Vasectomy in Rhesus Monkeys

IV ELECTRON MICROSCOPIC STUDIES OF THE SEMINIFEROUS EPITHELIUM

E. S. CHAPMAN,' P. M. HEIDGER, JR.,' R. M. HARRISON,' J . A. ROBERTS,2
G. J. DOMINGUE AND J. U . SCHLEGEL
I Department of Anatomy, University of Iowa, Iowa City, Iowa 52242; and Department
of Urology and Delta Regional Primate Research Center, Tulane University School of
Medicine, New Orleans, Louisiana 70112

ABSTRACT Studies were undertaken to assess t h e fine structural effects of

vasectomy upon t h e rhesus monkey testis, using 27 monkeys subjected to one of
three surgical techniques: control sham operation (COS), unilateral silk
vasoligation (USV), or unilateral clasp vaso-occlusion (UCV). The monkeys
were sacrificed from 1 t o 66 weeks after surgery; tissues were fixed in glutaral-
dehyde in collidine buffer and processed for electron microscopy (EM) using
routine techniques. No alterations were noted in t h e seminiferous epithelium of
any COS animal; only focal lesions were found in vasectomized animals. These
changes correlated with length of time post-vasectomy, a finding consistent
with earlier light microscopic studies (Heidger et al., '78). Even a t t h e longest
post-operative periods studied, spermatogenesis appeared in most animals to be
normal. One animal exhibited areas in which t h e seminiferous epithelium con-
sisted of Sertoli cells and spermatogonia, only. The seminiferous epithelium of
approximately one-third of other animals, both on t h e vasectomized and contra-
lateral sides, exhibited such alterations as development of extensive infoldings
and duplication within t h e basal lamina, and presence within t h e basal Sertoli
cell cytoplasm of late spermatids and sperm tails. These alterations were ob-
served in both UCV and USV animals. The presence of such late germ cells in
t h e basal Sertoli cell cytoplasm was noted by light microscopy; however, lu-
minal spermiophages encountered in one animal at t h e light microscopic level
were not detected in our EM study, which underscores t h e limitations of EM as
a survey technique. All UCV animals developed spermatic granulomas of t h e
vas, whereas only 3 of 13 USV animals developed such granulomas. I t does not
appear t h a t the alterations exhibited by t h e animals in this study would
necessarily dispose toward impaired spermatogenesis following vasovasostomy.

Its relative surgical ease, apparent low in-
cidence of clinical complications, and overall
effectiveness have contributed to t h e current
popularity of vasectomy as a means of ster-
ilization. Although as many as one million va-
sectomies are performed each year in t h e
United States alone (Derrick et al., '73), few
comprehensive studies of t h e effects of t h e
procedure have been conducted; even these
offer no uniformity of surgical technique and
underscore t h e considerable species variation
in response t o vasectomy. Studies at t h e elec-
tron microscopic (EM) level, where more sub-
tle alterations might be detected, have been
particularly limited (reviewed by Heidger and
Sawatzke, '77); other studies in primates have
emphasized t h e response of t h e excurrent duct
system, rather t h a n t h e testis (Bedford, '76;
Alexander, '75).
Accordingly, a multidisciplinary investiga-
tion of t h e immunologic and morphologic ef-
fects of vasectomy in rhesus monkeys was un-
dertaken. The surgical procedures, observed
effects at t h e gross anatomical and light mi-
croscopic (LM) levels, and results of immu-
nologic studies have been published earlier
(Harrison e t al., '77; Heidger e t al., '78; Dom-
ingue et al., '77). The present paper describes
at t h e electron microscopic level t h e response
of t h e seminiferous epithelium of t h e rhesus
monkey testis t o vasectomy.
Received Jan. 23, '78. Accepted Mar. 29, '78.

ANAT. REC. (1978) 192: 41-54. 41

42 E. S. CHAPMAN ET AL.

MATERIALS AND METHODS (c) Unilateral silk vasoligation (USV). The

The monkeys used in this study were all exposed vas was ligated with 2-0 silk suture
mature Macaca mulatta (rhesus) male mon- but was not sectioned.
keys weighing 5.2-12.0 kg, purchased from Vasectomies were performed a t three dif-
commercial dealers and other primate centers. ferent time periods; this schedule was necessi-
Details of the animal population and its care tated in part by efforts to evaluate possible
were as previously reported (Heidger et al., seasonal influences upon spermatogenesis and
'78). The monkeys were randomly assigned to thus upon testicular morphology (Harrison et
undergo various surgical procedures including al., '77). The animals were followed from 1 to
the following: 66 weeks after surgery (table 1); control and
(a) Sham-operated control (COS). Following experimental animals were killed on the same
standard presurgical preparation and with the date in an attempt to determine any influence
monkey maintained on halothane, a 2-cm inci- of seasonal variation upon spermatogenesis,
sion was made in the scrota1 skin lateral to the and thus upon interpretation of results from
base of the penis. The tunica vaginalis was ex- animals killed a t various times throughout
posed by blunt dissection and then cut. The the year. Tissue samples were taken from the
vas deferens was exposed in a similar manner superficial, anterior, equatorial portion of the
and a 2-0 silk suture was passed around i t a t a testis.
point just proximal t o the upper pole of the For electron microscopy the tissues were
testis. The suture was then removed; the fixed in 5% glutaraldehyde in collidine buffer
tunica vaginalis and skin were closed with 2-0 with post-fixation in osmium. This step was
chromic sutures. The monkeys were carefully followed by dehydration in ethanol series,
observed post-operatively and were returned solvent substitution with propylene oxide, and
to their cages in 24 hours. No antibiotics were embedment in Epon 812. Thin sections were
given. Except as noted below, all monkeys re- cut on a Sorvall MT-2b Ultramicrotome; tis-
ceived this same surgical procedure and post- sue contrast was enhanced with lead citrate
surgical treatment. and uranyl acetate, and/or bismuth (Barrett
(b) Unilateral clasp vaso-occlusion (UCV). e t al., '76). Microscopy was performed on a
Doctor T. H. Clewe prepared silastic-covered JEOL lOOB electron microscope.
stainless steel clasps which were placed over
the vas, perpendicular to the lumen; these RESULTS
clasps were designed to occlude the lumen The histology and fine structure of the semi-
without contusion and tissue damage. niferous epithelium from unilaterally vasec-

TABLE 1

Summary of experimental groups

Type of Time studied Number of Month Number of animals
surgery post-operatively animals sacrificed with vas granulomas

usv 66 weeks 2' November 2

48 weeks 32 November (1) 1
February (2)
18 weeks 2 June 0
4 weeks 3 February 1
1 week 3 March 0
ucv 66 weeks 2 November 2
36 weeks 2 April 2
18 weeks 1 May 1
cos 66 weeks 3 April (1) 0
November (2) 0
48 weeks 2 February 0
18 weeks 2 May 0
1 week 2 March 0
~~ ~

Note: Four additional animals were originally included in this study but were not used for morphological analysis for various
reasons: death, disease. or use of the animal for active immunization experiments.
' At the LM level, one of these animals exhibited tubules containing only Sertoli cells and spermatogonia.
'At the LM level, one of these animals exhibited tubules containing only Sertoli cells and spermatogonia; such affected tubules
contained spermiopbages within their lumina, and late spermatids were noted within the basal Sertoli cell cytoplasm.
 VASECTOMY IN RHESUS MONKEYS
tomized animals was compared with that of
monkeys subjected to sham operation; in addi-
tion, the seminiferous epithelium from the li-
gated side was compared with that of the
unoperated side. Attention was directed to-
ward certain features which seemed most
susceptible to change after vasectomy as re-
ported in studies involving other species. Mi-
crographs were selected to illustrate these
points and to indicate the general quality of
the tissues studied.
Sham-operated control animals
No consistent fine structural alterations
were found in the testes from COS animals,
nor was any seasonal variation in tubular
morphology noted.
Unilateral silk vasoligation animals
Changes in the morphology of the semi-
niferous epithelium in monkeys sacrificed 66
weeks after vasectomy were found within the
basal lamina and the basal Sertoli cell cyto-
plasm. In one animal, in which focal areas of
bilateral aspermatogenesis were noted a t the
LM level, several late spermatids and sperm
tails were found in the basal and perinuclear
Sertoli cell cytoplasm on both sides. This was
also the case on the operated side of a second
animal; this latter animal had exhibited no
changes a t the light microscopic level, how-
ever. The germ cells and acrosomal complex
appeared to be normal in all cases, and the spe-
cialized Sertoli cell-germ cell and Sertoli-Ser-
toli cell junctions appeared to be intact (fig.
1). The basal lamina displayed prominent
infoldings and projections into the Sertoli cell
cytoplasm bilaterally in one of these animals
and on the operated side of the second. Few
spermatocytes or early spermatids appeared
t o be present on the side contralateral to va-
sectomy in one animal; however, the signifi-
cance of a reduced number of developing germ
cells is difficult to ascertain. LM examination
of this animal revealed areas of epithelium
consisting of Sertoli cells and spermatogonia
only (Heidger et al., '78). Several instances of
residual bodies located in the perinuclear Ser-
toli cell cytoplasm were noted; this was never
encountered in the testes from control-oper-
ated animals.
The seminiferous epithelium of the three
animals examined 48 weeks postvasectomy
exhibited few alterations when compared to
the control animals sacrificed a t the same
time. The Sertoli cells typically contained nu-
43
merous, small, electron-dense mitochondria;
lipid droplets were plentiful, as were dense
bodies. Rough endoplasmic reticulum and
multivesicular bodies were also notable com-
ponents of the Sertoli cell cytoplasm. Germ
cells were present at various stages of matura-
tion, suggesting no apparent disruption of
spermatogenesis (fig. 2). Acrosomal develop-
ment appeared to be normal. Typical junctions
between Sertoli cells and germ cells, composed
of the Sertoli cell plasma membrane, the clas-
sical parallel array of electron-dense fibrils,
and a discontinuous layer of cisternae of the
endoplasmic reticulum, were frequently en-
countered. The morphology of the myoid cells
was consistent with that present in the con-
trol animals. These plaque-shaped cells were
characterized by overlapping cell processes,
numerous microfilaments, and an abundance
of pinocytotic vesicles. In two of three ani-
mals, the basal lamina appeared to be dupli-
cated and folded upon itself in certain areas of
testis from the vasectomized side, similar to
that cited above in the 66-week animals (figs.
3, 4). Interestingly, the animal in which LM
studies revealed isolated areas of epithelium
consisting of Sertoli cells and spermatogonia
only, and with spermiophages present in the
lumen (Heidger et al., '781, did not show any
evidence of testicular alteration in the sam-
ples studied a t the EM level.
Both animals sacrificed a t 18 weeks post-
vasectomy exhibited generally normal mor-
phology, with only a few exceptions. The Ser-
toli cells contained the usual array of lipid
droplets, dense bodies, endoplasmic reticulum,
and normal mitochondria. However, late sper-
matids and sperm tails were occasionally pres-
ent in the basal and perinuclear Sertoli cell
cytoplasm on the operated side of one animal
(fig. 5). In addition, spermatids and sperm
tails were often seen in a similar position in
the contralateral side of the same animal.
Some of these spermatids exhibited the usual
array of membranes: the spermatid nuclear
membrane, the inner and outer acrosomal
membranes, the spermatid plasma membrane,
and the abutting Sertoli cell plasma mem-
brane, while others were surrounded by rem-
nants of the usual investing membranes. In
control animals, late spermatids were often
seen embedded within the luminal cytoplasm
of the Sertoli cell, but never within the basal
cytoplasm. In the second vasectomized ani-
mal, residual bodies were seen more basally
within the seminiferous epithelium than were
44 E. S. CHAPMAN ET AL.
similar bodies within the control tissue. In
addition, fewer developing germ cells ap-
peared t o be present in some tubules, but it is
difficult to quantitate such an impression due
to the small size of the tissue samples used in
electron microscopy and the focal nature of
such lesions. Our LM studies failed to reveal
any such changes in the number of developing
germ cells in these 18-week animals. While
the myoid layer in the vasectomized animals
was indistinguishable from that of controls
sacrificed a t the same time, the basal lamina
in some areas exhibited convolutions and dup-
lications extending into the Sertoli and germ
cell cytoplasm (fig. 7 ) . In control animals, the
basal lamina was usually flat and even, or
somewhat wavy, but never as highly infolded
as in these tubules. Several examples of un-
usual, abnormally formed spermatids were
encountered in these animals, but small num-
bers of abnormal spermatids were also a fea-
ture of the testes from COS animals.
Few alterations were noted in the semi-
niferous epithelium of the three animals
killed four weeks after vasectomy. The Sertoli
cells contained the usual complement of or-
ganelles - mitochondria, dense bodies, rough
endoplasmic reticulum, lipid droplets. On the
vasectomized side of one animal, late sper-
matids and numerous sperm tails were seen in
the Sertoli cell cytoplasm near the basal lami-
na. The remaining germ cells and their germ
cell-Sertoli cell junctions appeared t o be nor-
mal, as did the Sertoli-Sertoli cell junctions.
Some infoldings of the basal lamina were seen
in certain tubules on the vasectomized side in
two monkeys; such infoldings were not nearly
as pronounced as in the longer term animals.
Three animals were sacrificed one week
after vasectomy; in general, the seminiferous
epithelium of these animals appeared normal.
On the vasectomized side of one animal, bun-
dles of filaments were observed within the
basal Sertoli cell cytoplasm (fig. 6 ) . These
bundles were of variable length and diameter;
the filaments comprising them were approx-
imately 65 A in diameter and closely packed.
Unilateral clasp vaso-occlusion animals
Few alterations were present in the semi-
niferous epithelium of the two monkeys sac-
rificed 66 weeks post-vasectomy. On the va-
sectomized side of one animal, the basal cy-
toplasm of some Sertoli cells contained, in
addition t o the normal complement of organ-
elles, late spermatids enveloped in a loose
configuration of membranes (figs. 8, 9).Germ
cells and their junctions with Sertoli cells ap-
peared to be normal; the structure of the Ser-
toli-Sertoli cell junctions also was apparently
unaltered by vasectomy. The basal lamina and
myoid layer in both animals exhibited normal
morphology.
Two UCV animals were sacrificed 36 weeks
after vasectomy. On the vasectomized side of
one animal, sperm tails were found in the Ser-
toli cell cytoplasm in some tubules. Testes
from the vasectomized side of both animals
contained some bundles of fibrils in the basal
Sertoli cell cytoplasm, similar t o those de-
scribed in the results from the 1-week USV
animals. In addition, the basal lamina dis-
played folds similar to those mentioned previ-
ously. The side contralateral t o vasectomy in
the first animal was normal except for the
presence of a degenerating late spermatid in
the perinuclear Sertoli cell cytoplasm. In the
second animal, late spermatids, sperm tails,
and residual bodies were present in the basal
Sertoli cell cytoplasm in some tissue speci-
mens from the contralateral side.
The animal sacrificed a t 18 weeks post-va-
sectomy displayed few alterations in its semi-
niferous epithelium. In several instances on
the side contralateral to vasectomy, observa-
tion of the basal and perinuclear Sertoli cell
cytoplasm revealed late spermatids and sperm
tails (fig. 10).The basal lamina of both sides
exhibited areas of infoldings and duplications,
but not to the degree seen in some other ani-
mals. The remaining germ cells and their
junctions, as well as the Sertoli-Sertoli cell
junctions, exhibited normal morphology.
DISCUSSION
Although others (Bedford, '76; Alexander,
'75) have examined the excurrent duct system
of the rhesus monkey following vasectomy,
and have given brief attention to testicular ef-
fects, the present study is the first t o empha-
size the fine structural response of the rhesus
monkey testis to vasectomy. In assessing the
effects of vasectomy upon the testis of any
species, several variables must be considered.
In particular, these include: (1) well-con-
trolled surgical procedures which avoid such
secondary effects as post-operative cryptor-
chidism, infection, and the compromise of the
testicular blood supply; (2) the inherent dis-
tensibility of the duct system of the species
studied; and (3) the development of gran-
ulomas of the vas or epididymis following va-
sectomy. The results of our present study are
 VASECTOMY IN RHESUS MONKEYS
discussed below with reference to these con-
siderations.
Concerning t h e first variable, t h e authors
a r e confident t h a t the surgical procedures
used were carefully performed and controlled
such t h a t t h e effects seen in t h e post-vasec-
tomy seminiferous epithelium were due t o
vasectomy alone. Support for this view lies in
t h e complete lack of any effect of t h e control
operation upon t h e testes of t h e COS animals.
It should be noted also t h a t differences in sur-
gical procedure may preclude direct com-
parison of results from different laboratories
even utilizing t h e same species.
Correlation of t h e distensibility of t h e duct
system and t h e role of pressure damage with
t h e testicular response to vasectomy h a s been
attempted by many investigators. Studies at
t h e EM level in t h e rabbit by Flickinger ('75)
and Alexander and Tung ('77) have shown
t h a t t h e rabbit testis is apparently unaltered
up to six months post-vasectomy; they re-
ported t h a t t h e rabbit duct system exhibits a
remarkable degree of distensibility. Corre-
spondingly, t h e dog, which exhibits little dis-
tention of t h e excurrent duct system after
vasectomy, h a s been reported to show altera-
tions in t h e testis at both t h e LM and EM
levels as early as one month post-operatively
(Heidger and Sawatzke, '77; Grewal and Sa-
chan, '68; Derrick et al., '74). From his short
term studies of t h e rhesus monkey response t o
vasectomy, Bedford ('76) suggested t h a t tes-
ticular changes in this species may be pre-
vented for a short time after vasectomy by t h e
distention of t h e vas and the presence of low
epididymal cysts. Certainly our findings of lit-
tle or no alteration in t h e rhesus testis up to
18 weeks post-vasectomy are consistent with
those of Bedford. Longer term studies in all of
t h e above species show alteration in t h e testis
following vasectomy, implying t h a t there is a
limit to t h e protection from pressure afforded
by distensibility.
Pressure upon t h e testis which may develop
after vasectomy may be minimized not only by
distensibility of t h e excurrent duct system,
but also by t h e formation of spermatic gran-
ulomas of t h e vas. Kuwahara and Frick ('75)
noted t h a t r a t s which developed large gran-
ulomas exhibited normal testicular morphol-
ogy three weeks after vasectomy; in their
study, tubular degeneration was evident in
those animals which did not develop gran-
ulomas. Granuloma formation is rare in t h e
dog, in which tubular alterations occur early
45
post-vasectomy (Heidger and Sawatzke, ' 7 7 ) .
In t h e present study, 8 of 26 animals de-
veloped granulomas a t t h e site of vasectomy;
these included 3 USV animals and all 5 UCV
animals (table 1).However, there appears to
be no correlation between alterations in the
seminiferous epithelium and t h e presence or
absence of a spermatic granuloma; i.e., the
role of granuloma formation in the develop-
ment of testicular alterations after vasectomy
remains unclear. With respect to granuloma
formation, t h e present study suggests t h a t
clasp-ligated animals are more likely to de-
velop such granulomas and t h a t granulomas
may develop as early as four weeks post-vas-
ectomy in t h e monkey. Further studies on
the spermatic granuloma of t h e vas - t h e
most common complication of vasectomy
in humans (Alexander and Schmidt, '77;
Schmidt and Morris, '73) - are in progress.
Our evaluation of spermatic granulomas of
t h e epididymis, which may also afford relief of
pressure on t h e testis, has not progressed suf-
ficiently a t this time to permit correlations
between epididymal granuloma formation and
possible testicular alterations.
Our studies demonstrate t h a t t h e altera-
tions which may accompany vasectomy are
more prevalent a t t h e longer post-operative
times evaluated. No remarkable changes were
present in animals followed for one week. Few
alterations were present in animals followed
for four weeks, with a higher frequency of al-
terations found in animals vasectomized for
longer periods. These results are similar to
those reported in t h e rabbit (Flickinger, '75)
and rat (Bedford, '76); however, they clearly
differ from findings in t h e dog (Heidger and
Sawatzke, '77; Grewal and Sachan, '68; Der-
rick et al., '74) in which alterations developed
shortly after vasectomy and later subsided.
We would stress t h a t alterations detected in
t h e seminiferous epithelium were focal in
nature. In this respect, our findings corre-
spond t o those in t h e dog (Heidger and
Sawatzke, '77) and rabbit (Alexander and
Tung, '77) and a r e distinctly different from
those reported in t h e vasectomized guinea pig
(Alexander, '73). This is noteworthy in t h a t
other studies (Alexander, '75) have utilized
testicular biopsies to evaluate t h e testicular
response t o vasectomy; our d a t a suggest t h a t
restricted tissue sampling may fail to detect
focal alterations. Furthermore, we would em-
phasize Flickinger's ('73) point t h a t electron
microscopy is limited as a survey technique
46 E. S. CHAPMAN ET AL.
due to t h e restricted tissue specimen size; it is
best utilized in conjunction with light micro-
scopic studies of the larger tissue samples per-
mitted. Illustrating this point in our studies is
our failure to detect alterations a t t h e EM
level in the testis of a 48-week animal which
under LM examination exhibited luminal
spermiophages within seminiferous tubules
containing only Sertoli cells and spermato-
gonia.
As mentioned in RESULTS. no morphological
alterations were seen in the complex Sertoli-
Sertoli cell junctions, t h e principal locus of
the blood-testis barrier in this species, as de-
scribed by Dym ('73). However, studies by
Heidger and Sawatzke ('77) of t h e dog blood-
testis barrier following vasectomy and by
Neaves in the r a t ('73) have shown t h a t the in-
tegrity of these junctions cannot be deter-
mined by fine structural examination, alone.
When lanthanum, a tracer of t h e intercellular
space (Revel and Karnovsky, '67) was in-
cluded in the fixative in the above studies, nu-
merous junctions which appeared to be mor-
phologically intact were shown t o allow t h e
penetration of this tracer over t h e entire
length of t h e Sertoli-Sertoli cell junction,
reaching the spermatid compartment of t h e
seminiferous epithelium. I t is hoped t h a t stud-
ies in progress in our laboratory will clarify
this question in t h e rhesus monkey.
The fine structural effects of vasectomy
upon the seminiferous epithelium of this spe-
cies are focal in nature and include changes
within the basal lamina and the presence of
late spermatids and sperm tails in basal and
perinuclear Sertoli cell cytoplasm. The latter
finding suggests a possible increase in phago-
cytic activity of t h e Sertoli cells, which are
known to be capable of phagocytosis (Carr et
al., '68; Kingsley-Smith and Lacy, '57). Alter-
natively, such placement of intact spermatids
could reflect alteration in the usual compart-
mentation of t h e seminiferous epithelium.
Impaired fertility following vasovasostomy,
despite patency of t h e vas, has been reported
in men (Montie and Stewart, '74). In a recent
study by Alexander ('77) 15 rhesus monkeys
which had been vasectomized for six months
were subjected t o vasovasostomy. After a 4-
month mating period, 2 of 15 monkeys were
classed as infertile; three others were classed
as subfertile. It does not appear t h a t t h e rela-
tively few alterations observed in the semi-
niferous epithelium of t h e animals in our
study following vasectomy are of such se-
verity as to be responsible for t h e reduced
fertility in these primate studies. In certain
animals, changes were noted on t h e side
contralateral to vasectomy; if these changes
have a n immunological basis, t h e responsible
antibody was not amenable to detection by t h e
techniques used in our earlier study (Dom-
ingue et al., '77). Further studies, both basic
and clinical, are necessary t o understand more
fully t h e effects of vasectomy - the most
common surgical procedure in adult males
(Schmidt, '68) - upon t h e male reproductive
tract so t h a t t h e reversibility of this procedure
can be assured.
ACKNOWLEDGMENTS
The authors gratefully acknowledge the ex-
cellent technical assistance of Mrs. Rita
Azzam, Mrs. Mary Donnell, and Mrs. Pearl
Gervais.
This investigation was supported by NIH
Contract NO 1 HD 3 2758, NIH Grant HD 1
0571, and a Sigma xi Research Award to
E.S.C..
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 VASECTOMY IN RHESUS MONKEYS
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Heidger, P. M., Jr., and C. L. Sawatzke 1977 Fine struc-
tural effects of vasectomy upon the male reproductive
system. In: The Male Reproductive System. R. D. Yates
and M. Gordon, eds. Masson, New York, pp. 131-153.
47
Kingsley-Smith, B. V., and D. Lacy 1959 Residual bodies of
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941-947.
 PLATE 1

EXPLANATION OF FIGURES

1 Electron micrograph of adjacent Sertoli cells showing a Sertoli-Sertoli cell junction;

66-week USV animal, unoperated side. These specialized junctions are characterized
by numerous tight junctions (arrowheads), arrays of electron dense fibers, and sub-
jacent parallel cisternae of endoplasmic reticulum. X 32,200.Inset: Higher mag-
nification of figure 1 showing the junction in greater detail. X 45,000.

2 Electron micrograph of a portion of t h e apical seminiferous epithelium from a 48-

week USV animal, operated side. The presence of normal late spermatids exhibiting
characteristic spermatid-Sertoli cell junctional complexes is indicative of active
spermatogenesis. Note the manchette formation in a n adjacent spermatid sectioned
a t a different level. x 10,000.

3 Electron micrograph of the periphery of a seminiferous tubule from a 48-week USV

animal, operated side. Note the prominent infolding and duplication of t h e basal
lamina (star). The myoid layer exhibits normal structure. X 22,600.

4 Electron micrograph of the basal region of a seminiferous tubule from a 48-week

USV animal, operated side. Processes of the basal lamina (star) project deeply into
the cytoplasm of this Sertoli cell. Note the presence of numerous lipid droplets and a
large dense body in the Sertoli cell basal cytoplasm. X 12,000.

48
 PLATE 1
VASECTOMY IN RHESUS MONKEYS
E. S. Chapman et al.
 PLATE 2
EXPLANATION OF FIGURES

5 Electron micrograph of the basal Sertoli cell cytoplasm from an 18-week USV ani-
mal, unoperated side. The presence of sperm tails (arrowheads) and a developing
spermatid (arrow) in the basal Sertoli cell cytoplasm reflects a disruption in the
usual Sertoli cell-germ cell spatial relationship within such tubules. X 9,000.

6 Electron micrograph of seminiferous epithelium from a I-week USV animal, oper-

ated side. These bundles of parallel filaments of unknown origin (arrows) were occa-
sionally present in t h e basal Sertoli cell cytoplasm. X 25,000.

7 Electron micrograph of t h e seminiferous epithelium from an 18-week USV animal,

operated side. Infolding8 of the basal lamina (star) project deeply into the Sertoli
cell cytoplasm, which contains numerous lipid droplets. X 8,300.

50
 PLATE 2
VASECTOMY IN RHESUS MONKEYS
E. S. Chapman e t al.

51
 PLATE 3

EXPLANATION OF FIGURES

8 Electron micrograph of the seminiferous epithelium of a 66-week UCV animal,

operated side. A spermatid head (arrowhead) is present in the basal Sertoli cell cy-
toplasm; its usual surrounding membranes appear to be lacking. The sper-
matogonium (sg),lipid droplets, the junctions, and t h e deeply indented Sertoli cell
nucleus all reflect the condition observed in COS animals. x 9,400.

9 Electron micrograph of the basal Sertoli cell cytoplasm from a 66-week UCV ani-
mal, operated side. Note the presence within the Sertoli cell cytoplasm of a late
spermatid (arrowhead) with its enclosing membranes in disarray, suggestive of
phagocytosis by t h e Sertoli cell. Several extensive Sertoli-Sertoli cell junctional
complexes are cut in oblique section (arrows). X 8.000.

10 Electron micrograph of the basal portion of a Sertoli cell from the 18-week UCV
animal. Note the presence of a late spermatid (arrowhead) with intact Sertoli cell-
spermatid junctional complex in t h e Sertoli cell cytoplasm. Sc, Sertoli cell nucleus.
X 24,700.

52
VASECTOMY IN RHESUS MONKEYS PLATE 3
E. S. Chapman e t al.