A REVIEW OF ANATOMY AND PHYSIOLOGY
OF THE STALLION
R. P. Amann, Ph.D.
INTRODUCTION
An understanding of the structure and function of the
reproductive organs is desirable when developing a
management strategy to maximize reproductive efficiency
or for understanding results of research. Knowledge of
anatomical terms and of the location, shape and relative
size of each reproductive organ is essential when evaluating
a stallion. Malfunction of one or more reproductive organs
is not rare. An understanding of the process of
spermatogenesis, the importance of epididymal function
and secretions from the accessory glands in influencing
semen quality, and the nature of the ejaculatory process is
essential for developing a prognosis and a plan for possible
treatment of any stallion suspected of having a
reproductive problem.
The male reproductive organs consist of the two testes,
each suspended by a spermatic cord, the two epididymides,
the two deferent ducts each with an ampulla, the paired
vesicular glands, the prostate gland, the paired
bulbourethral glands, and the penis (Figure I)." The
vesicular glands, prostate gland, and bulbourethral glands
often are termed the accessory sex glands. The tract is
supported within the pelvic cavity by the hammock-like
genital fold and externally by the scrotum and prepuce.
Several muscles, including the urethralis, bulbospongiosus,
ischiocavernosus, and retractor penis muscles are of
importance.
THE SCROTUM AND ITS CONTENTS
The Scrotum
The scrotum encloses the testes as an outpouching of the
skin in the inguinal region. The scrotum is globular in form
with a distinct longitudinal groove or raphe where the two
scrotal sacs were fused. One of the two scrotal sacs lies on
either side of the penis (Figure 2). If one testis is larger than
the other, the scrotum may appear asymmetrical.
The scrotum consists of four layers. The outer-most layer
Author's address: Animal Reproduction laboratory. Colorado State
University. Fort Collins. 80523. Portions of this review are included in a
bulletin "Management of the Stallion for Maximum Reproductive
Efficiency". The illustrative skill of John Daugherty is gratefully
acknowledged.
aThe general anatomical description is based on the author's observations
and texrs by Banks.to Gcrty.z« Krolling and Grau," Montane et "I. ll and
Nickel ct "I...• References for newer anatomical observations and for
specific physiological principals are supplied in the text.
May/June 1981
is the skin, which contains sebacous glands and an unusually
large number of sweat glands. Underlying the skin and
associated connective tissue is the tunica dartos muscle. The
tunica dartos is a layer of smooth muscle that contains some
fibroelastic tissue. This muscle layer is closely adherent to
the skin and the scrotal septum so that it forms the outer-
most component of each of the two scrotal sacs. .
The third layer is the scrotal fascia. This layer consists of
loose connective tissue that lies between the tunica dartos
and the vaginal tunic. The looseness of the scrotal fascia
allows the testis great mobility for vertical or horizontal
movement within the scrotal sac. Normally, the scrotal
fascia prevents 180 rotation of the testis within the scrotal
0
sac, although in certain stallions such rotation docs OCCUr.~9
If horizontal rotation of the testis occurs, permanent
correction is unlikely unless surgery is used to form
connective tissue bridges to assist the stretched or
weakened scrotal fascia in preventing rotation. During
castration, the scrotal fascia can be easily torn with the
fingers so that the underlying vaginal tunic and testis can be
freed from the scrotal sac.
The inner-most layer of the scrotum is the parietal
vaginal tunic. The parietal vaginal tunic (also called the
common vaginal tunic) is a membranous sac which extends
from the abdominal cavity, through the inguinal canal, to
the bottom of the scrotum. The outer aspect of the parietal
vaginal tunic is only lightly fused to the scrotal fascia. The
vaginal tunic forms a covering for the spermatic cord, testis
and epididymis. The parietal vaginal tunic is formed by an
evagination or ourpouching of the parietal peritoneiurn of
the abdominal cavity.
In a normal stallion, the parietal vaginal tunic is not
fused to the inner or visceral vaginal tunic that directly
covers the testis or epididymis. Rather, a vaginal cavity is
present between the two layers of the vaginal tunic (Figure
2). The space within the vaginal cavity contains a watery,
serous fluid which serves as a lubricant and facilitates
movement of the testis, covered by the visceral vaginal
tunic, within the sac of the parietal vaginal tunic. In some
older stallions, adhesions may develop between the parietal
and visceral layers of the vaginal tunic. Such adhesions
impede mobility of the testis and may reduce the
effectiveness of temperature control mechanisms described
below.
The visceral layer of the vaginal tunic directly covers the
testis and epididymis. This layer is only a few cells thick and
is fused with the tunica albuginea (the outer covering of the
testis) from which it cannot be separated. The visceral
83
 Deferent duct
Rectum
Genital fold
Urinary
bladder
Testicular
artery and
Internal inguinal ring--T:~m:~~~~~~~
within
canal
Glans penis
Prepuce
Figure I. Drawing of the reprodecrive tract of the stall ion as seen in a left later al dissection, From Pickert
vaginal tunic covered the testis before its descent from the
abdominal cavity into the scrotum. Thus, it was carried into
the outpouching of the parietal layer of the vaginal tunic
(vaginal process) during the descent of th e testis from the
abdominal cavity into the scrotum. The visceral layer of the
vaginal tunic actually is part of the test is ' rather than the
scrotum.
The Testis
The testis (plural =testes) is the male gon ad and is the
site of production for both sperrnarezoa and the
predominant male sex hormone, testosterone. The testes
ate ovoid or walnut shaped structures, slightly compressed
from side to side and with their long axis almost horizontal.
When the testis is retracted, however, the long axis
becomes almost vert ical. The testes of a post-pubertal
stallion have a considerable range in size, but they might
average 11 to 12 ern in length by 5 to 7 ern in width and
weigh about 225g per testis. The testis is reta ined within
the parietal vaginal tunic by the mesorchium and by the
proper ligament of the testis, that attaches the testis to the
84
Ampulla
gland
gland
Pelvic urethra
Bulbourethral
gland
Retractor penis
muscle
'H:;;;'~r-- Penile urethra
Bulbospongiosus
muscle
spongiosum
Corpus cavcrnosum penis
Epididymis
ellll. 6 1
tail of the epididymis which in turn is attached to the
parietal vaginal tunic by a ligament.
If the testis is exposed, as during open castration, the
visceral vaginal tunic is the outer-most layer. The visceral
vaginal tunic is inseparably fused to the tunica albuginea
(Figures 2 and 3). The tunica albuginea is a thick, dense,
connective tissue capsule of collagenous, elastic and smooth
muscle fibers from which septula extend to divide the testis
into lobules . The inner surface of the tunica albuginea
conta ins the tortuous tes ticular artery, as well as numerous
smaller arteries and veins (Figure 3).
The parenchyma, or noncapsular part of the testis, is
fairly firm and orange-brown in color. The parenchyma is
lightly pigmented in a young , postpubertal stallion,
moderately pigmented in a 4-5 year old stallion, and darkly
pigmented in an aged srall ion .t? A cross section through the
parenchyma (Figure 3) reveals the conspicuous, central
vein that runs cranially near the dorsal surface. The
testicular parenchyma consists of seminiferous tubules and
interstitial tissues .(Figure 4). Leydig cells are the major
component of the interstitial tissue " and they produce
EQUINE VETERINARY SCIENCE
hormones. The ratio of Leydig cells ro seminiferous tubules Corpus cavernosum penis
Penis
increases from 1:12 [Q 1:4 as a horse grows older (Table D.
The total length of seminiferous tubules in the testis also
increases by one-third. "

Table I. Age-Related Differcnces in Testicular Composition a

of epididymis
2-3 yr 4-5 yr 13-20 yr
duct
Testicular weIght ( g) Central vein
Parenchyma 105 J. 146 c 184d
Tunica albuginea 12 h 15 c 29°
Parenchyma I
composition (%)
Leyd i g ce I I s 6b 12' 18 d
Other Interstitial dartos
c
tissue nb 16 10 d
Seminiferous
tubules 72 72 72
Seminiferous Testicular
parenchyma Visceral vaginal tunic
tubule
Diameter IIJm) 219 b 230' 242'
Length/testis (m) 2040 h 2390 b c 2790' Figure 2. Drawing of the penis. scrotum and testes of the stallion as seen
Dally sperm in a vert ical section. From Picken et al.G1
production
(l09/testis) 1.3 b 2.7 c 3.2 c Both ends of a convoluted seminiferous tubule continue
l9 as transitional zones leading to straight tubules ," The
a Modified from Johnson and Nca\'es.
b ,e,d Means in th e same row with diffcrent superscripts differ transitional zone of a seminiferous tubule is tapered and
(1'< 0.05). lined by cells similar in appearance to Sertoli cells, if they
are nor actually Serroli cells. The straight tubule is a
The seminiferous tubules are arch-shaped (Figure 5) and continuation from the transitional zone and leads to the
are surrounded by a basement membrane which is a rete testis (Figure 5); it serves as the first component of the
laminar layer of collagenous and clastic fibers interspersed passageway through which sperm pass from the germinal
with myoid (smooth muscle-like) cells. A seminiferous epithelium to the epididymis. The narrow straight tubules
tubule consists of three zones. The major portion of the vary greatly in length and arc lined by a cuboidal
seminiferous tubule is highly coiled and is the site of sperm epithelium.
production. This portion is termed the convoluted Straight tubules con verge and an astomose in an area
seminiferous tubule. A convoluted seminiferous tubule is around the central vein in the cranial two-thirds of the
lined by the germinal epithelium which contains Serroli testis in an area termed the rete testis (Figures 3 and 5 ).6
cells (somatic cells) and several types of germ cells. 66Serrol i This area is analogous to the rete testis of the bull or
cells line the basement membrane and extend toward the human, although a distinct mediastinum is not evident. The
lumen in a radial pattern ro occupy 15 to 25% of the anastomosing tubules in the rete testis area (Figure 6) are
germinal epithelium in an adult stallion. They arc tall cells lined by a cuboidal epithelium identical to that of the
with a complex basal nucleus. Although it is conventionally straight tubules. The tubules of the intra testicular rete
accepted that Scrroli cells do not divide after puberry.P:" testis penetrate the tunic albuginea at the dorso-cranial
recent research " has shown that proliferation of Sertoli pole of the testis and continue as an extraresticular rete."
cells may occur as a stall ion approaches the breeding Additional straight tubules join the anastomosing tubules
season . A detailed consideration of the germinal epithelium of the rete testis just prior to their penetration of the tunica
and the process of spermatogenesis is in preparation. albuginea. Eventually, each rete tubule fuses with one of the
13 to 15 efferent ducts that lead to the epididymal duct . n
The testis has two primary functions: production of
Table 2. Development of thc Stallion Testis a spermatozoa by the germinal epithelium within the
Weight (g) of seminferous tubule and production of testosteroneband
Age (mo) Left Testis Right Testis Total other steroid hormones by the Leydig cells of the interstitial
<12 29 24 53
13- 16 54 47 101
17- 20 67 61 128 bTrivial names used for steroid hormones are test ost erone =-l-androsrcne-
21- 24 106 103 209 17 -01- 3·one; androstenedione = Ij-androstene- 3.17-dio ne;
15- 36 113 119 232 androstencdiol = 5-androstcne-3 , 17 -diol; dihydrorestosreronc = 5 -
36-120 156 161 317
>120 203 207 410 androstan-I? -ul-3·one; 3 -a nd rosranediol = 5 -androsran-S ,17 -
diol ; 3 -androsranediol = 5 -andros rane-y , 17 -d iol; progest erone =
-l- p rcg ncnc -y, 20·dionc; estrone = 1,3,5( IO)·eslratrienc-3-ol -17·onc;
a From Pickert and Voss.(l) estradiol = 1.3.5( lO)estratriene ·3.17 -diol,

May/June 1981 85

Figurc 3. Cross sections through a fixed testis. The upper view (1.2X)
shows the testicular artery (A), central vein (circle), tunica albuginea with
overlaying visceral vaginal tunic (TAl, blood vessels within rhc tunica
albuginea, and senrela radiating from the tunica albuginea to divide the
testis into lobules. The lower view (lOX) shows the distribution of
convoluted scminiferous tubules and straight scminifcrous tubules (black
arrows) leading to the rere tubules (black arrows) that anastomose around
the central vein (CV). The Iobular organization is evidcnr. From Amann ct
,".6
86
tissue. During the first 15-20 months after birth, the testes
are not producing sperm and the animal is incapable of
reproduction. Starting near 18 months.]H.H.(,() rapid growth
and development of the testes are initiated (Table 2) and
the testes gradually begin to produce sperm.":"
Concurrently, there are changes in the production of steroid
hormones by the Leydig cells. This prepubertal
development is culminated in puberty when a stallion
produces sperm and would be fertile if allowed to cover a
mare of appropriate size. After puberty, the quantity of
sperm produced and the quality of the sperm both increase
for some time. With bulls, fertility of semen increases for
about 6 months after puberty.':"
In the adult stallion, billions of sperm are produced daily
in the convoluted seminiferous tubulesY·16.2J Although
about 70,000 sperm are produced each and every second
during the breeding season by the two testes of an adult
stallion, production of each individual sperm requires about
55 daysy"·67 When sperm are liberated from the germinal
epithelium (Figure 7), fluid (presumably produced by the
Serroli cells) carries the sperm from the convoluted
seminiferous tubules into the straight seminiferous tubules
and the tubules of the rete testis (Figure 6) where additional
fluid may be added. The suspension of sperm is moved
rapidly through the efferent ducts into the proximal
epididymis.
The number of sperm produced by a normal stallion is a
function of testicular size. For living stallions, testicular size
can be estimated accurately by measuring scrotal width."
Thompson et al. 69 found that measurements of total scrotal
width were highly accurate, especially if the average value
for measurements made by two individuals was considered.
They also found that scrotal width was correlated (P < 0.01)
with daily sperm production (r=0.75). When evaluating the
normalcy of testicular development in a stallion, his age
must be considered. Table 3 provides norms for testicular
development. The ranges for ±1 standard deviation and±2
standard deviations represent about 67% and 95% of the
population. Thus, a 5-year-old stallion with a scrotal width
of 93 mm would be considered normal, bur a 7-year-old
stallion with the same scrotal width would be considered
subnormal and probably should not be used for breeding.
The number of sperm produced by a stallion also is
influenced by age (Table 4).7.39 The increase with age is
largely a matter of the seminiferous tubules becoming
longer and the testes larger.v The daily sperm production
per gram of testicular parencyma is reasonably similar,
during the breeding season, for stallions 4 years of age or
older." Since the daily sperm production rate is one factor
influencing the number of sperm available for ejaculation,
more sperm usually will be ejaculated by an older stallion
(possibly 5 to 16 years old) than a 2- to 4-year.old stallion.
This fact should be considered when evaluating or
managing a stallion.
The Leydig cells, within the interstitial tissue, produce a
variety of steroid hormones (Figure 8). Although
testosterone" is the major androgen secreted by the stallion
testis, the testosterone precursors androstenedione and
androstencdiol and the metabolites dihydrotestosrerone,
EaUINE VETERINARY SCIENCE
Table 3. Norms for Scrotal Width 3
Age (yr) ±1 SO
t-lean
J steroid content of jugular vein blood.'
b 88-103
2-3 96
4-6 100 b 93-107
27 10g e 102-117
a Modified from Thompson et af. 69
be ~fcans with different supcrscriprs differ (I' < 0.05).
3X·androstandiol, 3B-androstandiol and estradiol also are
secreted.':":" The exact site(s) of production of estradiol or
other estrogens is unknown.
The Leydig cells constantly produce a basal level of
testosterone. Periodically, however, at fairly random
intervals through the day and night, the Leydig cells are
stimulated to increase their production of testosterone and
episodic (2- to 4-fold) elevations of testosterone in
peripheral blood result."?' There appear to be 4 to 8 of
these episodic or transitory bursts of testosterone
production each day. Consequently, if a single blood sample
from a stallion is analyzed for its testosterone content,
Surprisingly, concentrations of steroids in testicular vein
±2 SO blood accounted for less than 16% of the variability in the
81-111
Injection of human chorionic gonadotrophin (hCG) is
85-115
95-124 followed within 1 or 2 hr by an elevation of the 17 -
hydroxy-androgen content of jugular vein blood (Figure
13) and testicular vein blood (Figure 11).4 Peak production
of testosterone induced by hCG appeared to occur only over
a short interval, but the effect of hCG treatment on
testosterone production 3 to 48 hr later has not been
studied. Surprisingly, the concentrations of total estrogen
in testicular vein or jugular vein blood were similar
throughout the 5-hour sampling period of this study.'
Obviously, the concentration of total estrogen in testicular
venous blood was independent of testosterone production

Table 4. Effects of Age on Daily Sperm Production"

Testes Oai Iy sperm production
parenchyma 9
Age (yr) wt (g) 106/ g testis 10 /stall ion
2-4 265 b 15.7 4.2 b
5-9 371 17.3 6.3
10-16 416 16.0 6.6
3 From Amann ct al,'
b Means for 2- 10 4-year-olds differed from those of 5- ro 9- and 10- ro 16-
year-old stallions (I' < 0.05); differences between 5- 109- and 10- ro 16-
year-olds were not significant.

occasionally an unusually high value may be obtained that is

not typical of peripheral concentrations in blood of that -'~::'+'*-l,- Ly m p hat ic
horse. v ess el
The relationship among concentrations of androgen and
estrogens in the blood leaving the testis through the
testicular vein or obtained from the jugular vein have been
compared.' As shown in Figure 9, testosterone accounted >;:.,.-,.;.:;.-.;~~..:..,..~:.-n- Fi bra b la s t
for over 70% of the total 1713·hydroxy-androgen found in
testicular vein blood. The concentration of total 1713-
hydroxy-androgen in testicular vein blood averaged 62
ng/ml (Table 5) or about 45 !imes the concentration of 1.3
sig] ml in blood taken concurrently from the jugular vein of
these anesthetized stallions. Surprisingly, the correlation
between the concentrations of 17B-hydroxy-androgen in
testicular vein blood and in jugular vein blood was very low
(r=0.32; P>0.05). The concentration of total 17B-hydroxy-
androgen in testicular vein blood was similar for stallions 2 Leyd ig ce lls
to 3,4 to 9 or 11 to 14 years old,' although the total mass of
Leydig cells is 3 to 5 times greater in old stallions than in 2
to 3 year olds (Table I). \? Testicular vein blood contains
extraordinary concentrations ot total estrogen (estrone Se mind e ro s
plus estradiolj.! The concentration of total estrogen in
Figure 4. A scanning electron micrograph (upper; X200) and drawing of
testicular vein blood (Table 5) was influenced by age of the stallion testis (lower) showing the relationship among the blood vessels,
stallion (P<0.05) and averaged 20,700, 36,000 and 17,700 lymphatic vessels and Leydig cells of the intersririal tissue and the
pg/rnl for stallions aged 2 to 3,4 to 9 or 11 to 14 years. seminiferous tubules. Upper figure from Johnson ct uf. 11

May/June 1981 87

figurc 5. Drawing showing the location of the slraighl tubules and the
rete resris and also the vascular pattern in the stallion testis, The straight
tubules converge in a group of inrcrconnccung rete tubules that penetrate
the tunica albuginea. The testicular artery becomes highly coiled in the
pempiniform plexus. After emerging from the pampiniform plexus, the
testicular artery passes along the dorsal aspect of the testis to the caudal
pole where it starts to branch to vascularize the parenchyma. Venous
drainage of rhc parenchyma is via the central vein and superficial testicular
veins . After leaving the testis, the veins form an anastomosing plexus of
veins.j crmcd the pampiniform plexus, that is in intimate contact with the
testicular artery, About 7-10 cm above the testis, the veins converge into
the testicular vein . From Pickert et ,,1. 61
88
as represented by the venous concentrations of total
androgens (r=0.03). Amann and Ganjarrr' concluded that
concentrations of total 1713·hydroxy-androgen (primarily
testosterone) and total estrogen in blood sampled from the
jugular vein bore little relationship to their concentrations
in blood leaving the testis a few minutes earlier.
Considering all sample pairs in their study (N=84), the
concentrations of total 1713·hydroxy-androgen and of total
estrogen in testicular venous blood accounted for only 50
and 10% of the variation in concentrations of these
hormones in jugular vein blood .
The actual production rate of testosterone was reported"
for one stallion as 2.2 ~g/min/testis . Using the average
blood flow of 26 mlJmin reported by Lidner,18 Amann and
Ganjarrr' estimated that the 20 stallions used in their study
secreted about 2.3 ~g of testosterone and 0.71 ug of total
estrogen per minute per testis. By extrapolating from data
for hCG-treated stallions, they speculated that the
concentration of testosterone in testicular vein blood might
approach 700 ng/ml during an episodic peak in production
rare.'
Figure 6. A scanning electron micrograph (X(,O) of the anastomosing
tubules wirhin the rete testis ncar the central vein (CVI. Rete tubules (R)
and smaller blood vessels (8) arc comm ingled in this area . From Amann et
.11.'
EaUINE VETERINARY SCIENCE
 A·· PREGNf:NOL.ONf: PROGESTE.RONE

~/o(1
c-o
HO~·OH

1711· HYDROXYPREGN~NDLOI'4E" 171l1-HYOROXYPROGESTERC,E

j
HO~ O~ -HO~
O(HYORO[PlANCROS1E:ROf,;E .t.··J...-.rOROST£~EorC~E tSTRONE

I
Ha~ o~ -HOroP
TESTOSTERO~E ESTRAOIOl

DIHYDROTESTOSTERONE

10M

Ha• .q5P
H

~.-ANCROSTA~-3•• 17/! DIOL

Figure 7. A scanning electron micrograph (X5,200) of stallion

spermatozoa just as they complete spermatogenesis and are about to be
released from the germinal epithelium into the lumen of a convoluted
seminiferous tubule. After their release, sperm are washed into the
straight seminiferous tubules by flowing fluid. From Johnson ct 011. 1 1
From the studies reported 'above, it is evident that
testicular vein blood, and presumably testicular tissue,
typically contain about a 45-fold higher concentration of
testosterone than peripheral blood and that during episodic
production of testosterone the concentrations of this
steroid within the testis may be further elevated,
Consequently, the seminiferous tubules are continuously
exposed to a high concentration of testosterone. This very
Figure S. The pathway for steroid biosymhesis in the sra llion testis.
Cholesterol is converted by one of two pathways to androstenedione or
androstenediol which are the two immediate precursors for testosterone.
Testosterone, in turn, can be converted to dihydrotcsrosterone or 3 -
androsrancdiol (androgens acrive in many tissues) or to estradiol
(generally considered 3S 3 female sex hormone although it also may be
important in the male). Note the structural similarity of many of these
hormones. Synthetic compounds, or analogues having a similar structure,
may affect a stallion by blocking or mimicking an action of a natural
hormone. From Picken et al.6 1
z 80
w
c;
0
II:
0
z

11
« 70
high concentration of testosterone around the seminiferous )-
x
tubules probably is essential for normal spermatogenesis as 0
;r
has been shown with other speciesYM.61 0 :: .1
)-
:I: 60
Descent of the Testes
A complete discussion of the embryology of the equine I
...J
reproductive system is beyond the scope of this review. «
~
However, an understanding of the process by which the 0
~
restes normally descend from the abdominal cavity into the 20
LL.
scrotum is desirable. In the normal fetus or colt, both testes 0
W
should descend into the scrotum between 30 days before l!l
«
birth and 10 days after birth. 14.27 Failure of normal testicular
z 10
~

descent is common in horses and is termed cryptorchidism. w

The gonad in a male fetus differentiates into a testis at
about day 40 of gestarion.!' The testis is suspended from the
ventral surface of the mesonephros by the cephalic
ligament of the testis, the mesorchium, and the caudal
gonadalligamcnt (Figure 12). The mesonephric duct,which
later gives rise to the epididymis and deferent duct, leads
caudally into the pelvic area. A narrow evagination from
the caudal end of the abdominal cavity forms around the
U
II:
C1.
0
Figure 9. The profile of androgens in stallion testicular vein blood. The
percentage contributions of individual steroids to the total 17S·hydroxy-
androgen pool for two resras are shown. 5·D = androsrenediol, T =
testosterone. DHT = dihydrorestosteronc, 3 -D = 3 -androsranediol, 3 -
D = 3 -androsranediol and U = unidentified steroids. From Picket ct 011. 6 1

May/June 1981 89
Table 5. Hormone Concentrations in Testicular Vein and Jugular
Vein Blood."
.....E ..
-E
<,
Testis Jugular 0 10 0 ... 0
61.5 1.3 c: c:
17a-hydroxy-androgen (ng/mll ..
Total estrogen (pg/ml) 25,400 58 30
z z
w w
aFrom Amann and Ganjarn.! (!)
0
o
a: 20 0
0
z 5
.. ....cr
en
<t w
...J 10 ...J
<t 250 <t
6 r t-
O 0
E
......
HCG ....
0 ., .. 0
-- l
0'
c:
I hr 2 hr
Z
1LI
4 HOU S AF E CG
(!)
oa:: Figure II. Differential effect of hCG on concentrations of 17/3.hydroxy
o androgen (left) and total estrogen (right) in testicular vein blood. From
z Amann and Ganjarn.!
< 2
...J
<!
b
r
vaginal process. From about day 70 to day 120 of gestation,
the testis and gubernaculum increase in size and the testis is
O~---.l"--_......r. _ _......._ _......._ _....&..-_ _
drawn closer to the inguinal ring.
By day 150, the junction between the gubernacular cord
:::: 60 and the developing tail of the epididymis is drawn to or just
E within the internal inguinal ring (Figurel S)." This results
......

--
0'
C. in a reflection of the deferent duct around the urethra
(Figure 13). The caudal gonadal ligament is quite short so
~ 40 .1 that the testis is in close proximity to the internal inguinal
~t
C)
o ring. The mesorchium (the double layer of peritonewn that
a:: previously attached the dorsal aspect of the testis to the
r
en abdominal wall) is bordered anteriorly by the cephalic
1LI
...J 20 gonadal ligament that connects with the peritoneum. The
~ gubernacular bulb increases in size, but the gubernacular
~ 0 cord remains short (Figure 13).The tip of the gubernaculum
-2 -I 0 +1 .2 is connected by fibrous threads to the base of the developing
HOURS FROM HCG INJECTION
scrotal folds.
Between day 150 and day 275, passage of the testis
Figure 10. Effect of hCG on concentrations of 17/3-h}'droxy-androgen through the inguinal canal is impossible." The equine fetal
(upper) and total estrogen (lower) in jugular vein blood. An injection of testis weighs about 20g at 150 days, 50g at 250 days and 30
saline (e) or 1500 IU of hCG ( ) was given 2 hr before castration. From
Pickett et al. 61
at 300 days." Thus, the equine testis at 250 days of
gestation 14 is more than twice the size of a typical, adult
human testis.' At 250 days of gestation, the caudal pole of
lateral and ventral borders of the gubernaculum around day the testis, closely attached to the tail of the epididymis by
43 of gestation. This evagination is termed the vaginal the caudal gonadal ligament, is about 3 em from the internal
process and progressively develops to form an internal inguinal ring. The tail of the epididymis is within the
inguinal ring through which the gubernacular cord is inguinal canal, while the testis remains within the
suspended. By day 65, the mesonephrous has degenerated peritoneal cavity.
so that the testis is suspended directly from the dorsal Entrance of the testis into the inguinal canal typically
abdominal wall. The gubernaculum extends from beyond begins between 270 and 300 days of gestation. 14 This occurs
the caudal end of the vaginal process, through the lumen of only after the vaginal process and internal inguinal ring
the process and the inguinal canal, to the connection have been enlarged by the combined mass of the
between the caudal gonadal ligament and the developing gubernaculum, tail of the epididymis, and caudal gonadal
tail of the epididydmis (Figures 12, 13 ). The gubernacular ligament. The internal inguinal ring is stretched by the
bulb is an increasingly large mass developing within the enlarging tail of the epididymis (Figure 14) until it is of
differentiating muscle layers of the pubic region. The sufficient size to allow entrance of the testis that is
future external cremaster muscle develops around the diminishing in size. Eventual expansion of the internal

90 EQUINE VETERINARY SCIENCE

 Cephalic ligament
Mesonephric of testis
duct
Gubernacular bulb
~ Dorsal body wall
~Mesorchium
~~~\~~~;;;;~;~;;o~:::g
A Tunica albuginea
of testis
Invaginated
peritoneal lining
\\:.3JI.---I- Vaginal processes
B B
Figure 12. Drawing of a horse Ictus 31 75 days of gestation, The resris is
suspended wirhin the abdominal cavity by a th in double-layered band of
fused lis sue termed the mesorchium. The caudal gonadal ligamcnr,
extending from the testis 10 its point of fusion with the mesonephric ducr
(future epididymis and deferent duct), is conrinuous with the gubernacular
cord that runs 10 the gubernacular bulb . The vaginal process (future
parietal vaginal tunic) encloses about one-third of the gubernacular bulb.
From Pickell et al.6 1
inguinal ring allows passage of the testis through this
upper ring and into the inguinal canal. Pressure from the
peritoneal fluid, and possibly the abdominal viscera, forces
the testis down through the inguinal canal. During passage
through the inguinal canal, the testis is stretched into a
narrow cylinder rather than its characteristic ovoid shape.
Mav/June 1981
Epididymal Mesorchium Cephalic ligament
duct
Deferent duct of testis
~~~~~
Vaginal processes
A
Mesorchium
Figure 13. Drawing of a horse fetus at 175 days of gestation. The vaginal
process extends almost 10 rhe scrotum and has virtually enclosed the
gubernacular bulb. A thin band of fus ion, or double-layered mesorchium, is
formed between the gubernaculum and the vaginal process and is
conrinuous with the mesorchium supporting the testis. The epididymis
and deferent ducr have formed from the mesonephric duct and the future
tail of the epididymis will form where the duct is sharply reflected, From
Pickell et al.61
The fine fibrous strands connecting the tip of the
gubernacular bulb to the developing scrotum may provide
directional guidance for the vaginal process, but there is no
evidence that tension pulls the gubernaculum into the
developing scrotum."
Bergin et al," reported that the earliest complete descent
91

Caudal gonadal
ligament
A (c) insufficient growth of the gubernaculum and tail of the
Parietal layer of
vaginal tunic
Mesorchium
Deferent duct
surrounded by
mesoductus deferens
r Vaginal cavity
Blood vessels +
nerve surrounded
B by visceral layer
of vaginal tunic
Parietal layer of
vaginal tunic
Figure 14. Drawing of a horse fetus near term. The testis has passed
through the inguinal canal, but is not fully within. the scrotum. The
of both testes was at 315 days of gestation; about 25 days
before parturition. In 32 fetuses between 9 months of
gestation and birth, they found that descent of the right
testis was further advanced than the left in 78% of the
fetuses, while the left testis was more advanced in only 3%.
Of 12 fetuses collected at term, 42% had completely
descended testes, 25% had both testes within the inguinal
canals and equally distant between the internal and external
92
inguinal rings, 17 % had one testis in the scrotum and one
in the inguinal canal, and 17% had both testes within the
abdominal cavity. Five of nine colts less than one week old
had complete, bilateral descent of the testes into the
scrotum.
In contrast to descriptions in texts of veterinary
anatomy, Bergin et al. ll concluded that the vaginal process
originates as an evagination of the peritoneum in early
rather than in mid gestation. Furthermore, the testis passes
through the inguinal canal rather than retroperitoneally to
the canal. Consequently, the vaginal process directly
becomes the parietal vaginal tunic following entrance of
the testis in the scrotum. Furthermore, Bergin et al."
contend that the testicular mesorchium of the adult
represents the fold of tissue already suspending the testis
from the dorsal abdominal cavity at day 55 of gestation.
Movement of the testis into the vaginal process simply
pulls the mesorchium into the sack-like vaginal process. As
mentioned in the preceding description of the vaginal
tunic, the visceral layer of the vaginal tunic simply is that
portion of the visceral epithelium that covered the tunica
albuginea of the testis prior to the initiation of testicular
descent.
As reviewed by Bergin at al.," failure of the testes to
descend has been attributed to abnormalities of the testis,
development of adhesions between the testis and adjacent
structures, or an abnormal outpouching of the vaginal
process. Bergin et al.,11 discount these factors as causes of
cryptorchidism and suggest that the most obvious reasons
for the testis to remain in the abdominal cavity include (a)
stretching of the gubernacular cord, (b) insufficient
abdominal pressure to properly expand the vaginal process,
epididymis so that they are unable to expand the inguinal
ring sufficiently to allow entrance of the testis, and (d)
displacement of the testis to a position where the pressure
of the viscera prevents gubernacular tension from pulling
the testis into the vaginal process.
The truly cryptorchid testis, one retained within the
abdominal cavity, apparently occurs when the testis fails to
enter the inguinal canal before closure of the internal
inguinal ring during the first 2 weeks after birth. The
preponderence of failure of the left testis to descend" might
result from the relatively slow rate of descent of the left
epididymis and testis.
Diagnosis of cryptorchidism should include careful
external palpation of the scrotum and of the external
inguinal ring as well as rectal palpation of the internal
inguinal ring and pelvic area. During the first several weeks
after birth, the gubernaculum may be quite large; it should
not be confused with a testis. At birth the weight of each
testis is 5 to 109 and testicular size does not change through
10 months of age.Y'There is a slight growth between II and
16 months of age and rapid development of the testes
generally starts around 18 months of age. It is important to
recognize, however, that there are great differences in the
age when rapid growth ofthe testes begins. ' R.H .m Diagnosis
of cryptorchidism might be aided by analyses of blood
concentration of testosterone and esrradiol.»
EaUINE VETERINARY SCIENCE
 From a structural point of view, the epididymis consists
M esorchi m of three compartments (Figure 17): intertubular
connective tissue, epithelium, and luminal contents. From a
functional point of view, the epididymis has three
segments." The efferent ducts plus the initial segment of
the head are involved in the resorption of fluid and solids
Duc tu s from the ductal lumen and probably secrete several
de fere ns compounds. The middle segment is comprised of the major
portions of the head and body of the epididymis and is
V isc eral involved in sperm maturation. The terminal segment is
vag i na l comprised of the tail of the epididymis and proximal
t ic Test icu lar artery. vei ns. deferent duct and is involved in the storage of fertile sperm.
an d n er ves Most aspects of epididymal function have not been
studied in the stallion. However, based on data for other
Figure 15. Drawing showing a cross section through the right spermatic
cord above the level of the parnpiniforrn plexus. From Pickett et ,,1.
61 species.Y" it seems certain that most of the fluid, protein
and other material that enter from the testis are reabsorbed
in the proximal head; spermatozoa, however, are not
The Epididymis resorbed.v'? There is an obvious increase in the
The epididymis (plural = epididymides) is divided concentration of sperm as one views successively more
anatomically into three parts: the head or caput, the body or distal cross sections through the epididymal duct in the
corpus, and the tailor cauda. The head curves from the head of the epididyrnis.?'
dorso-medial aspect of the testis, around the spermatic cord
(Figures I and 5) and returns caudally around the lateral
aspect of the spermatic cord. The head is rather flat, has aJ-
shape and is closely attached co the cranial end of the testis.
The body of the epididymis appears grossly as a narrow,
cylindrical structure loosely attached to the dorsal surface of
the testis and it lies on the lateral aspect of the mesorchium
(Figure 2). The tail of the epididymis is large, bulbous, and
loosely attached to the caudal pole of the testis by the
proper ligament of the testis and to the parietal vaginal
tunic by the ligament of the tail of the epididymis.
The epididymis is covered by a dense, connective tissue
capsule termed the saccus epididymidis which is similar to
the tunica albuginea covering the testis. The superficial
aspect of the saccus epididymidis is covered by the same
thin, visceral vaginal tunic that covers the testis.
Careful examination of the epididymis reveals that the
proximal head actually contains the distal ends of the 13 to
15 highly coiled, efferent ducts that lead from the tubules of
the exrrarcsricular rete testis. 6,l S Within the head of the
epididymis, the efferent ducts fuse" into a single, duct
termed the epididymal duct. This single duct, possibly 45
meters long, is folded in pleats and continues in a tortuous
pattern through the head, body and tail of the epididimis
and is continuous with the deferent duct. The epididymal
duct has a smooth muscle coat that increases in thickness
distally.
The 13 to 15 efferent ducts have an epithelium (Figure
16) that contains columnar cells with true, motile cilia and
interspersed, non-ciliated columnar cells." The epithelirn
probably is resorptive in some areas but secretory in other
areasY·5ij The epididymal duct (Figures 17 to 19) is lined by
a pseudostratified, columnar epithelium (Figure 20) that
contains tall principle cells, small spherical basal cells,
wandering rnacrophages, and possibly other cell types. Figure 16. Cross section through an efferent duct (upper) showing the
Based on epithelial morphology," six regions can be ciliated and nonciliared cells in this pseudosrrarified epithelium (X800).
The luminal surface of an efferent duet and a spermatOzoon with a
distinguished in the stallion epididymis and probably the
proximal cytoplasmic droplet (normal for a spermatozoon from this
function of each region is different. region) also are shown (lower; (X.1200). From Johnson et "IY

May/June 1981 93
 Based on data for several species other than the horse, it within the epididymis, as well as testis size and daily sperm
is evident that sperm entering the epididymis are infertile, production, are influenced greatly by age of the stallion.7.l 9.(/)
whereas sperm recovered from the' tail of the epididymis The paired tails of the epididym is of a typical adult stallion
arc fertile. ~8 The process by which sperm develop the (5 to 16 years old) contain about 54 billion sperm (Table 7)
capacity for fertilization is termed sperm maturation. It has or 61 % of the total number found within the excurrent duct
been shown in several species" that sperm maturation is system."
dependent upon the availability of testosterone co the Transit of the sperm through the epididymal duct
epithelium of the initial or middle segments of the primarily is by continuous parastaltic contractions of the
epididymis (head and body) . Evidence for sperm smooth muscle in the wall of the duct within the head and
maturation includes the acquisition of fertilizing capacity, body of the epididymis. In the rail, however, the ducr
acquisition of progressive motility, changes in sperm normally is quiescent except when the smooth muscle is
morphology, changes in characteristics of the pl asma stimulated co contract. Consequently, the time required for
membrane of sperm, and changes in sperm metabolism. transit of sperm through the head and body of the
Simple retention of sperm within a given segment of the ep ididymis is not altered by ejaculation and averages about
epididymal duct, however, is insufficient co induce sperm 4.1 days in the stallionY ·2l.67
maturation." Secretions by the epididymal epithelium arc Since the rate of sperm transit through the head and body
directly involved in modifying the spermatozoa in some of the epididymis, those areas where sperm maturation
unknown way. occurs, is not influenced by ejaculation," it is reasonable to
As an initial step toward understanding the process of assume that the fertility of sperm should nor be depressed
sperm maturation in the stallion epididymis, sperm from even in males ejaculating frequently. Extensive data for
. four regions of the epididymis were evaluated by criteria stallions are not available to support this conclusion, but
considered co reflect the maturational st ate of sperm. H Data comprehensive studies have shown that the fertility of
for epididymal sperm also were compared with those for
ejacul ated sperm. Since evaluation of fertilizing capability
was impractical, characteristics of sperm associated with
maturation in other species were studied. It was found
(Table 6) that temperature shock induced by rapid cooling
to DOC did not decrease the percentage of sperm that
excluded the dye eosin. Thus, sperm from all regions of the
epididymis were considered to be resistant to cold shock.
Ejaculated sperm, however, were altered by rapid cooling co
DOC. Consequently, when handling ejaculated stallion
semen, precautions should be taken co eliminate sudden
changes in temperature (other than when thawing frozen
semen).
Equine sperm from the head or proximal body of the
epididymis were immotile when released into a
physiological salt solution (Table 6) .H The percentage of
motile sperm in samples from the tail of the epididymis,
however, was similar co that in ejaculates collected from the
same stallions. Thus, as measured by progressive motility,
maturation of stallion sperm is completed before sperm
enter the tail of the epididymis." Based on these
observations, and on the structural stability of stallion
sperm from different regions of the epididymis,johnson et
alY concluded that maturation of sperm in the stallion is
not completed until the sperm leave the body of the
epididymis. From this research and the limited data on the
fertility of stallion sperm from the tail of the epididymis," it
is reasonable to assume that sperm from the tail of the
epididymis of a valuable stallion could be used co
inseminate mares provided such sperm were recovered
within a few hours after death or castration of the stallion.
Although sperm are found throughout the epidi-
dymis,7·'?·2~
the tail of the ep ididymis and the deferent duct
are the major sperm storage areas (Table 7) . Since sperm
Figure 17. Cross and sagittal sections through the head of the ep ididym is
within the tail and deferent duct have completed their showing the epithelium and th in layer of smooth muscle (upper; X!!O)and
maruration," sperm scored in these regions should have the surface of the epithelium and spermatozoa within the duct (lower;
normal fertility. However, the number of sperm scored <X375).FromJohnson et ",.~2

94 EQUINE VETERINARY SCIENCE

Table 6. Change In Sperm During Passage Through the not ejaculated at copulation or voided by masturbation, arc
Ep ididymis a voided periodically during urination."? We speculate that in
. a normal, sexually inactive stallion, sperm intermittently
Unstained sperm (%)
p ass from the deferent ducts into the pelvic urethra and arc
Not Cooled to Progressive voided during micturition. The direct cause for and interval
Sperm source cooled 0 0 In s 2 min moti Iity (%)
Central head 96 95 0 between such emissions are unknown, but their occurrence
Proxi mal body 97 95 0 in normal stallions is likely. Certain sta llio ns accumulate an
Distal body 93 93 11 c abnormally large number of sperm in the epididymis." In
Central tai I 90 87 b 56 d
EJ aeul ated 68 27 64 such a horse, it is likely that spontaneous emissions do not
occur and that sperm accumulate in the epididymis until the
a Modified from Johnson et <I1.4l limit of distensibility of the epididymal duct is reached. As a
b The percentage of unstained sperm was reduced (P < 0.01) by cold shock. consequence of this accumulation, the storage interval of
c Sperm from two of five stallions were progressively motile.
sperm in the tail of the ep ididymis of such a stallion is much
d The percentage of rnorile sperm was similar (P >0.05) to that in
ejaculated semen. longer than 7 to 10 days and the sperm may undergo
marked alterations. The motility of sperm in the first
several ejaculates collected from a stallion accumulating
sperm from bulls ejaculating daily, or at a similar high sperm might be lower than usual because of the prolonged
frequency, is equivalent, if not slightly superior, to that for storage interval.
bulls ejaculating once weekly.2.1I.10 When seven successive
ejaculates were collected from beef bulls and the semen was
used to artificially inseminate cattle, fertility did not differ
among the seven successive ejaculates. II Fertility of stallion
semen used for artificial insemination probably is similar
when stallions are collected daily or every other day since
conception rates of mares inseminated daily or every other
day are similar.
Although transit time of sperm through the head and
body of the epididymis is not altered by ejaculation
frequency, the interval that sperm spend in the tail of the
epididymis is influenced by ejaculationY The number of
sperm in the tail of the epididymis is maximal in sexually
rested stallions (Table 7) and reduced by 20 to 30 % in
males ejaculating daily or every other day.' Since fewer
sperm arc present in the tail of the epididymis of a stallion
ejaculating regularly than in a sexually inactive male, the
transit time for sperm through the tail of the epididymis of Figure 18. Sagittal sectio n through the body of the epid idymis. Th e
undul arions result from differences in cell height (X250). FromJohnson et
a sexually active stallion is reduced to 7 or 8 days from the
al. 4l
10 days characteristic of a sexually rested stallion.'
Sperm are produced continuously at a high rate
regardless of ejaculation frequency. Since sperm enter the
epididymis at a constant rate, they also must leave the The Spermatic Cord
excurrenr duct system at a relatively constant rate, although
this rate is altered by ejaculation. Based on work with The spermatic cord extends from the abdominal inguinal
several species.v" it is likely that all sperm that enter the ring to its attachment on the testis. It serves to suspend the
excurrent duct system of the stallion leave through the testis in the scrotum and as a passageway for the deferent
urethra. It is unlikely that sperm resorp tion occurs within du ct, nerves and blood vessels associated with the testis
the excurrent duct system.l.1·4?Consequently, sperm that are (Figure 15). One of the most obvious components of the
spermatic cord is the external cremaster muscle. This is a
striated muscle that encloses the lateral and caudal aspects
Table 7. Sperm Reserves in the Adult Stallion a of the spermatic cord and runs from the abdominal muscles
to its attachment within the parietal vaginal tunic near the
Sperm In Relative
Segment of both sides distribution head of the epididymis. A continuous band of connective
exc ur r ent duct (10 9) (% ) tissue also runs the entire length of the spermatic cord. This
Head of epididymis 12 13 band is formed prior to descent of the testis, through fusion
Body of epididymis 17 19 of the parietal and visceral layers of the vaginal tunic. 14 This
Tall of epididymIs 54 61
band is termed the mesorch ium. The relationship of the
Deferent duct 4 4
Ampulla 2 2 mesorchium to the tunica vaginalis, external cremaster
muscle, deferent duct and blood vessels is evident in
a Modified from Amann et <If using data (or stallions 5·16 years old. Figure I').

Mav/June 1981 95
Figurc 19. Cross and sJgittal sections through the rail of the epididymis. In the proximal tail (I 9-A) short papillae of epithelium radiate from the wall of the
duct and the smooth musclc layer is of moderate thickness (X80). Details of the papillae (P) and smooth musclc (SM) arc shown in 19-B(X ·100). In the distal
tail (21,Cl. the papillae are much longer and thc smooth muscle layer is very thick (XGO). From Johnson £'1 dl. 4!

96 EOUINE VETERINARY SCIENCE

 Principal cell
Basal cell
Figure 20. Drawing of a cross section through the cpididymis showing its
organizat ion into luminal, epithelial and intertubular compartments. The
epirheliurn contains tall , columnar principal cells and apical cello; and also
small spherical basal cells. Occasional rnacrophages (possibly leucocyrcs)
arc found with in the ep ithclim.
The spermatic cord includes the hi~hly coiled testicular
artery (see Figure 5) from which branches lead to
vascularize the head and body of the epididymis. The veins
draining the testis from an intimate network of small veins
around the highly coiled artery. " This network of veins is
called the pampiniform plexus and is part of a heat
transfer mechanism. Autonomic nerves parallel the
testicular artery and 4 to 8 lymphatic vessels accompany the
testicular veins.
REGULATION OF TEMPERATURE
WITHIN THE TESTIS AND EPIDIDYMIS
The scrotum serves to cover and protect the testes, but its
primary function is regulation of temperature in the testis
and the tail of the epididymis. The testes of a stallion must
be below body temperature for normal spermatogenesis.
Based on research with other species,34.6l.71.71 we assume that
if the intraresricular temperature is elevated to ~ <10.5°C
(105° F) for 120 min, then certain germ cells (B-
spermatogonia and pachytene spermarocytes) developing
within the seminiferous tubules will be affected and die .
Consequently, a transitory decrease in the number of sperm
produced or ejaculated would be expected about 50 days
later. The influence of elevated temperature on sperm
within the epididymis has not been studied with stallions.
In other species, however, prolonged elevation of
intrascrotal temperature docs affect the quality and
functionality of sperm within the epididymis.s"
Although it generally is assumed that thermal regulation
of the testis and epididymis evolved as a mechanism to
reduce inrraresticular temperature and allow normal
May/June 1981
spermatogenesis, it has been suggested that maintenance of
the tail of the epididymis at a tempera ture <1 to 5°C below
body temperature may be even more important than
maintaining a low testicular temperature." The low
temperature within the tail of the epididymis may be one
mechanism involved in maintaining the fertilizing capacity
of sperm stored within this organ.
Thermal regulation of the testis and epididymis is
dependent upon the combined anion of the scrotum,
pampiniform plexus, tunica dartos muscle and external
cremaster muscle. Based on data for sheep," we assume that
blood within surface veins of the testis is cooled by removal
of heat through evaporation of moisture from the scrotal
skin. Thus, the temperature of testicular venous blood is
similar to that beneath the scrotal skin. The cooler venous
blood enters the pampiniform plexus which serves as a
counter-current heat exchange area. Heat is transferred
from the warm arterial blood to the cooler venous blood so
that the arterial blood leaving the parnpiniforrn plexus is
several degrees cooler (4.5°C in the ram) than that in the
testicular artery just above the pampiniform plexus. The
extent of cooling of the arterial blood passing through the
pampiniform plexus is governed by the temperature of the
venous blood.
The tunica dartos muscle, within the scrotal wall,
Contracts in cold weather co raise the testes close to the
abdominal wall and minimize heat loss. In warm weather,
the tunica dartos relaxes to make the scrotum more
pendulous and maximize cooling of the testicular venous
blood by evaporation from the scrotal skin. The external
cremaster muscle can reflexly contract to raise the testis for
a short interval.
THE REPRODUCTIVE TRACT
The Deferent Duct
a
The deferent duct is continuat ion of the epididymal
duct and runs from the tail of the epididymis, through the
spermatic cord co the pelvic urethra (Figure 1). Near its
origin, the deferent duct lies medial and parallel to the body
of the epididymis on the dorsal surface of the testis. The
deferent duct of the stallion has an extremely thick wall of
smooth muscle so that the proximal deferent duct can be
palpated readily through the scrotal skin. The lining of the
narrow lumen is a pseudostrarified, columnar epithelium
and is folded longitudinally. The epithelium probably has a
secretory function.
As the deferent duct approaches the pelvic urethra, it
widens into a structure termed the ampulla of the deferent
duct (Figures I and 21). The ampulla is abour 18 mm in
diameter as compared co 4 to 5 mm for the major portion of
the deferent duet . The increased diameter in the ampullary
region is primarily due to a thickening of the wall associated
with the presence of crypts and glands, although there is
some increase in luminal diameter. After the ampulla
passes under the body of the prostrate (Figure 21), the duct
joins with that from the ipsilateral vesicular gland to form a
shore ejaculatory duct that empties into the urethra within a
small prorruberance termed the colliculus serninalis. The
colliculus serninalis contains the ducts leading from both
sides of the tract.
97
 Urinary
bladder - - - T . - - - - - - - : ; . : - - - - - - . . . '
Gen ital fold --ET~+.t_~~'<:""""_;.~,.....::;::....--
Ischium-~~
~""";~~i-:-+--- Urethralis
Retractor
penis muscle---n-~-~~rr:~
l@a~~~~F--~k'\--Isch iocavernosus muscle
Figure 21. Drawing showing a dorsal view of the pelvic portion of the reproductive traer, The fascia and most of the genital fold, that support the pelvic
porti on of rhe tract , were dissected away. From Pickell et al.61
The Vesicular Glands
The two vesicular glands (previously termed seminal
vesicles) are long, tubular-alveolar gl ands lying lateral to
the ampulla and dorsal to the neck of the bladder (Figure
21). The portion of the reproductive tract lying within the
pelvis, including the vesicular glands and ampullae, is
suspended by loose connective tissue and a sheet of fascia
termed the genital fold. Essentially, the vesicular glands are
elongated, hollow pouches about 15 to 20 cm long and 5 cm
in diameter. Their smooth muscle wall is lined by a folded
mucus membrane containing numerous branched tubules
with grape-like, secretory, alveolar pockets. A single
excretory duct leads from each vesicular gland, through the
colliculus seminalis, to the ejaculatory duct and an orifice in
the pelvic urethra.
In sp ite of the historic use of the term seminal vesicles to
describe these glands, the vesicular glands do not serve as a
storage organ for sperm. Sperm normally are not found in
the vesicular glands.
The fluid secreted by the vesicular glands contributes a
major portion of the seminal plasma in an ejaculate. The
gelatinous material found in seminal plasma, especially in
April to july,?' is secreted by these glands." Their fluid also
contains fructose, citric acid and proteins.
The Prost ate Gland
The prostate gland is a single, firm and nodular gland
(Figure 21) with tWO narrow lobes (each 7 x 4 x 1 cm)
98
~rm:l~T;;rrm~r--Deferent duct
:;;;::'~--Ampulla
~;;;J<.r-.:r.:-~~~;"";:'!I---- Ves i c ular gland
--.-:-:,:,.,..:.;:;;;,:.;;m7r.T.~~~~-- Lobe 0 f prost ate g Iand
Isthmus of prostate gland
muscle
Bulbourethral gland
Bulbospongiosus muscle
connected by a thin, trans verse isthmus (about 3 cm long).
The lobes are somewhat prismatic in shape and lie over the
dorsal aspect of the vesicular glands. A series of 15 to 20
ducts lead from each lobe of the prostate into the pelvic
urethra . The prostate has a tubular alveolar, grape-like
structure and is interspersed with smooth muscle and
connective tissue.
The secretion of the prostate gland is thin and watery."
This secretion may help to cleanse the urethra during
ejaculation and also contributes to the seminal plasma.
The Bulbourethral Glands
The two bulbourethral glands (previously termed
Cowper's glands) are compound , tubular alveolar glands
that lie on either side of the pelvic urethra near the ischial
arch (Figures 1 and 21) . These glands have a simple
columnar epithelium and produce a mucoid secretion. A
series of 6 to 8 small ducts lead from each bulbourethral
gland to open in the pelvic urethra caudal to the openings
from the prostate gland. The bulbourethral glands are
covered by a dense capsule of fibrous tissue and are overlain
by the bulboglandularis muscles and bulbospongiosus
muscles. These muscles contract during emission and
ejaculation to force the bulbourethral gland secretion into
the pelvic urethra where it contributes to the semen.
Collectively, the vesicular glands, prostate gland and
bulbourethral glands are termed the accessory sex glands.
The accessory sex glands contribute most of the fluid to the
EaUINE VETERINARY SCIENCE
ejaculate, but are not essential for normal fertility of the
sperm. Normal function of all the accessory glands is
dependent upon the availability of testosterone in the
peripheral blood.
The Urethra
The urethra is a long mucus tube that extends from the
bladder to the free end of the penis. The proximal portion,
within the pelvic cavity, is lined by a transitional epithelium
which changes 10 a stratified squamous epithelium (similar
to that of the skin) distally . The pelvic portion of the
urethra is overlain by a thick, powerful, striated muscle
running circularly around the cylindrical tube of the urethra
(Figure 21). This muscle is termed the urerhralis muscle
and contracts vigorously during ejaculation.
The ischial urethra extends from the bulbourethral
glands around the ischial arch to the root of the penis and
crura penis . This part of the urethral tube is surrounded by a
small amount of cavernosus tissue and covered by the
bulbospongiosus muscle which contracts during
ejaculation. The extrapelvic or penile urethra runs the
length of the penis and terminates in a free extension
termed the urethral process (Figure 22). The penile urethra
is surrounded by the corpus spongiosum (Figure 23) which
is an area of cavernous, erectile tissue. The urethra serves
the joint excretory canal for both urine and scmen.
The Penis
The penis is the male organ of copulation and consists of
three parts (Figure 22). These are the root or the bulb
which is attached at the ischial arch by two crura; the body
or shaft which is the main portion of the penis and extends
from the crura to the glans; and the glans penis which is
External pudendal artery
Cranial artery of
Dorsal process of glans
Corona glandis
Corpus cavernosum penis
of glan s
Urethral
Figure 22. Draw ing showing a left lateral view o f the penis and its attachment to th e ischium. From Pickell et ,,1. 6 1
May/June 1981
the enlarged, free end of the penis . The penis extends
forward from the ischial arch to the umbilical region and
lies on the exterior aspect of the abdom inal wall supported
by the penile fascia and skin (Figure 1). The stallion penis is
of the musculocavernosus type as compared to the non-
erectile, fibroelastic penis of the bull. The stallion penis
contains a large amount of erectile tissue that surrounds the
urethra and is enclosed in a connective tissue capsule, the
tunic albuginea (Figure 23).
The root of the penis is formcd by the two crura which
arc extensions of the corpus cavernous urn penis that
terminate on either side of the ischial arch . The root of the
penis is attached to the pelvis by two strong, suspensory
ligaments and the paired ischiocavernosus muscles (Figure
22). When the body of the penis is viewed in cross section
(Figure 23), it is evident that the corpus cavernosum penis
forms the major component. Thecorpus cavernosum penis
is spongy, erectile tissue which is continuous with the veins
draining the penis. This erectile tissue is supported by
trabeculae arising from the tunica albuginea. The corpus
spongiosurn is a small area of spongy, erectile tissue that
immediately surrounds the urethra, It also becomes
engorged with blood during erection. The glans pen is has a
thin covering of skin that is devoid of glands. The glans
penis is richly endowed with nerve endings and becomes
engorged and erect during sexual excitement. A short
urethral process (Figures 22 and 24) extends beyond the
glans penis.
When the smooth muscle of the trabeculae within the
body of the penis loose tonus, the flacid penis will drop.
Erection involves several structures. At times of sexual
excitement, the smooth muscle within the trabeculae relax
Obturator
artery Rectal part of
r et ra ct or penis
.~\\ muscle
~~.
Tuber ischii
'
Suspensory
ligament of
penis
artery of penis
penis muscle
Bulbospong iosus muscle
99
 the corpus cavernosum penis results in erection. Arterial
veins of penis blood then passes into the cavernous spaces of the corpus
spongiosum [0 bring about dilation or flowering of the
glans penis. Loss of the erection, or detumescence, occurs
nerve of penis
after sympathetic nerve impulses induce contraction of
smooth muscle in the trabeculae and blood is forced out of
the cavernous spaces .
When not erect, the penis is about 50 cm long by 2.5 to
5.0 cm in diameter. About 15 to 20 ern lie free in the prepuce
(Figure 24). Erection increases the length and diameter of
albuginea
the penis by about 50% while the glans penis increases 300·
400 % in diameter through engorgement.

cavernosum The Muscles

Retractor pen is muscle
Figure 23. Drawing showing a cross section through the body of thepenis.
From Pickell " I ,,1.61
and the walls of the helicine arteries of the penis also relax
so that blood enters these tortuous arteries. Dilation of the
helicine arteries allows blood to enter the cavernous spaces
of the corpus cavernosurn penis. Simultaneously, drainage
of blood from the penis is blocked . The resulting dilation of
The paired ischiocavernosus muscles are short, strong,
striated muscles that enclose the crura penis and insert on
the lateral surface of the penis to connect the root of the
penis with the ischial arch (Figures 21 and 22). They arc
flattened laterally. During erection, the ischiocavernosus
muscles pull the penis against the ischial arch thereby
compressing the dorsal veins and preventing outflow of
blood from the penis. This results in engorgement of the
cavernosus spaces of the penis and erection.":"
The urethralis muscle is a circular, straired muscle which
surrounds the pelvic urethra (Figure 21). It is more or less
continuous with the bulbospongiosus muscle . The
urethralis muscle serves to empty the pelvic urethra during
urination or ejaculation.
The paired bulbospongiosus muscles arc powerful,
bulbous striated muscles that cover the ischial portion of
the urethra and the root of the penis and extend most of the
way through the body of the penis (Figures 21 and 22). A
medial raphe separates the two muscles and they arc

venous plexus

cavernosum penis

Urethra

Urethral

Fossa glandis
spongiosum

External prepuce
Internal prepuce

Figure 24. Drawing showing a left lateral view of the glans penis and prcpucc. From Pickell "I al.6 1 .

100 EaUINE VETERINARY SCIENCE

covered by dense connective tissue. An extension of each
bulbospongiosus muscle covers one bulbourethral gland
and is termed the bulboglandularis muscle. The
bulboglandularis muscles help to evacuate the fluid from
the bulbourethral glands during emission. The
bulbospongiosus muscles empty the extrapelvic pan of the
urethra by pulsitile contractions during ejaculation.v-'-
The paired retractor penis muscles arise at the first two
coccygeal vertebrae, divide and pass on either side of the
rectum, to a point of insertion on the penis below the crura
(Figures 1 and 22), and continue on the ventral surface of
the penis. The retractor penis muscles are smooth, rather
than striated, muscles and by contracting, serve to retract
the f1acid penis back into the prepuce or sheath.
EJACULATION
The process commonly considered as ejaculation actually
involves three sequential processes: erection, emission and
ejaculation. Erection is the lengthening and stiffening of
the penis resulting from engorgement of the corpus
cavernosum penis with blood. Emission is the movement
and deposition of sperm and fluid from the deferent ducts
and tails of the epididymis, together with fluids from the
accessory sex glands, into the pelvic urethra. Ejaculation is
the actual expulsion of the semen out through the urethra.
In the stallion, ejaculation occurs as a series of strong
pulsitile contractions so that several successive "jets"
(typically eight) of semen are ejaculated."
HORMONAL CONTROL OF MALE
REPRODUCTIVE FUNCTION
Function of the reproductive organs is controlled by the
neuroendocrine system.v'-" Although the autonomic
nervous system has a role in controlling function of the
reproductive organs; transport of sperm from the testis
through the epididymis and deferent duct; and the
processes of erection, emission and ejaculation; the
maintenance of normal function of reproductive organs is
dependent more upon the neuroendocrine system. In
addition to the reproductive organs considered earlier, the
hypothalamus and anterior lobe of the pituitary gland have
crucial roles in the neuroendocrine system.
The hypothalamus (Figure 25) is a pan of the
diencephalon of the brain, but its exact boundaries have not
been critically defined in the horse. Based on work with
other species, it is known that the hypothalamus is involved
in regulation of appetite and thirst, body temperature,
vasomotor activity, emotion, use of body nutrient reserves,
activity of the intestinal tract and bladder, the state of sleep
and wakefulness, sexual behavior, and release of tropic
hormones. This latter role, controlling the release of tropic
hormones, is the role of primary importance in controlling
reproductive function.
The pituitary gland is connected to the hypothalamus
and extends downward from it (Figure 25). The pituitary
gland consists of three parts or lobes: the infundibulum,
anterior lobe and posterior lobe. The anterior lobe secretes
May/June 1981
Figure 25. Diagram showing the hypothalamus and pituitary gland in the
skull of a horse. From Pickell et ,,/.61
and discharges a number of hormones that control
reproductive processes in the stallion and mare." Several
cell types are found in the anterior pituitary and each cell
type produces one or more hormones. The posterior lobe,
on the other hand, does not secrete hormones, but simply
serves as a storage reservoir for hormones produced by
neural tissue within the brain. These hormones, oxytocin
and vasopressin, come from the brain to the posterior lobe
of the pituitary by flowing down several large nerve tracts
that extend from the hypothalamic area, through the
infundibular stock, and terminate in the posterior lobe of
the pituitary gland. 17.10
The anterior lobe of the pituitary gland is not connected
to the hypothalamus by nerves. Rather, the hypothalamus
and anterior lobe of the pituitary gland are linked by portal
vessels which run directly from the neural tissue of the
hypothalamus to the glandular tissue of the anterior
pituitary. 17 Within this portal vascular system, blood
normally flows directly from the hypothalamus to the
anterior pituitary without going to the heart. The portal
vessels are the only. direct link between the hypothalamus
of the central nervous system and the anterior pituitary
lobe.
Under appropriate neurochemical stimulation, the
hypothalamus synthesizes and discharges a number of
"releasing hormones".17.30 The releasing hormone most
directly involved in controlling reproductive function is
gonadotrophin releasing hormone (GnRH). GnRH is
discharged by the hypothalamus in short, pulsatile bursts.'?
This fact, coupled with its rapid clearance from the blood,
results in a pulsatile discharge of hormones produced by the
anterior pituitary in response to stimulation by GnRH.
GnRH has been isolated and contains 10 amino acids. A
large number of similar compounds, called structural
analogs, have been synthesized and used in research aimed
at studying and controlling reproductive function. Two
types of analogs to GnRH are available: antagonistic
analogs that block the action of the natural hormone and
agonistic analogs that are more effective in evoking a
response than the natural GnRH.
The anterior lobe of the pituitary gland produces at least
six trophic hormones, but only two or three arc of direct
101
 - - ---- ------1
I
I
I
I
~--0~~~)~*:~-------- --~
I
I
oI
Accessory
sex glands I
I
I
I

Leyd ig ce lls in Sertoli cell in

intersti t ial t issue sem iniferous tu bule

Figure 26. Diagram showing inrerrelarionsh ip of hormone production in the Leyd ig cells and in the seminiferous tubules and the Icedback control of
gonadal hormones on the hypothalamus and anterior lobe of the pituit ary gland. An increased level of testosterone in peripheral blood. cirher as J result of
increased production by the tesres or following injection of exogenous hormone, feeds back on the hyporhal arnus and anterior pituitary to suppress discharge
of LH and, therefore, the Leydig cells produce less testosterone and the concentraticn of resrosterone around the semin iferous tubules also drops. GnRH =
gonadotrophin releasing hormone. LH = luteinizing hormone, FSH =follicle srirnularing hormone. PRL =prolacrin, T = testosterone, E =estradiol or orhcr
estrogens, AnI' = androgen binding pro tein, and I = inhibin. From Pickel[ et .11.61

102 EaUINE VETERINARY SCIENCE

importance in male reproduction.":" The anterior
pituitary lobe produces luteinizing hormone (LH) and
follicle stimulating hormone (FSH) in direct response to
stimulation by GnRH. LH and FSH are termed
gonadotroph ic hormones because they act on the testes and
stimulate them to produce steroid hormones. Both LH and
FSH are glycoprotein hormones with a molecular weight of
about 32,000 daltons. Among the other hormones produced
by the anterior pituitary gland, only prolactin (a protein
hormone) has been shown to have a direct effect on
testicular function . Although prolactin influences testicular
function in the rat and human," its role, if any. in
controlling function of the stallion testis has not been
established.
The concepts for endocrine control of reproductive
function in the stallion are based mostly on data for other
species. 17•10,46.H Between birth and puberty. circulating levels
of testosterone are low. Blood levels of FSH probably are
elevated at some interval before the onset of
spermatogenesis and FSH probably is important for
development of Sertoli cell function. As puberty
approaches. blood levels of LH tend to increase and the
circulating level of testosterone increases rapidly in parallel
with the increase in testicular weight that starts ncar 18
months ? f age and the concurrent onset of spermatogenic
activity.
In the adult male (Figure 26). production of testosterone
is controlled by LH. In the stallion, as in other species, LH is
released in episodic bursts which result in blood levels of
LH that temporarily arc far above the basal value.
Consequently, the basal level of testosterone production is
augmented by episodic bursts of testosterone
producrion.n -» Testosterone, produced by the Leydig cells,
passes through the venous blood dr aining the testis, into
the general circulation, and to the hypothalamus and
anterior pituitary gland. In this manner, testosterone feeds
back via a "long loop" and modulates the discharge of
GnRH and LH. If the concentration of testosterone
reaching the hypothalamus and pituitary gland is relatively
high. the discharge of GnRH by the hypothalamus is
suppressed and the response of the anterior pituitary gland
to available GnRH also is suppressed. Because of this
negative feedback. the concentration of LH would be low in
blood entering the testis so that the Leydig cells would be
exposed to a low concentration of LH and. consequently,
the Leydig cells would only secrete testosterone at a basal
rate. As the concentration of testosterone in peripheral
blood declines, the negative block is removed and episodic
discharge of GnRH from the hypothalamus can occur. This
is followed by a discharge of LH from the pituitary gland, an
elevated concentration of LH in blood flowing to the testis,
and rapid stimulation of the Leydig cells to produce and
discharge testosterone. Thus, the Leydid cells of the testis.
the hypothalamus. and the anterior lobe of the pituitary arc
involved in a circular feedback loop which regulates the
concenrrarions of LH and testosterone in peripheral blood
(Figure 26).
Although secretion of FSH also is stimulated by GnRH,
secretion of FSH and LH must be controlled by different
May/June 1981
mechanisms. Bursts of LH secretion frequently are not
accompanied by discharge of FSH . Based on research with
non-equine species, it appears that FSH acts exclusively on
Sertoli cells within the seminiferous tubules." Although
stimulation by FSH is necess ary for initiation of Sertoli cell
function , it is not clear if normal function of Serroli cells
requires coneinuous or intermittent stimulation with FSH.
or if their function can be maintained solely by high
concentrations of testosterone. In any case. the basal region
of the Serroli cells is exposed to FSH.
The Serroli cells produce a compound termed inhibin that
passes through the blood stream to the anterior lobe of the
pituitary gland (Figure 26) and also may be taken up by the
hypothalamus. The concentration of inhibin in blood
apparently reflects the function of the germinal
epithelium." If the blood concentration of inhibin is
normal, there is a partial suppression of FSH synthesis and
discharge. If spermatogenesis is disrupted, however, the
concentration of inhibin in peripheral blood declines and
FSH production and secretion increase; the ratio of LH to
FSH is altered.in peripheral blood. This feedback loop for
controlling Serroli cell function , which apparently involves
the action of inhibin (or possibly a steroid) on the anterior
pituitary gland, is depicted in Figure 26.
As discussed in the section on testicular function, the
stallion resris produces un iquely high concentrations of
estrogens.' It remains to be established if the estrogens are
produced in the Sertoli cells. Leydig cells or some other
component of the testis. Figure 26 incorporates the
speculation that testosterone produced by the Leydig cells
enters the Sertoli cells and is transformed, within the Serroli
cells, to estradiol or other estrogens that then return to the
interstitial tissue and enter the blood draining the testis.
Clear proof of this concept is not available for any species.
In any case, blood draining the stallion testis is rich in
estrogens and high concentrations of estrogens in blood
flowing to the hypothalamus and anterior pituitary may
suppress discharge of GnRH or of LH and FSH
In several species. ll.62 the Sertoli cells also produce a
compound termed androgen-binding protein (ABP).
Androgen-binding protein is thought to serve as a transfer
protein to carry testosterone to the germ cells developing
within the germinal epithelium or, alternatively. to provide
a relatively high concentration of testosterone to the
epithelium lining the initial segment of the epididymis.
The physiological importance of either of these two roles,
for the stallion or for any other species. has not been
established.
From the preceding discussion of feedback loops
involving the hypothalamus, anterior lobe of the pituitary
gland, and the testis. it should be evident that injection of
hormones will alter this delicate balance and may
profoundly influence reproductive function . Injection of
GnRH or another compound that induces a discharge of
LH from the anterior lobe of the pituitary gland usually will
induce (by action of LH) secret ion of testosterone by [he
Leydig cells. Typically. this release of testosterone will be
similar in magnitude to those occurring naturally
throughout the day or night and no prolonged alteration of
103
Table 8. Effect of Anabolic Steroids on Testicular Characteristics a
Testis No.
spermatids
parenchyma
Treatment wt (g) Cl06/g testis)
b spermatogenesis would be retarded in a colt receiving
Control 257 b 36
Equipoise (1.1 mg/kg) 151 c 27 b
Equipoise (4.4 mg/kg) 102~ 24 c
c but it is likely that both FSH and testosterone levels within
Deca-durabol in 93 c 24
Cl.l mg/kg)
a From Todter. 7o Equipoise (boldenone undecylenate) or Deca-durabolin
(nandrolone decanoate) was administered intramuscularly every third
week over 15 weeks and stallions were castrated 16 weeks after the
initial injection,
b,c Means with different superscripts differ (P < 0.05).
reproductive function would be anticipated.
Injection of hCG would mimic the action of LH and cause
synthesis and secretion of testosterone by the Leydig cells.!
In many instances, the induced secretion of testosterone
would continue longer than secretion following a natural,
episodic discharge of LH. If injection of hCG induced
prolonged secretion of testosterone, the normal episodic
discharges of LH might be suppressed for several hours or
days following a single injection of hCG. There is no
evidence for the stallion to prove or refute this suggestion.
Based on work with other species, repeated injection of
hCG is likely to induce decreased responsiveness of the
Leydig cells to either LH or hCG and a decline in
testosterone production might result even in the face of
relatively high blood concentrations of LH or hCG.
Consequently, repeated injections of hCG probably would
be unsuccessful in maintaining elevated levels of
testosterone in a stallion thought to have impaired
reproductive efficiency associated with a low concentration
of testosterone in peripheral blood. Repeated injection of a
stallion with hCG is not recommended.
Injection of testosterone, a similar androgen or an
anabolic steroid is conrraindicated.v-":" Injection of
testosterone may maintain or raise the normal peripheral
level of testosterone reaching the hypothalamus and
anterior pituitary and suppress LH discharge by the
pituitary gland. Consequently, the Leydig cells will produce
little, if any, testosterone. Although the testosterone
concentration in peripheral blood may equal or exceed the
normal level, failure of testosterone production by the
Leydig cells results in a decline in the concentration of
testosterone within the interstitial tissue (Figure 26) from
a concentration possibly 45 times that in peripheral blood
(see Table 5) to a level equivalent to that normally found in
peripheral blood. The Serroli cells and developing germ
cells then would not be exposed to the high concentration
of testosterone normally found in fluids bathing the
seminiferous tubules. Since a high concentration of
testosterone is essential for normal spermatogenesis,
sperm production typically will decline in a stallion treated
with testosterone or other androgens (Table 8).16.17 The
severity of the decline is at least partly dose dependent and
damage to the germinal epithelium induced by repeated
injections of androgens can be very severe." Repeated
104
injection of a colt or stallion with any androgen or anabolic
steroid is contraindicated.
It is reasonable to assume the prepubertal development
of the germinal epithelium and the onset of
exogenous androgens. Experimental evidence is lacking,
testicular tissue would be abnormally low and the onset of
puberty would be retarded.
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