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Testicular development in the rhesus monkey.

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TESTICULAR DEVELOPMENT I N T H E
RHESUS MONKEY
GERTRUDE VAN W A G E N E N AND MIRIAM E. SIMPSON
Department of Obstetrics and Gynecology, Y a l e University School of Medicine,
N e w Haven, Connecticut;
Institute of Experimental Biology and Department of Anatomy,
University o f California, Berkeley
TWENTY-EIGHT FIGURES
An analysis of normal testicuIar development in the rhesus
monkey, Macaca muZatta, was undertaken in order to provide
basic data f o r the interpretation of changes brought about
experimentally by the action of gonadotrophic hormones in
the immature animal. An obstacle to the effective use of the
monkey for this purpose has been the lack of exact knowledge
of the developmental status of the testes of animals of known
ages, so necessary as a foundation for any experimental work.
Even in isolated experiments, normal control animals of the
same age raised under the same conditions at the same time
are rarely available for the monkey. No colony is large enough
to supply such paired animals in sufficient numbers. Rarely
is it possible even to obtain animals of known age, while littermates are out of the question since no male twins have thus
far been reported. Purchased animals, which are usually of
unknown history, although of the same body weight and approximate appearance, may be very different in age and
sexual development. It is true that certain observations help
This investigation was supported in p a r t by research grants from the National
Institutes of Health, Public Health Service, t o Yale University School of Medicine
and to the Institute of Experimental Biology, University of California; by grants
from the Research Board of the University of California, and by grants from the
Committee on Research, Council on Pharmacy a n d Chemistry, American Medical
Association.
231
cr,sLE I
Sertoli nuclei move basally ; occasional spermatocytes
appear.
Sliermatids appear and a few differentiate to sperm;
Leydig cells mature.
Sperm ; tubules filled with debris.
S~i~rmatogenc~sis
conil)lete. Ta1)oles orderly and free of
de1)ris.
E’ully acti\e.
70-1 (I0
100-150
1x-200
200-2.70
2,j~J-:$IlO
2.0
2.2
3.00
3.25
4.00
ca. 2 yr. 9 mo.
2 yr. 10 mo.-3 yr. 2 mo.
3 yr.-3 yr. 5 mo.
3 yr. 5 mo.-4 yr.
from 4 yr.
-~
Leydig cclls epitlielioid, not mature ; Sertoli cells increaqe in numbcr.
30- 60
2.0
Relatrcely lattle advance,
Inciease in spermatogonia.
50- 60
50- GO
Regression of Leydig and Sertoli cell differentiation ;
Scrtoli nuclei no longer basal.
50- 60
Few spermatogonia p..rsent; Sertoli nuclei basal; Leydig rells differentiated.
TESTICULAR D I F F E R E N T I A T I O N
ca. 2 yr. 7 mo.
1.10-1.60
1.10
8 1110. to 1yr. 3 mo.
1 yr. 3 m0.-2 yr. 7 mo.
0.75
Birth to 3-4 mu.
micru
70- 80
cni
0.04
0.06
SEMINIFEROUS
TUBULE
DIAMETER
TESTIS
LENGTH
Fetal life : 7 2 d.
154 d.
AGE OF MONXEY
1)ctxlopnicnt of t c o t i s o f the monkey, Macacu aiiilutta. Sequence of clrunges correlated w i t h age
TESTICULBR DEVELOPMENT I N MONKEY
233
in establishing the probable age. Body weight and body
length (sitting height) may be compared with known growth
curves, and roentgenograms with the normal standards of
skeletal differentiation. Dental age is somewhat more easily
determined and applied (Hurme and van Wagenen, '53).
However, both dental and osseous development are of value
only when significant steps in differentiation coincide with
those parts of the life span which are of interest in a particular experiment.
It remained essential for the work with gonadotrophins to
establish the age at which critical changes occur in the testis,
and the sequence and duration of each developmental phase.
3Iaterial suited for this study has accumulated since 1935
from the monkey colony established in the Department of
Obstetrics at Yale University. The colony is an inbred group
of rhesus monkeys (Macncn w t k i t t a ) now in its sixth generation. Histological preparations from biopsies and autopsies
were studied from 40 male monkeys of known age, from fetal
life to eleven years. Table 1 gives in outline form the critical
periods in growth and differentiation of the testes.
Figure 1 (described below) is based on serial observations
of body weight and testis length of animals born and raised
in the colony. Of a total of 24 normal untreated animals
studied during the first year, 15 were followed through the
second year, 14 through the third year and 12 to the end of
the fourth year. The smaller numbers in succeeding yeass
means only that these normal monkeys were allocated to
experiments.
OBSERVATIONS
Fetal development. The testicular tubules before birth are
still short, are only slightly coiled and are separated by much
interstitial tissue. However, both tubular and intertubular
tissue show evidence of early differentiation. The degree of
development exceeds that attained for months or even years
after birth, being almost comparable to that of the preadolescent testis. At 90 to 110 days of fetal life the tubules
are widely separated by abundant interstitial tissue (figs. 2,
234
G. VAN WAGENEN AND M. E. SIMPSON
19 and 20). This tissue consists for the most part of small
epithelioid cells, which, though not so large as in the adult,
are recognizable as Leydig cells. The tubules of the testes
a t this time are short and only slightly coiled but are larger
in diameter (70-80 p ) than after birth. The Sertoli cells have
acquired morphological features which are here interpreted
as differentiation. The cytoplasm is abundant and filamentous
and extend across the lumen; the nuclei occupy a peripheral
or basal position in the cytoplasm.
The presence of abundant interstitial tissue (identified as
Leydig cells) has been described in fetal testes of both the
human and the horse (Bouin and Ancel, '03 ; Cole et al., '33 ;
Gillman, '48) and has been correlated with the increased
amounts of gonadotrophin present in the maternal blood of
these species. Since chorionic gonadotrophin has been reported in the urine of the pregnant monkey (Hamlett, '37),
it is reasonable to attribute to this agent the Leydig cell growth
and differentiation in the fetal monkey testis.2 The descended
testis characteristic at birth of both man and monkey has also
been correlated with intrauterine action of gonadotrophin
(Engle, '32b ; Wislocki, '33).
Post-natal regressiow. Actual regression in the development of the testes occurs after birth (figs. 3 and 21). At birth
the Sertoli cell differentiation is still present and nuclei are
basal. The interstitial cells have decreased in size and probably in number. Figure 21 shows, especially by comparison
with the fetal picture in figure 20, the more closely packed
tubules at birth. This regression is most pronounced within
the first 3 months and resumption of development may not
become evident before 9 months to 1 year 3 months (figs. 4,
5, 6). The diameter of the seminiferous tubules decreases to
50 p and does not increase appreciably during the first year,
still measuring between 50 t o 60 p at 1year 3 months (fig. 7).
'Evidence will be given in a paper devoted to the effects of interstitial cell
stimulating gonadotrophins in the prepuberal male monkey t h a t differentiation in
the Sertoli cells is also due t o the chorionic gonadotrophin. Engle ( '32) shows
this change in his illustrations.
TESTICULlR DEVELOPMEST IX MOSICET
235
The tubules grow in length, h o m r e r , and become more convoluted, accounting f o r the increasing size of the testis. The
Sertoli cells lose the cytoplasmic differentiation attained
during intrauterine lifc, and the nuclei of these cells again
come to fill the lumen of the tubules. These and a relatively
fern basally situated spermatogonia are the only cells present
in the tubule. Differentiated Leydig cells disappear. Within
3 or 4 months (figs. 3, 4 and 5) after birth, the intertubular
spaces have become nanotv and the tubules are closely packed
together. The only intertubular cells which remain, except
for a few flattened cndothelial cells, are inactive Leydig or
capsule cells with small dark staining nuclei, which lie very
close together and sunound the tubules in 1 or 2 concentric
layers.
B e g i m i w g d i f e r e 12 f Lotion. The regression in differentiation of tubular and intertubular tissue may be attributed to
the withdrawal at birth of maternal hormone influences. There
supervenes a period which lasts through most of the first
year, during which tubules are not elaborated cytologically,
though they lengthen and beconie more convoluted. The tubules at this time characteristically contain only Sertoli cells
and scattered sperniatogonia. Intertubular tissue is scanty
and Leydig cells are not distinguishable. Early in the second
year spermatogonia become more numerous, forming a continuous basal layer in some tubules. The multiplication of
spermatogonia sometimes becomes obvious at 11months, and
is more evident still between 1 year 3 months, and 1 year 9
months (figs. 7 and 8 ) . These cells are larger, more rounded
and possess more abundant cytoplasm than the previously
seen primitive sex cells.
Although the tubules had begun to increase slightly in size
by 2 years 2 months to 2 years 4 months, they still measured
between 50 and 6 0 p , and even by 2 years 7 months they
reached only 70 to SO p, corresponding to the diameter in late
fetal life (compare fig. 3 with figs. 9, 15 and 16). It is more
difficult to assess the increase in tubule length which un-
236
0. V A N WAGENEX AND M. E. SIMPSON
doubtedly occurs. Some indication of this is the gradual
overall increase in testis size (table 1 and figure 1).
F o r a n entire year, extending through most of the third
year of the individual’s life, no appreciable further advancement in diff erentiatiori of the germinal elements of the tubules
takes place (compare figs. 8, 9 and 11, 12). The next phases
in development concern the Sertoli and the Leydig cells.
Grams
7000
I
I
I
I
2
3
4
Age in Years
Fig. 1 Testis length correlated with body weight and age of norm:il rontrol
Ihesus iiionkeys from the colony a t Yale University School of MedicGne. Thc
serial observations were based on twenty-four niiinials through the first year.
Alloeatioii t o experiments reduced this number t.0 twelve at end of the fourth Tear.
The histological observations were based on serial biopsy and autopsy of 40 male
n i o n k e p of knoxm age.
Diffwentiation of Sertoli cells. At 2 years 7 months, when
the tuhules have reached 70 to 80 I.I in diameter, multiplication
with differentiation, of Sertoli cells occurs. The increase of
Sertoli nuclei may be very conspicuous. The enlarging tubules
a r c filled with these nuclei before the differentiation becomes
pronounced (fig. 9). The nuclei enlarge and become pear&aped with indentations, the chromatin is scattered, and
there a r e one or more prominent nucleoli. The cytoplasm
increases in amount, becomes filamentous and stretches across
TESTICI'LSB DEVELOPMENT I N MOXKEY
237
to fill the lumen of the tubule. The nuclei come to lie more
and more basally (figs. 10, 13, 15 and 16).
Differentiation of L e y d i g cells. Accompanying the differentiation of Sertoli cells, between 2 years 7 months and 2
years 9 months, the tubules become less crowded within the
testis. The vascularity of the intertubular tissue is increased
and the tissue fluid increases. At this time, differentiation of
interstitial cells occurs. They enlarge and become rounded
(epithelioid). Although not fully mature, at 2 years 9 months,
they a r e now definitely i*ecognizable as Leydig cells (fig. lo),
apparently differentiated from cells hitherto arranged in
layers around the tubules. Proliferation from encapsulating
cells can be seen in illustrations from the fetal testis (fig. 20).
This subject is further developed and illustrated by Simpson
and van Wagenen ( ' 5 3 ) . The small, elongated, dark-staining
nuclei of capsule cells enlarge and become rounded; the
chromatin scatters and the nucleus therefore heconies lightstaining. One or sometinies two prominent nucleoli appear.
Abundant vacuolatioii in the peripheral part of the cytoplasm
is characteristic of the 1,eydig cells of the mature testis but
is not comnionly seen iii animals that have recently matured.
The number of Leydig cells recognizable hy these criteria in
normal mature animals is surprisingly small ; they form
clumps usually coiitaining 110 more than 2 to 5 cells and rarely
10 to 20.
D i f f e r e i L t i a t i o n of p
~
~ eleincizts
k
l of t h e tubules. The
appearance of priinurjj s p w m c i t o c y t e s marks the onset of
activity in the germinal elements of the tubule which will
lead to sperm formation. Between 2 years 7 months and 2
years 9 months (figs. 9 and 10) the tubule diameter has increased to about 1 0 0 ~ .A few primary spermatocytes, distinguishable by their size, position and distribution of chroinatin in the spirenie, appear in the meshes of the Sertoli
cell cytoplasm. No open lumen is present in the tubule at
this time. The first formed spermatocytes undoubtedly desquamate, or degenerate, and their number does not increase
immcdiately (figs. 13, 15, 16). Only a few scattered primary
238
G. \'AS
W A G E S E N A N D 31. E. SIMPSOX
spermatocptes were found in tubules of four animals between
2 years 9 months and 2 years 10 months of age. After 2
years 10 months, spermatocytes begin to accumulate in great
numbers in the tubules and succeeding stages of development
then follow rapidly (figs. 14, 17).
A series of four biopsies from the testes of ZIIm 642 illustrates the intratubular development from the quiescent prepuberal state to the formation of spermatocytes (figs. 11 t o
14). Adequate material is not available to support general
statements. However, it is seen in this animal that 4 months
elapsed between the assumption of a basal position lip the
Sertoli nuclei and the formation of the first spermatocptes.
Three weeks later the number of spermatocytes had increased
hut no further differentiation of cell types had occurred. I n
a second series of biopsies (figs. 15 to 18) from another maturing animal, hlni 252, the first testis sample was taken after
migration of the Sertoli nuclei and thc accompanying tubule
enlargement; the last sample was taken when fully formed
sperm were present. The entire period covered 10 months.
At the end of the first 4; months the only advance t o be seen
was an occasional germinal cell, lying central to the Sertoli
cell layer, in which a spireme was forming. The final organization of the seminiferous tubule with clearing of the lumen
is shown in 3 stages f o r the monkey Mm 686 in figures 23 to
25, and the age span covered was from 2 years 10 months to
3 pears 5 months.
S p e r m a t i d s and s p w n were first seen in the testicular tubules at 2 years 10 months, but in other animals this degree
of development was not noted until 3 years or 3 years 6 months
of age. Spermatids appeared when the tubules had attained
a diameter of 100 to 1 5 0 ~ .A considerable time appears t o
be consumed in the final ripening to form the sperm. The
slender sperm tails, even after they have become elongated,
remain for a time difficult to stain. During this terminal
ripening phase, from the end of the third year to the middle
of the fourth, desquamation of cells in all the later stages of
derelopment is occurring and the tubule lumen is packed
TESTICULAR DEVELOPMENT I N MONKEY
239
with debris (figs. 23 t o 25). By the time the tubules are opened
and cleared of all debris the diameter of the tubules has increased to 200 p and the cells of the stratified epithelium are
arranged in more orderly fashion (figs. 26 to 28). Fully
mature tubules a t the height of activity measure 250 to
300 p.
Lacking monkeys of known ages, experimentalists heretofore have classified males simply as immature or mature.
Body weight has been the useful index in such classifications,
and where the reports have been accompanied by illustrations
of the testes, the histology and body weights have agreed
well with the data here presented (Engle, '32a; Smith, '38
and '44). Hartman ('32)' who maintained males of known
age, reported sesual maturity as occurring within the upper
limits here assigned. Figure 1 shows the correlation between
body weight, testis size and age of the monkeys used in this
study. I n all instances the measurements of the testes were
found to give a reliable index of the tubular diameter and
differentiation, and such measurements, correlated with the
finding summarized in table 1 and figure 1, are useful in
selecting male nionkeys f o r experimental work with gonadotrophins.
DlSCl~SSIOS
Testicular development during childhood is described to
be as slow as that observed here f o r comparable periods in
the monkey, an interval of 3 years in the monkey being considered roughly equivalent to 15 years in the human. The
diameter of the testicular tubules is surprisingly similar.
Clharny et al. ( ' 5 2 ) give the following measurements from a
developmental series (mostly from biopsies) in the human :
tubule diameter 6611 from 3$ weeks to 4 years; an increase
only to 72 p at 10 years and 85 p at 12 years; an increase to
100-150 p a t puberty, and an average adult measurement of
150-180 p. (Compare measurements for the monkey as given
in table 1.)
The time when dedifferentiation or regression of Leydig
tissue occurs in the human has not been ascertained accu-
240
G. VAN WAGENEN A X D M. E. SIMPSON
rately but it is generally assumed that the hypertrophy of
the fetal period, being dependent on maternal hormones,
probably disappears soon after birth. Albert ( ’ 5 2 ) , though
finding it somewhat variable and uncertain, assumes that
regression may occur within a ~ e c l after
i
birth. C‘harny et
al. (’52) find no Lcydig cells distinguishable 3; weeks after
birth.
Theoretically one might expect that a t puberty differentiation, or even hypertrophy of Leydig cells would precede
differentiation of the tubules, judged from the ability of
testosterone to stimulate spermatogenesis and the supposed
production of male hormone by Leydig cells. I n this study,
Leydig cells were found to be distinguishable from indifferent
intertubular cells at about the same period a s the carliest
differentiation occurred in the tubules. The appearance of
the fully functional cells (on basis of size, rounded cell body
arid granular or vacuolated cytoplasm) was not seen, however, until spermatogenesis was f a r advanced. Albert et al.
(’52) in ii recent revie\\. of the developmental stages in the
human testis havc stressed the fact that the changes leading
to maturity at puberty definitely occur in the tubules before
they do in the Leydig cells. Sniffen ( ’ j 2 ) also describes the
appearance of tlie first spermatocytes in human testes, e.g.
a t 13 years of age, before differentiation of the Leydig cell
has occurred. T o quote: “ I n early puberty, when the secondary sex characteristics a r e beginning to develop and tubular activity is well established, the interstitial cells seem to
lag lnehiiid and remain uiidiff erentiated. ” When they appeared during the next few years, however, say by 17 years,
by differentiation f roni mesenchymal cells, more Leydig cells
were observed thaii were characteristic in the adult. Hooker
(’44) also reports that in the bull the differentiation of the
tubule occurs earlier than that of the Leydig cells. He states,
“ S o striking change in either the Leydig cells or the androgen content of the testes was evident at puberty which apparently occurred duriiig the second 6 months of life. The great
changes in androgen content after 8 years of age were ac-
TESTICULAR DEVELOPMEKT IR- N O N K E Y
241
companied by changes of comparable degree in vacuolation
and numbers of Leydig cells.”
I t may be noted that in descriptions of the development
of the human testis by Charny et al. ( ’ X ) ,Albert et al. (’52)
and Sniffen ( ’ 5 3 ) , these workers are conservative about the
differentiation of the spermatogonia from Sertoli cells in the
testis early in childhood, referring to the presence of a few sex
cells and a synctium. I n the description given here of the development of the testes of the monkey the enlarged, rounded,
basally located cells seen from birth onward have been designated spermatogonia. Sertoli cells have been described, rather
than designating a syncytium, though it is realized that the
boundary of individual Sertoli cells is difficult to distinguish
either in the immature or the mature testis. The opening
of a lumen in the seminiferous tubule also is described here
as occurring relatively later than it is given in the accounts
of human development; the potential lumen is considered to
he filled successively by the nuclei of the Sertoli cells, then
by their filamentous cytoplasm, and later by successive generations of desquamating unripe and finally ripe germinal
elements, before a truly patent lumen is established in the
adult.
SUMMARY
Fetal life. During late fetal life the testicular tubules of
the rhesus monkey attain a diameter of 70 to 80 p and reach
a stage of development not seen again until prepuberal changes
appear. Although only Sertoli cells and a few spermatogonia
are present, the nuclei and cytoplasm of the former show
differentiation. The nuclei come t o lie near the basement
membrane and the cytoplasm fills the lumen. Interstitial cells
are abundant, filling the wide intertubular spaces and many
are differentiated and identifiable as Leydig cells.
Post-na,tal regression and subsrqzcent slow growth. Definite
regression of the testes occurs after birth. The tubuIes decrease to a diameter of 50 to GO p and remain so for the first
year while increasing slowly in length. The interstitial cells
decrease in number, dedifferentiate, and are not again clearly
242
G. V 4 S W A G H S E S A S D 31. E. SIMPSON
distinguishable as Leydig cells until late in the third year.
Only a few scattered spermatogonia are seen in the post-natal
period, but they become more numerous, appearing enlarged
and rounded at the end of the first year.
Adolescence a w l m at urit y. After this prolonged period of
slow development, lasting until the end of the third year,
changes which lead to maturity follow in rapid succession.
Leydig cells differentiate. Sertoli cells increase in number
and become differentiated, with filamentous cytoplasm filling
the lunieii of the enlarging tubules and with nuclei lying
basally. Primary spermatocytes appear and the changes
leading to formation of spermatozoa take place rapidly. The
earliest appearance of spermatozoa was observed at 2 years
11 months, the latest at 3 years 5 months.
LITERATURE CITED
ALRERT,A., L. 0.UXDERDAHI.,
L. F. GREENE A X ~ IN. LORENZ1953 Male hypogonadism: I. The normal testis. Pioc. Staff Meetings Mayo Clinic
$8: 409-40'7.
EOVIIS,
I'., A N D P. A S ~ L 1903 Reclierclles stir lcs cellules interstitielles du
tcsticule des nianimif8res. Arch. de zool. esp6r. et g h . , I : 437-523.
CI-ISKNY, C. W.,A. S. CONSTON .WJI D. R. MERANZF. 1952 Testicular developmental liistolog~. Annals X. Y. Acad. Sci., 5 5 : 597-608.
COLE,H. H., G. I*. H A R T , W. R. LYONSAXD H. R. CATCIIPOLE 1933 The development and hormonal content of fctal horse goiiads. Anat. Rec., 56:
'775-293.
ESGLE,E. T . 19321 The action of extracts of anterior pituitary and of pregnancy urine on the testis of immature rats and monkeys. Endocrinology,
1 6 : BOG-512.
183'7b Expcrin~entallyinduced descent of the testis in the macacus
monkex I)? hornioues from the anterior pituitary and pregnancy urine.
Endorrinology, 1 G : 513-520.
GIzLaras, J. 1048 The derelopment of tlie gonads in man, with a consideration
of the rolr of fetal endocrines and the histogenesis of ovarian tumors.
Carnvgie Inst. Wnsli., Contvib. to Enibryol., 3 2 : 81-131.
HAIILETT,0. W. D. 193i Positive Fricdinan tests in the pregnant rhesus monkey, .Wacncn nmlattn. Am. J. Pliysiol., 118: 664-666.
HARTMAS,
C. G. 1933 Cited by Scliultz in The Anatomy of the Rhesus Monkey
Ch. I T , Grolvtli and Development. Edited by C. G. Hartman and W. L.
Strnus, J r . The 1Villi:ims and Wilkins Co., Baltimore.
HOOKER,
C. Tit'. 1944 The 1)ostnat:tl history and function of the interstitial cells
of tlie testis of the buli. Ani. J. Anat., 7 4 : 1-37.
TESTICULSR DEVELOPMEXT I N MONKEY
v. o.,
243
A X D G. E AX WAGEXEN 1953 Basie d a t a on the emergence of
deciduous teeth in the monkey (,Macaca midatta). Proc. Am. Phil Soc.,
9 7 : 291-315.
S r ~ r s o x?if.
, E., A'KD G. V A X X ~ G E X
E NPersistent nodules in testis of the
1963
monkey associated with Leydig cell hyperplasia induced by gonadotrophins. Canecr Research (in press).
SsIITII, Y. E. 1938 Comparative rffcets of hypophgsectomy and therapy on the
testes of monkeys and rats, in Les Hormones Sexuelles, Edit. L. Eroulia,
Hermann ct Cic., Paris.
-___
1914 Mainteiiance and restoration of spermatogenesis in hypoph)-sectoinizcd rhesns m o i i k e ~ sbv androgen administration. Yale J. Biol.
Med., 1 7 : 2 8 1 - 2 8 i .
S s r r ~ c R.
~ , C'. 195;' H i e t n l n p of tlir normal 2 n d a b n o r m a l trfitifi n t pnhcrty
Annals K. 1'.S c a d . Sci., 5 5 : 609-618.
Wr s r . o c~ r ,G. E. 1933 Observations on the descent of the testis i n the macaque
and in the eliimpanzce. A4nat. Rec., 5 7 : 133-148.
HUEME,
Testis of Mm 601 at birth (174 da. gestation). Testis biopsy. Tubule diameter GO-80 p . Nuclei of Sertoli cells arc basal
and cytoplasmic strands fill tubule lumen. Spermatogonia are sparse. Intertubular tissue is abundant but undifferentiated.
Testis of Mm 630 a t 3 mo. More coiling of tubules is present; diameter 50-60 p. Sertoli cytoplasm is developed and
still fills the lumen. A few spermatogonia are present. Intertubular spaces are narrow and interstitial cells have regressed.
3
4
Testis of Mm 570 :it 2 ~ r 7. mo. 6 (la. Note that durillg : i n c n t i i c year tlieie has been no appreciable advance in
development except in ninltipiicntion of Sertoli cell nuclei ; ~ n d:I slight inrreasc iu di;imetcr Of tubule (60-70 P). S n l l l C
of the interstitial cells are now lighter staining.
10 Testis of Mm 665 a t 2 yr. 9 mo. 21 da. Tubules are now definitely larger (90 p). Sertoli cells have moved t o the
periphery of the tubule. Spermatogonia are numerous and rounded. Rounded Leydig cells are now frequently seen.
9
lumen. Intertubular tissuc is undifferentiated.
~ the~
Testis of Mm 5.50 a t 1 yr. 3 mo. 24 da. Tubules are still sm:ilI, diaineter 40-50 p . Spermatogonia are no\v iiicreascd
in number and size. Nuclei of undifferentiated cells fill tlie lumen. 0111) dark stained nuclei in rows around tubules arc
seen in narrow peritubular spaces.
r
8 Testis of Mm 385 a t 1 yr. 8 nio. 7 da. Tubules are still small (diameter 30 p ) . Sertoli nuclei continue t o c
Testis of Mm 619 a t 4 inn. 24 da. Tubules are sinall and closely paelted. The size has not changed, 50 6 0 ~ . Sertoli
nuclei fill the lumen, only occasional spcrinatogonia being seen.
G
5 Testis of Mm 668 a t 3 ma. 25 da. Considerable growth in lciigtli n i t l i coiling of tubules has occurred. Tubules are siiiall,
50-60 1, compact, filled with Sertoli nuclei. There are oeensional slic,rmatogonia. The peritubular arrangement of dark
stained nuclei of intertubular tissue is clearly seen.
Testis of Mm 486, 96 day fetus. Tubules are short and straight, diameter 60-i0 p . Only Sertoli cells and a few spermatogonia are present in tubules. Sertoli cells are large and fill the lumen. Intertubular spaces are wide and contain mally
cells, some epithelioid.
2
Pigs. 2 t o 1 0 Developmental stages in Macaque testes, from biopsies and autopsies. H.E. X 237.
ESPLAN~\TION O F F I G U R E S
PLATE 1
d
C . V A N WAC4EXhS A S D .\I. h S I J I P b O X
TESTICULAR DEVETAIPNEYT IN >IONKEY
PLATE 3
PLATE 2
CSPLIXATIOK OF FIQL-EES
Fign. 11 to 14 Series of rlerrlo~inientalatages hi trntis of an indiridunl inonkcy
(Jfm 642). x 87.
11 Right teatir biopsy a t 2 yr. 4 1110. 7 1 dn. H and E:. Tubulcs long, coarolutcrl,
clowlg packed 50-70 p in dinmeter. Rcrtoli crlls poorly clifforentint&, but
cytoplasm is increasing niid nuclri hiire partirilly moved basnlly. Iiitrrstitia11
eells sintill and sparse.
12 Riglit testis biopsy rtt ! yr. 9 ino. I6 &I. 11 m d E. The changes nrc sliglit.
Tiibulcs are aomc\vliat lrtrgrr (70 p ) nod Rrtoli iiuclei are Nore bnlial. SO
rliffcwntinting of Leydig wlls accii.
13 Right trstis biopsy :tt 3 yr. 1 mo. 26 dsi. H rind 1:. Tubulcq hare incrcnwd
slightly in cliiimeter (scI-no p ) . 8rrtoli nuc*lc.i nrr now definitely basal. A few
crlln, recognimlile ns spcrnintocytcr I Iiy tlir spirriiic, nre Iircseut. Intrigtitiril
cells still not clearly rwugniuib!c.
14
I p f t trntin rt 3 pr. I! 1110. 14 cln. 1f:iIlnry stnin. Tubiilrn nirnsurc 100-150 p.
3Inny aprnintocytra nrc now Iirescnt. Cnnnlizntion is seen in n few tnliiilcw.
A few cpithclioid Lcyclig rclln, niiigly or in Iinim, :ire p r m n t .
Figs. 15 to 18 Scrim of dcre1opiuciit:tl stagrn in iiiacnque testcs n t higher
power, X 237.
16 Testis liiupsp (Jim 362) at 9 yr. 7 i i i c i . 14 (la. I€ tind $2. Tubules mrnniirr
70 p. Rertuli nuclei :irc btimil. Thrrc urr n few desqu:im:tting cells in the
nicslics uf Sertoli rytuplasiii. Occ~siunrrlLryclig crlls nre reeogniuiblc.
16 Testis biopsy (Mm ,“x;”)
tit
3 yr. 0
itin.
IT (la. TI nnd E. Tnbulcs inenailre
70-80 p. Rcrtoli nuclei arc Iiriwl. Ynsrukirity h:ta inrreawd, ttnd tlirrc itj
iiiore space betwen tnliulm. E:pitlieliuid 1,cylig cells itre present though iiot
of mature size. Sprmatoertcs arc appearing.
1i I d t teetin (MN 663) u t 3 yr. 2 mo. 14 chi. (Hec fig. 13.) Jhllory rtnin.
Tubules meamrc 100-120 p. Bprrmatayte I foriit:ttion is here nbnndnnt.
(Mm .Y’8,0) n t 3 yr. 5 inn. 28 rln. I1 nnrl E. Tubulrs nicnsiirc
130-15.0 p. Mniiy spcniinticls aiid some sperm are present. Some desqn:iiiiritiuii
of immature cells is d i l l prcsrnt in the tubular lumen.
18 Tentir biopsy
240
C . V A N WAC4EXhS A S D .\I. h S I J I P b O X
TESTICULAR DEVETAIPNEYT IN >IONKEY
Litc de~-elopn~cnt:ilstagcs i n testis of
Jfiii
686. Iron HIICIII.x 85.
Riglit twtii sniii~ilc after niiother oiuntli, tit 3 yr. 0 1110. 1 &I. Tubulc dianictcr 120-150 p. Tu1)ulcs :ire tightly packcvl
and it is difficult to find Leylig cclls. Cellular debris 1ias been clcnred from most tubules :tiid :I new generation of
speriixitids with orderly ;irr~ingeinentis present.
Riglit testis s:inililc five niontlis Inter, n t 3 y.3 ino. 1 &I. Tubule tliainctcr 350 p. Leylig ~ c l l s:ire very rare. Tlicro
arc ni:iny prcsperrn. nnd R few inaturc spenn free in the lumen.
94
25
23 Riglit testis s:iniplc one iiioiitli Ititor, :it 1 yr. 11. 1110. 1 d;i. Tiiljule diaiiictc*r 100-140 p. Slicriii are piweiit.
22 Riglit twtis snmplc a t 2 yr. 10 nio. 3 (la. Tubule dirineter 100-120 p. Spcriiiutids Ii:ive differentinted to lircsperni
or :iliiiost ni:iture sperin. Great aniounts of cellular dcbris fill l11nien. Few Leytlig cclls :ire differeiitititecl.
Figs. 22-25
Testis lfni 601, Iiirth, 1 i 4 tlnys' gest:itioii, I1 and E x 85. Slioivs Scrtoli cell nuclei rcinriining in a. bas;il position and
a persistent witle iiitcrtubu1:ir space ; however, Leylig cell size 1i:is decreased.
21
11;
c
of tlw interstitiill tissue
Testis M i 1 1 497, 110 i1:i. fetus, 11 niid
x 235. Shows tiil)iilcs in C r o S S spctioii. ~ i uricnt:itioii
coiirciitri~:illy :iroiinil tlic tul.iulcs, :ilso tlic enltlrgeiiieat of the cells within tlic coiieentrie rings, is sliowii.
"0
19 Testis M m f07, 110
&I. fetus, 11 and 15 x 87. Slio\vs short, uncoi!(~dtuliiilrs, di:iinetcr iO-80 p. Scrtoli cell cytoplas1ll
is well dcvelolied rind largely fills I U I I I C ~ I ~ . Tilc h o d i l l t ~ ~ r t l l ~ J lYl ~~IaCrC S contain abuad:int ceIIs, nian? of which arc
enhtrged and rounded.
ESPIANATIOX OF FIGURRS
PLATE 3
C . V A N WAC4EXhS A S D .\I. h S I J I P b O X
TESTICULAR DEVETAIPNEYT IN >IONKEY
b
EXPI,.IX.\'l'IO?I
EX?L.\I.\rnOs OF
OP FIGCUBS
m1:nUI
PLATE
PLATK 4
C . V A N WAC4EXhS A S D .\I. h S I J I P b O X
TESTICULAR DEVETAIPNEYT IN >IONKEY
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