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Response of epididymal duct to the temporary depletion of spermatozoa induced by testicular irradiation in mice.

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THE ANATOMICAL RECORD 207:17-24 (1983)
Response of Epididymal Duct to the Temporary Depletion of
Spermatozoa Induced by Testicular Irradiation in Mice
KAZUHIRO ABE, HIROKO TAKANO, A N D 'I'AKASHI I T 0
Dc~part t n c w t of A natomy, Hok ka d o Utr LOCV ally Sc h 001 of M d r L I 11 c, Sapporo
060, Japan
ABSTRACT
The mouse epididymal duct can be histologically divided into
five segments (I-V), and the principal cells in segment I1 appear to secrete
periodic acid-Schiff (PAS)-positive material into the lumen. In this study, male
dd-mice received one, two, or four 800-R doses of radiation beginning a t age 50
days. Mice receiving multiple doses were irradiated a t 1-week intervals.
After irradiation, marked depletion of spermatozoa, or aspermia, occurred in
the epididymal duct for 2 to 16 weeks after a latency period of 3 to 4 weeks
according to the times of irradiations. During oligospermia or aspermia, PASpositive inclusions appeared in the principal cells in segment IV. The inclusions occupied a supranuclear position and appeared as round granules and
globules measuring 2-15 pm in diameter, and increased in number, size, and
staining intensity with time. They disappeared after reappearance of spermatozoa. The findings suggest that PAS-positive material may bind to spermatozoa and, if not bound, is reabsorbed by the principal cells in segment IV and
deposited as intracellular inclusions, and the principal cells in segment IV are
capable of digesting the accumulated PAS-positive material.
The mouse epididymal duct is divided into
five segments (I-V) according to the morphological differences of the principal cells (Takano, 1980; Abe et al., 1982a, 1983a). The
principal cells in segment I1 have PAS-positive cytoplasm and appear to secrete PASpositive material into the lumen (Abe et al.,
1982a). Such PAS-positive material may contain the specific epididymal glycoprotein
which binds to spermatozoa and renders
spermatozoa capable of fertilization (Lea et
al., 1978; Olson and Hamilton, 1978; Faye et
al., 1980; Moore, 1980).The principal cells in
segment IV acquire PAS-positive cytoplasmic inclusions after ligation of the efferent duct or epididymal duct proximal to
segment 11(Abe et al., 1982a). These findings
suggest that the PAS-positive inclusions in
segment IV appear under the following conditions: 1) release of PAS-positive material
into segment IV; and 2) interruption of flow
of spermatozoa from the testis into segment
IV. Thus we assumed that, unless bound to
spermatozoa, the specific epididymal glycoprotein is absorbed by the principal cells in
segment IV.
Ligation at the efferent duct or epididymal
duct proximal to segment I1 stops the flow
0 1983 ALAN R. LISS, INC
not only of spermatozoa but also of secretions
from the testis into segment Tv. In this study,
to understand more clearly effects of the
presence of spermatozoa on the function of
the principal cells in segments I1 and IV, we
attempted to eliminate spermatozoa temporarily by testicular irradiation. In addition,
to demonstrate the association of the intraepithelial PAS-positive inclusions in segment
IV with the intraluminal PAS-positive material from segment 11, the epididymal duct a t
the junction between segments I11 and IV
was ligated after irradiation and before disappearance of spermatozoa in the epididymal
duct. The changes induced in the epididymal
duct were analyzed by qualitative and quantitative morphology.
MATERIALS AND METHODS
In this experiment involving 115 male ddmice, 102 mice received one, two, or four 800R doses of radiation beginning at age 50 days.
Mice receiving multiple doses were irradiated at 1-week intervals, Five mice received
two radiation doses and underwent unilatKrct,ivrd September 20, 1982; iicceptcd April 20, 1983
18
K. ABE, H. TAKANO, AND T. IT0
era1 ligation of the epididymal duct between
segments I11 and IV, according to the previous procedure (Abe et al., 1982b), 2 weeks
after the second irradiation. Eight mice, 60
to 90 days old, nonirradiated, served as controls.
During irradiation, the mice were anesthetized with pentobarbital sodium (Nenibutal)
injected intraperitoneally, placed supine, and
covered with wet cotton and a 6-mm thick
plastic board in order to achieve the optimum effective depth (see “build up” in the
textbook by Jones and Cunningham, 1971).
The wet cotton filled the space between the
plastic board and the lower body of the
mouse. Cephalad to the umbilical level, the
body was shielded by placing a 5-cm thick
lead slab on the plastic board. From a 6oC0
source, 800 R was administrated at a dose
rate of 125 R per minute to the lower half of
the body. After irradiation, the mice were
killed a t various intervals and the epididymides and testes were removed. They were
fixed in Bouin’s solution for 3 hours and then
embedded in paraffin. Serial 6-pm longitudinal sections of the epididymis and testis were
cut, stained with periodic acid-Schiff (PAS)
and hematoxylin, and examined by light
microscopy.
For quantitative observations, in one section among the serial sections from each epididymis, the one ductal cross section showing
the most marked appearance of the intraepithelial PAS-positive inclusions plus the three
neighboring sections were selected. The minimum and maximum inner diameters of
these ductal profiles and the number and size
of the PAS-positive inclusions were rneasured. The ductal circumferences were calculated using the minimum and maximum
diameters. The appearance of the intraepithelial PAS-positive inclusions in each epididymis was graded using the equation C d,
2 C d + + iL x 100, where d+ = diameters
of inclusions staining more weakly than the
luminal material, d, + = diameters of the inclusions staining more intensely than the luminal material, and L = the total circumferential length of the ducts.
spermatocytes were regenerated. Spermatozoa disappeared a t 5 weeks and reappeared
at 7 weeks postirradiation.
Two of four 800-R irradiations depressed
the number of type A spermatogonia and
accelerated the sequential disappearance of
the other spermatogenetic cells. Recovery of
spermatogenesis began 3 weeks after the last
irradiation but was more protracted. Thus,
two and four irradiations resulted in disappearance of spermatozoa from the seminiferous epithelium from 4 to 10 weeks and 4 to
16 weeks after the first irradiation, respectively,
Epididymal Duct
In the normal epididyniis, the epididyinal
duct in all segments except for segment I
contains spermatozoa and PAS-positive material (Figs. 1, 2a, 3a). In segments IV and V,
the lumen is extremely distended with abundant spermatozoa. The principal cells in segment IV were 15-20 pm high and about 8 pin
’I
,111
II’
-lV
+
RESULTS
Testis
The seminiferous tubule epithelium
showed disappearance of type B spermatogonia and leptotene and zygotene spermatocytes 1week after a single 800-R irradiation.
Cells in all stages of spermatogenesis disappeared and reappeared sequentially. At 3
weeks postirradiation, spermatogonia and
Fig. 1. Longitudinal section of the testis and epididyinis of the mouse. The epididymal duct is divided into
five segments (1-V). Scgmcnt I1 appears dark because
the epithelial cells are stained with periodic acid-Schiff
(PAS).The duct in segments IV and V contains abundant
spermatozoa (black content of the lumen). PAS-hematoxylin. x7.
EPIDIDYMAL PAS-POSITIVE MATERIAL AND SPERMATOZOA
Fig. 2. Segment IV of the mouse epididymal duct. a)
Normal. The duct contains numerous spermatozoa and
PAS-positive material. b) Segmciit IV 10 weeks after the
first of the two irradiations. The duct contains strongly
PAS-positive material and little spermatozoa. PAS-positive inclusions (arrows) are seen in the cpithelial cells. c)
19
Segment IV in the duct which h a d been ligated between
segments 111 and IV 2 weeks after the second irradiation,
is observed 4 weeks after ligation. The duct is decreased
in diaimtcr, and shows no spermatozoa and little PASpositive matcrial in the lumen and no inclusions i n the
epithelium. PAS-hematoxylin. x 170.
wide and had round nuclei, about 6 pm in appeared in the epididymal duct (Fig. 3b).
These cells disappeared soon after spermatodiameter (Fig. 3a).
After irradiation, aspermia or marked oli- zoic regeneration. The luminal material was
gospermia occurred in the epididymal duct more strongly PAS-positive in the spermatofor a time of duration after a latency period, zoa-depleted ducts when compared with the
as shown in Fig. 4. Spermatozoa disappeared controls (Figs. 2b, 3b,c).
During aspermia, PAS-positive inclusions
sequentially from the proximal to distal segments of the duct. In segment IV, after one appeared in the principal cells in segment
irradiation, spermatozoa were normal in IV, especially in the proximal region (Figs.
number until 4 weeks, disappeared at 5 2b, 3b,c). They usually occupied a supranuweeks, began to reappear a t 6 weeks, and clear position and appeared as round granbecame normal in number again a t 7 weeks ules and globules measuring 2-15 p m in
postirradiation. After two and four irradia- diameter (Figs. 3b,c). The inclusions intions, spermatozoa in segment IV were nor- creased in number, size, and staining intenmal in number until 3 weeks but disappeared sity with time.
Figs. 5 and 6 show the relationship bea t 4 weeks, and aspermia persisted until 10
weeks and 16 weeks after the first irradia- tween the disappearance of spermatozoa from
tion, respectively. Spermatozoa in segment the epididymal duct and the appearance of
IV became nearly normal in number a t 15 the PAS-positive inclusions in the principal
weeks and recovered incompletely at 20 cells in segment IV. After one irradiation,
weeks after the first irradiation of two and the epithelium in segment IV was normal for
the initial 4 weeks while spermatozoa were
four irradiations, respectively.
As spermatozoa disappeared, spermatoge- present. Then, as spermatozoa were depleted
netic cells, mainly consisting of spermatids, from the duct, the PAS-positive inclusions
20
K. ABE, H. TAKANO, AND T. IT0
Fig. 3 . Segment IV of the epididymal duct. a ) Normal.
The lumen has many spa-matozoa and PAS positive material. The epithelial cells have no inclusions. b, c) Ten
weeks after the first of four irradiations. The lumen
contains strongly PAS-positive material and has spermatids (ST) and few spermatozoa. The principal cells
contain PAS-positive inclusions varying in size and
staining intensity (arrows). PAS-hematoxylin. ~ 6 5 0 .
Fig. 4. Changes in number of spermatozoa in the cpididymal duct of segment IV after one (1 x ), two ('2 x I, and
four (4 x 1 irradiations (arrows).The normal range (standard deviation) is shown by parallel lines. Bars represent
standard deviations. The initial changes after two irradiations are almost t h c hiltlie as those after four irradiations.
21
EPIDIDYMAL PAS-POSITIVE MATERIAL AND SPERMATOZOA
.
!X
'X
.
0
.
.
.
-
.
.
.*
:
-.*.
t
we.
6
-&e-
a
-.-
.*
rn
r-10 12-15
7-9
*
20
Weeks
Fig. 5. Intraepithelial PAS-positive inclusions in segmcnt IV various weeks after onc (1x1 irradiation and
after the first of two ( 2 X ) and four (4X ) irradiations.
Each dot represents the grade of appearance of the inclusions in each epididymis. Dots in (-) exhibit epididymides which had no inclusions (0%in grade). Bars among
dots represent average values. Black belts show the duration in which the duct contains many spermatozoa.
Note that PAS-positive inclusions appeared and increased in grade with time during aspermia and that
epididymides had little or no inclusions after reappearance of spermatozoa.
IOpm : Diameter
25 : Number/mm
30% : Grade
t
t
4 1 4 . 4
Irradiation
0
Spermatozoa ( - )
4
8
Weeks
Fig. 6. Possible changes, summarized from the results, in diameter, number, or grade of PAS-positive inclusions in segment IV after one (1 X), two (2 X), and four
(4x1 irradiations (arrows). Thick lines represent the periods of aspermia. The diameter and number of the inclu-
12
16
20
sions showed similar changes to that in the grade. During
the phase of the most significant changes, the averaged
inclusions were about 10 pm in diameter, numbered
about 25 per mm of the lateral aspect of the epithelium,
and 30% in grade.
22
K. ABE, H. TAKANO. AND T. IT0
appeared a t 5 weeks and became more significant 6 weeks after the irradiation. The inclusions disappeared after reappearance of
spermatozoa. After four irradiations, the
PAS-positive inclusions were not observed for
initial 3 weeks after the first irradiation
while spermatozoa were still present. Following disappearance of spermatozoa from the
duct, the inclusions appeared 4 weeks after
the first irradiation, increased in grade until
about 10 weeks, and began to decrease in
grade at 16 weeks (Figs. 5 , 6). During the
phase of the most significant changes, the
averaged PAS-positive inclusions were about
10 pm in diameter, numbered about 25 per
mm of the lateral aspect of the epithelium,
and about 30% in grade (Fig. 6). In the
group irradiated twice, the radiation-induced
changes were of intermediate severity between those after one to four irradiations.
The inclusions were still present in the duct,
which showed incomplete recovery of spermatozoa 15 weeks and 20 weeks after the
first of two and four irradiations, respectively (Fig. 7). Segments I, 11, 111, and
V showed no distinctive postirradiation
changes.
In the mice with ducts ligated 2 weeks after
the second irradiation, the diameters of the
50-
40-
-<
c
30( 5 "
''
0
C
o
o
-5
20-0
"C
._
--2+
n
0
.=
10-
0
0
'OD
0
0
100
moob o 800
. .
.*
200
300
Spermatozoa : Number/lO>ml
Fig. 7. Relationship between the number of spermatozoa a n d the PAS-positive inclusions during recovery of
spolmntozoa. Dots a n d circles represent thc valucs 12 to
15 weeks a n d 20 weeks after t h e first of two a n d four
irradiations, respectively.
duct and the lumen in segment IV were decreased, and neither luminal PAS-positive
material nor intraepithelial PAS-positive inclusions were seen on the ligated side 4 weeks
after the operation (Fig. 2c). The contralatera1 nonligated epididymis showed the PASpositive inclusions.
DISCUSSION
Radiation induced disappearance of spermatozoa from the epididymal duct after a
latency period. This may be related to the
sensitivity of spermatogenic cells to irradiation and the duration of the spermatogenetic
cycle in the seminiferous epithelium. Oakberg (1956) observed the mouse testis histologically after a 100-R radiation dose and
estimated the duration of spermatogenesis
from type A spermatogonia to spermatozoa
is 34.5 days. The present study showed that
spermatozoa in the seminiferous epithelium
disappeared 5 weeks after 800-R irradiation.
Thus, the interval from irradiation to
aspermia in the testis corresponds to the duration of spermatogenesis and indicates that
only the earliest cell types in the spermatogenic series were destroyed when irradiated
with the larger 800-R dose, as is the case
with the 100 R used by Oakberg (1956). However, the destructive effect on the young spermatogenic cells seems to be exaggerated by
multiple irradiations, because spermatozoa
disappeared earlier and showed a more retarded recovery after four irradiations than
after one irradiation.
In contrast to the radiosensitivity of spermatogenic cells, the interstitial cells and
Sertoli cells showed morphologically no detectable damage. It is also indicated that the
interstitial cell androgen production and Sertoli cell function were almost unaffected by
radiation. Because the segmental differentiation of the epididymal epithelial cells is dependent upon androgen, the segmental
differences of the epididymal duct disappear
after gonadectomy (Cavazos, 1958; Alexander, 1972), but after irradiation the segmental differences of the epididymal duct are
maintained. In addition, because differentiation of the initial segment also requires the
intraluminal androgen-binding protein produced by Sertoli cells, the initial segment
shows decrease of epithelial height-dedifferentiation-after efferent duct ligation (Fawcett and Hoffer, 19791, but shows no such
changes after irradiation. Thus, the present
study suggests that flow of the testicular fluid
into the epididymal epithelial cells retain
EPIDIDYMAL PAS-POSITIVE MATERIAL AND SPERMATOZOA
their functions under the influence of androgen after irradiation.
It has been suggested that the biochemical
response of the epididymal epithelial cells is
affected by irradiation (Ito, 1966; Gupta and
Bawa, 1971). Morphologically, however, despite damage of spermatogenic cells after irradiation, the epididymal duct did not show
any changes during the initial postirradiation latency period. In addition, because
changes in segment IV after the latency period were prevented by ligation between segments 111 and IV and similar changes could
be induced by ligation of the efferent duct
without irradiation (Abe et al., 1982a), the
changes in segment IV after radiation are
definitely considered due to no radiation
damage and to be concerned with a functional activity of this segment. Thus, it is
considered that irradiation induces a temporary disappearance of spermatozoa from the
epididymal duct, leaving the ductal epithelium functionally active and the flow of the
testicular fluid uninterrupted.
During the period of ductal aspermia, the
intraepithelial PAS-positive inclusions, such
as shown in the ligation experiment (Abe et
al., 1982a1, depended upon the release of the
luminal material from the segment 11. In
segment 11, the cytoplasm of the principal
cells stains with PAS, the apical portions of
the cells are strongly positive, and the stereocilia are embedded within strongly PASpositive material; and the lumen from segment I1 distally contains PAS-positive material together with spermatozoa (Takano,
1980). These findings suggest that the luminal PAS-positive material is secreted by
the principal cells in segment 11. The ligation
between segments I11 and IV obstructs the
flow of the luminal material to segment IV
and prevents the appearance of the PAS-positive inclusions (Abe et al., 1982a). Thus, the
PAS-positive inclusions in the principal cells
in segment IV appear when the duct in segment IV received the PAS-positive material
from segment I1 but no spermatozoa. Duration of the appearance of the PAS-positive
inclusions clearly depends on the duration of
ductal aspermia.
Recent studies in the rat, rabbit, and hamster demonstrated that the principal cells
in the epididymal duct distal to the initial
segment secrete specific epididymal glycoproteins or proteins which bind to the spermatozoa membrane (Lea et al., 1978; Olson
and Hamilton, 1978; Faye et al., 1980; Moore,
1980). In the mouse, Vernon et al. (1982),
23
using a monoclonal antibody, demonstrated
immunohistologically that a n antigen binding to sperm tails is secreted from the principal cells in a short segment of the distal
epididymal head. Such substances have been
hypothesized to be responsible for rendering
spermatozoa capable of fertilization (see reviews by Bedford, 1975, and Hamilton, 1975).
Because of the stainability and the site of
secretion, the luminal PAS-positive material
in the mouse may contain such a specific
epididymal glycoprotein. The epididymal
principal cells are known to be absorptive as
well as secretory (Moniem and Glover, 1972;
Moore and Bedford, 1979; Fain-Maurel et al.,
1981). It is therefore likely that the specific
epididymal glycoprotein is reabsorbed by the
principal cells in segment IV and deposited
as intracellular inclusions if not bound to
spermatozoa.
The present study showed that the inclusions appeared 1 week after the disappearance of spermatozoa in the epididymal duct.
In the previous study, we observed that PASpositive inclusions develop in segment IV beginning 1 week after efferent duct ligation
(Abe et al., 1982a). Vernon et al. (1982) also
examined the sperm tail antigen of the mouse
epididymal duct 10 days after efferent duct
ligation, but found no labeling of the luminal
material and no intracellular deposits in the
principal cells corresponding to segment IV.
Thus, the site of the antigen secretion seems
to correspond to segment 111, but the sperm
tail antigen demonstrated by Vernon et al.,
(1982)may be different from the PAS-positive
material demonstrated by our group.
The PAS-positive inclusions disappeared
soon after the reappearance of spermatozoa.
This suggests that the principal cells are capable of digesting the accumulated PAS-positive material. The principal cells contain
dense bodies and multivesicular bodies which
contain acid-phosphatase and are lysosomal
in character (Friend, 1969; Moniem and
Glover, 1972). We have observed that the
principal cells in segment IV in normal mice
have specific dense bodies and peculiar
inclusions, both of which are considered
to represent various stages of digestion of
accumulated materials (Abe et al., 1983b).
The particular inclusions which contain a
bundle of tubules occur only in segment IV,
and they may be concerned with a specific
function such as absorption of the glycoprotein in this segment. Thus, even under normal conditions, the principal cells may absorb
and digest the glycoprotein which is un-
24
K. ABE, H . TAKANO, AND T. I T 0
bound to spermatozoa. Although the sperm
coating antigen secreted from the proximal
epididymis has been shown only on the stereocilia of the principal cells without cytoplasmic labeling in the distal epididymis,
absorption of the antigen has been also suggested from the distinctive cytoplasmic labeling of the clear cells in the distal epididymis
(Vernon et al., 1982).
As demonstrated above, the principal cells
in segment IV can absorb the luminal PASpositive material coming from segment 11.
The intracellular PAS-positive inclusions are
formed by the absorbed luminal material and
depend directly on the absence of spermatozoa from the lumen. The luminal PAS-positive material could be bound to or used by
spermatozoa in the lumen. In this study, temporary depletion of spermatozoa in the epididymal duct induced by testicular irradiation
clearly demonstrated the relationship between spermatozoa, PAS-positive material
secreted in segment 11, and the function of
the principal cells in segment IV. The luminal PAS-positive material also may play a
role in the differentiation of the principal
cells in segment IV, as shown previously (Abe
et al., 1982b).
ACKNOWLEDGMENTS
We wish to thank Professor Goro Irie, Department of Radiology, Hokkaido University
School of Medicine, for advice on the radiation method and Libby Cone, S.B., University of Massachusetts Medical School, for
correction of English in the manuscript. The
irradiation was performed at Central Institute of Radioisotope Science, Hokkaido University.
This study was supported by a grant from
the Ministry of Education, 1981 (5670002).
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Ahe, K., H. Takano, and T. Ito (1983a)Ultrastructure of
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Abe, K., H. Takano, a n d T. Ito (198313)Tubule-containing
inclusions in the epithelial cells of the mouse epididyma1 duct. Arch. Histol. Jpn., 46: (in press).
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Bedford, J. M. (1975) Maturation, transport and fate of
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Cavazos, L. F. (1958) Effects of testosterone propionate
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Fain-Maurel, M.A., J. P. Dadoune, and M. F. Alfonsi
(1981) High-resolution autoradiography of newly
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Fawcett, D. W., and A. P. Hoffer (1979) Failure of exogenous androgen to prevent regression of the initial segments of the r a t epididymis after efferent duct ligation
or orchidectomy Biol. Reprod., 20:162-181.
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Gupta, G. S., and S. R. Bawa (1971) Phosphatases in
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Hamilton, D. W. (1975) Structure and function of the
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5 , D. W. Hamilton and R. 0. Greep, eds. American
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Ito, H. (1966) Histochemical observations of oxidative
enzymes in irradiated testis and epididymis. Rad. Res.,
28.266-277.
Jones, H . E., and J. R. Cunningham (1971) The Physics
of Radiology, 3rd ed., Charles C. Thomas, Springfield.
Lea, 0. A,, P. Petrusz, and F. S. French (1978) Purification and localization of acidic epididymal glycoprotein
(AEG): A sperm coating protein secreted by the r a t
epididymis. Int. J . Androl., ISuppl.], 2t592-607.
Moniem, K. A., and T. U. Glover (1972) Comparative
histochemical localization of lysosomal enzymes in
mammalian epididymides. J. Anat., 111:437-452.
Moore, H. D. M., and J . M. Bedford (1979) The differential absorptive activity of epithelial cells of the r a t
epididymis before and after castration. Anat. Rec.,
193t313-328.
Moore, H . D. M. (19801 Localization of specific glycoproteins secreted by the rabbit and hamster epididymis.
Biol. Reprod., 22705-718.
Oakberg, E. F. (1956)Duration of spermatogenesis in the
mouse and timing of stages o f t h e cycle of the seminiferous epithelium. Am. J. Anat., 99:507-516.
Olson, G. E., and D. W. Hamilton (1978) Characterization of the surface glycoproteins of r a t spermatozoa.
B i d . Reprod., 19126-35.
Takano H. (1980) Qualitative and quantitative histology
and histogenesis of t h e mouse epididymis, with special
emphasis on t h e regional difference. Acta Anat. Nippon, 551573-587, (In Japanese with English Abstract.)
Vernon, R. B., C. H. Muller, J. C. Herr, F. A. Feucher,
and E. M. Eddy (1982)Epididymal secretion of a mouse
sperm surface component recognized by monoclonal
antibody. Biol. Reprod., 26:523-535.
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