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Sperm formation and growth of accessory reproductive organs in hypophysectomized ground squirrels in response to substances from blood and human urine.

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SPN€i,\I FOR,14dTION AXD GROWTH OF AC~CESROR,Y
REPRODUCTIVE ORGANS I N HYPOPHYSECTO3IIZED GROUND SQUIRRELS I N RESPONSE
TO SUBSTRN('ES FBQAT FLOOD AND
HUMAN URTKE
ONE TEXT FIOUR.E BWD TWO PLATES (EIGHTEEN FIGUF~ES)
INTRODUCTION
Gonadotropic materials known to cause the testis of the
ground squirrel (Citellus tridecemlineatus ) t o produce spermatozoa and secrete testis hormone are : pituitary substance,
Antuitrin-S and blood serum from pregnant mai*es (Wells,
'35 a, '37; Wells and Moore, '36; Baker and Johnson, '36).
Anterior hypophysis implants in totally hypophysectomized
ground squirrels have resulted in sperm production (Wells and
Gomez, '37). It was deemed desirable to see if gonadotropic
materials other than pituitary substance would affect the reproductive organs in hypophysectomized males of this species.
The present papcr reports the response of reproductive organs
in liypophysectomized male ground squirrels t o administration of the following substances: 1) blood serum from pregnant mares, 2) an extract of urine from pregnant women
(Antuitrin-S) and 3 ) an extract of human male urine (Prospermin). Xost of our results have been abstracted elsewhere
( Wclls and Overholser, '38).
This study was aided by n grant to L. J. Wells from the National Research
Council, Committee for Research in Problems of Sex.
231
232
L. J. WELLS A N D RI. D. OVERHOLSER
Tlie writers are greatly indebted to the following individuals: Dr. F r e d F. McKenzie, Dr. A. J. Durant, and Dr.
Victor Berliner of the University of Missouri for preparing
blood serum from p r e p a n t mares; Ilr. A. ,J. Morrell and
E.R. Xquibb ancl Sons for furnishing Prospermin; Dr. Oliver
Kamm and Parke, Davis and Company f o r supplying Antuitrin-S; and Dr. Carl R. Moore of The TJriiversity of Chicago
who has given valuable advice.
XETHODS
Reproductive organs of eighty-six male ground squirrels
(Citellus tridecemlineatus) were studied grossly arid microscopically in this investigation. Fifty-four of the males T.r;ere
hypophysectomized and they survived the operation for
periods varying from 8 clays to 6 months.
Hypophysectomy was accomplished by an oral approach,
using both Nembntal and ether to induce surgical anesthesia.
The details of hypophpsectomy technique, post-operative care,
periodic examination of reproductive organs by manual palpation and observation, diet of the animals and autopsy
routine Tiwe identical with those published elsewhere (Wells
and Gomez, '37), and €or this reason they are not repeated
in this paper.
Hypoplij-sectomized animals tolerated iiijections poorly,
especially during the first few days of treatment. Ignorance
of this fact in the earlier experiments resulted in many deatlis
within the first week of injection. Siicli animals exhibited
local cutaneous swelling at injection sites and also symptoms
resembling thosc of hypoglycemia. Greater toleration was
secured in the later eAqerimentsby giving small closes f o r the
first few clays. Tolerance to injection was successfully established in most cases by giving 25% of the desired dose (no
more than 0.15 cc. of aqueous substance) once dailj- on the
first arid second clays, 50% on the tliird day, 75% on the fourth
and, finally, the desired dose twice daily.
In all experiments an injection period of 25 to 30 days was
planned. Sixteen treated animals tolerated injections so
I N D U C I N G SPERM AFTER H Y P O P H Y S E C T O M Y
2 33
poorly that it was necessary to terminate the treatment at an
earlier date than that wliich had been scheduled. Thirteen of
these died and autopsy was performed as soon as death was
discovered ; the interval between death and autopsy varied
from a few minutes to a maximum of 10 hours. Tlie other three
were killed when it appeared definite that they had no chance
of survival.
At autopsy the utmost care was taken in preserving possible
pituitary remains. The brain was cautiously removed without destruction of tlie dorsal dural wall of the pituitary. In
each hypophysectomized animal, the sphenoid bone was removed without disturbing the sella turcica and fixed in Bouin’s
fluid. This entire block of tissue was decalcified, sectioned
serially at 10 p, stained with hematoxylin and eosin and examined microscopically f o r fragments of pituitary tissue.
Most of the hypophysectomized animals were fourid to have
fragments of pars intermedia and pars tuheralis, and lack
pars nervosa with the exception o f tlie stalk wliicli almost
inrariably remained. Some anirrials showed no microscopically detectable trace of anterior lobe tissue, and such animals
are referred to in this paper as ‘completely hypophysectomized, ’ while others exhibited srnall fragments.
I n the cases that x-ere incompletely hypophysectornized, the
amount of anterior lobe tissue remaining was calculated in
percentage of tlie entire anterior lobe by the following method.
Every third section of anterior lobe fragments was marked
on the didp with an ink spot. The marked sections were then
projected by an Edinger projection apparatus at a magnification of 20 and the outline of thc sections drawn. The total
area of these drav-hgs was then clctermiried in square centimeters by means of a planimeter. A complete anterior lobe
from a normal animal was sectioned and measured by the
same method. Actual percentages of remaining tissue in the
experimental animals could thus be calculated. Figure 2
shows a section through a normal intact hypophysis. Figures
3 and 4 show sections through the sella tarcica from experimental animals in which the percentages o f anterior lobe fragments were 4.6 and 5.474 respectively. Figure 5 is a similar
234
L. J. \VELLS A N D M. D. OVEBHOLSER
section from a series in which no anterior lobe fragments
were found.
RESULTS
I. Normal malPs
Twenty-two normal males were killed during the August to
November period. Each had non-fmictional accessory reproductive organs and no spermatozoa (tables 1 and 2). The
testis was invariably abdominal in position and small.
The above results are in harmony wit11 those previously
reported (Johnson, Foster and Coco, '33 ; Moore, Simmons,
Wells, Zalesky and Nelson, '34; Wells, '35 a ; Wells and Moore,
'36). During the past 7 years, only six males exhibiting
spermatozoa within the aspermatic period shown in figure 1
have been discovered by one of us (L.J.W.). These six exceptional animals were killed near the extreme ends of the
aspermatic interval.
Hypophysectomized contl-ols
Nine immature males mere hypophysectomized during the
aspermatic period of the year and autopsied after intervals
of 44 to 93 days. Autopsied testes wcre small and in an abdominal position, and accessory reproductive organs were
similar to those of normal controls. There were great variations in testis weight; however, in most cases, the testis
weighed less than that of unoperated controls (tables I and 2)
and showed less advance in spermatogenesis (figs. 10 and 12).
Spermatozoa were not found in any case.
No consistent differences between the reproductive organs
of completely liypophysectomized controls and those of partially hypophysectomized controls were noted.
2.
3.
Injectioias
Pvospermim (extract of human male urine) was administered to twenty-two hypophysectomized males for periods of
6 to 31 days. I n most cases it was dissolved in distilled water
and injected subcutaneously twice daily in doses of 1.0 to
10.0 mg.
IXDUCTRG SPERM AFTER HYPOPHYSECTOl\fY
235
Fourteen of the animals proved to be completely hypphysectomizecl, eight immatwe and six adult. Each of five immature males injected for periods of 19 to 31 days showed
large scrota1 testes Ti-ith spermatozoa (table 1, fig. 14), and
accessory organs similar to those of normal breeding adults
(fig. 15). The average weight of accessory organs was approximately 27 (bulbar gland) to 63 (seminal resicles) times
greater than that for untreated controls, and the height of the
prostate epithelium was about 2 to 3 times greater. Three
othcr immature males injected f o r periods of' 10 to 18 days
exhibited no spermatozoa. Each of six adults possessed scrotal testes with sperm following an injection period of 30 t o
ezZa Sperrnotozoa
Young
k
;
;
:
:
Jun. Jul. Aug. Sept. Oct. Nov. Dec. Jan. Feb. Mar. Apr.
m:
1
;
;
May
rn/////////fyfQ@////J/m.j
Fig. 1 Occurrence of spermatozoa in 511 normal males autopsied (232 cases)
or castrated (278 cases) during the period 1931 t o 1938.
31 days. Onc of these adults was killed during the normal
breeding season ; however, at laparotomy preceding injection
one testis was removed and fourid to be small, abdominal and
aspermatic.
Eight males had anterior lobe fragments. Six of these had
spermatozoa at autopsy, ~ h i l etwo injected for intervals of
6 to 10 days had none. The presence of spcrmatozoa in one
animal killed on January 29th is not considered significant,
due to the fact that it had a large fragment of the anterior
hypophjsis and was killed during the season of sperm expectancy in unoperated controls.
0
W
N
,
-.
4
5
45
3
2'
1 2.4
1 4
4
2
34
Days
I
XOlsTH
KILLED
3140
19-26
Nov.
5.0
26
Nos.
38
5.0
30-37
10-18
Jan.
1.0-2.0
92
JSn.
3.0
31
3 w 7 3.0-10.0 6-10
Jan.
Jan.
31
92
5.0
None
49-93
Ok-Jan.
4443
None
Aug.-Jan.
Normal
None
Nov.
___ .- -.
179-185
5.0
38-185
5.0
140
31
5.0
Normal
None
grams
daily
Yilli-
--.
CBEATMEXT
-I-I
I
BODY
119
160
107
146
159
190
142
135
m.
50.0
0.0
0.0
8.7
4
5
.0
.~
0.0
TIWIIE
BIS
HYPOPHP
PEB C E N l
.~
WnIGIHT
I
0.6358
0.7055
0.3491
0.7546
0.4040
0.8550
-0.1858
0.1343
0.2522
.0.7346
0.8654
0.6626
0.3033
Urn.
Weight
+
+
+
0
0
0
_ -
+
+
0
+0
+
sperm 6
TIPSTIS
"I
~
.-
0.0033
0.0036
0.0042
0.1904
0.2372
0.2591
0.0135
__.
__.._
11.0
20.5
21.7
-
Proatata gland
m.
0.1684
0.7233
0.9692
0.2067
-0.0258
0.0252
0.0266
0.7506
0.7779
0.4802
0.1209
-
Bulbar
gland
weight
.
0.0054
0.0059
0.0060
0.4001
0.4624
0.3508
-0.0271
0.0728
0.3065
0.3378
0.0894
Pa.
Cowper'a
glands
weight
ACOESSOBY OBGAXS
4.3
1 2
0
'Data expreseed aa averages for each group.
'Autopsy date and presence of pituitary fragment make sperm presence not aigniflcant.
One testis surgically removed on April 19th before injection. It weighed 0.1695 gm. and contained no spermatozoa.
Prospermin suspended in olive oil.
Sixteen of eeventeen spermatic males showed spermatozoa in fresh epididymia smears, and eight exhibited apcrm motility.
Adult
Immature
I
I
PIXYSEOTOYIZED
-0-
DAY#
TABLE 1 1
Effects of Prospermin on testis and accessory gtalazaa of hypophyseotomiaed males
0.7371
1.0397
0.1566
0.8806
0.1172
2.0051
0.0094
0.0117
0.0090
0.9326
1.0718
1.3184
0.0375
--.
Urn.
weight
Seminal
vesiclee
I
NO.
~
1 -
DAYS
HYPOPHYSECTOXIZED
Suhstance
R.U.
daily
REATMENT
~
~
Pays
MONTH
KILLED
um.
117
112
120
BODY
WEIQHT
sls
T1ssuE
IYPOPHY-
'ER CENT
Urn.
I I
Weight
+
Sperm 6
TESTIS
Urn.
Weight
1
P
tz&,,
Prostate gland
um.
Bulbar
gland
weight
m.
um.
!owper's Seminal
glands vesicles
weight
weight
ACCESSORY CRGANS
4
2
5
16-19
51-55
56-59
17-30
53-88
63
44
58
Normal
Normal
1 1
+
+
0.1257
M.S.
40
dug.
0.0
0.1234
0
14.5 0.1421 0.0695 0.1470
9-12
0.2886 0.2226 0.5032
0.1394
Oet.
0.5119
19.7
29-33
0.0
40
M.S.
0.2584 0.2063 0.5257
13.7
0.4232
419.6
0.1244
2040
30-3 7 Sept.-Oct.
M.S.
0.1230 0.0687 0.1370
0.0419 I 15.5
Aug.
100-200 10-16
0.0
0.1197
0
105
Ant.
0.4128 0.3483 0.9027
0.1990
18.6
105
0.0
0.3950
Ant.
50-75
31-42 Sept.-Oct.
0.4950 0.3226 0.6200
Oct.
115
0.1860
24.0
3.9
0.3850
75
37
Ant.
0.0179 0.0061 0.0097
121
0.0038
None
Aug.
0.0956
0
10.7
9.0
0.0216 0.0046 0.0094
0.0034
None
Oct.
0.0
0.1625
0
133
8.6
2
0.0224 0.0068 0.0101
0.0038
Normal 0.0448
0
134
10.6
None
Aug.
6
0.0048
Normal 0.1905
174
None
11.1 0.0240 0.0066 0.0112
Oct.
0
2
0.4136 0.4426 0.3994
125
17
Ant.
50
13
Aug.
0.1545 0.1185 0.1275
6
190
100
8
Ant.
Aurr.
0.8181 1.1800
0.6006
M.S.
40
16
Aug.
170
19
1.5337
1.0757
0.4579
M.S.
25
30
140
0.2825
139
May
- 1
:4
0.1606 0.0396 0.0414
223
Normal None
Aug.
5.8
0.0193 1
1 8
Data expressed as averages for each group.
a A t laparatomy on August 8th before injection, testis size was 9.0 X 4.0 mm. Testis a t autopsy 12.0 X 6.0 mm.
a A t laparotomy on July 31st before injection, testis size was 12.0 X 6.0 mm. Autopsy testis 14.0 X 7.0 mm.
'One testis surgically removed on April 19th before injection. Testis weighed 0.1959 gm. and contained no spermatozoa.
S T ~ of
o four Antuitrin-S spermatic males had motile sperm in epididymia. Seven of eight mare serum males with testieular eperm also
had spermatozoa in the epididymis; sperm motility observed in four of these.
AGE
TABLE 2 1
Effects of mare serum and Antuitrin-X on testis and accessoru .a, l a d s of. hvvo. usectomized males
238
L. J. WELLS A N D M. D. OVEK€IOLSJ!X
Some of the average weights in animal groups of table 1
are someivhat greater f o r injected animals with hypophysis
fragments than are the average weights f o r treated males
with no detectable anterior lobe tissue. Comparison of
protocols f o r indiridual animals showed that this difference
in weight was related t o a variation in the length of the injection period and not to the presence or ahserice of an antcrior lobe fragment.
Ten non-hypophysectomized immature males were caastrated
bilaterally f o r 7 1 days. Nine of these were injected twice
daily for 30 days with doses ranging from 0.5 to 10.0 mg.
suspended in olive oil. The nine injected animals and one
untreated castrate were killed 017 the same day. The following data represent averages for the nine injected castrates,
and these data are compared respectively with those in the
nninjected castrate : body weight, 206 and 210 gm. ; prostate
gland, 0.0036 and 0.0037 gm. ; height of prostate epithelium,
7.3 and 8.6 p ; seminal vesicles, 0.0079 and 0.0094 gm. Since
no constant significant difference was found between accessory
organs of injected and untreated castrates, it is concluded
that prospermin in the above dose range exhibited no androgenic potency.
Blood serum f r o m pwqrzavzt rnarP.9 was given t o eleven immature and two adult hypophysectomized males in two daily
subcutaneous injections for periods of 9 to 37 days. The
daily dose ranged from 20 to 40 rat units.
Six immature and two adult males were completely hypophysectomized. Two immature males following 29 to 33 days
of injection (table 2, figs. 16 and 17), showed large scrota1
testes with spermatozoa and functional accessory reproductive organs. Average weight of accessory organs was approximately 1 2 (bnlbar gland) to 44 (seminal vesicles) times
greater than that for untreated controls. The remaining four
immature males exhibited lesser responses to shorter injection periods which varied from 9 to 12 days. These short
periods failed to cause testicular descent and sperm formation, but they resulted (probably indirectly through the testis)
INDUCIA-G SPERM AFTER H P r O P H P S E C T O M Y
231)
in a great stimulation of accessory organs. One completely
hypophysectomizetl aclnlt liad suiall, abdominal and aspermatic testes, one of which was removed at laparotomy; treatment f o r 30 days caused the remaining testis t o enlarge,
descend into the scrotum, and produce spermatozoa ; accessory
organs attained a size equal to that for nornial breeding
adults. In the other adult, 16 days of injection caused a
stimulation of all reproductive organs but no sperm production.
Five immature males had anterior lobe fragments. Treatmeiit for 30 to 37 days caused a stimulation of the entire reproductive tract in each case. At autopsy, testes were large,
scrotal and spermatic, all accessory organs mere large, and
the epithelium of the prostate gland was higher than that of
control animals (table 2).
A9efuitrii2-S as given subcutaneously to eight immature
and ti%-oadult liypophysectomized males twice daily i n doses
of 23 to 100 rat units for periods of 6 to 42 days.
S e w n immature animals aiid oiie adult were found to be
completely liypophysectomiz~d. Threc immature males injected for 31 to 42 days had large, scrotal testes with spermatozoa (table 3, fig. 18). Average weiglit of their accessory
organs was approximately 8 (bulbar gland) t o 80 (seminal
Twsicles) times greater than that for untreated controls. The
arerage heiqht of the epitlicliurn of their prostate glands was
about two times greater than that for controls (fig. 19). F o u r
otlier immature males ~ w i - etreated f o r only 10 to 16 d a p .
Their testes x7ere abdominal in position arid aspermatic, and
their accessory organs n-cighecl about 6 to 10 times more than
those of untreated males. Tlie adult male receil-ing only 13
days of treatment liad reproductive organs that mere definitely larger than those of control adults. No spermatozoa
x e r e fourid in this animal.
One immature and oiie adult male had anterior lobe fragments. All rcprocluctive organs of the immature male were
greatly stimulated after 37 days of treatment. The testes had
descended into the scrotum and produced spermatozoa. The
240
L. J. W;ET,LS A N D AX. D. QVNRHOLSER
adult was injected f o r only 6 clays. I t s rcprocluctire organs
were larger than those of uninjected adult controls. This
adult clid not have spermatozoa.
DISCUSSION
Formation of spermatozoa in response to these substances
in the absence of the hypophysis is sigriificarit mainly i n the
fact that sperm production W R Y induccd in immature as well
a s adult males and at a period of the year whcn spermatozoa
are normally absent in this species. Ailatomically mature
spermatozoa with o r without motility wei*c found to be abandant i n the epididpmis in most cases. Also, it is a n interesting fact that Prospermin, an extract of hnman male urine,
was found t o have gonadotropic and no androgenic properties.
Immature ground squirrels h a w been previously reported
to undergo precocious sperm production, and adults showing
seasonal aspeimia have been stimulated to renewed sperm
formation in respotisc to Antuitrin-S and mare serum (TVells,
733
r a, '35 b ; Wells arid &Ioore, '36 ; Kalicr and Johnson, '36).
Tlie ferret, another seasonal breeder, whcn subjected to an
increase in the length of day and injections of pregnancv
urine, produced sperm earlier in thc year than untreated coiltrols (Allanson, Romlaacls and Parkes, '34). Illorre11 ( '33)
reported that precocious spermatogenesis was induced in immature rats by injections of Prospermin combincd with either
a p o w t h preparation of beef pituitaries or pituitarF liebin.
Rloore ( 3 6 ) has reviewed the worlir of investigators ~.7'ho
haw
reported upon the effects of these sulsstances on spermatogcncsis in immature mice and ruts ; Moore found that these
materials (rnare serum, ilntnitrin-S arid Pmspermin) did not
hasten spermatogenesis i n immature r a t s to the 'sperm-head
stage,, this stage proving to be a dependable and effective end
point. Cole ('36) reported no sperm production in rats following pregnant mare serum injections. In adult liuman
liypogonaclisni, Antnitrill-S has been found to cause the
appearance of spermatozoa i n condom samples of semen
(Rrosias and Schaffer, ' 3 3 ) .
INDOCING SPEllM AF'TElt 1IYPOPHYSECTObTY
241
Smith and 1,eonard ( ' 3 3 ) have shown that pregnancy urine
extracts will maiiitairi o r repair spermatogenesis in liypophysectoniizetl rats. Other urinary and pituitary substances have
a1so been found capable of regulating spermatogenesis in
hypoplq-seetomized r a t s (Smith, Engle and Tyndale, '31 ;
E r a n s , Pencharz and Simpson, '31 ; Greep aiicl Fevold, '37),
Altbough it seems clear from the above reports that sperm
formation can he readily inclaced in adult rats, spermatogcnesis in rats immature at the time of hyp'ophysectomy has resulted rarely, if ever, in the formation of fully mature spermatozoa with motile mpacity such i l s we are able to report
for immature li~popliysectomizeclground squirrels.
It has been our experience that the length of tlie injection
period is prohahly more important tliaii the size of the close,
since 19 days was the shortest period of treatment resulting
in spermatozoa formation a i d llie substances were injected
in large overdoses in most cases. W e feel that the optimum
lcngth of treatment should he about 30 days.
SUMMARY
1. Immature ground squirrels normally do not produce
spermatozoa before late January, at \vhich time they a r e
about i.5 months old. Normal adults have small, abdominal
and aspermatic testes f o r R period of about 6 months hetween
Jnlj- a i d December.
2. IIypophysectomy was performed during months of
aspcbrmia, arid surgical success was chcckecl hy a study of
serial sections of the seila tnrcica. Volame of anterior hypophysis fragments in cases of incomplete hypoplq-sectomy
was mcasured by means of an Edinger projection apparatus
and a planimeter.
3. Adequate treatment of lij-popliysectomized males witli
extracts of iiniriari male urine, pregnancy urine and pregnant
mare serum caused the testis to enlarge, clescend into tlie
scrotum and produce spermatozoa. ,Accessory reproductiw
organs of some aiiimalq reaclied a size and histological state
characteristic of noymai breeding adults.
242
L. J. WELLS AND
Sf. D. OVEBHOLSEB
4. Both immature hypophpsectomized males a i d hypophyscctomizcd adults were stimulated to produce sperm, which i n
most cases T1-el.e present in the epididyrrlis arid in some cases
were motile.
5. Sperm production occurred equally as well in the absence of the antcrior hypophysis as i n animals with aiitcrior
lobe fragments.
6. The substances were deliberately administered in overdoses ; therefore, our data have more qualitative than quantitative value.
LITERATURE CITED
ALLAXSON,>I., I. TIT. ROT7T,A4NDS AND A. 8. P ~ R K 1931
E ~ ~Induction of fertilitS
and pregnancy in the anoestrns ferret. Proc. Roy. Soc., B,vol. 115,
pp. 410431.
BAKFR,
R. L., BND G. E. J o m r s o N
1936 The effect of injections of Antuitrin-8
oil the sexually inartioe male ground squirrel. Endoerinoloffy, vol. 20,
pp. 21 9-223.
BROSIUS,
TT. L., ~ N D12. L. SCHAFFER1933 Spermatogenesis following therapy
with the gonad stimulating extiact from the urine of pregnancy.
J. A41U.3Iea. ~ ~ s ( ] 1c70. 1,.101,p. 1227.
CQLE, H. H. 1936 On the biological properties of mare gonadotropic hormone.
Am. J. Anat., vo!. 59, pp. 299-331.
EVAKS,11. IN.,X. I. YFNCHARZAND 3.1. E. SIMPSON1934 Maintenanec and
repair of the reproductive system of hypopb~sectoniizedmale rats by
11) pophyeal s p c r g i s t , prcgriancv prolan and combinations thereof.
Endoerinologp, vol. 18, pp. 607-618.
GXEEP, R. O., ~ K H.
D L. FEVOLD
1937 The spermatogenir a n d secretory function
of the gonads of hFpophysectomized arhilt rats treated with pituitary
FSH and LII. Endocrinology, rol. 21. pp. 611-618.
JOIIVSOK,
G. E., $1. E. F o s ~ r wA K D R. &COCO
I.
1933 The sexunl cycle of the
thirtecii lined ground squirrel i n t h e laboratory. Trans. Kansas Acad.
Sei.. vol. 36, p. 250.
A~OORE, C. R. 1936 Responses of immature rat testcs to gonadotropir agents.
Am. .T. Anwt., vol. 59, pp. 63-88.
RIOORE, C. R., G. F. SInmoNs, I,. J. WELLS,11. ZALESKYAND W.0. NELSON 1934
On the control of reproductive activity in an anniial breeding mammal
(Citellus tridecenilincatus). l i n a t . Rep., vol. 60, pp. 279-289.
&~OREFTL, J. A.
1933 The effert of a new snhstance 'prospermin,' occurring in
noriiial male nrine, on the gonads of the innnatnre male rat. Anat.
Rcc., 7701. 5.5 (Ruppl.), pp. 29-30.
SMITII,P. E., E. 'r. EKGLE
-4ND H. €1. TPm.4LE
1934 Gametokinetic action of
extracts of folliclc-stirnulating urine. Proc. Roc. Exp. Biol. and Med.,
vol. 31, pp. 745-746.
1NL)LClK-C: SPERM AFTER HYFOPHYSECTOAIY
2.1::
1!133 Rcsponses o f t h e rt,pro(luctivc systenl
SMITH,P. E., APID W. L. I,FONARD
of 1I;cI’opliSsectonnzed and normal rats to injections of pregiianry urine
extracts. I. Tlic male. Anat. Xrc , vol. 5$. pp. 145-174.
WELLS, I,. J. 1935 a Stnsonal scxual rh) tlini and its expernnental modification
in the n ~ l eof tlrc thirteen-lined ground squirrel (Citellus tndeeemlineatns). Anat. Rcc., 101. 6 2 , pp. 409-147.
____
19:+,5b Stimulation of the reproduetlve system of male glound
squirrels hp injections of blood serum f r o m pregnant inzres. Anat.
Rec., vol. 61 (Siippl.), p. 49.
1937 ltc~~rodiictire
organs of t n o manimxliaii herniaphrodi
their response t o injections of pregnant marc’ serum. Anat. Rec.,
101. 67, ~ 1 > 233-251.
.
WELLS, L. J., A K D F. T. (:UUIW 1937 Hypopliysectoniy and its effects on male
reproductiTe organs iii a m l r l mammal with ~ n n u a lrut (Citclluu).
Anat. Rec., rol. 69, pp. 213-227.
WELLS, L. J., ~ K I IC. R. M O O K ~ 1936 llorinonal stimnlstion of spcrmatogenrsis
111 the testis of the ground squirrel.
Anat. Rec., vol. 66, pp. 181-2110.
WELLS, L. J.. AXD M. D. OVERIIOLSER 1938 Production of sperm in hypophpsectomizcd ground syuiircls (Citcllus) injected with substances from
urine a n d 1)lood. Anat. Rrc., vol. 70 (Huppl.), p. 81.
PLATE 1
EXPLANATTON OF F I G U R E S
Sectiun tlirougli solla tureica of riorrnal ground squirrel showing liypophysis.
N, pars nervusa. (posterior lobe) surrounded by liars intermcdirt. I), pars distalis
(anterior lob-). Pituitary &ilk and pars tuberalis lie t o tlic right of the anterior
lobe. X 15.
3 Section t,hrough s d : t turcica of hppoI'hyscctornizcd ground squirrel no. 1405.
Volume of t,be anterior lnhe fragment, l), as found to be 4.6%. X IS.
4 Section t h r n n g h the sella, tiircica of hypopli~acc.torrii~~d
ground squirrel no.
1406. Volume of anterinr lohe fragment, D, mas 5.4%. N, pars IiervosiL 8111'rounded by pars intermedia. X 15.
5 Section through tlic sella turcica of hq-poPhysectomi~edground squirrel no.
1403, which was ' completely hypopliysectorriized ' so f a r as anterior lobe is c o w
cerned. N, pars nervosa, surrounded by pars iritcrmcdia. X 15.
6 Seetion o f testis f r o m normal adult killed I)wc:ml)er 3rd. Notc :thence of
spermatids and spermatozoa. X 200.
7 Section of prostate gland from ahovc animal. X 200.
8 Section of testis f r o m adult liy~io~~hyscctortiizcd
malc no. 1407 which received Prospwniiii end was killcd Kovember 28th, 181 days after hypopliysectomy.
This :mima1 received 5.0 nig. Prospermin daily f o r 26 days. S o aritcrior lobe
tissue remained. Many mot,ile spermatozoa foiind in rpididymis. x 200.
9 Section of prostate gland of ahove animal. X 200.
2
244
INDTJCINB SPXRM AFTER HYPOPHYSECTOIY
1.J. W R U S A N D Y. D. UVEWNOJ~S&IC
PLATE 1
PLATE 2
EXPLAVATION OF P I O U & E S
1 0 Section o f testis from normal immature male killed Novembcr 19th. Note
ah-ence of spermatids niid spermatozoa. X 2000.
11 Section of prostate g h n d from :ik)ovc~aniirial. X 200.
1 2 Section of tcstis from uninjected hyp(ipIi\ sectvrnizcd iinmatnre m a l e nu.
1424 which was killed on Xnveinhcr 28th, 46 d:i~s after liypophywctoin;.
No
X 20G.
sprririatida or spermatozoa. Tolunie o f anterior lolir fragrrieiit \\as tiS7..
19 Srctioii of y i o s t n t e glaiid froin abme .rnininl. x 200.
14 Section of testis frorn irrirnaturc lippopli? ~eetoini7edmale no. 1412 \vlirch
rereired 5.0 nig. Prosycrrriiii dailb for 26 daps and as killed So\crub;r 28th.
40 daps after bypophysectomy. No anterior lob:) tissue. Many sperin:~to/oa ;ri
tlic tcstis arid in the epididymis. X 200.
15 Secation of prostate glxiid nf nhuvr animal. X 200.
fioin iriirii:Lture li4popliyscctoiiiized inalc no. 1279 nhic.11
received 40 rat units of inare wriini daily for 33 d n k y arid nas killed on Octohci
27th, 5.5 days aftcr hypophysectomp. No anterior lobe tissue. Xany spermato7oa
in testis. Many motile sperm f r o m epididymis. x 5 0 0 .
1 7 Scetiori of prostate gland from ahove animal. X 200.
I S Section of testis f r o m i m m a t u r e h y ~ n ~ ~ h ~ s e c t o m imale
7 c ~ d no. 15.58 which
rewived 75 rat units of Antuitriii S daily f o r 42 d a y and mas killed on Septcmbcr
24th, 70 days after EipEioIiiivsrh.tonip. No anterior lobe tissue. Many sy~rmato7oa
in testis and epididjniis. F e w rpidictyniiq speirri showed motility. x 200.
19 Section of prostate gland from :tbovr xiiinial. x 200.
237
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grounds, growth, formation, urine, sperm, human, organy, blood, accessory, response, substances, squirrel, hypophysectomized, reproduction
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