the controls of the first group; those uticc received an injection
of 0.2 ml sunflower-seed oil 10 days prior to irradiation. Animals of
the first and second group were kept under observation for 10 days
prior to irradiation and for 47 days thereafter. The third group (lii
animals) was used to study the effects of diethylstilbestrol on the
blood in the absence of a subsequent irradiation. The fourth group
(16 animals) were the controls of the third group, and these animals were
given an injection of 0.2 ml olive oil without subsequent irradiation.
The animals of the third and fourth groups were kept under observation
for 28 days after the injection.
During the course of the experiment, records were kept of the
total amount of lenkocytes and erythrocytes, and the percentage of
hemoglobin content was deter?ined. Blood for analysis was withdrawn
from the caudal vein, and each time in six animals of the first and
second groups, and in four animals of the third and fourth groups
. 3o ..
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(ani exception were the ti- rtoonth and twenty?? f i rst day? following
i.rraclintion when three An mnls were used in each o1 the nxperirrlentai
l
and the control group). The hematological chungee in animals of all
four groups are ropi'es& ;tod rti,y- curves (Figures M,
data are shown in the addendum (Tf~ble, 3 and 4).
t, 3) ; r;umer.cai
ecause of the fluctuations of the blood-index values in the
nor,irradiated control animals, it is difficult to make a definite state-
~.aent concorning fire effects of die4hyltilbestrol on the blood of non-
.rradiated ai1iirrils, but the fact is apparent tit ' thr.~ blood indices of
animals whici hari been g vcn the cstrogeii are it all caso.a iowcr thm
those of t-ie control anir,; 1s . Maximum decrease ref the total nuxa er of
leukocytes in the ir-radi ited animals was observed on the fifth day
following the exposure, and i1iis decrease was less in mice which had
been given diethylstiloestrol) than in those of the control group.
:1rtximum decrease in the number of erythrocyte: was observed on the
ninth day following i.rradi;ati.on, and was also somewhat greater in the
control group than in the oxperir'iental group. A somewhat-increased
number of erythroc;? tes on the thirteenth day following irradiation n
the control group, ns comp sec with the experimentaai, may be of a forte toes
nature, since at this time only a few animals were investigated (three
mice in each control and e7:primentai grour,, as was stated above).
Restoration of the number of erythrocytes occurred more rapidly
i~ the experimental group than in the control group, attaining its
normal level on the twenty-first day following irradiation. At that
time the controls still revealed a lagging-behind in relation to the
normal values,
?
-3l
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Lowcr:.n~y then 11 uo .~oL,J it cotuat wa& (xsc)rved o~i tilc) ninth to
ti.irtevnth (buy foiiowirg ~.rradxatl.or., ad it waof a les:-or etdut ia;
Sao P.1-~i~7r Wt3;ktal group, wiicro:ix ..tom roatorati +a; also occurred i ore rapidly
t:la11 among tyre corn: u
In summing up the results of the effects of diethylstilbestrol on
the total amount of ioukocyt?s, erythrocytes, and the liemoglobin per-
centage in the irradiated :nice:, it can be stated that the lowi~ring c
the eaumorMatod indices dui g g he coursc " r?.sd .ation injury was 1s
cnuals protected by UL.C. ~vstrog ;n than ..: to cc k rol ax i;na].u.
Restoration occurred more r idly ii the oxp rinental group than ;:
cr n ; rots (;pOCICliy acCi:CUrfCd the w b4?r (~i k i'ytrocyto ~ a.ids l:e -
(labin peccontage). Thus tiro data obtainou leave uo doubt concerning ti1L
pos...t vc effort.; i estrogens subs tance:-, and especially of diet::y.w
stilbestrol, on the coarse of radiation it jury.
Correlation Between the Protewti ve infect and tine Dosr ge a$ Dieth i_
stilbestrol
To deter~nine th corn?elat. on betwee l tlio survival rate of tine
animal subjected to Xrny ia'radiation and the amoun 4 of estrogenic sab-
st nce adr~inistored to ttwii, ws have tested the a ct:ion of the foiiowikig
dosages of diethylsl ilbestrol. 0.025, 0.05, O.1, u.2, 0.8 mg. The
diethylstilbestrol was dissolved in olive oil and administered to the mice sub-
cutaneously l0 days prior to the Irradiation (each mouse was given 0.2 ml
of the oil solution of the estrogen). The control animals were injected
with a corroaponding amount of olive oil. The dosage of :Pray irradiations
was 500 r. The conditions of irradiation, the keeping of the experimental
animals, the nature and time intervals of observations were similar to those
described above. The results of the experiments are shown in Table 2,
32 -
0
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3
?rABLE 2
THE EPF tD DIETHYLSTILBESTROL P'OLIAWING IRRADIATION ON T1. SURVIVAL OF MICE
~'1' ~tI ' VARIOUS DOSAGES O
Goi00 r.
f
Illl ,`l~ ~/sz: V
--
c / ; - - l . / / \___-
?g"1? 1~ / /
o ,90 \ I' /
E \ '~
d ~ J
-!0 1 13 full 23 4
Days
,Xylr
ft: ' ~ lahin cortt,ent of the blood of irradiated and non-
irrr:vIiatcd mice on acininistration diethylstilbestrol.
1, otl; 2, di.rtbyisti1b' stro1; 3, oil 4 irradiation
5O.) r; i4, di.c thytilbcstral + i.rradta.tion 5O) r.
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)~. C11f1:'i3 in the
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00
I _- L
0 10 l0 X
Days of experiment
,
`-------?Z
0 18 20 Jo
Days of experiment
Figure 6. Life duration of mice following irradiation.
I - upon administration of 0.025 mg diethylstilbestrol;
II - upon administration of 0.0~ mg diethylstilbestrol;
III - upon administration of 0.1 mg diethylstilbestrol;
IV - upon administration of 0.2 mg diethylstilbestrol;
V - upon administration of 0.8 mg diethylstilbestrol.
1, experiment; 2, controls
0 10 l0 30
Doys of expprlment
I,
if
v D ro 28 X
`- Days of exreriment
i' w
o
'-`--------2
0 50
!1 ID /0
Days of experiment
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no 1 1 Fl, D ,t 110
.~
~:t
.
_rn, .~ _,%'
urd0
N loo
. -'
C !0
p
'N Ii 7M f Mt11f 1! IY-~ Oil 7 tcft Mfr 1311 -a 0g 7 p $ f sad0
D a y s of e x p e r t men t
IF
"0
n
d
u 9!D
;1B SAO
G
c~ -70~
l 1 , . 1
P-% ,z 1 lJ n i,1f 1~ -8 81 7 /I / r tr 2719
Days of experiment
Figure 7. Changes in weight of mice following irradiation.
I - upon administration of 0.02 mg diethylstilbestrol;
II - upon administration of 0.0~ mg diethylstilbestrol;
III - upon administration ot'0.1 mg diethylstilbestrol;
Iv ? upon administration of 0.2 mg diethylstilbestrol;
V . upon administration of 0.$ mg diethylstilbestrol.
r ,
1, experinient; 2, control.
. 65 ..
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 a?
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50
/0 20 30
Doys
N
to :o jo ? a' '9 1y
Days Dove
Figure 8. Life duration of mice following irradiation.
I . upon administration of diethylstilbestrol one day
prior to X-ray irradiation; II - upon administration of
diethylstilbestrol 3 days prior to X-ray Irradiation;
III - upon administration of diethylstilbestrol 5 days
prior to X-ra.y irradiation; IV upon administration of
diethylstilbestrol la days prior to X-ray irradiation;
V - upon administration of diethylstilbestrol 12 days.
prior to X-ray irradiation; VI - upon administration
of diethylstilbestrol 15 days prior to X-ray irradiation.
1, experiment; 2, control.
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9 !0 ZO
Days
foe
50
00 0
foeC.
'1
2
-_________
if
1
,
---..t
_L_.__
10 20 30
Days
' `-.
50
0 a Po 30 0
-01 043R0002001 80025-9
^
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1OO
?, ~:=-=~~t 100
05
117
_j__ L a.. Ll~t _. I
r0 15 10 23 30 Doys 10 S 0
100
80
_1._L__4___L- I--
5 10 15 ?fl t5JB Days
~l._, 1 11 1 i 1 t i 1
58 5 /0 15 20 25 X Doys 15 l0 5 b 3 r0 15 t0 ZS J0 Doys
10
18
,17
vi
.
3 0 5 10 1s 20 25 30 Days is 10 J 0 3 /0 /5 20 83 30 Days
Figure 9. Changes in weight of mice following irradiation.
I - upon administration of diethylstilbestrol one day
prior to X-ray irrad:iat,ion; Ii - uOon a(tmi.nistration
of diethylsti 1bestro] I day~3 prior to X-ray irradiation;
ill - uoon administration of diethy1sti1he5troi 5 days
prior to X-ray irradiation; IV - upon administration of
diethylst,i.ibestrol. 10 days prior to X-ray Irradiation;
V - upon administration of diethylstilt,e ,trod 12 days
prior to X-ray irradiation; Vi - upon adminic;tration of
diethylstilbestrol 15 (Jays prior to X..ray irradiation.
1, exp?. riment; 2, coritr. olo
07'.
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'Co
p ,, ZO JO Do y s
a SOD
C
'`-----~
.- 50 _---~,__ Z
r
7
0(10 )J ZO 30 Doy*
111
v 0 j 20 Doys
Figure 10. Life duration of mice following irradiation.
I ? upon introduction of pellets contaning diethylstil-
bestrol. 10 days prior to X-ray irradiation; II - 20
days prior to X-ray irradiation; III - 30 days prior
to X-ray irradiation.
1, experiment; 2, control.
'loo
.. 50
0
Figure
11
0 10 ZO 30 Days
p
'---------2
Lire dust: c n
10 ZO 37 Days
of lUC?? foilowing irr,s.~i tic~n upon repeated
a:dmiriistratinrt of (1iethyl5tllhf stro1 (series I and II ).
a., experiment; 2, cont,roi.
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P
C
S *1 0 m IS
Dore of experiment
Figure 12. Life duration of mice following irradiation upon adminstra.
Lion of diethylstilbostrol in conjunction with progesterone.
1, diethyllstilbestrol; 2, diethylstilbestrol + progesterone;
3, progesterone; Li, control.
O
C
-_-- --.----
a
--------------
a S XI 20 15 7
Days of expert me nt
Figure 13. Life duration of mice following irradiation upon adndnstra.
tion of diethylstilbestrol in conjunction with pregnenolonr
1, diethylstilbestrol; 2, diethylstilbestrol + prenenolon;
3, pregnenolon; 14, control.
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,2of
Figure 11j.
ftO
W
-to -5 D 5 !O 15 ZO 25 50
Dots of experiment
Chant;:; in weight of mice following irradiation upon
administration of diethylstilhe trot in conjunction
with pregnenolon,
1, diethylstilbestrol; 2, diethylstiibestrol 4
prepnenolon; 3, pregnenolon; 11, control.
tic,
70
-18 -s O s ra l5 Zo 2 3O
Days of experiment
Figure 15. Chan~Yes in wei cht of mice foL owing; irradiation upon
administration of d.iethy1st1lbestro7 in conjunction
with proest,erone.
1, dlethvlsLi.i1 estrol; 2, diethylstilbestrol +
prole:>terof( ~, pro :teror~e; 14, control.
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THE HOLE OP THE PHYSIOLOGICAL STATE OF TN$ ORGANISM ON
UTILIZATION OF PAOTNCTIVE RMDIES AGAINST TAE DAMAGING
ACTION OF PENETTMTII'N~- RADIATIONS
N.
I.
A.
Shapiro,
M. Bu$in
and N.
I.
Nushdin
Among the contributions concerned with the biological action
of penetrating radiations, there are a sufficiently large number of
investigations which indicate a correlation between the radioaensi-
tivity of organisms and their physiological state. This correlation
has been repeatedly demonstrated in the case of plant as well as of
animal forms (Dugger, 1986). In some researches the role of the phy-
siological state has been revealed upon the study of individual varia-
tions in the radiosensitivity of the irradiated objects, in others on
observation of organisms at different stages of embryonic or postembryonic
development, and finally in still other researches by means of alterations
of the state of animals or plants due to the effects of environment. The
facts which show a correlation between radiosensitivity and the physio-
logical state of the organism are of exceptional importance primarily as
substantiation of the posaibility:of an active intervention in the reaction
of the organism to radiation exposure. In other words, the facts which
show a correlation between the radiosensitivity of an organism and its
physiological state constitute a certain experimental substantiation of
the possibility of finding substances which protect the organism against
the damaging action of penetrating radiations. From the ascertaining of
these facts also follows still another most important consequence, the
Significance of which is at the present ti?e clearly underestimated.
Since the physiological state of the organism affects its radiasensitivity,
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It is quite natural that this state must also affect the action of many
protective agents inasmuch as it is uuquestionable that many of these
substances exercise their effects by altering the physiological state of
the animal or plant being protected.
All of the foregoing apparently indicates the necessity, in searching
for and undertaking the study of substances which protect biological objects
from the damaging action of radiations, of taking into consideration the
physiological state of the organism. It is entirely probable that effect-
iveness of many protective remedies will depend upon the physiological
characteristics of the animals undergoing exposure. Yet in the overwhelming
majority of investigations conducted in this field, the physiological
state of the experimental animals is not taken into account. Such an
abioiogical approach constitutes perhaps one of the most characteristic
features of the current researches which are being conducted abroad on
the study of the protective action of various chemical substances.
The abiologism is further heightened by the nature and dosages of
chemical substances which are being tested in this work. At the present
time intensive studies are being carried out on the protective action of
such strong poisons as, for example, the cyanides (Herve and Baoq, 1949), or of
huge doses of less harmful substances such as, for instance, cysteine,
reaching the limit of tolerance (Smith, Patt, Tyree, and Straube, 1950).
Thus, the researchers operate with means of such potency in the presence
of which it is not always, by far, the case that the physiological state
of the experimental animals can be of any significance. The action of
substances of this kind has no relationship whatever with a utilization or
furtherance of the protective mechanisms of the organism itself and con-
stitutea rather something extraneous to the irradiated obect.
.72.
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Without denying the importance of the work on uncovering
protective means, the mechanism of the action of which is not associated
with the protective reactionsinherent to the organism itself, we believe
that a limitation of the scope of researches to such strongly acting
means would be erroneous. There is no doubt that means which enhance the
protective characteristics of the organism itself should be of no lesser,
but possibly of even greater, significance.
in this connection, we consider it of special importance not only
to substantiate theoretically the necessity of taking into account the
physiological state of the organism, but also to show experimentally
the correlation between this factor and the protective action of any
given preparation. We have selected diethylstilbestrol which, according
to previously derived data,was found to exercise a good protective action
upon the irradiatiu:. of mice (ace preceeding communication of the present
symposium (Shapiro, Nuzhdin, Kuzin, 1955)).
Although the mechanism of the protective action of diethylstilbestrol
as yet remains uncertain, there is no doubt that it is associated with some
specific effects of estrogenic hormones on individual organs and tissues
of the animals.
According to previously conducted experiments diethylstilbestrol
increases approximately twofold the rate of survival in mice exposed to
lethal dosages of Xrays (Shapiro, Nuzhdi, guzin, 1955). Table 1 shows the
data on the survival rate of mice following a single Xray irradiation with
a dosage of 500r. The conditions of irradiation (in this instance, as well
as in all subsequent experiments) were as follows. Voltage 160 kv; current
intensity 5 ma; filters 0.75 mm Al +0.5 mm Cu; focal distance 40 cm; dosage
- 73 M
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intensity 15.3r/min. The irradiation technique has been described in
detail in one of our papers (Shapiro and Nuahdin, 1955).
Ten days prior to the Xray irradiation, each mouse was given a
subcutaneous injection of 0.05 mg diethylstilbestrol dissolved in 0.2 ml
of refined vegetable oil. The control animals received only the oil.
Sexually mature mice of strain A, not yet having propagated, aged 2-3 months,
were used in this experimental series. The initial weight of the animals
varied between 20 and 24 g. Observations of the experimental mice were
continued (in this instance and in the other experiments) nor 30 days.
The data of Table 1 leave no doubt concerning the protective action
of diethylstilbestrol which manifests itself to an equal extent in female
and male animals. Thus, the results obtained appear to indicate that the
action of this preparation can hardly depend extensively upon the physio-
logical state of the organism, since even such a great difference as exists
between males and females does not alter its effectiveness. Nevertheless,
we have made an attempt to determine the conditions which change the results
of the action of the preparation under study. Since we are dealing with
an estrogenic substance, it could be expected that its efficacy should
depend upon the general hormonal level of the experimental animals and
the associated therewith of specific changes in the functional state
of the different organs. Considering that the factor which alters
drastically the physiological state of the entire organism and in particular
its hormonal conditions is the begetting of offspring in females, a compar-
ison was made of the protective action of diethylstilbestrol in virgin
females and those which had previously propagated. A preliminary determina-
tion was made of the radiosensitivity of these two types of animals,
A
irrespectively of any supplementary influence.
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rv-
TABLE 1
&FPFCTS OF ? PROPHYLACTIC ADMINISTRATION OF DIETHYLSTILBESTROL Old 3IIRVIYAI. RATS
OF g~J-i,$ AND MALE ![ICS Ole STRAIN A FOLIAWII~ TOTAL PRAY IRRADIATION (DOSAGE
OOx)
Object
Treatment Total ntsber Survived Died
Itsaa lire
duration
of animals
Number Percent Number Percent
(days)
]P'easles
Diethylstilbestrol 45 39 64.4?9.1 16 35.6#7.1
ia.s
Control 48 14 29.2?i.4 34 ?0.8?6.4
9.8
Males
Diethylstilbestrol 89 55 62.915.1 34 37.115.1
10.3
Control 102 29 28.514.4 73 71.5_*4.4
e.s
c
I
TABLE 2
RADIOSEN3ITIVITY OF VIRGIN AND OF PARENT FEMALE ffiZCE OF STRAIN A (DOSAGE OF EXPOSURE, 500r)
Object
Total nwbes
of animals
Survived
Died
Ydift mdif
(967L)
!less life duration
(days)
V>;rgin
peygeg
48
Number
14
Percent
29.26.6
Number
34
Percent
70.816.6
9.8
29.1?10.5
Bred ye~ales
36
21
58.318.2
ZS
41.7?8.2
13.1
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The results of experiments on the survival of virgin female mice
and those which had propagated (twice), of strain A, following Xray
irradiation (dosage 500r) are shown in Table 2, from which it is apparent
that the radiosensitivity of virgin females is greater than that of the
females which had propagated. There are reasons for assuming that the greater
physiological stability of females which had given birth is not selective,
as concerns the harmful consequences of Xray irradiation, but can be revealed
also as concerns other unfavorable conditions.
The testing of the protective action of diethylstilbestrol in virgin
females and those which had propagated was effected in the following manner.
Ten days prior to Xray irradiation (dosage, 500r) each female of the
experimental group was given a subcutaneous injection of 0.05 mg diethylstil-
bestrol dissolved in 0.2 ml of vegetable oil. The control animals received
the same amount of oil. Table 3 shows the results of this experiment.
The tabulated data show that the protective action of diethylstil-
bestrol, which is so clearly manifested in the case of virgin females, is
practically absent in the case of the females which had previously propagated.
Thus, the relatively high resistance of the females which had previously
propagated cannot be enhanced to any appreciable extent by an administration
of diethylstilbestrol. From the data presented, it is also apparent than
an administration of diethylstilbestrol to virgin females increases their
radioresistance only up to the level which is characteristic of the females
which had previously propagated. In this connection it is also important
to bear in mind the fact that the protective action of diethylstilbestrol
in males is quantitatively similar to that in virgin females (See Table 1),
and that the level of radioresistance in mt_les after they have received
diethyistilbestrol is also proximate to the level found in females which
had previously propagated.
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TABLE 3
SPP&.T3 OF DI~THYL4TILBSSTR4L (0.05 mg ) ON THS SURVIVAL RATS OF VIRGIN 1QCS AND THOSE WUII HAD
PROPAGATED, OF STRAIN A, FOLiAMIHG TOTAL ERA7f IRRADIATION
(DOSAGE 500r)
Object Treatssnt Total number Survived Died
of animals
virgin
female Diethylstilbestrol
Control
Females
which had
propagated DiethylstilbeStl'Ol
E:ontro'
r Percent Number percent
Nuimbe
'bi:t Halt
moan life
duration
(days)
45
29
64.437.1
16
35.617.1
12.9
35.219.7
48
14
28.216.6
34
70.816.6
9.8
35
23
65.718.0
12
34.318.0
12.6
7.4111.5
36
21
58,3?8,2
15
41.718.2
13.3
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On the basie of the comparison of all the data obtained itia
natural to aeeume that the difference in radiosensitivity between virgin
females and those which had propagated, on the one hand, and the feaal~e
which had propagated and males, on the other, moat be attributed to a
difference in their hormonal (estrogenic) level. This assumption can
also be confirmed by facts well known in experimental oncology (Grinshteyn,
1831). The development of the cancer of maimaary glands in mice requires
on the one hand a cancer virus (the so-called lactary factor), and on the
other the action of estrogen hormone. Only the simultaneous presence
of both these components determines the development of neoplasms. In
this connection, in mice of strain A which are characterized by the presence
of the lactary factor, the breast tumors occur only in females which have
propagated. In order to induce the occurrence of tugs in males or females
which have not propagated, of this strain of mice, it is necessary to ad-'
minister additional estrogenic substances.
Thus, these data also indicate that in females of strain A which
have propagated the level of estrogenic hormones is much higher, not only
as compared with the males but also as compared with the females which had
not propagated. Since, after receiving estrogen, the males and virgin females
become similar in radiosensitivity to the females which had propagated, it must
be assumed that in this instance increased radiostability under the action
of an estrogenic hormone is governed to a certain extent by the so-called
law of "all or nothing." In other words, the attainment of a definite
level of resistance requires a definite minimum of the hormone; additions
thereof in excess of this amount are practically useless. This standpoint
is supported by the results of our previous experiments on ascertaining the
protective action of different dosages of diethylstilbestrol. Ten days
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prior to Yray irradiation (dosage 500r), male mice of strain A, 2 to 3
months of age, were given subcutaneous injections of different amounts
of diethylstilbestrol dissolved in vegetable oil. The data thus obtained
are shown in Table 4.
On examining the data of this table it is necessary to bear in mind
that, since in our investigations a fairly large variation was observed in
the death rate of the irradiated animals from one experiment to another,
controls were used concurrently with all the experimental series. To
facilitate a comparison of the protective action of various doses of
diethylstilbestrol, an index was computed, which we have designated as
the survival index and which constitutes the ratio of the percent of
animals that survived in the experimental group to the percent of animals
which survived in the control group.
The data listed in Table 4 show that a tenfold or more increase of
the diethylstilbestrol dose does not result in corresponding increase of the
resictance of the mice. A somewhat higher survival rate of mice following
the administration of 0.8 mg diethylstilbestrol is not statistically reliable.
Thus, on the basis of these data we also arrive at the conclusion that the
protective action of diethylstilbestrol is governed in practice by the
socalled law of "all or nothing,"
All the above cited data leave no doubt that the protective action
of diethylatilbestrol depends on the physiological state of the organism
subjected to irradiation, and as we have shown, in particular on its hormonal
conditions. The question thus arises as to whether the presence or absence
of tfe protective action of diethylstilbestrol is associated only with the
hormonal conditions.
.. 79 ..
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TABLE 4
EFFECTS OF DIFFERENT DOSES OF DIETHYLSTILBESTROL ON THE SURVIVAL RATE OF MICE OF STRAIN A POLI.OAING
I
~o
C
1
Dosa of
diethylstilbestrol
(1n ag}
0.2
0.8
IRRADIATION (DOSAGE 500r)
Experiment
Control
Survival
Total Survived
number
Died
Total Survived
number
Died
Index
of ant-'
of ani-
orals
Number
Percent
Number
percent
male Number percent
Number
percent
[2]
[3] [4]
[5]
[s]
[7]
[s]
[a]
(lo]
f11l [12I
63
45 71.4f5.6
18
28.6?5.6
54 19
35.2t6.5
35
64.8?6.5 2.03
88
56 62.915.0
33
37.115.0
102 29
28.5?4.4
73
71_5?4.4 2.20
40
31 77.5t6.6
g
22.5#g,g
40 14
35.0?7.4
26
65.Of7,4 2.22
89
69 79.5?4.4
20
22.514.4
92 36
39.1?5.1
56
60.9?5.1 1.98
33
24 72.7?7.7
9
27.3?7.7
34 8
26,5?7.6
25
73.517.6 2.74
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aw,
A certain answer to this question is provided by the results
of our szperiments concerned with the determination of the effects
of diethylstilbestrol on mice of strain Cg7 (black). The mice C57
(black) are extensively used in experimental ontology. They are character-
ized (in contrast to strain A) by the absence of the so-called lactic
factor which induces the occurrence of cancer of the mammary glands.
In addition, a number of researches have established numerous physio-
logical and biochemical differences between these two strains of mice
(Grinshteyn, 1951), of special interest to us is the investigation which
has shown that screening of the spleen during total Xray irradiation sharp-
ly increases the survival rate of the animals of strain A and, to a much
lesser extent, that of strain C57 (black) (Kaplan and Janice, 1952).
Studies of the strain C57 (black) were initiated with a determina-
tion of its radiosensitivity and a comparison with the radiosensitivity of
mice of strain A. In the investigation, use was made of mice, 2 to 3
months old, which had not yet propagated, weighing from 20 to 24 g. The
results relating to the survival rate of mice of both strains during the
30 days after a total Xray irradiation (dosage 500r) are shown in Table 5.
-81-
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TABLE 8
BADIO:fBlSITIVITY OF MICE OF STRAIN A AND Cgq (BEACH) (DOSAGE OF EXSURE, 500r)
Str,* i
r
Total
amber
Survived
Died
Mdi4?mdif
~II lift
duratim
of sai-
('X9L)
(days)
Fatales
eels
48
Number
14
Percent
29.2?6.6
Number
34
Percent
70.8?6.6
8.8
20.519.7
Cgs (black)
48
24
50.0t7.2
24
50.017.2
15.8
R
Melee
47
15
31.916.8
32
88.1?6.8
9.6
24.229.4
(black)
57
32
56.116.6
25
43.8t6.3
9.0
I
Males and
95
29
30.514.7
66
69.5?4.7
8.7
i
Cyr (black)
females
105
56
53.3?4.9
49
46.714.4
12.4
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As is readily apparent frog the data listed in Table 5, the
radiosensitivity of male and female mice of strain C37 (black) is
lower than that ;of the axles and feaalea of strain A.
Tests of the protective action of diethylstilbestrol on mice of
strain C57 (black) were started with a dose of 0.05 mg. Dissolved in
vegetable oil, the prparation was adainistered subcutaneously 10 days
prior to irradiation. The data of Table 6 show that administration of
this amount of diethylstilbestrol had only a fairly slight protective
action. At the same time, upon administration of a larger amount of
diethylstilbestrol (0.2 mg), its effects were clearly manifested.
Thus, diethylstilbestrol protects mice of strain C57 (black)
from the damaging action of Xray irradiation upon utilization of somewhat
greater doses than in the case in regard to animals of strain A. Evidently
the physiological characteristics of mice of strain C57 (black) are such
that, in spite of their higher radioresistance, the protective action
of diethylstilbestrol is manifest in them to a lesser extent. It is
possible that the reverse relationship observed between the natural
radioresistance of mice and their ensitivity to diethylstilbestrol is
far from being fortuitous.
Adverting to the que$tton concerning the causes of the different
effects of the estrogen on mice of strains A and C57 (black), it can only
be. assumed that these causes are different from those which we have en-
countered in studying the action of the preparation on virgin females and
those which havo propagated.
g,2 ..
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TABLE 6
nCfS OF D2ETHY7.STILHBSYibf3L ON THE SHRVIVAL RATE OF YALE TICS OF STAINS A-ABiD C57 (BIdCE)
gpLLpfING TOTAL SsAY IRRADIATION (500x)
Strain
Dose of
diethyl-
stilbestrol
'Dotal
n~ber
of sai-
Survived
Died
ydiY?mdif
(sssc)
11eau life
duration
(days)
awls
Number
Percent
Number
Percent
0
05
sa
56
62.915.0
33
37.1?5.0
10.3
.
34.436.7
Control
loa
29
28.5?4.4
73
71.b#4.4
9.6
a
3
10
0
2
89
69
77.519.4
20
22.514.4
.
.
38.4?B.7
Co
Control
92
36
39.115.1
56
60.83.1
11.2
0
05
37
27
73.017.3
10
27.0t7.3
12.1
.
11.2?9.6
C57 (black)
Control
34
zi
61.8t6.3
13
38.2?6.3
10.5
0
2
22
is
si.s?s.z
4
18.2?8.2
18.7
.
34.0113.3
Control
23
12
47.8?10.4
12
sa.s?io.4
7.4
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Theresa in the latter case, the difference can be attributed to the hormonal
(estrogenic) levels of the two groups of animals referred to, this cannot
be said sa conoerns the mice of strains A and C57 (black). Sven on assuming
that the estrogenic level of females of the C57 (black) strain is higher than
that of females of the A, we cannot possibly consider that the estrogenic
level of males of the C57 (black) strain is higher than that of virgin
females of the A. Thus, the specific features of the reaction of animals
of strain C87 (black), as compared with the animals of strain A, to the
administration of diethylstilbestrol must be attributed to some physiological
differences which are not directly related to the hormonal level of the animals.
In other words, the means whereby the protective action of diethylstilbestrol
is effected evidently depend on the Ahysialogicsl characteristics of the
organism in the sufficiently wide meaning of this term.
In connection with the establishment of the facts which indicate
that diethylstilbestrol increases the radioresistance of the organism within
the limits of the existing physiological norms, it is important to discuss
the question concerning the maximum dosage of Bray irradiation at which the
estrogen retains its protective properties. It was natural to assume that,
if the preparation does not increase the radioresistance of the animals be-
yond that which is encountered under uormal conditions, it should also not
protect the organism in cases of exposure to absolutely lethal dosages.
As was shown in another paper (Shapiro and Nuahdin, 1955) the minimum absolute-
ly lethal dosage of Brays in the case of male nice of strain A is 700r.
Three small series of experiments were carried out with a dosage of 700r,
which yielded fully conclusive results. In the first series 18 male mice. of
strain A, which, 10 days prior to the exposure, had been given subcutaneously
0.05 mg of diethylstilbestrol each, were irradiated. At the same time
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five control aaia*la;were irradiated. All the mice ?- the experimental
as Nell as the controls -- died (the mean life duration of the mice was
11.3 days in the experimental group and 6.5 days in the control group). In
the second series the dose of diethylstilbestrol was increased to 0.4 mg.
The irradiation was carried out on 12 males which had received diethylstil-
bestrol and 12 control males. As in the first series, all the animals died
and at approximately the same intervals of time (the mean life duration
of the mice was respectively 6.2 and 5.5 days). Finally in the third series
use was made of the most resistent group of animals, the females of strain A
which had previously propagated. Ten days prior to irradiation, 12 females
were administered 0.2 mg of diethylstilbestrol, and 12 females wore irradiated
without any preliminary treatment. In this instance, as well, all the
animals died (there was a mean life duration of 9.0 days in the control
group, and of 8.2 days in the experimental group).
Thus, on summarizing the results of these experiments, it can be
stated that diethylstilbestrol does not increase the resistance of the
animals to such extent that they are capable of withstanding the deleterious
effects of irradiation in the presence of an absolutely lethal dosage.
The experiments have shown that dosages of Xray irradiation at which diethyl-
stilbestrol exercises its protective action do not exceed the limits of
those at which, even under normal conditions, i.e., without an administra-
tion of the hormonal preparation, a sash1 portion of the mice survive.
It is possible that diethylstilbestrol, while protecting some organs of the
animals, has no effect on other organs, and therefore death occurs as a
result of the damage to the latter. Thus, all the data cited in the present
paper indicate that diethylstilbestrol is s preparation the action of which
is aasociated with the protective mechanisms of the organism itself; there?
from follow all its positive and negative properties.
..
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The materials which we have considered, relating to the role of the
physiological state of the organism upon the use of diethylstilbestrol, evoke
a number of general questions which are significance, in principle, as
concerns the search for and the study of substances which protect the organism
from the damaging action of penetrating radiations. First of all, it is
desirable to provide an answer to the question as to what significance may
be attributed to the study of substances the action of which is directly
associated with the protective mechanisms which are inherent to the organism
itself. Evidently in such instances we will encounter, as a rule, an in-
crease of the radioresistance of the organism only within the limits of lethal,
but not absolutely lethal, dosages of exposure. In addition, the efficacy of
such preparation may depend to a large extent upon the physiological state
of the objects subjected to irradiation.
The above-stated, seemingly unfavorable circumstances notwithstanding,
we believe that the study of substances intended to enhance the protective
properties of the organism itself is of great interest not only theoretically,
but also practically. It is unquestionable that in the course of investiga-
tions on this kind of preparations there may be ascertained the concrete
physiological nature of the so-called radiosensitivity of the organisms.
Up to the present time, the widespread use of this concept in radiobiology
notwithstanding, it is to a considerable extent devoid of a concrete physio-
logical meaning. The importance of a cognizance of the nature of radio-
sensitivity is further emphasized by the fact that this constitutes a
concomitant cognizance of the concrete means of the action of radiation
upon the organism. Of no lesser importance may also be the study of sub-
stances of this kind in the practice of safeguarding the organism against
the damaging action of penetrating radiations. it is quite natural that one
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must learn how to protect the organism from radiation daaage not only
on exposure to absolutely lethal dosages but also to ordinarily lethal
dosages. At the ware time we believe that protection of the organism on
exposure to absolutely lethal dosages also requires the discovery of
preparations which enhance the natural radioresiatance of the organise.
Radiation injury in animala, arising on their exposure to large dosages
of radiation, constitutes in practice a result of deaagee to ail organs
and tissues. Hence, both preventive methods and treatasnt procedures in
regard to this disorder will be of a complex, composite nature. There
are no reasons for hoping that in this instance it will be possible to
find some kind of single, universal therapeutic remedy. Thum, the
composite control of radiation injuries requires the utilization of the
most diversified preparations. AmOng these not the least position, by
far, will be allocated to those preparations which enhanca the protective
properties of the organisms itself.
BIBLIOGRAPHY
GrunshteYn, Dzh., Biokhimiya raka [Biochemistry of Cancer), 1951, Moscow,
Inoizdat, 1-394
Shapiro, N. I.. Nuzhdin, N. I., Buzin, A. M., Deyatviye edtprogennykh veah?
chesty na luchevu u reaktsi sbe (The Effect of Estrogenic Sub-
stances on the Radiation Reaction in Mice), cOllectian of works on
radiobiology, 1966
Shapiro, N.
zbivae~oat' tEffect of Various Doses of xray
Irradiation on the Duration of Life of Mice), collection of works on
radiobio1og7, 1965
Nuzhdin, fit. I., V1i i e ramlichn kb dos ,rent ohovak
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Duggar, B. L, Biological Effects of Radiation, 1936, McGrai-Hill,
I, 1-676; II, 6771343
Hervs, A., Hacq, Z., cyanure et dose lethale de rayons X, Compt. Rend. Soc. de
Biol., 1949, CXLIII, 881.883
Kap1$, U. S., Janice, P., Genetic Modification of Response to Spleen Shield-
ing in Irradiated, Proc. Exp. Biol. and Med., 79 (4), 670.672
Smith, D., patt, H., Tyrre, S., Straube, a., f,~uantitatiye ASPects of the
protective Action of stein Against x-Radiation, Proc. Exp. Bid. and
Med., 1930, 73, 198-200
CONCERNING THE ROLE OP DAMAGE TO HEMATOPOIETIC ORGANS IN THE
COURSE OP RADIATION REACTION
N. I. Shapiro,
N. I. Nuxhdin,
M. A. yolkovich,
Ye. N. Kolodiy
INTRODUCTION
One of the most characteristic features of the damaging action of
penetrating radiations inflicted to mammals under conditions of a total
irradiation in the disruption of the structures and functions of all the
systens of the organs and tissues of the animal. However, it is well known
that, in spite of such total damage, not all the tissues of the organism,
by far, are equally radiosensitive. In other words, different organs and
tissues of the irradiated animal by far do not become damaged to the same
extent. The urgent necessity arisasof ascertaining the role of the damage
to individual organs in the course of the radiation reaction. This analysis,
in addition to its theoretical significance in determining the nature of the
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radiation reaction in mammals, aty be also of a more direct practical value
in connection with the necessity of developing effective procedures for the
treatment of radiation daaages.
At the present time there are two principal methods for ascertaining
the role of the damage to individual organs and tissues during the course
of the general radiation reaction in mammals. The first method is a local
irradiation of the aniaais (and correspondingly a screening of individual
organs on general irradiation), and the second is the implantation of in-
dividual organs and tissues to animals having been exposed to radiation.
By mutually supplementing each other, both these methods permit the deriving
of the correct solution of the postulated problem.
In studying the role of the damage to different organs in the course
of radiation reaction, the researchers devoted special attention to ascertain-
ing the significance of damage to the hemopoietic system. This concern is
due on the one hand to the exceptionally important role of the blood and
heaopoietic organs in the vital activities of the animals, and on the other
to their high radiosensitivity. As was shown by numerous investigations,
the heaopoietic tissue is one of the most radiosensitive tissues of the
organism (Yegorov and Bochkarev, 1950).
Leaving out of consideration the numerous morphological investiga-
tions concerned with the study of the changes in the blood and hemopoietic
organs under the influence of ionizing radiations (see the synopsis of this
work in the book of Yegvrov and Bochkarev "Krovotyoreniy'e i ionixiruyunhchaya
(Eematopoiesia and Ionizing Radiations], 1950), we will mention the
contributions in which a study was made of the role of the heaatopoietic
organs in the radiation reaction by a screening of these organs during the
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general irradiation of the aniallis or by the implantation of heaaopoietic
organs of nonirradiatad animate to the irradiated animals (Jacobson,
Marks, Robson, Gaston, and Zirkie, 1949; Mandart, Lambert, and Maisin,
1982s, 1952b; Barnes and Isnatit, 1953; Laugendorff, Koch, and Sauer, 1954).
The overall result of these investigations, the objects of which
were in most instances small rodents (mice, rats), was a conclusaion as to
the very important role of damage to the hemopoietic organs in the course
of the radiation reaction in mammals, In spite of the unquestionably cone
clusive nature of this deduction, it is necessary to note that a number of
features involved in the damage to the hemopoietic organs and their corre-
lations with the most important aspects of the course of the radiation re-
action have remained unelucidated.
One must take into account the procedural difficulties which are
encountered in the study of the problems under consideration. It is
prmcisely these difficulties that are responsible to a considerable extent
for the great variations in the experimental results of different researchers.
The latter applies to the data secured on screening the hemopoietic organs of
the animals during irradiation (Kaplan and Janice, 1952), as well as on the
transplantations of these organs (Barnes and Loutit, 1953; Laangendorff,
Koch, and Sauer, 1954), Variations in the results were found to be so prom
nounced that they have been the subject of a special research in which the
author has studied the influence of the hereditary features of sex and age
of the experimental animals on the curative effects of transplanted spleen
(Cole and Ellie, 1853).
In carrhifg out the study of the regularities of tht diction re-.
action in mats, aid in particular, while determining the significance of
the damage to specific organs as concerns the course of thin reaction,
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we also had to investigate the role of the hsaopoietic organs. The
present paper is devoted to the description of the resulte of experiments
carried out in this direction.
Material and Method
The material consisted of male and female mice of strain A, aged
from 3 to 3 months. The mice were subjected to Xray irradiation under
the following conditions; voltage 180 kv; current intensity 5 ma; filters;
0.5 Cu + 0.75 A1; focal distance 40 cm; dosage 18.3 r/min. Observations
of the irradiated aice were continued for 30 days following the exposure.
Determination was made of the time of death of the animals, changes in
their weight, andte general course of the radiation reaction. In a nusber
of cases observations were conducted on the peripheral white blood compon-
ents. Weighing of the experimental mice was done every fourth day, and
blood samples were taken once every 5 days.
The investigation consisted of three parts. In the first, a coapari-
son was made of the course of radiation reaction in the totally irradiated
wales and fealales with screened spleen or bone marrow gear extremities).
Screening was done with lead plates 3 thick, in the manner shown in
Figure 1.
Withdrawal of the spleen was effected without narcosis of the animals
operated upon. The mouse was fastened supinely to the board. At the left
upper portion of the abdomen an incision about 1 cm in lemgtheM asde.
Through this incision the spleen was withdrdwn, pulled aside, and placed
in a sterile gauze bandage moistened with physiological solution and held
in a sasll lead cell. During this operation the spleen is not damaged and
its conaeation with the organism is not disrupted, in particular all of the
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blood vessels remaining intact. This operation was perioraed on the
aniaale of the experiaental series (on screening of the spleen), as
well as on the controls (without screening of the spleen). On coapletion
of the irradiation, the spleen was put back into the abdoeainsl cavity and
the incision was sutured. Screening of tho bamr marrow was effected by the
covering of one or both rear extremities.
In the second part of the investigation the transplantation of the
spleen from nonirradiated mice to the irradiated was carried out, In these
c9503 the irradiation was carried our, using special containers which could
hold 12 animals. In so doing, the focal distance was reduced to 20 cm.
The dosage was 7.4 r/min. Total dose of exposure was JOOr.
We have tested a numter of procedures ~yyleon transplantation.
In some cases, a spleen taken from an adult mouse of the same strain was
transplanted during the first hour following irradiation to male and
female mice of strain A. In transplantation the vessels of the spleen of
the donor were tied up, and the transplanted organ was sewed to the epiploon
of the recipient. In the animals of the control series pieces of the epiploon
of the donor mice were sewed on. In other cases, from one to three spleens
of 8-22 day old mice were transplanted to the males during the first hour
following irradiation. The operation technique is the same as in the pre-
ceding oases. The use of these procedures of transplanting the spleen did
not yield the expected result, viz., a higher survival rate of the irradiated
animals. In this connection, a new procedure was tested. The ppincipal
variant of the experiments, the reswlts of which are given in the present
paper, was carried out in the following manner. As recipients, female mice
of strain A which had given birth to a litter were used, while the donors
were the offspring, at the age of 1-5 days, of each female. To each female
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there were transplanted four to five spleens. By this procedure we ensured
the greatest consanguinity between the tissues of donor and recipient which,
as is known, provides the moat favorable conditions for the adoption as
well as the survival of the transplanted organs and tissues. By this
transplantation procedure we excluded to the maximally possible extent those
difficulties in the adaption of the transplanted organ which are connected
with tissue incompatibility. Thus, the procedure utilized makes it possible
to form an objective opinion on the basis of relatively limited experimental
material concerning the prospects of the effected transplantation of any
given organs or tissues.
In our experiments the spleens for transplantation were prepared half
an hour before irradiation. The removed spleens were kept in a sterile
Ringer solution. During the first hour following irradiation the transplanta-
tion of the spleens to mice of the experimental group was carried out.
For this purpose the females were fastened to a special board. No narcotic
was used during the operation. At the left side of the abdomen a small
incision of the skin was made, and the abdominal cavity was exposed. The
spleens being transplanted were inserted into the abdominal cavity in such
manner as to place them in the region of the epiploon (recessus lienalis)
which adjoins the spleen of the recipient mouse. The vessels of the trans-
planted spleens were not tied, and the spleens were not attached to any
organs or the epiploon. After tae insertion of the spleens the incision was
sutured. The control group of animals consisted of female mice which had
given birth at the same time as the animals of the experimental series.
The control animals were operated on in the same manner, but without the
insertion of spleens. Finally, in the third part of the investigation a
study was made of the effects of on intravenous administration ofhomologous
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bone marrow upon the course of the radiation reaction in mice. Male mice
of strain A, aged 2-3 months, were used both as donors and as recipients
in these experiments. The conditions and technique of exposure to Brays
were the same as in the studies of the effects of spleen transplantation
upon the survival rate of the irradiated animals.
During the first hour following the gray irradiation, the mice of
the experimental group were given an intravenous tail injection of 0.6 ml
of a bone marrow suspension in a buffer solution. The injected volume
of the suspension contained the bone marrow isolated from the two femurs of
a nonirradiated animal. Mice of the control group were injected with a
corresponding amount of the buffer solution. The buffer solution contained
in one lit of distilled water NaCI (6.8g), KC1 (O.4g), CaC12 (0.2g),
MgSO4 (O.lg), NaH2P04 (0.125g), NaHCO3 (2.2g), and glucose (O.lg). The
pH of the solution was 7.4. The suspension of bone marrow was prepared
in the following manner. Femurs thoroughly cleaned to remove muscles and
fasciae were placed (in pairs from each donor) in vessels containing
sterile Ringer solution. To remove the marrow, the upper end of the
femur was cut oft, and the needle of a syringe was inserted in the opening.
The bone was only slightly incised at the lower tip, and this end was
immersed in a small vessel holding 0.6 ml of the buffer solution. y
means of the syringe, this solution was drawn 2 or 3 times through the bone,
thereby removing the marrow therefrom. The bone was then removed from
the needle, and the same procedure was repeated with the second femur of
the donor. As a result, the concentration of the thus prepared suspension
corresponded to the bone marrow of the two femurs of the donor.
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Effect of ti, screening of iieaatopoietic 0 ans upon the Course of
Radiation Reaction in Mice
To study the role of the damage to hematopoietic organs in the
course of the radiation reactions in the animals, three fundamental
kinds of experiments were first carried out. In one (control) experi-
ment a total Xray irradiation of the animals was carried out. In the
second, a total irradiation with screening of the spleen. Finally,
in the third a total irradiation was combined with a screening of the
rear extremities of the animals. In the different variants of the
experiments different doses of xray exposure were used. The purpose
of this portion of the work was to determine the changes in the nature
of the reaction of the animals under conditions of a complete preserva-
tion of different hematopoietic organs. Securing of these data was not
only of interest to use per se but was also of great importance in the
carrying out of subsequent experiments.
The results of a total irradiation of the mice with Xrays and of
the irradiation under conditions of a screening of spleen and bone marrow
are shown in the synoptic Table 1.
(See Table 1 on Page 97)
This table shows data relating to the survival of the animals and
the mean duration of life of the mice that died.
Examination of the tabulated data shows that screening of any of the
studied hematopoietic organs increases sharply the survival of the animals.
In moat cases, in spite of the relatively small number of animals investigated,
the results relating to experimental and control series show statistically
reliable differences.
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TABYS 1
SURVIVAL OF MICE FaLLON12B1G X&AY IRRADIATION CARRIED ODT UNDER CONDITION OF A SCREENING
OF TUE H7lATOPOISfIC ORGANS
Ezperi..ntal
Dose
Group
Total
Survived
Died
Mean
series
(r)
number
of ani-
mals
Number
Percent
Number
Percent
duration
of life
(days)
soo
Experiment
27
19
70.4?8.8
8
29.618.8
14.&
Control
24
10
41.6?10.1
14
38.4t10.1
11.4
Experiment
17
is
64.7111.6
6
35.3#ll.6
9.5
Screening
600
of spleen
Control
17
11.717.8
15
88.3?7.8
8.6
Experiment
21
33.4#10.3
14
66.8?1Q.3
6.0
800
Control
Screening of one
20
0.0
20
100.0
4.6
extremity
34
24
?0.617.9
10
29.4?7,8
11.0
600
Screening of two
Screening of
bone marrow
extremities
34
30
88.2?5.5
4
11.8}5.5
19.2
Control
Screening of one
34
17.6?6.v
28
82.416.5
9.6
extremity
23
30.4t9.G
16
69.6?9.6
6.3
700
control
24
0.0
24
ion
5.6
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As was to be expected, a more complete protection of bone marrow
(screening of two extremities) ie more effective than a partial (screening
of one extremity), although a direct proportionality is not observed in
these instances.
On comparing the protective effect resulting from screening of
the spleen and bone marrow, it is noted that screening of the spleen yields
very similar results to those obtained by a partial screening of bone
marrow and is somewhat less effective than protection of both rear extremities.
Worthy of attention are the results of the screening of the spleen at dif-
ferent doses of exposure. In theme cases the greater the irradiation dosage
(within the range of ordinarily lethal but not absolutely lethal dosages),
the greater the relative protective action of the screening. Thus, the
survival of the animals following an irradiation dosage of 500r under
the conditions of a screening of the spleen is only 1.7 times greater than
that of the controls; whereas, with a dosage of 6001', it is more than 5
times greater.
Evidently the death of the animals following relatively small
irradiation dosages is associated to a lesser extent with a damage to hema-
topoietic organs. It should be noted that this conclusion is in good agree
ment with literature data and also with our own unpublished data showing a
relatively alight damage to hematopoietic organs following chronic exposure
to small doses of radiation (Zirkie, 1984).
From the data shown in Table 1, still another unquestionable con-
clusion foli.ows; damage to hematopoietic organs plays an important,
possibly a decisive, part in the death of the irradiated animals upon
application of the minimal absolutely lethal dose or even a somewhat higher
dose). This is evidenced by the relative increase in stature of the
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role of the screening of hematopoietic organs in the survival of the
animals upon increase of the irradiation dosage, as well as by the high rate
of survival of the screened anlm*ls (30-33%) following exposure either to
a minimal absolutely lethal dose (lOOr) or a higher dosage (800r). In
other words, it may be assumed that determination of the level of ex-
posure (irradiation dosage) at which a 100% death rate of the irradiated animals
occurs is connected primarily with a damage to the hematopoietic organs.
On comparing our results with the literature data concerning the role of
the damage to individual systems of organs in the radiation reaction follow-
ing different doses of xray exposure, the following assumption can be made.
Upon application of relatively small dosages, death of the animals results
from a general weakening of the irradiated organism which involves no
predominant damage of any one specific system of organs or tissues. Death
of the animals following exposure to a dose of the order of X80_100/30
is due primarily to a disruption of hematopoietic functions. Finally,
upon application of considerably higher dosages of the order of LD100/3,
the determinant role in the course of the radiation reaction appertains
to a damage of the digestive system and above all of the intestines
(Quastler, Lanzl, Keller, and Osbone, 1951). The beneficial effect of a
screening of the hematopoietic organs on he course of the radiation
reaction can be ascertained not only from the final result, viz., the
survival of the animals, but also from the course of this reaction. A
relatively milder course of the radiation reaction is evidenced primarily
by the behavior and appearance of the irradiated animals. This is also
indicated, in particular, by the mean values of life duration of the mice
that died, which are always higher in the experimental than in the control
series. Finally, this is made still more apparent upon analysis of the
data which characterize the dynamics, in time, of the death of irradiated
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animals. Figure 2 shows a graph which represents the time of death
of irradiated animals in regard to cases involving acreening of the spleen
and those without such screening (the data on the basis of which this and
the following graph are plotted are shown in the addendum of the present
paper. Figure 3 shows curves of the same nature which reveal the effects
of a screening of the bone marrow.
Herein, as well as in the case of a safeguard of the spleen, the
shape of the animal-survival curves relating to the experimental series
differs sharply from those relating to the control series. This dif-
ference is made especially evident upon comparison of the curves, showing
the survival of animals which were subjected to irradiation as well as
screening of both rear extremities, with the curves relating to the controls.
In this instance death occurrences among animals of the experimental series be-
gin at a time when such occurrences completely ceased among those of the
control series.
A good index of the beneificial effect of the screening of hema-
topietic organs upon tie course of the radiation reaction of the animals
is provided by the dynamics of their change in weight. Figure 4 shows
the curves of weight chaa es in surviving mice during the 30 days following
irradiation. The irradiation was carr#+id out with and without screening
of the spleen.
To facilitate a comparison of the curves, the weight of the animals
is expressed in the form of indices (ratio of mean weight of the mice at the
time of weighing to the mean weight at the beginning of the experiment,
which is taken as being equal to 100).
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Analysis of the curves shown in the graph reveals a lesser decrease
in weight among animals irradiated under the conditions of a safeguard
of the spleen, as compared with those which were not so protected. This
is especially manifest on examination of the data relating to irradiation
with a dosage of 500r. Most characteristic in the weight changes of the
animals of all the experimental groups, in comparison with the controls,
is the acceleration of the recuperative processes. It is also of interest
to note that the changes in weight of the animals irradiated with a dosage
of 800r while their spleen was screened, are about equivalent to the
changes in weight observed in animals following a total irradiation with
dosages of 500-600r.
Figure 5 shows the curves of the changes in weight of mice which have
been subjected to irradiation with and without screening of the extremities.
No particular differences are found in the shape of the weight variation
curves relating to animals irradiated with screening of one and both
extremities. Both these curves are characterized by a relatively rapid
drop followed by a sharp increase which contrasts with the curve relating
toy the controls.
Examination of the shape of the curves which characterize changes
in weight of the experimental animals and of the controls, permits, on the
whole, arriving at the following general conclusion. Screening of hemato-
poietic organs at the time of Bray irradiation manifests itself not so much
by a decrease of be primary overall damage to the organism as by the
rapidity and intensity of the recuperative processes.
Effect of Spleen Imp1antst ons Upon the Course of ldidtion- Ruction in Mice
The beneficial action of spleen screening upon the course.of radiation
reaction in mice naturally suggested the possibility of attaining the same
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effeot not by screening of this organ but by means of it$ transplantation
from nonirradiated to irradiated animals. Experiments of thia nature,
as rats mentioned hereinbefore, have been carried out by a number of
researchers and have yielded positive results (Jacobson, Simmons,
Marks, and Eldredge, 1951; Barnes and Loutit, 1953, et al.). At the
same time, up to now the mechanism of the positive influence of the
implantations has not been determined. The fate of the transplanted
spleens has not been followed, and an effective transplantation procedure
has not even been developed, i.e., a procedure which would make certain
a positive action of the transplants upon the course of the radiation
reaction. One of the characteristic features of the work carried out in this
direction is the very great variability of the results. All this made
necessary further study of the effects of a transplantation of the spleen
of nonirradiated animals upon the course of the radiation reaction.
As was stated be'einbefore, the principal variant of our experi-
ments on determination of the effects of spleen implantation upon the
course of the radiation reaction consisted in a transplantation of the
spleens of the offspring to their mothers. To ascertain the fate of
transplanted spleens, the mice of the experimental series were dissected
2 to 2 1/2 months thereafter. Thus, it was found that, in addition to
their own spleen, about 60% of the animals also had 3 to 4 spleens which
had undergone adaption. The adapted spleens are smaller in size than
normal (but are considerably larger than their size at the time of
transplantation) and have the appearance of normally functioning organs,
situated most frequently at the inner wall of the ab ominal cavity. A
ramified network of blood vessels extends to the transplanted spleens.
The picture observed during one of the dissections is shown in the
photograph of Figure 5.
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A histological analysis of the implanted spleens has been carried
out. Fixation of the material (pieces of the tissues of the implanted
and,the natural spleon) was conducted in a Zenker solution. The prepara-
tions were stained by hematoxylin with eosin (for an examination of the
general microscopic structure of the orgaus), according to the Mallory
method (to reveal the extent of development oz connective elements), and
with azure-eosin (for a differential staining of the cellular blood ele-
ments).
Mention should be made of the great similarity in the microscopic
structure of the natural and implanted spleens. The adapted spleens, as
well as the natural, are enclosed in a connective-tissue capsule. Both
kinds of spleen show normally developed white and red pulps; there is
present a reticular syncytium in the meshes of which are found free cells
including erythrocytes, megakaryocytes, very numerous lymphocytes, a small
number of neutrophiles and eosinophiles. The enumerated cellular forms
are found at most diverse stages of development. The implanted and natural
spleens are permeated with Mood vessels and in the I~lpighian bodies
there is invariably found an eccentrically disposed central artery. Stain-
ing with azure-eosin clearly reveals the presence in the balpighian bodies
of more lightly stained areas, the centers of cell proliferation.
A detailed comparison of the adopted and natural spleens also as-
certains certain differences in their microscopical structures. As was
stated above, both kinds of spleen have a well developed connective tissue
capsule; nevertheless, while in the natural spleen numerous fairly coarse
and thick trabeculae extend from the capsule into the depth of the tissues,
in the implanted spleens the trabeculae are few, very thin, and delicate,
and it is only at the places where the vessels enter the spleen that the
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trabecular are of a fully developed form (see Figures ? and 8).
In the implanted apleens the Malpighian bodies are more uumeroue
than in the natural spleen. Moreover, in the adapted apleens the Mal-
pighian bodies have more definite contours (see Figures 9 and 10). In
the Malpighian bodies of both the natural and the implanted spleens there
are observed a large number of cells undergoing division; nevertheless,
in the implanted spleens the mitoses proceed with greater intensity (see Figure
11).
within different areas of the adopted spleens there can be observed
a considerable amount (greater than in the natural spleen) of eosinophiles
(up to 12 to 14 within the field of vision) and aaegakaryocytea (up to
8 to 9 within the field of vision) at different stages of development.
Some of these cells are in process of undergoing division (see Figures
12 and 13).
Many of the differences noted between the implanted and the natural
spleens are probably due to age differences between the organs being
compared. As was stated in the part of the paper relating to the pro-
cedures, the spleens for implantation were taken from mice one to 5 days
of age, whereas the recipients were adult mice aged 2 1/2 to 3 months.
All the above-presented observations permit reaching the conclusion
that in the irradiated mice the adopted, as well as the natural, spleens are the
sites of active hematopoiesis.
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TABLE 2
SURVIVAL OF THE 11NIl1At.4 POLi.OIiI*G IRRADIATION. EXPERIMENT: IRRADIATED IfICS WITH ThANSPLANTED
SPLEENS, OONPROL: IRBADIAT~ lIICB. HAVIBiIi i!!IDffiifONS IUD 3PLS?3N I1a+IJlttfATION
1bt+a1
aiaber
of ani-
mals
Ntmtber
Survi vied
Percent
Number
Died
Percent
Yosn duration
of life (days)
sspanaeac
71
53
74.6f5.1
18
25.4?5.1
8.5
control
63
2?
42.8?6.2
ss
57.2?6.2
10.8
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we note in addition that the histological analysis data constitute
still another confiraatioo of the previously made aaeus+ption concerning
the great importance, in relation to the results of the investigation, of
the procedure utili$ed in the experiments involving a study of the in-
fluence of spleen transplantation on the course of radiation reaction. Thus,
in particular, the data obtained by us are of an opposite nature in compari-
son with those recently published by Langendorff, et al. The procedure
used in the latter research did not ensure an adaption of the transplanted
spleens (Langendorff, Koch, and Sauer, 1954).
The results which we have obtained indicate a greater effectiveness
of the procedure of transplanting organs and tissues Prom newborn animals
to their mothers. This method may be of value also in other experimental
work concerned with the study of the mechanism involved in the effects
produced by transplantation of various organs on the course of the radiation
reaction.
Table 2 shows data which characterize the survival of irradiated
females with and without implanted spleens. As is apparent from the
tabulated data, the survival of the animals of the experimental series
exceeds almost twofold that of the controls. The differences observed are
statistically reliable. Conspicuous is the fact that the mean life duration
of the mice which died in the experimental series is less than that of the
controls. Usually, under the conditions of a safeguarding, the animals
live longer than the controls. Later on we will give further consideration
to this somewhat unexpected fact.
The beneficial effect of spleen implantation is most clearly mani-
festod upon examination of the survival curves shown in Figure 14.
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The survival curve of the animals of the experimental series
is characterized by a more elevated level as compared with the control
series. We note in this connection that the death rate of the two
groups being compared is almost the same up to the eighth day. There-
after, the death instances among animals with implanted spleens occur
at a somewhat slower rate, and cease almost entirely after the thirteenth
day. Death instances among mice of the control series subside only on the
seventeenth day, but isolated instances of death occurrence are encountered
up to the last (thirtieth) day of observation. This nature of the death
rate among experimental and control animals is precisely the reason which
brings about the fact that mean life duration of the mice which died in
the experimental group is found to be shorter than in the case of the
controls.
The observations of animals with implanted spleens have shown that
a transplantation of the spleen renders milder a number of symptoms of
the radiation reaction. The general appearance of the experimental mice
is better than that of the controls, as are the weight indices and the
data which characterize the dynamics of changes in the number of leukocytes
found in the peripheral blood. Figure 15 shows the curves which characterize
the changes in the number of leukocytes in the animals of the groups being
compared over the 3Q-day period of observation.
The group of experimental animals appears to be subdivided into
two subgroups, one of Which includes the mice in which (according to the
results of the subsequent dissections) the spleens have become adapted,
While the other comprises those animals in which an adaption of the spleens
has not taken place,
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Accordingly, the experimental group of animals is represented in the graph,
in addition to the swmaative curve (curve 1), by still two other curves.
In all cases a decrease in the number of leukocytes occurs during the 5
days following irradiation. The greatest difference between experiment
and control manifests itself in the recuperatooy process. In the mice
of both experimental groups the recuperatooy process takes place more
intensively (especially in mice with adopted spleens) than in the control
animals.
On examination of the data relating to the changes in the weight
of the mice, represented in the form of curves in Figure 16, we also per-
ceive that in the animals with adapted spleens the decrease in weight is
somewhat less than in the controls, and, what is of special interest, that
in the former the recuperative processes are effected much more rapidly
and to a fuller extent. Here, as in the case involving the leukocytes, we
are in a position to ascertain that the adaption of spleens manifests it-
self essentially by an accelerated progress of recuperation. Mice in which
the spleens have not become adapted constitute a separate instance. Their
weight after definite intervals of time following irradiation is found to be
even below that of the controls. The cause of this is obscure. If this
instance is omitted from consideration, all the above-presented data indicate
that implantation of the spleen, carried out by the above-described pro-
cedure, has a most beneficial effect on the irradiated animals. This
manifests itself by a greater survival and a less pronounced nature of
the radiation reaction. The latter is objectively evidenced by ;the dynamics
of changes in the number of leukocytes and weight among the irradiated
animals.
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Effects of Intravenous Administration of Bone Marrow on the Course of
Radiation Reaction in Mice
The beneficial action of the screening of bone marrow (as well as
of the screening of the spleen) on the general irradiatior of the animals
suggests the possibility of its administration to irradiated animals with
the view of rendering milder the radiation reaction in these animals. Tile
study of this question by means of experiments on mice is of unquestionable
interest, primarily for the reason that it is precisely the 'Done marrow that
constitutes the principal hematopoietic organ in highly davloped ma?,mals.
Attempts to administer bone marrow derived from nnn'.rradiatxd mice ?t,c
irradiated mice have been reported in the literature. In some instances az.
unquestionably positive effect was o'Userved, with increasing survival of
the irradiated animals (Rekers, Coulter, and Warran, 1950; Lorenz, Congdon,
and Uphoff, 1953); in other instances no such effect: were noted (Tai'bot
and Pinson, 1951). The conflicting results thus obtained and also the
undetermined nature of the mechanism of the action of bone-marrow admini-
stration in those experiments which yielded positive results have led
us to undertake the study of this question. Therein are presented the
preliminary data which we have obtained on the effects of an intravenous
injection of bow marrow upon the course of radiation reaction in mice.
Table 3 shows the data which characterize the survival of irradiated
animals following administration of bone marrow (experiment) and without
such administration (control).
The higher percentage of survival among animals of the experimental
series, as compared with the controls, leaves no doubt as to the beneficial
action of an administration of bone marrow upon the course of the radiation
reaction in mice. The same is also indicated by the greater life duration
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of the animals which died.
Figure r7 shows carves which illustrate th death rate oi' the
experimental animals and that of the controls, following irradiatio.~.
In these cases, as in those involviri; implantation of placn, the ui-
ference In death rate values among ep rire ltai aid control aiii aal ;
becomes apparent only after tao eight day.
TABLE a
SUfVIVAL OF ANIMALS FOi..LOWIG IRc,,ADIATNN. EXPERIMENT:
.;iZC~a
TO WUICH BONE MARROW AS ADMINISTER INTA` ENOUSLi kTE)
RADIATION. CONTROL; IRRADIATED ?!ICE TO WUICH NO DO.1E MARR0
WAS ADMINISTERED
series Total Survived Died Mean
number duration
of ani_ of life
orals N,arter perceart Nunh r percent (days)
Ercperi-
went 104 (j5
Control 104
'49 C7.ll4.
1l r U~ r' 64.5?4.7
355.5+_'. d v
Starting from this point of time, death occurrences among control
mice take place at a considerably more rapid rate than among the experimental
animals, and last up to about the twentieth day. Thereafter, only isolated
instances of death occur among animals of both series.
As in the preceding experiments on screening of hematopoietic organs
and implantation of spleen, an investigation was also made of the other
objective indices of the course of the radiation reaction. These indices
include first of all the amount of leukocytes in the peripheral blood of
irradiated animals. Figure 18 shows the curves which characterize the
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changes in the number of 1oukocytes . i na~ce injected with Ions marrow
after irradiation, and L'i the controls.,
As is apparent #ro~;l those curves lnukopeia occurs at -the sarw
time irl the animals oa both groups being compared, and it roachcs an
Identical level a A the earn time the recup r atory process ta'~ : piece
much more intensely in mice whi&:Y have riser, giv :n a i s octio=, of one
marrow. ThuaG the lumber cf leukocytes In the blood of tiro coz.tro
a ib is reachos dr.1;.i tgi tw ltieth day the
.t :rya attaiae
on tho f tcenth day tie mice of they expor:irueryt. at ser.ioz
Not 1esi:7 asld!eat Ve iJ tc ma1Jif G9ta 'n c'1. I+dS byne.P6ic1a1 act.'.o
o bone-marrow l'1 jOC tii ii. upon e a ninatioz: oz data slating to changes
in the weight of the irradiated animals. Figure 19 shows the pertinent
data,
in mice cif ti:O experim sat d group, l px r'ux! decrease is noted
on the fourth day a ttiar- irradiaiot which i foliowcd by a gradual in-
crease. By the sixtenti9 day the animals reach tho?r initial weight.
Mice of the control group o9 sc ` quite a different picture 1n these
cases decrease iweight continues up to the tweii rh day following
irradiation. The titia]. weight is reached only on the twenty-fourth
day. Thus all the data presented in this sectio:.indicate the beneficial
action of an intravenous injoction of bone marrow on the radiation reaction
in mice.
Discussion
It appears of unquestionable interest to compare the date character-
izing the course of radiation reaction in mice which have been exposed to
-ill-
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radiation under conditions of screening of hematopoietic organs, with
the course of reaction in animals which have had a spleen implantation or
an injection of bone marrow after the irradiation. Of no losser interest
is a comparison of the data obtained on the study of the radiation reaction
in animals with protected or implanted spleen, with analogous data relating
to animals w.th screened bone marrow or injected with bone marrow.
Notwithstanding the tact that these experiments were carried out riot
at the sane time, their comparison is fully permissible. Their corlpara-
Ality i.~ determined Ly the sirailarity of conditions both physical (Xray
irradiation) and bio1o~:Lca1 (the keeping of the animals) maintained on con-
ducting the: o experirnentM. 'ri.e presence of such similarity ir, conditions is
evidenced in particular by the values which characterize the survival of the
animals of the differeszt o.~:rol series.
In comparing the data obtained, it i~ necessary first of all to
point out the great sirglilarity iri the effectiveness of all the tested pro-
tection methods. For a comparison of the efficacy of the different
protection methods, it is better to utilize not the percent cf survival
of the anirr:als in any giver: series of experiments, but rather the so-
called survival index, i.e., the ratio of percent of surviving animals
in the experiment to the percent of surviving animals in the control.
The survival index on exposure to a dose of 500r, with screening of the
spleen, is 1.7, on implantation of the spleen it is also 1.7; and,
finally, with injection of bone marrow it is 1.5 (see data showl in Tables
1, 2, and 3). Such coinciding values of the survival index can hardly
be considered a fortuitous occurrence.
An especially important feature which is common to all the tested
protection forms is in our opinion the predominant influence of the pro-
tection upon the course of the recuperatory processes and to a lesser
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extent upon the inanifestations of the primary damage. This has been
sufficie~Ytly streissed hereinbefora. It i.s true that in these instances it
is probably more correct to speak not of a predoriinant influence of the
different forms of protection upon the course of the recuporatory pro-
cesses, but merely of a better^-defined man.ifestatior of the protective
action during this period of the radiation reaction.
In addition to the noted features of similarity, there asp also
found certain differences in the nature of the protective action of the
screening of an organ during irradiation and that othe protective
action due to subsequent repiacemenL of the organ. such a, difference wGs
ascertained by us, in particular, in the case of a comparison of the
results of spleen screening and spleen transplantation. \'Vheroas, ii tip
absolutely lethal dosages of irradiation, the screening o the spiee~z
produces a protective effect (see Table 1), the transplantation of the
spleens does not affect in these instances the surviv-01 rate. We havo
carried out a special i mited?sca1e expertent (with 15- animals o: the
experimental series and 14 control animals) in which irradiation was
effected with a minimum absolutely lethal dosage (70th') and the implanta-
tion of spleens did not produce any beneficial effect. The cause of the
observed difference in the effects of screening and transplantation is
in our opinion the fact that implantation of spleen begins to exercise its
beneficial effect upon the irradiated animals at a much later time than
is the case with screened organ. Since with absolutely lethal dosages
the radiation reaction proceeds at a much more rapid rate and is more
intensive than following lethal dosages, the effects of an implanted spleen,
which manifest themselves unquestionably subsequent to the transplantation
operation, do not have sufficient time to alter the course of this reaction.
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However, the i.,ieffectiveness of spleen.. transplantation upon application
of absolutely lethal dosages is due not only to this reason. Another
cause, in our opl;tion, is the fact that spleen implantation produeea a
beneficial effect primarily by enhancing the resistance of the irradiated
organism to secondary infectionti (which will be considered in detail here-
inaftor), whereas the death oi animals irradiated with absolutely lethal
dosages .a riot due to info ~tio zS aCu:a~:~ .
The facts under d isucs ion are directly cor~nocted with the
question concerning the atur . of the protective ation of irnplar ted
spleen and injected bone marrow. s it duct to clluiar structures or to
some humoral factor produced by tic retmlving introduced t:LSSUCS? Our
data, which show the importance of an adaptiox, of the spieens for the
effectuation of a protective action, indicate that it is associated with
cellular structures. At the same time, the fact that in the case when
no adaption of the implanted organ has takem place, a certain beneficial
effect is still oroducecl :indicates the possibility that a hurnoral factor
may also be involved it the protective effect. We beleve that the same
is also indicated by the data relating to injection of bone marrow, where-
in it can hardly be assumed that the protective action is associated with
adaption of the cells of this tissue. Thus, we are inclined to assume
a dual nature of the protective effects of spleen transplantation anal bone
marrow injection.
in conclusion, let us consider briefly the possible mechanism of the
beneficial action of spleen transplantations and bone marrow injections.
The nature of the effect of spleen transplantation upon the irradiated
animals is determined by the functional significance of this organ. The
role of the spleen as a hematopo1eti4 organ in mice is very great; therefore
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there is every reason for assuming that the adapted spleens rormai:.zo
the heuatopoiesis in animals having been nubjeetoi to irradiation. We
assume that of no lesser impartaiico in the protective effect are the
adaptod spleens as a source o: onhaiicere it o : the protective funct .or
of the organism in its cant ?Oi of secondary -infoctio-nsy The role of '- ~ ;
spleen. 1='. this coimoction is wo11 'cnown; it is associated with the pagocyt.c
aeti'Vity o this urgar aid id.:"_l t3 capab.-ii ty to I'OdUCO aitibodie.
Our data provi 2 a ew?ta ft ii.dicat on that the posi. ti vo ofSec,.s
of spieer i7np1antatiori is associated 'with decreased probabii ty of the
occurrence of izifectioudiaoas 3 it is precisely to thim t ?ct then
can be attributed the above>mc:ntio iod divergence to survival curves
rcla?t n to the experimental and the contra. ,routs ox mice, only 8 day,.
after the irradiation, aria tine fact assited therewith of a greatoa ::e
duration of the mice which died in the control group rather than those
which died in the experimental group.
It is known that dc;ath of the animals duri g the 2 to 3 week;
after irradiation and late :is usually due to a development of bacteriemia.
It is also possible that .ii plantation of spleen has a stthulating effect
on the leucopaietic functiox. of the natural spider of the recipie.rt. The
latter is indicated by the large amount of leukocytes not only in to :food
of mice with Adapted spleensut also in the blood of mice which have under-
gone implantation but in which adaption of the implanted spleen has not
yet occurred. An increasing !c-umber of leukocytes within the cir'uiating
blood enhances the defensive capabilities of the organism in its control
of infections.
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On considering tlao qution of the pot~u1L1o rechaizism oi' the
protoctivo action of the bone marrow, it should be stated that in a
numbor of feature. it is probably similar to the mechanism of the
action of splc oi~ itpIa t ;. It i#R true h this ?;istanet: th3
tin; acticn of the Lntroduced bone marrow upon the hematopoiesiW of the
irradiated aixii;ialbecomes of primary importar~cc. Increased hematopo r~s s
i ?. r t 1t, (.,i +"''M?1i fir. r1. ,,. LE. ..', 'j, lk
..a... e .., .:. ~ ?aLi ata~l a: , a,.c; , :os tit:. z~.1 a .~. L.L.. 4:.~.a: ..,
z .'i,.t.'i:rai lt+ciF.oL,; 3si~t, lab part tcalai 9 to :J~i~ eta! saKb'' .ri:cc,,..,1..3. .
+tu: w d:,1.Vi~dd:14tW.dicsonsi-
tivity of the testes and also the fact that recuperat+_ve processes
occur therein more slowly than those upon which the general condition
of the animal depends,
Histological Analysis oi` i Testicular Structure of Irradiates Animals
Known at the prese.d t, ire a 1 u'go ?i nber of researches eonceraied
with cufficientiy-detai1od citudies of the damages to he testes of
mammals (especially of rothmts) induced by ionizing radiations (vedgenidze,
Kotik, larionov, et al., i936; Bloom, 1945; Sschonbrenner and Miller, 1E4
Fogg and Cowing, 1952; Shaver, 1953), The histological analysis of the
testes of irradiated aniiais which we have carried out is limited to
the general picture of the :addation damage of this organ. Such an
analysis was indispensible for a comparison (at the selected time in-
tervals) of the damage pi?txes revealed in the microscopical structure
of genital glands, with those damages disclosed by means of a hybridour
analysis of fertility.
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In connectiolx with the task involved fixation of the testes
of animals was carried out at the followiiu points of fir o
(i)
after 12 hours, (2) after 24 hours, (3) after 15 days, (4) after one
month, and (5) after 3 months, following irradiation. The first,
second, fourth, and fifth time intervals gat' fixatyon coincide with the
time intervals of the fertility tests of the irradiated males, which
makes it possible to form an opinion concerning tho course of spermatogenesis
at the time of the tests. At each of the time intervals two animals wore
killed in both the experiment; l end control series. The testes were
subjected to a fixation procedure in Zerikor's solution, as modifio
by MaksimovL The sections were 4 to G ' thick. staining was done with
iron hematoxylin, according to the Haydenhein method,.
(a) Histological structure of Testes 12 Hours After irradiation
Twelve hours after the exposure, the general microscopic picture
of the structure of the tastes of animals subjected to an irradiation of 200r,
as well as to one of 400r, does not differ essentially from that of the
control animals. The tubtzii are well filled-?out, and their diameter does
not differ from that of tho seminal tubuli of the control animals. The
tubuli contain seminal cells at different stages of spermatogenesis.
Some spermatocytes in the testes of irradiated animals are in the process
of isitosis.
On a more detailed study of the sections of the testes of
irradiated mice it is possible, even during this period, to detect the
action of radiation. Thus, among the cells adjoining the basement mem-
brane, there are cells having pycnotic nuclei which possibly constitute
apermatogonia which were in the state of division at the time of exposure.
Pycnotic alterations also affect the nuclei of sperlaatocybes. In these
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ifBtanceg they acquire a uniform, fairly intensive stain coloration.
Sometimes the chromatin is conglomerated into a cluster and is dis-
tributed in the shape of a crescent along the periphery of the nucleus.
Found fairly frequently at the locations of spermatogonia and sperm-
acytos are rounded particles of unstained plasma which apparently con-
stitute the remnants of degenerated cells. Also found in the testes of
irradiated a 4mals are gigantic multinuclear spnrmatides. Some of them
bear clear traces of degeneration. Tho described alterations are en-
countered somewhat more frequently in the testcs of animals subjected
to an irradiation of 400r than in those subjected to 200r.
(b) Histological Structure of Testes 24 Hours After Irradiation
At this time, the testes of animals irradiated with a dose of
400r show an appreciable decrease in the vaunt of spermatogonia. Other-
wise, the histological picture of the testes of animals previously sub-
jected to exposure does not differ from that described above.
(c) Histological Structure of Testes After 15 days Followin
Irradiation
After 15 days subsequent to exposure there is noted in the testes
of irradiated animals a decrease in the number of germ cells. At the
same time the degree of devastation of seminal tubuli indicates a direct
correlation with the dosage of exposure. With a dosage of 200r, de-
generation is not as extensive as in the case of a 400r dosage. The
testes of animals irradiated with a dose of 200r are characterized by
the presence of a small number of cells constituting the initial stages
of spermatogenesis, viz., speranatogonia and spermacytes. The principal
mass of germinal tissues is composed of spermatides and spermatozoids,
which is adequately illustrated by the microphO ograph shown in Figure 4.
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With a dosage of 400r, the devastation of seminal tubuli has
proceeded considerably further. Conspicuous is the lack of uniforaaity
in the degree of damage to different seminal tubuli (Figure 5). Tubuli
are found, the histological structure of which is similar to that observed
in the case of a dosage of 200r. In those the principal mass of germinal
cells consists of spermatides and spermatozoids. Present on the other
tubuli are essentially Sertoli cells and only a small amount of spermatides
and spermatozoids. Spermatogonia and sperrnacytes are very rarely encountered.
Figure S shows a microphotograph illustrating the structure of a tastes of
an animal of the same age which had undergone no Xray irradiation.
(d) Histological Structure of Testes One Month After Irradiation
An interesting picture is that of testes subjected to fixation one
month after irradiation. The diameter of seminal tubuli in irradiated
animals is less than in the controls. At the same time there is manifested
therein a relationship with the dosage, since the diameter of the tubuli
is less in testes of animals subjected to a larger dosage of xrays (400r),
as compared with the other experimental group subjected to a lesser dosage
(200r). The different composition of the germinal cells in the testes of
animals irradiated with different dosages also emphasizes the considerable
difference in the effectiveness of these dosages. With a dosage of 200r
almost all the seminal tubuli are uniform in cellular composition (Figure 7).
They contain cells in all stages of spermatogenesis, but the amount of
spermatozoids is very small. The picture observed indicates that in
the testes of animals subjected to 200r an intensive process of spermatogenesis
restoration takes place one month after exposure.
In the testes of animals subjected to 400r a sharp lack of uniform
ity is observed in the structure of seminal tubuli (Figure 8). Some tubuli
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are filled only with Sertoli cells, the nuclei of which are clearly
visible at the basis of the membrane. Vacuolation of the plasma of these
cells is clearly manifested. The structure of other seminal tubuli
recalls the tubuli of testes of animals irradiated with 200r. In these,
restoration of spermatogenesis is in progress. Not infrequently these
tubuli have a reserve of spermatogonia. Spermacytes are not disposed in
two or three rows as in the controls, but are scattered at random. A
number of tubuli show a structure that is of an intermediate nature:
a varying number of spermacytes are scattered among the Sertoli's cells.
Testes of animals irradiated with 400r are characterized during this
period by an almost complete absence of spermatides and ~ermatozoids.
(e) Histological Structure of Testes After 3 months Following
Irradiation
The histological structure of testes of animals irradiated with
200r (Figure 10) approximates that of the testes of the control animals.
The seminal tubuli bear almost no traces of damage due to irradiation.
Only occasionally are there encountered gigantic multinuclear spermatides.
In animals irradiated with 400r there also takes place by this time a
restoration of spermatogenosis (Figure 11). However, the histological
picture of the structure of testes in these animals differs somewhat
from that of the controls (Figure 12). Thus, the lumina of seminal
tubuli in the experimental animals are larger, which is apparently connected
with the lesser number of cells in the late stages of spermatogenesis, viz.,
spermatides and spermatozoids.
Thus, the above-presented data show that by action of the irra-
diation a sharp disruption of spermatogenesis occurs in the male mice.
The disruption is associated on the one hand with damage to the spermatogenetic
cells at different stages of spermatogenesis, and on the other with its
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temporary cessation. The utter is apparently caused by the fact that
the spermatogonia retained by the irradiated animals temporarily lose
the capability of undergoing division; since in all the cases which we
have investigated spermatogonia were present in the testes, although some-
times in a very small amount, the restoration of spermatogenesis is
apparently the result ci a restoration of the capacity of spermatogonia
to undergo division,
The lesser degree of damage following irradiation with 200r
results in a situation wherein recuperative processes begin at a time when
the available reserves of germinal cells have not yet been fully depleted,
and therefore a period of total sterility, if it occurs in these animals,
is of only brief duration. With a dosage of 400r this period is more pro-
longed, which is evidenced by the incomplete restoration of spermatogenesis
even after 3 months following exposure.
Offspring Derived from Irradiated Males After the Restoration of their
Fertility
Since the histological analysis has shown that Xray irradiation
results in damage to germinal cells present at different stages of
spermatogenesis, the question arose concerning the radiosensitivity of
these cells from the standpoint of the effects upon the offspring. In
this connection, it was of particular interest to trace the effect produced
by the radiation on genital cells which at the time of irradiation are i
the stage of spermatozoids or at stages close thereto, and also whether
or not irradiation affects genital cells formed during the process of
recuperation.
Known are a large number of researches conducted mostly on insects,
in particular on Drosophila, in which it was shown that hay irrro'liation
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dasages primarily the nuclei of speraatozoids or of cells which are at
stages close to these. Nuclei of cells present at earlier stages of
spermatogenesis (spermatogonia) Buffer to a considerably lesser degree
from the radiation damage (Shapiro, 1831; Serebrovskaya and Shapiro,
1935). Similar data were subsequently secured in regard to mammals
(Hertwig, 1535). Thus, every reason existed for assuming that the viability
of offspring derived from irradiated males after restoration of their
fecundity will be substantially greater than that of the offspring derived
from the same males directly after their irradiation.
The data which we have analyzed hereinbefore provide in part an
answer to the question concerning the effects of an irradiation of sperma-
tozoids on the viability of the offspring. It is known that the supply of
sperms contained in the testis appendage of the male mouse is sufficient
for only four to five copulations (Hertwig, 1935). Hence, it was of
interest to determine the size of the first litters sired by irradiated
males and the number of stillborn. In testing the fertility of males,
females were brought together with them successively at different time
intervals following the irradiation, and in every instance records were
kept of the effectiveness of the mating. Thus, we were in a position to
segregate the first litters irrespectively of the fact whether they were
sired immediately after irradiation or at a later time. Data showing
the average size of only the first litters and the number of etillbirhp.
occurring therein are shown in Tables III and IV of the addendum. A
comparison of these data with those shown in Tables 4 and 5 indicates
that the number of offspring in tht first litters is lees, and the number
of stillbirths is greater by comparison with the subsequent litters. The
data of the above-mentioned tables also permit reaching the following
conclusions the average size of the first litter and the number of still-
births do not depend upon whether this litter was sired immediately after
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irradiation or at a later date. These data indicate the high senaitivity
of spermatozoids, and the stages close thereto, toward irradiation (as
concerns the effects upon the offspring), which is in good agreement with
the large number of facts concerning the action of ionising radiation on
alteration of hereditary characteristics of animals and plants (Shapiro,
1531; 9erebrovskaya and Shapiro, 1935; Hertwig, 1935).
To provide an answer to the question concerning the effects of
irradiation on the genital cells formed during the process of reparation,
a special investigation was carried out. Each of the irradiated males
(dosage 4001') sired up to seven to eight litters. Records were kept of
the size of the litters and the number of stillbirths. It was assumed
that if the first litters are produced by genital cells which at the
time of irradiation are present in the form of spermatozoids, the sub
sequent litters will be produced by germinal cells which at the time of
irradiation were at earlier stages of the spermatogenesis. U the damaging
action of Xraya affected only the mature genital cells or the ells
approximating them in their stage of development, then, in the presence
of an increasing consecutive number of litter, its size will increase
and the stil4births will become iess numerous. If, on the other hand,
the damaging action of Xrays affects to the same extent germinal cells
which were at earlier stages of gametogenesie all the litters sired by
irradiated males will be about the same size and will include the same
proportion of stillbirths. The results of these experiments are shown
in Table 10 and Figure 13.
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TABLE 10
AVERAGE NU!lggS OF OFFSPRING AND STILLBIRTHS IN A CONSECUTIVE SERIES
OF LITTERS
f Average number of offspring Stillborn offspring
b
consecutive
nunab?rs of
litters
number of
litters
er o
Total num
per litter (Miji)
offspring Numbe3
9."7?0.30 18
Percent
6.511.98
1
59
277
4
3.411.64
:i.3t0.38
a
zs
122
1
1.011.02
410.61
6
15
.
96
2
2.831.96
5?0.81
G
11
,
71
411.03
7
,
3?
1
2.212.16
i
6.6?0.43
46
N
m
7,4?1.40
37
i
8.0
Control
1-8 182
1240
s,sto.i7
13
1.030.28
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Table 10 includes data showing the average number of offspring
and the number of stillbirths in the consecutive litters. Figures
characterizing the average size of the litter show that, from the first
to the fourth litter, the average number of offspring per litter (being
relatively small) increases and remains thereafter at about the game level
as is observed with the controls.
The graph of Figure 13 shows the distribution of litters sired by
the irradiated and the control males, depending upon their size (numerical
data are shown in Table V of the addendum).
The graph shows clearly that the size of the first two litters sired
by the irradiated males is smaller than those sired by the control animals
2.1f0.34 and Mdi; 2
(the differences are statistically reliable: ~dif 1
?.3?0.41), whereas the size of the subsequent litters approximated that
of the controls. It should be noted that the first two litters sired by
irradiated males are not only less numerou$ in comparison with the sub-
sequent litters but the stillbirths therein are encountered oftenest.
This permits arriving at the following two conclusions. (1) The
most sensitive stage of spermatogenesi$ as concerns the effects upon the
offspring is the stage of spermatozoids. It is precisely from cells,
which at the time of the irradiation were at the stage of spermatozoids or
aloes thereto, that there could be produced the first two litters in
reduced and the greatest number
which thta number of offspring is daarply
of stillbirths occur. (2) Mature genital cells formed during the
roce$S do not carry gross traces of the damaging action of
re'-aration p
i n- This is evidenced by the normal Size of the eub5e1uent
,rradiatio
litters mad a number of stillbirths approximating that sired by the con-
r
trols
These conclusions are impo*tant not only froaa a theoretical standpoint;
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possibly they are of no lesser practical significance. Although
we are not justified to apply directly to humans the data secured
by means of investigations of laboratory animals, we should still take
them into account. If subsequently, these data will be confirmed in
other mammals, the following conclusion of practical importance can be
arrived at; in cases of radiation therapy applied to the region of testes
there is less to be feared from the damaging action of the irradiation
upon the issue of the patient if this issue is derived from genital cells
which were produced after the recovery of fertility.
postembryonic Development of Offspring Sired by Irradiated Males
As was previously stated, decreased number of offspring derived from
males subjected to irradiation is due to the death of the embryos at
different stages of their development. In this connection, the question
arose as to the normalcy of the course of postembryonic development of
mice sired by these males, and whether the traces of the damaging action
of penetrating radiation affect the postembryonic development of the
offspring. To provide an answer to this question a follow-up was
carried out on the postembryonic development of offspring resulting from
the mating of males subjected to an irradiation of 240 and 400r with
nonirradiated females. Investigations were conducted not only on the off-
spring of litters sired soon after the irradiation (first and second
litters), in which by analogy with the embryonic development a maximum
effect of the irradiation could be assumed, but also on litters sired 3
months after the irradiation (sixth and seventh litters), in which the
number of offspring and the percentage of stillbirths did not differ from
those of the controls, and hence a damaging effect of irradiation could
hardly be assumed to be present. 4baervation of the animals were carried
out through the first month and a half of their postembryonic development,
i.e., from the time of birth to sexual maturity. As a result, it was
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ascertained that the survival of the members of the first generation eired
by the irradiated males does not differ from that of the controls (off-
spring of first and second litters, as well as those of sixth and seventh
litters).
In processing the materials, separate account was kept of the
mortality of the offspring of different sex. No differences in the sur-
vival rate of males and females were detected. A determination was also
made of the ratio of sexes among the offspring of irradiated males. The
literature contains conflicting data on this subject. According to some
investigators (Parkes, 1925; Kalmus, Metrakos, and Silverberg, 1952) the
ratio of sexes among the offspring of irradiated males deviates from the
normal. According to the data of other researchers, no deviations from
the normal are found in the numerical ratios of the sexes (Trasher and
Metrakos, 1953). In our experiments, determinations were made of the
number of females and males in litters sired at different intervals
of time following the irradiation of the males. Table 11 shows the
data relating to the ratio of sexes in litters sired immediately after
irradiation of the males. These data show that the offspring of irradiated
and of control males contained equal numbers of either sex. Similar re-
suits were obtained upon analysis of the offspring sired one month and
3 months following irradiation.
Notwithstanding the large number of litters analyzed, we have found no
morphological deviations in the development of the offspring of the animals
of the experimental groups, as compared with that of the controls.
we have also studied the growth of animals sired by the irradiated
males. It is known that growth regularities are usually studied by
aeons of the weight change curves of the developing animals.
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TABLE 11
SEX RATIOS OF THE OFFSPRItIG OF IRRADIAT&D MALES (LITTERS SIRED
II~tiDIATELY AFTER IRRADIATION)
Irradiation
dosage
Total number
of offspring
Including
% of males
(r)
200
148
males
79
females
69
53.4?4.10
400
110
00
50
54.5?4.75
Control
150
74
7t;
49.3?4.10
The mice were weighed at the following time intervals: first, fifth,
thirteenth, twenty-first, twenty-eighth, thirty-fifth, and forty-second
day alter birth. These time intervals were selected for the following
reasons. The fifth day coincides with the appearance of fur, the thirteenth
with the opening of the eyes, twenty-first with beginning of independent
food intake, and the twenty-eighth with separation from the dam. Since
the growth characteristics of mice depend upon the sex and number of
animals in the litter, a corresponding grouping of the data was carried out
in the processing data. Thus, the average weight of the animals during
the different postembryonic periods of development was calculated separately
for the females and for the males. Separate processing was carried out
on the data relating to litters containing from one to three, from four
to six, and from seven to nine offspring. Data were derived which
characterize the growth of offspring sired immediately after irradiation
of the males, i.e., of the first two litters and also of that sired 3 months
after the exposure, i.e., the sixth and seventh litters. Numerical indices
relating to the changes in weight of the animals are shown in their
entirety in Tables VI, VII, VIIIQ and IX of the addendum.
Here we confine ourselves to a consideration of data limited to
offspring sired immediately after irradiation. Figure 14 shows the curves
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of growth of the animals, which reveal upon examination a great similarity
between experimental and control groups in all three series being compared.
The data cited leave ro doubt that the postembryonic development of mice
sired by the irradiated males does not differ from that of mice derived
from nonirradiated parents.
On comparing the data on postembryonic development of mice sired
by the irradiated males with analogous data relating to their embryonic
development, we note the following interesting regularity. While the
embryonic development of offspring sired by irradiated males bears the
mark of the damaging action of the radiation, the postembryonic develop
ment is free from these effects. Evidently the gross damage arising in
the genital cells due to the action of irradiation manifests itself during
the process of embryogenesis, and the individuals which pass through this
peculiar filter are found to be practically normal.
Conclusions
1. Xray irradiation of males has the following results.
(a) Decreased mating capability of the males
(b) Decreased number of offspring in litters resulting from
mating with nonirradiated females.
2. The offspring of irradiated males includes a large number of
stillbirths.
3. Decrease in the number of offspring of irradiated males occurs
due to the death of the embryos during different stages of their develop-
merit. Thus, the problem of the sterilizing action of Xrays is partially
a problem of the development of the offspring of irradiated males.
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4. ~hauges in weight of the testes can serve as a qualitative
index of the sterilizing action of gays.
5. The greater the exposure dosage of Xrays, the greater their
effect on the fertility of the irradiated males.
6. After the lapse of one to 3 months following irradiation, the
fertility of the males becomes restored. This recuperation proceeds at
a rate which increases with decreasing dosage of exposure.
7. Restoration of testes proceeds slower than the overall re-
cuperation of the irradiated organism.
8. Mice resulting from the fertilization of ova by spermatozoas
developing from regenerated germinal cells do riot differ in viability
from the controls. In other words, the regenerated genital cells bear
no gross traces cf the damaging action of radiation:
9. Postembryonic development of the offspring sired by the irradiated
males reveals no deviation from the normal.
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Eschenbrenner, A. B., Miller, E., Effect of Roentgen Rays on the Testis,"
Arch. Path., 1950, 50, 738749
Fogg, L. C., Cowing, R. F., "Effect of Direct X-Irradiation on Mammalian
Testicles," Exper. cell. Research, 5, 1952, 19-32
Geller, F. C., "Der Brunstcyclus der weissen Maus each Sterilisationsbestrahlung
nebst ailgemeinen Betrachtungen ueber den Brunsteyclua ueberhaupt,"
Arch. f. Gynaeck., 139, 1930, 530-536
Halberstaedter, L., "Die Binwirkung der Roentgeustrahlen auf Ovarien,"
Berl. Kiln. Wochenschr., 42, 1905,64-66
-174-
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 Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Hertwig, p., "8teritaetserscheinungen bet roentgenbeatrahlten Meeueen,"
Zeitschr. Abst. Vera Vererb. Lehr., 70, 1935, 517.523
Hertwig, P., "Unterschiede in der Entwicklungsfaehigkeit von F1 Maeusen
nach Roentgen-Bestrahlung von Spermatogoni.en, fertigen and un-
fertigen Sperastozoen," Biol. Zbl, 58, 1938, 237-301
Kalmua, H., Metrakos, J., Silverberg, M., Sex Ration of Offspring from
irradiated Male Mice," Science, 116, 1952, 274-275
Neuhaus, M., "Sterility Mutations in D. Melanogaster," Dros. Inform.
Serv,, 7, 1937, 91-92
Parkes, A. S., "The Effects on Fertility and the Sex Ratio of Substerility
Exposure to Xrays," Proc. Roy Soc. Biol. 98, 1925, 415-438
Russell, w. L., Maatualian Radiation Genetics, Symposium on Radiobiol.,
1952, New York, 427-440
Schugt, P., "Untersuchungen ueber die Wirkung abgestufter Dosen von
Roentgenstrahlen vershiedener Wellenlaenge auf die Struktur and
Funktion der Ovarien," Strahlentherapie, 27, 1928, 603-662
Shaver, S. L., "X-Irradition Injury and Repair in the Ger~ainal Epithelium
of Male Rats, I. Injury and Repair in Adult Rats, "Am. Journ. Anat.,
92, 1953, 391-433
Snell, G. D., "Xray Sterility in the Kale House Mouse," Journ. Exp. Zool.,
65, 1933, 425-441
Snell, G. D., Bodemann, E., Hollander, E., "A Transiacation in the House
Mouse and Its Effect on Development," J. ~cp. Zool., 67, 1934, 93-
104
Trailer, D. O., Metrakos, J. D., Sex Ration of Offspring froa Irradiated
Male Mice," Genetics, 38, 1953, 697
- 175 M
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 1
N
EA
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ADDENDUM
TABLE 1
DISTRIBUTION OF ANIMALS THAT DID BY HAYS OF OBSERVATION
gsFosyte Mbar of
dosage animals
(s*) that died
- N N LC) T
1 1 1 1 I
+-I N M q~ u~
Dsys of observation
e?. ca m ~
1 1 i 1 -4
N t) '+ L.) tD
-4 pd *i r4 rl ri
1 ( I I I I
2 N M
.i ~ ??1
ri N M '~ it: t4 h aQ Q
9-4 I N C N N N N N N C44N M
1 I 1 I 1 I : 1 i
M i' N
t* i CA C N N N Ci N N tL'
;.~ ri ri N N N N N
200 7 - .a - Z 3 1 1 - _ - .- 1 r w - 3 w
400 48 - - - - - .+ 2 6 7 5 8 1 4 - 2 Z - 1 - 11.4
Control
Mean duration
of life (days)
400 11.5
Control
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S S
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TABLE II
AVERAGE N[AB&R OF OFFSPRING IN LITTERS WHICH IICLIIDBD OR DID IT IEI.IS
STILLBIRTHS
Exposure
Litters Including Stillbirths Litters Not Including Stillbirths
dosage
(r)
Number of
litters
Number of
offspring
Average number
of offspring
per litter
Number of
litters
Number of
offspring
Average nuaber
of offspring
per litter
aoo
5
32
6.4
50
296
5.9
4010
10
58
5.8
49
219
4.5
Control
2
16
80
60
424
7.1
N
y
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I
AVERAGE NUMBER OF OFFSPRING IN LITTERS SIRED BY IRRADIATED MALES. O3ASED
Exposure
dosage
(r)
(Wd1W'
Time
Ismediately After 1 month
Number Number Average Number Number
of of off- number of of of off-
litters spring offspring litters spring
per litter
(gym)
TABLE III
After 3 months
Number Number of Average
Average
of of offspring number
number
litters of off-
offspring
Per litter spring
per litter
(M?m)
total
Number Number Averago
of of off- number
litters spring of off-
spring
per litter
(M?m )
200
Zy
143
5.7?0.32
21
133 6.3?0.45 9
52
5.8?0.78
55
328
6.010.26
400
39
172
4.4t0.29
:>
16 3.2?1.5 1G
gg
5,9?0.62
59
277
4.7?11.30
Control
36
263
7.310.33
19
119 6.3?O.0 '1
58
8.3?1.06
fit
440
7.110.30
I
Excluding Stillbirths
w
o~
9
32
~.8?O.73
55
320
5.913.27
i
200
25
136
5.4?0.33
71
132 0.3?0.46
400
39
161
4.1?0.30
5
14 2.8?1.21 15
$4
5.6?0.63
59
259
4.4?0.29
Control
36
262
7.3?0.33
19
116 6.1?0.uO 7
56
3.3?1.OE3
62
436
7,0?0.30
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TABLE IV
Exposure
dosage
(r)
NUMBER OF STILLBORN MICE IN LI1T&RS OF IRRADIATE YAI.SS (BASSO ON THE FIRST LITTERS)
Time Following Irradiation
Immediately After 1 month After 3 months Total
Stillborn Stillborn Stillborn Stillborn
Offspring Offspring Offspring Offspring
investigated Number Percent investigated Number Percent investigated Number percent investigated Number Percent
200 143
400 172
Control 263
Exposure
dosage
T 4.9f1.81 133 1 0.710.72 52 - - 328 8 2.410.85
11 6.411.87 16 2 12.530.27 89 5 5.632.44 277 18 6.5?1.48
1 0.4?0.40 119 3 2.511.43 58 - - 440 9. 0.910.45
TABLE V
SIRED BY IRRADIATED HALES, ACCORDING
DISTRIBUTION OF LITTERS
,
TO THEIR SIZE
Number of offspring per li1;ter
Litters
1
2
3
4
S fi
7
8
9 10
11
12
13
Total number
of litters
1 and 2 Number
7
5
8
16
18 9
9
5
3 2
-
-
-
SZ
96
8.5
6.1
9.7
19.5
22.0 11.0
11.0
6.1
3.7 2.4
-
-
-
100.0
From 3 Numlaer
-
4
1
2
5 A
10
11
3 2
2
-
-
44
to 8 %
-
9.1
2.3
4.5
11.9 9.1
22.8
2~.0
6.8 4.5
4.5
-
-
100,0
All Humber
2
10
6
10
15 3G
33
38
20 11
5
Y
1
182
q,
1.1
5.5
3.3
5.5
8.3 16.5
18.1
29.8
11.0 &.u
2.7
0.E
0,6
100.0
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00
0
TABLE VI
CHAtRrffi IN WEIGHT OF OFlSPAI*'i SIRED IAI1RDIATffi.Y AFTER IRRADIATION OF
THE MALE
Litters numbering 1 to 3 oYYspring,~Q
Exposure Average number Days of weighing
dosage Number of of offspring Number
(r) litters per litter 1 5 13 21 28 35 42
aoo i
400 10
Control 7
2.0 2 1.5 3.8 6.9 10.5 13.9 16.4 17.5
2.9 14 1.5 3.4 7.6 10.5 15.1 18.7 20.5
2,8 10 1.6 3.6 6.3 10,1 13.5 16.7 18.6
Litters numbering 4 to 6 offspring,
200 14 4.9 93 1.5 3.3 6.6 9.7 13.5 17.0 18.1
400 10 5.3 23 1.6 3.3 6.6 9.3 12.5 16.2 18.8
Control 12 5.6 37 1.4 2.9 6.1 8.5 12.1 15.3 16.8
Litters numbering 7 to 9 offspring,??
gpp 7 7,3 25 1.3 3.1 5.3 7.7 10.9 14.3 15.8
400 3 7.3 10 1.6 3.4 5.8 8.5 10.1 15.8 16.7
Control 13 7.7 45 1.5 2.8 5.4 8.2 11.7 14.7 15.9
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 TABLE VII
CHAI9GSS IN WEIGHT OF OF!'SPRI*'r SIRS IMMEDIATELY AFPER IRRADIATION OF THE MALE
d
Litters numbering 1 to 3 offspring,
i
N
Co
1
Exposure Average number
dosage Number Qf of offspring
(r) litters per litter Number,
I~
200 3
400 12
control 5
xpp 13
400 31
Control 12
200
400
Control
3
13
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Days of weighing
1 5 13 21 28 35 42
2.7 3 1.8 2.8 5.9 8.6 12.6 16.6 18.1
2.6 17 1.5 3.3 7.2 10.6 15.2 19.1 22.1
2,8 g 1.7 3.4 7.9 11.0 14.4 17.7 20.9
Litters numbering 4 to 6 offspring, d^ b
4.9 32 1.5 3.3 6.6 9.8 13,2 17.6 19.3
5,2 Zq 2,6 3.3 7.2 9.7 13.8 18.2 20.7
5.6 30 1.4 2.9 6.1 9.1 18.3 16.9 18.8
Litters numbering 7 to 9 offspring,
7,5 34 1.3 3.1 5.3 7.8 11.9 15.3 17.2
7,3 12 1.6 3.4 6.0 9.1 11.5 17.3 19.5
7,7 52 1.4 2.8 5.4 3.4 12.3 15.8 17.2
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TABLE VIII
CHANGES IN IYSIGIiT OF OFFSPRING SIft~ 3 MONTHS AFTER IRRADIA'7I4'iT
Litters numbering i to 3 offspring,~ ~
Exposure
dosage pwber of Average number Number Days of weighing
Cr) litters of offspring
per litter 1 5 13 21 28 35 42
200 2 2,0 3 1.9 3.5 10.3 11.4 14.5 16.9 23.0
400 1 3.0 1 1.8 3.3 8.4 13.6 - 19.E 21.7
Control 1 3.0 1 1.2 3.2 6.6 - 10.0 11.9 14.7
Litters numbering 4 to 6 offspring, ??
Zpp 8 5.5 21 1.6 2.9 6.4 8.1 11.6 14.8 17.3
400 5 4.8 15 1.7 3.6 6.1 9.2 12.2 16.1 17.1
Control 7 5.4 19 1.5 2.7 5.6 8.3 11.1 14.6 17.6
Litters numbering 7 to 9 offspring,
gpp g 7,5 27 1.4 2.6 5.5 7.3 9.7 21.4 13.6
400 g 8.0 24 1.3 2.7 5.7 7.6 10.6 12.8 14.8
Control 6 7.8 22 1.3 2.5 4.7 6.5 9.3 12.0 13.7
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87~posurs
dosage
(r)
Number
of
litters
200
2
400
1
Control
2
200
9
AOQ
5
control
T
oc
w
i
200
8
400
8
Control
6
TABLE IX
CHANGES IN WEIGHT OF OFFSPRING SIRED 3 MONTHS AFTER IRRADIATION
Litters numbering 1 to 3 offspring,
cT~
Average number Days of weighing
of offspring
per litter Number 1 5 13 21 28 35 42
3.0 4 1.3 2.3 2.4 8.4 12.4 18.9 21.0
3.0 2 1.6 3.3 7.9 12.3 - 18.7 21,6
2.0 4 1.7 4.3 8.7 10.9 14.8 19.7 23.4
Litters numbering 4 to 6 offspring,
5.4 28 1.5 3.1 6.0 8.5 12.2 15.3 17.6
4.8 9 1.7 3.6 6.3 9.6 12.6 16.0 18.9
5.4 17 1.5 2.7 5.6 8.1 12,2 14.9 18.3
Litters numbering 7 to 9 offspring,
7.5 27 1.4 2.6 5.3 7.5 10.3 12.4 14.8
8,0 24 1.3 2.7 5.8 7.7 10.5 12.8 15.1
7.8 22 1.3 2.5 4.9 6.6 11.3 12.5 14.4
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 Iuul
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. U u d 12 1. L5
Doy~ of observation
Fi pure 1. Ch &rr es i..n ;:ht of survivin,; rn:-1.e :i'oii c~winr irr t i i J, i on.
1, control; 2, doe ?OO r; 3, (1() f;!' ).100 r,
'
_. s-----"--
\
1/ ,> 49 Z
Days of observation
Ffture 2. Survival of race fol]owinF; irradiLion.
1, control; 2, rjo+Fe 200 r; 3, dosage li00 r.
/001
'`'z
!
\ti 1' /T T
1,
\V ,
Time following irradiation, in days
Figure 3. Changes in wei ht of body, tt!Sts'D, and acceory genital
glands of irradiatr d males.
Lody wci ght: 1, dosage 200 r; 2, dosage Lt00 r. Weight
of testes: 3,, dos-e 200 r; h, dosai P LtOO r.
Weight of accessory genital glands: 5, do:,age 200 r;
h, dos i o 1100 r,
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Figure !1. Microphotograph of testa. of an animal irradiated with
200 r. Fixation 15 days after irradiated. 1).O X.
Figure ~. Microphotograph of testis of an animal irradiated with
1400 r. Fixation 15 days after exposure. l!40 X.
Figure 6, Microphotograph of testis of a control animal of the
same age. 1140 X.
a. 188
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Figure 7
: ?.s~
Figure 8. Microphotograph of testis of an animal irradiated with
1430 r. Fixation 30 days after irradiation, 1)O X.
Ma crophotof''r )h of test,i.s of an animal irradiated with
200 r' Fixtcd pr,b1eii, youurrg,
sezually mature, nonparent females of strain C57 (black) (24 animals) were
subjected to a single total irradiation with a dosage of 100r. The condition
of irradiation were the same as those utilized with animals of strain A.
The controls consisted of 25 fem lea of strain C5, `black), of the same ag~
Results of the experime-,ts on Xray irrad.ation of females of strain
C57 (black), were compared with analogous data obtained with mice of strain
A. Table 5 shows the percent ratios and duration of the individual stages
of the oestrous cycle in the mice of the strains under investigation. These
data testify to the similarity of the disruptions of the oestrous cycle
in irradiated mice of strain A and C57 (black), Experimental groups of
mice of toth strains show, beginning with the second month following irra-
diation, an appreciable decrease in the frequency of occurrence of the
stages of oestrus and proestrus, and increased frequency of the occurrence
of the dioestrous and metoestrous stages. A still more considerable de-
crease in the frequency of occurrence of the stages of oestrus and proestrus
was observed during the third and sixth month following irradiation. For
example, during the third month, in the females of strain A, 8.7% of the
total number of tests indicated a stage of oestrus and proestrus; while in
females of strain C57 (black) this figure was 10%, as compared with 57 and
50% in the controls. During the sixth month the data were 13.5 and 7.550
respectively for the experimental animals; 55.9 and 45.9% for the controls.
It should also be noted that with increasing disruption of tie oestrous cycle
in the irradiated mice there is observed a prolongation of the stages of
dioestrus and metnestrus, and this is more sharply manifested in females
of strain A. The mean duration of the stages of oestrus and proestrus in
experimental and control animals showed no difference. Drastic prolongation
of the oestrous stage in females of strain A during the sixth month following
irradiation is connected with a long-lasting oestrus in one of the females,
which also constitutes an indication of the disruption of the genital cycle
in the irradiated mice.
"218."
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the ore hand, and on the other of females which had or had not previously
given birth. To provide a solution of ;rust fcr iul>tcd pr,b1eii, youurrg,
sezually mature, nonparent females of strain C57 (black) (24 animals) were
subjected to a single total irradiation with a dosage of 100r. The condition
of irradiation were the same as those utilized with animals of strain A.
The controls consisted of 25 fem lea of strain C5, `black), of the same ag~
Results of the experime-,ts on Xray irrad.ation of females of strain
C57 (black), were compared with analogous data obtained with mice of strain
A. Table 5 shows the percent ratios and duration of the individual stages
of the oestrous cycle in the mice of the strains under investigation. These
data testify to the similarity of the disruptions of the oestrous cycle
in irradiated mice of strain A and C57 (black), Experimental groups of
mice of toth strains show, beginning with the second month following irra-
diation, an appreciable decrease in the frequency of occurrence of the
stages of oestrus and proestrus, and increased frequency of the occurrence
of the dioestrous and metoestrous stages. A still more considerable de-
crease in the frequency of occurrence of the stages of oestrus and proestrus
was observed during the third and sixth month following irradiation. For
example, during the third month, in the females of strain A, 8.7% of the
total number of tests indicated a stage of oestrus and proestrus; while in
females of strain C57 (black) this figure was 10%, as compared with 57 and
50% in the controls. During the sixth month the data were 13.5 and 7.550
respectively for the experimental animals; 55.9 and 45.9% for the controls.
It should also be noted that with increasing disruption of tie oestrous cycle
in the irradiated mice there is observed a prolongation of the stages of
dioestrus and metnestrus, and this is more sharply manifested in females
of strain A. The mean duration of the stages of oestrus and proestrus in
experimental and control animals showed no difference. Drastic prolongation
of the oestrous stage in females of strain A during the sixth month following
irradiation is connected with a long-lasting oestrus in one of the females,
which also constitutes an indication of the disruption of the genital cycle
in the irradiated mice.
"218."
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Figure 4 shows the curves whic} characterize the frequency of
occurrence of individual stages of the oestrous cycle in mice of strain A
and C57 (black). Frequency of occurrence of the oestrous stage (and also
of the other stages) is expressed on the curves in indices which indicate
the ratio of the percent of irradiateu
d .....
-ir\..... AM aa, Oho OAAt'OU St AtiEi~ tO
11C1Wwa r,uc wv .~~ a.~ ......,~...
the percent of such females in the controls, Expressed in an analogous
manner are the other data pertaining to the course of the oestrous cycle.
The curves show clearly the same nature of the changes of the oestrous
cycle in the irradiated females of both strains under study.
The similarity noted becomes even more apparent oz-i comparieon of
the data on the number of normal cycles per female and the number of females
in cycle. From Table 6 which shows these data it is apparent that dis-
ruptions in the course of the oestrous cycle in females of both strains
begin with the second month following irradiation and during the third and
sixth months the cycc process of the females ceases almost completely.
While in the control groups, during the entire period of investigation,
there occur 3.5 to 4 normal cycles monthly per female, with a mean dura-
tion of the cycle being 0-7 days; in the irradiated mice, on the second
month following exposure, there occur 1.2 cycle per female of strain A
and 2.2 cycle per female of strain C57 (black). On the other hand, during
the third and sixth months after irradiation, there are respectively 0.5
and 0.6 cycle per female, with a mean duration of the cycle being about
6 days. The difference between experiment and control as concerns females
of both strains is statistically fully reliable. Exceptions are the data
relating to the second month after irradiation in the case of mice of the
strain C57 (black), wherein the reliabil;ty of the difference is equal to
2.56. Similar data were also obtained on the percent of females in cycle.
- 220 -
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 * ~,,, ?~,
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For a more apparent comperisozt of the course of the oestrous cycle
irA mice of strains A and C57 (black), the above-stated data are presented
in the form of curves in Figures 5 and 6. In this ix~stancn, as beiura, the
average r:umber of cycles per female and the percent of fe iales irz cycle are
expressed iz3 indices (the ratio of experi.ent to control).
The results of the experiments peri,..t reaching the following G o:-
clusion. Disruptions of the oestrous cycle, induced by a single total
Xray irradiation with a dosage of IOor, are of a s{znilar nature th miee
of strains A and C57 (black). Thus, the latent period is both stra r3
lasts approxioately ox~e month, and disruption of the normal cyclic process
of females is observed only beginning with the second month following
irradiation, wring the third and sixth month after Irradiation there is
observed an almost complete cessation of the cyclic process.
To determine the question concernir3g the nat~ire of the reaction
to Xray irradiation in parent and nonparent females, a special experi&ient
was carried out. Upon reaching sexual maturityG a group of females of
strain A, of the same age, was kept with males of the same strain. Of
this group of mice 72 females were selected, which had given birth twice
in succession (without having suckled the offspring), of which 24 wore
subjected to a single total Xray irradiation dosage of 50r1 22 to s dosage
of IOOr, the 27 nonirradiated females serving as controls. In the enumeratod
females the oestrous cycle was studied according to the procedure adopted
by UL The results of this experiment are compared with the data obtained
on single total Xray irradiation (with the same dosages) of nonparent
females of strain A.
Tables 7 and d show the percent proportions and the duration of
individual stages of the oestrous cycle in parent and nonparent females.
The data of Table 7 show that with a dosage of 50r disruptions in the course
of the oestroua cycle in parent and nonparent females, begin with the second
month following irradiation, In the nonparent females the difference
221
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between experiment and control in the percent of occurrence of individual
s'agnn of the oestrcus cycle is $tUtisticaily reliable, whereas in the
case of the parent females these differences -? although they do not reach
a statistical reliability -- are of sufficiently high probability (reliability
of the difference as concerns the oestrus stage is 2.28; it is 2,46 as
concerns the proestrous stage). I the :iMVUr~ of disruptions of the
course of the oestrous cycle during the third and sixth months following
irradiation, the groups of females being compared show no differences,
It is possible that the later manifestation of disruptions in the course
of the oestrous cycle in the parent females as compared with the nonparent
constitutes an indication of their somewhat enhanced resistance to Xray
irradiation, sowever, no definite conclusion can be reached on the basis
of the data cited.
A comparison of the results concerning the Xray irradiation of parent
and nonparent females with a dosage of 100r (Table 8) shows that disruptions
in the course of the aestrous cycle in the animals of both groups occur
similarly. In the parent, as well as in the nonparent females, beginning
with the second month after irradiation, there is observed a statistically
reliable -- as compared with the controls ?- lowering of the percent of
occurrence of the stages of oestrus and proestrus, and an increase in the
stages of dioestrus and metoestrus. During the third and sixth months
the differences between experiment and control are manifested in animals
of both groups even more sharply.
For a more apparent comparison of the frequency of occurrence
of individual stages of the oestrous cycle in parent and nonparent females
the corresponding data are expressed in indices (ratios of experiment data
to the control data taken as equal to 1), and are represented as curves in
Figure 7.
. 222g.
*lpysm ?Ib
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
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6
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
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6
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
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6
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
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6
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
n?
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
n?
6
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
n?
6
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
n?
6
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
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6
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
n?
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
n?
6
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
n?
6
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
n?
6
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 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
n?
6
7
._ V..
 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
n?
6
7
._ V..
 TAHI.S 6
$lYSCT8 OF XRAY IRRADIATION ON THE COURSE OF THE OESTROUS CYCLE IN !LICE OF DIFFERENT STRI-IN9. (IRRADIATlbli
DOSAGE lOOr)
Time following irradiptloa 1 tenth g ti
Strain A CJ? (black) A Cyr (black)
$Xp@P1IDBAt Control Experiment Control Experiment Contr. Experiment Control
Total nuabsr of la~ales 24 24 24 25 24 24 16 29
I
,
96,0 79 .0
5
81
Derr nt nt ll~sstlu-~ it f nvni 1 f A A m k
96
0
--
.
Average nueber of noraal cycle
per }A~!l110 40?025 42
O3 ?
?
....3,3?0.3176?0. 231?041
.s nn ...., .... ..... ..,.. _
333 -.-? .-_ _.___.__..
.
3 6.52
6
seen duration of normal cycle
ffL YR 1 ' A # )
3 months 6 eont
h8
Total Auaue: of teaie@ Za 24 zt xz is as is 17
Percent of Ywalea is cycle 37.i 100,0 43.7 95.2 1X.1 95.5 25.0 100.0
Average nv^ber of noa~al cycles
per file 0.5t0.16 4.310.29 0.63r0.22 4.0530,22s 0.1?0.03` 2.4?0.24* 0,5?4.28 3.64t0.24
Yvan duration of normal cycle
(days) 8.4 5.7 5.52 5.62 8.0 6.9 8.17 7.48
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
Sanitized Copy Approved for Release 2010/04/02 : CIA-RDP81-01043R000200180025-9
n .. _ _._
n?
6
7
._ V..