Manual of Equine Reproduction, 3rd Edition

Visit the link below to download the full version of this book:

https://medidownload.com/product/manual-of-equine-reproduction-3rd-edition/

Click Download Now

About the Covers:

FRONT COVER BACK COVER

Photograph of American Quarter Horse champion race Photograph of American Quarter Horse performance
mare, Your First Moon, owned by Frank “Scoop” Vessels III, champion and elite sire, Shining Spark, owned by Carol
Vessels Stallion Farm, Bonsall, CA, with foal. The foal, Stray Rose, Carol Rose Quarter Horses, Gainesville, TX. Shining
Cat, owned by Lyle Lovett and Scoop Vessels, was sired by Spark’s foals have earned more than $7.3 million, and
the great Thoroughbred stallion, Storm Cat, Overbrook over 32,881 AQHA points. He is a $3 million dollar
Farm, Lexington, KY, after the stallion’s retirement from NRHA sire, and the only $3 million dollar NRCHA
Thoroughbred breeding. The successful breeding was sire. He is also the sire of 20 AQHA World or Reserve
achieved by transrectally-guided low-dose insemination of World champions, numerous NRCHA world champions,
semen that was first centrifuged though a silica-particle NRCHA Snaffle Bit Futurity champions, an AQHA World
solution, EquiPure (Nidacon International AB, Mölndal, Show Super Horse, and a NRHA Open Futurity cham-
Sweden) to enhance semen quality in the inseminate. pion. In the past 10 years, the stallion has sired foals
through transrectally-guided low-dose insemination,
and in recent years, with EquiPure-processed semen.

viii
 Reproductive Anatomy of the Mare

1
CHAPTER
O B J E C T I VES
While studying the information covered in this chapter, the reader should attempt to:
■ Acquire a working understanding of the anatomy of the reproductive organs of the mare.

■ Acquire a working understanding of how defects in anatomic development of the reproductive tract of the

mare, or changes in anatomy that occur with injury or age, may adversely affect fertility.
■ Acquire a working understanding of procedures to be used for safe examination of the reproductive tract of

the mare per rectum.

S TUDY Q U EST I ON S
1. Describe the normal nonpregnant equine repro- 4. List three major physical barriers to contamination
ductive tract, including location and shape of the of the mare’s uterus.
ovaries, ovulation fossa, oviducts, uterus, cervix, 5. Describe important guidelines for examination of
vagina, vestibule, and vulva. the mare reproductive tract per rectum, including
2. Describe normal equine ovarian structures, the pro- restraint, protective wear, lubrication, manure
cess of ovulation and oocyte entry into the oviduct, removal, guards against rectal perforation, and
the process of sperm entry into the oviduct, and anatomic orientation.
the site of fertilization. 6. Discuss congenital and acquired defects of the mare
3. List structures that are physically isolated from reproductive tract that may affect reproductive
the reproductive tract but play a central role in performance.
regulation of reproductive events in the mare.

The reproductive system is made up of two groups of body that communicates between the abdominal cavity
organs: (1) those structures that are intrinsic to the repro- and the external environment. More than half of the
ductive tract (ovaries and tubular genitalia) and (2) those reproductive tract lies within the abdominal cavity, with
structures that are physically isolated from the reproduc- the remainder confined to the pelvic cavity. When the
tive tract but play a role in the regulation of reproductive ovum is discharged from the follicle at ovulation, it is
events (e.g., pineal gland, retina, hypothalamus, pituitary received at the level of the ovarian bursa, which is
gland). thought to assist the passage of the ovum into the
The reproductive tract (Figures 1-1 through 1-7) con- oviduct. The oviduct is responsible for movement of
sists of two ovaries and a tubular tract, including the sperm and ova to a common site (the ampullary-
paired oviducts and uterine horns, and a single uterine isthmic junction) for fertilization. After fertilization, the
body, cervix, vagina, vestibule, and vulva. The lumen of developing equine embryo travels down the oviduct;
the female reproductive tract is the only channel in the and after 4.5 to 5 days, it secretes increasing amounts of

1
2 CHAPTER 1 ■
Reproductive Anatomy of the Mare

Left uterine horn Right uterine horn

Uterine body

Ovarian ligament Oviduct

Infundibulum
Left ovary

Ovarian artery
Broad ligament
Ovarian branch of ovarian artery (mesometrium)

Uterine branch of ovarian artery Ureter to bladder

Uterine artery Cervix

Vagina

Transverse fold
Urethral orifice

Vestibulum

Vulvar labia
Clitoral fossa
Glans clitoris
Figure 1-1 Dorsal view of the mare reproductive tract. Broad ligament attachments to the abdominal and pelvic walls are not
depicted, and the dorsal vaginal wall has been omitted to reveal the mucosal surface of the external cervical os, vagina, and
vestibule.

Ovary Ovary

Ovary Rectum
rn

Ut
ho

er
e

Infundibulum in
in

e Vagina
er

of the oviduct ho
Ut

rn
Uterine
Vestibule
horn

Uterine body Vulva

Uterine body Bladder Cervix

Figure 1-2 Frontal aspect of the suspended ovaries and

uterus in situ. The rectum and abdominal viscera have been
removed to facilitate visualization of the reproductive organs.

Figure 1-3 Lateral view of the reproductive organs and adja-

cent structures of the mare. Abdominal viscera are not
prostaglandin E2, which allows it to gain entrance into the depicted to facilitate visualization of the reproductive organs.
uterus for gestational support. The uterus provides the
proper environment for the embryo to develop further.
The cervix accommodates the expanded glans penis endometrial folds that line the uterine body. The cervix
of the stallion at estrus to allow intrauterine deposition secretes two types of mucus: a thin mucus to lubricate
of sperm and closes tightly during pregnancy to prevent the posterior genital tract in preparation for coitus and a
ascending bacterial or fungal infection from the poste- more viscid mucus to help seal the cervical lumen dur-
rior tract. The cervix also expands considerably at the ing pregnancy.
time of parturition to accommodate passage of the foal. The vagina is a potential space that expands to per-
The caudal portion of the cervix projects into the lumen mit penile and foal passage. A transverse fold (remnant
of the vagina (Figure 1-8). Longitudinal folds comprise of the hymen) overlies the external urethral orifice and
the lining of the cervix and are continuous with the is the anatomic division between the vagina, which is
 Reproductive Anatomy of the Mare ■
CHAPTER 1 3

Mesothelium

Medulla

Cortex

Germinal epithelium
in ovulation fossa
Figure 1-4 Schematic drawing of cortical and medullary A
areas of equine ovary. Germinal epithelium lines the surface
of the ovulation fossa, where all ovulations take place. The
oviductal fimbriae (not pictured) pick up the ovulated oocyte
and move it to the site where fertilization takes place.

Infundibulum

Ampulla B

Isthmus

Figure 1-5 Lateral surface of ovary, oviduct, and end of uter-

ine horn exposed through laparotomy site. The portions of the
oviduct (infundibulum, ampulla, and isthmus) are identified.

A B C

C
Figure 1-7 Corresponding histologic cross sections of the
Isthmus Ampullary- Ampulla dissected oviduct in Figure 1-6. Note the smaller lumen and
isthmus junction highly muscular component of the isthmus and the larger
stellate lumen/diameter of the ampulla.
Figures 1-6 Grossly dissected oviduct depicts the site of
fertilization near the end of the ampulla (ampullary-isthmic
junction). A to C correspond with photomicrographs A to C OVARIES
in Figure 1-7.
The ovaries of the mare are usually the most anterior part
anterior, and the vestibule, which is posterior. The of the reproductive tract in the nonpregnant mare (see
juncture between the vestibule and vagina is referred to Figure 1-3). The noncoiled and dorsally suspended uterus
as the vestibulovaginal ring. When closed, this ring of the mare (as compared with the coiled tract and more
restricts entry of air and debris into the upper tubular posterior [flank] broad ligament attachment in other farm
tract. The vulva is limited to the external opening of the animal species) accounts for the more cranial location
tubular tract. of the ovaries. So, in the mare, the ovaries are in the
4 CHAPTER 1 ■
Reproductive Anatomy of the Mare

cortical zone (which contains the follicles) is in the inte-

rior of the gland. The cortical tissue reaches the surface
only at the depression of the ventral or free border. This,
therefore, is the only area from which normal ovulation
occurs and is appropriately termed the ovulation fossa.
The ovulation fossa is covered by a layer of short
polygonal cells, which are a remnant of the primitive
germinal epithelium. The ovulation papilla of the cor-
pus luteum does not project from the convex surface of
the ovary as it does in other species but rather protrudes
into the ovulation fossa.
The ovary has both exocrine and endocrine functions.
The exocrine function is development of gametes, and
the endocrine function is production of hormones.

OVIDUCTS
The oviducts (Fallopian tubes, uterine tubes) are long
tortuous ducts that measure 20 to 30 cm in length when
Figure 1-8 Endoscopic view of endometrial folds within col- fully extended in horse mares. Cilia are present on
lapsed uterine horn.
the epithelium of the oviduct and produce a current
directed toward the uterus. The oviduct is divided into
cranial-most transverse plane, whereas in other farm ani- three parts: the infundibulum (funnel-shaped portion
mal species, a portion of the uterus is cranial to the ova- nearest the ovary), the ampulla (expanded middle por-
ries. The ovaries and the follicles are larger in the mare tion), and the isthmus (narrowed portion that connects
than in other farm animal species (see Figure 2-5). Equine the ampulla to the uterine horn). The cranial edge of the
ovaries are bean-shaped and vary in size according to infundibulum attaches to the lateral surface of the
ovarian activity; they are largest during the breeding mare’s ovary (see Figures 1-5 and 4-7). At ovulation,
season (spring and summer) and smallest during the this fan-shaped structure envelopes the ovulation fossa
nonbreeding season (winter anestrus). Average ovaries to facilitate ovum entry into the oviduct. Cannulation
are 6 to 8 cm in length, 3 to 4 cm in height, and 70 to 80 g of the infundibulum is possible through a flank
in weight. approach, which has been used for such clinical and
The ovaries are located in the sublumbar area (ventral research procedures as oocyte transfer to a recipient
to the fourth or fifth lumbar vertebra), suspended by mare, gamete intrafallopian transfer (GIFT) (instillation
long sheet-like broad ligaments, and are usually located of both sperm and oocyte into a recipient mare), and
several centimeters behind the corresponding kidney. flushing to determine oviductal patency. The ampulla is
The right ovary is typically more cranial (2 to 3 cm) than the middle part of the oviduct (see Figure 1-6) where
the left. Because the ovaries may be lifted by the intest- fertilization and early cleavage of fertilized ovum
ines, their actual location in the body is quite variable. occur. The highly muscular isthmus (see Figure 1-7)
Therefore, to facilitate locating the ovaries during palpa- serves both as sperm reservoir and as a conduit to trans-
tion per rectum, we recommend tracing the tip of each port sperm from the uterus to the ampulla and fertilized
uterine horn to the associated ovary (as the ovary may ova (embryos) from the site of fertilization into the uter-
be in contact with, to within 5 cm of, the tip of the ipsi- ine lumen (see Figure 1-6). The oviduct enters the uterus
lateral uterine horn). Each ovary consists of two surfaces just caudal to the blunt end of the uterine horn through
(medial and lateral), two borders (attached [dorsal] and a distinct papilla (oviductal papilla) that is easily visu-
free [ventral]), and two poles (cranial [tubal] and caudal alized from the uterine lumen (see Figure 4-35). Normal
[uterine]). The caudal border is connected to the uterine oviducts are not usually palpable per rectum.
horn by the ovarian ligament. Each ovary is kidney-bean Sperm gain access to the oviduct through the utero-
shaped, with a prominent depression on the free or tubal junction (UTJ), which is located in the center of the
ventral border. The convex dorsal border is sometimes oviductal papilla that projects into the uterine lumen near
called the greater curvature. the blunt end of the uterine horn. Deep edematous longi-
The ovarian surface is largely covered by peritoneum tudinal folds are present in the UTJ during estrus, and
except at the attached border where nerves and vessels numerous sperm can be found “bound” to epithelial cells
enter. The relationship between ovarian cortical and or “trapped” in these folds within 4 hours of breeding. The
medullary areas is unusual in the mare (see Figure 1-4). UTJ may play a role in the selection of morphologically
The ovary of the mare is “inside-out” compared with normal sperm (i.e., allowing only normal sperm to access
those of other farm animal species. In other words, the oviduct) and may also act as a storage site for sperm
the medullary or vascular zone is superficial and the awaiting transport into the oviduct. The muscular isthmus
 Reproductive Anatomy of the Mare ■
CHAPTER 1 5

is believed to contract rhythmically after breeding in a marked by a short uterine septum (see Figure 4-33),
fashion that propels sperm to the fertilization site in the and the body of the uterus is longer.
ampulla (near its end at the ampullary-isthmic junction). The vasculature of the uterus is supplied on each
Adhesion of sperm to epithelial cells in this area is thought side by three arteries and veins (see Figure 1-1) that
to prevent premature capacitation and increase the lifes- weave their way through the broad ligament: the uter-
pan of the sperm, resulting in a sperm reservoir awaiting ine branch of the vaginal artery and corresponding
the opportunity for release to fertilize the ovum. Because vein, the uterine artery (sometimes called middle uter-
mares may become pregnant (albeit at a lower rate) when ine artery) and corresponding vein, and the uterine
ovulation occurs as late as 6 to 7 days after breeding, the branch of the ovarian artery and corresponding vein.
sperm attached to the oviductal epithelium are thought to Rupture of these arteries or veins sometimes occurs
be gradually released in waves so as to continuously sup- during parturition in aged mares, leading either to
ply capacitated sperm that can fertilize the ovum. hematoma formation within the broad ligament or fatal
hemorrhage into the abdomen (see Chapter 10). The
UTERUS ovarian artery is located in the cranial portion of the
broad ligament and follows the course of the ovarian
The uterus consists of two horns and a singular body. vein and the uterine branch of the ovarian vein; in con-
The uterus has been described as T-shaped in the mare, trast to ruminant species, the ovarian artery is not as
but Y-shaped (see Figures 1-1 and 1-2) is probably a closely attached or applied to the ovarian vein. This has
more accurate description of the organ when viewed important functional considerations, namely for coun-
dorsally in its natural position in the mare. The uterus is tercurrent transport of prostaglandin-F2␣ (PGF2␣) from
suspended within the pelvic cavity and abdomen by the the veins draining the uterus into the ovarian artery.
broad ligament. These suspensory attachments also Because this countercurrent exchange is not efficient in
serve as sheaths that contain blood vessels, lymphatics, the mare, regression of the corpus luteum is induced by
and nerves; the serous layer is continuous with the PGF2␣ that reaches the ovary via the systemic circula-
serous lining of the abdominal cavity. The portion of the tion, as opposed to ruminants, in which PGF2␣ reaches
broad ligament that attaches to the uterus is called the ovary in higher concentrations via the ovarian
the mesometrium. In the mare, the mesometrium artery because of countercurrent exchange from the
attaches to the dorsal surface of the uterine horns, closely entwined uterine venous drainage.
whereas in the cow, the attachment is on the ventrolat-
eral surface. Therefore, in mares, the free (unattached) CERVIX
surface of the uterus is ventral to the broad ligament,
whereas in cattle, the free surface is dorsal to the broad The cervix is a dynamic organ in the mare. It is lined
ligament. This arrangement hinders digital evaluation internally by epithelium-containing secretory cells that
of the uterine body and likewise prevents retraction of produce a thin mucus to serve as a lubricant during
the uterus into the pelvic cavity during palpation of the estrus and a thick mucus to occlude the cervical lumen
mare per rectum, in contrast with the cow, in which these during diestrus and pregnancy so that it is less perme-
procedures are easily accomplished. The uterine horns of able to bacteria and foreign material. The longitudinal
the mare are entirely in the abdominal cavity and “float” folds of the cervix are continuous with the endometrial
on, or are intermingled with, intestinal viscera. folds present in the body of the uterus. The cervix
The serosal layer of the uterus and the vascular layer expands to accommodate the stallion’s penis during
plus longitudinal muscular layer are continuous with estrus and the foal during parturition, and it closes tightly
that of the broad ligament. The myometrium is com- during diestrus and even more so during pregnancy.
posed of an inner circular layer and an outer longitudi- The thick-walled cervix is usually identifiable via pal-
nal layer, with the outer longitudinal layer continuous pation per rectum, particularly during diestrus or preg-
with that in the oviducts. Finally, the innermost layer of nancy, and is typically 5 to 7.5 cm in length and 2 to 4 cm
the uterus consists of the endometrium, which is glan- in diameter. During estrus, the cervix is quite flaccid and
dular and secretory (see Figure 4-26). thus more difficult to feel via palpation per rectum.
The uterine lumen in the normal nonpregnant state The cervix of the mare has two features that differ
is nearly obliterated by the collapsed wall and promi- from the cow: (1) the cervical lumen greatly expands
nent endometrial folds (see Figure 1-8). The endome- and contracts during the estrous cycle as a result of a
trial folds are arranged longitudinally in the uterus and thick layer of circular muscle rich in elastic fibers; and
are usually palpable per rectum when the uterus is (2) the cervix has only longitudinal folds with no
“strummed” between the thumb and forefingers. The obstructing transverse cervical rings. Therefore, the
myometrium is quite thick and is responsible for varia- cervical changes are readily palpable throughout the
tion in uterine tone of the mare during estrus versus estrous cycle in the mare, and the uterus is more easily
diestrus or early pregnancy. In contrast with the cow, accessed through the cervix than in the cow. The exter-
the uterus of the mare is not coiled, the intercornual nal os of the cervix protrudes into the vaginal lumen
ligament is not prominent, the internal bifurcation is (Figure 1-9) and is surrounded by the vaginal fornix
6 CHAPTER 1 ■
Reproductive Anatomy of the Mare

Figure 1-9 Per vagina view of external cervical os protrud-

ing into vaginal lumen.

(i.e., the area of reflection of vaginal mucous membrane Figure 1-10 Persistent hymen in a maiden mare.
onto the cervix).

VAGINA VESTIBULE
The vagina is a tubular organ that extends horizontally The vestibule extends 10 to 12 cm from the transverse fold
for 15 to 20 cm through the pelvic cavity from the exter- overlying the external urethral orifice to the vulva. A ves-
nal os of the cervix to the transverse fold overlying the tibulovaginal ring exists at the junction of the vestibule
external urethral orifice (see Figure 1-1). In maiden and vagina and, because of vulvar and vaginal constrictor
mares, this transverse fold is often continued on either muscles that encircle this area, forms a seal, thereby mini-
side of the vagina and forms the hymen. Occasionally, mizing entry of foreign material into the upper tubular
the hymen completely encircles the vestibulovaginal junc- tract. This ring oftentimes is incompetent (weak or inca-
tion and is imperforate (persistent hymen) (Figure 1-10), pable of closing) when pneumovagina exists, allowing
precluding breeding until it is removed. The vagina entry of air into the vaginal space. The vestibule contains
continues caudally as the vestibule. vestibular glands ventrally that secrete mucus to provide
The lumen of the vagina is normally collapsed lubrication of the posterior tubular tract.
except during breeding and passage of the foal at par-
turition. The vagina, including its mucosa, is highly VULVA
elastic and expands considerably to accommodate the
passage of the foal. It becomes distended with air The vulva (Figure 1-11) refers to the external opening of
when the abnormal condition of pneumovagina exists. the female reproductive tract and the structures that sur-
The lumen of the vagina is covered with stratified round it. The vertical vulvar opening normally begins
squamous epithelium. The cranial vagina is covered 5 to 7 cm directly under the anus and is 12 to 15 cm in
with serosa and lies within the peritoneal cavity. The length. The dorsal commissure of the vulva normally is
posterior vagina is in a retroperitoneal position and is less than 5 cm above the ischium (floor of the pelvis). The
therefore not covered with serosa. Because most of the mare is prone to aspiration of air into the vagina (pneu-
vagina is retroperitoneal, vaginal injuries (such as movagina) if the dorsal commissure is greater than 5 cm
tearing during breeding) usually do not perforate into above the ischium, particularly if the anus is recessed
the peritoneal space, although this sometimes occurs. (sunken) and the vulvar lips are tipped horizontally so
Puncturing the vaginal wall near the cervix does pro- the vulva is no longer vertical (Figure 1-12). The labia
vide a surgical approach into the abdomen (colpot- of the vulva contain underlying musculature that func-
omy) for procedures such as ovariectomy. Unlike the tions to close the vulvar opening, providing a further
uterus, cervix, and vestibule, the vagina contains no barrier to the entrance of foreign material into the tubular
glandular structures. tract. The vulva contains much elastic tissue and expands
 Reproductive Anatomy of the Mare ■
CHAPTER 1 7

Clitoris

Fossa

Figure 1-13 The clitoris is located in a cavity just cranial to

Figure 1-11 Normal vulva and anus of a mare. the ventral commissure of the vulva. A wire is placed within
one of the clitoral sinuses, and the edges of the clitoral fossa
are retracted to expose the glans clitoris.

EXAMINATION OF THE REPRODUCTIVE TRACT

OF THE MARE PER RECTUM
Mare Restraint
The mare must be adequately restrained before exami-
nation of the reproductive tract. Such precautions pro-
tect both the mare and the examiner from severe injury.
Minimal but effective restraint is the key to a safe
examination and varies from mare to mare. The dispo-
sition of the mare should be determined before the
examination begins. Mares accustomed to frequent
examinations per rectum tend to need little restraint,
Figure 1-12 Sunken anus and tipped vulva in an aged whereas mares not accustomed to such practices often
mare. become anxious and sometimes explosive during the
examination. In any case, one should never be careless
and should always use caution. A strategically placed
kick can quickly terminate a career! If available, all
greatly during passage of the fetus at parturition. The mares should be placed in stocks (Figure 1-14) before
clitoris, a homologue of the penis, is located in a cavity the examination. Ideally, the stocks should be equipped
just cranial to the ventral commissure of the vulvar open- with a solid padded rear door to help prevent leg
ing (Figure 1-13). The glans clitoris is more prominent in extension if the mare does decide to kick and to afford
the mare than in other farm animal species. Three clitoral some protection against injury to the lower legs of the
sinuses are located on the dorsal aspect of the clitoris, mare. The height of the door should not be higher than
and a large singular clitoral fossa is located ventral to the the mid-upper gaskin region of the mare’s hindquar-
glans clitoris. The clitoral sinuses and fossa must be ters. Higher doors could damage the examiner’s arm
swabbed for bacteriologic culture to document freedom if the mare abruptly squats while the arm is in the rec-
from infection with Taylorella equigenitalis (formerly tum. If stocks are not available, the mare should be
Haemophilus equigenitalum), the causative organism of examined in a doorway with the hindquarters remain-
contagious equine metritis. ing 2 feet beyond the doorway. The examiner, when
8 CHAPTER 1 ■
Reproductive Anatomy of the Mare

The protected hand should be well lubricated before

use. A water-based lubricant such as methylcellulose
can be used. The lubricated hand should then be
shaped into a cone and gently inserted into the rectum
through the tight anal sphincter. Slow rotation of the
arm in conjunction with gentle forward pressure
enhances advancement into the rectum. Do not rush.
Rectal perforation is a common cause of malpractice
claims against equine practitioners. Use extreme cau-
tion, and never perform forceful manipulations in the
rectum against peristaltic waves or a tense rectal wall.
If necessary, products such as propantheline bromide
(Pro-Banthine, Schiapparelli Searle, Chicago, Ill.) or
N-butylscopolammonium bromide (Buscopan, Boeh-
Figure 1-14 Mare being palpated in a stock with padded
kickboard. ringer Ingelheim Pharmaceuticals, Ridgefield, Conn.)
can be administered to the mare to relax the rectum and
reduce straining.
standing to the side, is thus protected by the door As much fecal material as possible should be
frame. If a mare must be examined in an open area, removed from the rectum and distal colon before eval-
breeding hobbles can be properly secured to the mare’s uation of abdominal and pelvic structures. This task is
hind legs to reduce the range of limb motion, should accomplished by passing the coned hand past a small
she decide to kick. A leg strap can also be applied to a amount of feces and then cupping the hand to facilitate
flexed front leg. Lifting the tail directly over the back of removal when the arm is withdrawn.
the mare also helps distract her from the examiner’s Two established principles for examination of the
activities. If necessary a twitch can also be placed on female reproductive tract per rectum are (1) establish
the mare’s muzzle as an additional means of physical normal anatomic orientation and (2) follow a thorough
restraint. Tranquilization or sedation may be necessary methodical approach. Adherence to these principles
to adequately restrain an anxious mare. Remember, reduces the chance of error.
overrestraint can be as dangerous to the mare and Before emphasis is placed on detection of the repro-
the examiner as underrestraint. ductive tract, the examiner should first become prop-
Never develop a false sense of security through the erly oriented by identifying topographic landmarks
use of protective devices. Always take special precau- that are in a constant position, such as the outlines of the
tions when walking around the hindquarters of a mare. pelvic cavity (the pelvic floor, sacrum, wings of the ilia,
Approach the mare from the front and work your way and pelvic brim). Once the examiner is confident of the
back to the hindquarters while quietly talking to her so location in the pelvic and abdominal cavities, specific
she is aware of your location. Never surprise her. Mini- examination of reproductive structures may begin.
mize time spent directly behind the mare. Rather, stand Three initial landmarks for locating the reproductive
to one side when possible. Stand close to the mare to tract in the mare have been described by various
reduce concussion, should you be in the line of fire if authorities on the subject: the cervix, the leading edge
she decides to kick. of the uterus, and the ovary. All of the structures have
a relatively inconstant position. The cervix of the non-
Technique pregnant mare is located in the pelvic cavity, whereas
The first question that comes to mind is which arm to the leading edge of the uterus and the ovaries are
use to perform the examination. Ambidexterity may always in the abdominal cavity. To locate the cervix of
provide some advantage, but both hands are probably a nonpregnant mare, one should produce a side-to-side
equally clumsy at the onset. Therefore, a general rec- motion with the extended hand in the pelvic cavity,
ommendation is to teach the off hand to palpate per with downward pressure with the fingers held together
rectum. The more frequently used hand can then be and the palm facing downward (Figure 1-15). The cer-
used to perform other duties (i.e., operating an ultra- vix can be palpated in almost all instances but may
sound machine) during the examination. relax during estrus to the point of becoming barely
Shoulder-length rubber obstetric sleeves or dis- perceptible. However, even in this relaxed state, it can
posable plastic sleeves can be used to protect the arm be identified when the fingers slip caudally off the shelf
during the examination. Soft pliable sleeves increase formed by the external os. Uterine contents (e.g., fetus
sensitivity. Alternatively, the fingers can be removed or pyometra) may pull the cervix into an abdominal
from the plastic sleeve and replaced with a latex sur- position, making palpation more difficult. Palpation of
gical glove. To minimize horizontal disease trans- the dilated cervix per rectum is also difficult in the
mission, sleeves should be changed between mares. early postpartum period.
 Reproductive Anatomy of the Mare ■
CHAPTER 1 9

Figure 1-17 Palpation of an ovary per rectum. The ovary is

gently grasped with the finger tips, to enable the examiner to
identify raised or softened follicles. The rectum and abdominal
viscera have been removed to facilitate visualization.

Figure 1-15 Procedure used to locate cervix during palpa-

tion per rectum. A side-to-side motion is made with the pelvis, whereupon the hand is cupped and retracted.
extended hand in the pelvic cavity, with downward pressure The cupped hand “hooks” the uterus if the technique
with the fingers held together and the palm facing down- is properly applied and if the uterus is of a normal
ward. The rectum and abdominal viscera have been removed
to facilitate visualization. nonpregnant size and in the usual location. Enlarged
uteri that have begun descent into the lower abdomen
are difficult to identify with this method.
The leading edge of the uterus can be detected by The ovaries are located in the sublumbar area, cau-
first inserting the arm deeply into the rectum, cupping doventral to the corresponding kidney. By following
the hand downward, then slowly retracting the arm the leading edge of the uterus laterally and slightly
(Figure 1-16). Alternatively, the hand can be gently cranially, the ovary can usually be located with the
pressed downward as the arm is advanced into the rec- fingertips. Gently retracting the ipsilateral uterine horn
tum until a potential space is felt beyond the brim of the or broad ligament often helps bring the ovary to a
more readily palpable location. If the rectal wall is
relaxed, gentle grasping motions can be used to pick
up the ovary (Figure 1-17). The ovaries are pulled
downward and toward the midline when the uterus
becomes greatly enlarged (i.e., in the early postpartum
period, in advanced pregnancy [see Figure 7-11], or in
some cases of pyometra).
Once any of these landmarks is located, the practi-
tioner should proceed with a methodical examination
of the accessible reproductive tract. The examination
should include both ovaries, the uterus, and the cervix.
Structures caudal to the cervix do not lend themselves
to palpation per rectum but rather are evaluated via the
vulvar opening.

BIBLIOGRAPHY
Ginther OJ: Reproductive biology of the mare: basic and applied aspects,
ed 2, Cross Plains, WI, 1992, Equiservices, 1-40.
Greenhoff GR, Kenney RM: Evaluation of reproductive status of
non-pregnant mares, JAVMA 167:449-458, 1975.
Sisson S, Grossman JD: The anatomy of domestic animals, ed 4,
Philadelphia, 1953, Saunders, 606-614.
Figure 1-16 Palpation of uterine horns per rectum. The
cupped hand is used to grasp the leading edge of the uterine
horn. The rectum and abdominal viscera have been removed
to facilitate visualization.
 Reproductive Physiology of the
Nonpregnant Mare
2
C H A P TE R
O B J EC T I VES
While studying the information covered in this chapter, the reader should attempt to:
■ Acquire a working understanding of the physiology of the estrous cycle of the mare.

■ Acquire a working knowledge of the criteria used for staging the estrous cycle of the mare.

■ Acquire a working understanding of seasonal control of reproductive function of the mare.

S TUDY Q U EST I ON S
1. List the length of the estrous cycle, estrus, and 5. Describe the ultrasonographic changes that occur
diestrus in the mare. in the dominant follicles as ovulation approaches.
2. Explain the endocrine events that occur during 6. Describe the effects of long and short photoperi-
estrus, ovulation, formation of the corpus luteum, ods on control of reproductive function of the
and luteolysis in the mare. mare.
3. Explain why determination of the presence of a 7. Describe the seasonal effects on incidence of
large follicle on an ovary is a poor predictor of ovulation in mare populations in the Northern
estrus in the mare. Hemisphere.
4. List the criteria that are useful in staging the 8. Discuss effects of season on duration of estrus in
estrous cycle via palpation per rectum in the mare. the mare, including the first postpartum estrus
(foal heat).

THE ESTROUS CYCLE the mare is sexually receptive to the stallion, and the
genital tract is prepared to accept and transport sperm
The mare is a seasonal polyestrous animal. During the to the oviducts for fertilization) that involves the ovula-
breeding season (spring and summer), the nonpreg- tion process and a luteal phase (diestrus) (in which the
nant mare has recurring estrous cycles. The length of mare is not receptive to the stallion, and the genital
the estrous cycle can be defined as the period from one tract is prepared to accept and nurture the conceptus).
ovulation to a subsequent ovulation (interovulatory The diestrous period ends with regression of the cor-
interval). Each ovulation is accompanied by a period of pus luteum, initiation of the next follicular phase, and
behavioral signs of estrus while plasma progesterone onset of estrous behavior. The average length of the
concentrations remain low (⬍1 ng/mL). The estrous estrous cycle in mares during the physiologic breeding
cycle is divided into the ovulation process and an inter- season is 21 to 22 days (range, approximately 18 to
ovulatory period. The estrous cycle may also be consid- 24 days), with estrus comprising 4 to 7 of these days
ered to consist of a follicular phase (estrus) (in which (Figure 2-1). The length of diestrus remains relatively

10
 Reproductive Physiology of the Nonpregnant Mare ■
CHAPTER 2 11

Estrogen
FSH
LH
Progesterone
PGF2␣

Estrus Estrus

-6 -4 -2 0 2 4 6 8 10 12 14 16 18 20 1 3

Ov Ov

Figure 2-1 The estrous cycle of the mare averages 21 to 22 days, with 4 to 7 days of estrus (sexual receptivity) and 14 to
15 days of diestrus (during which the mare is not sexually receptive to the stallion). Ovulation generally occurs 1 to 2 days before
the end of estrus. (Modified from Irvine CHG: Endocrinology of the estrous cycle of the mare: applications to embryo transfer,
Theriogenology 15:85, 1981; and Neely DP, et al: Equine reproduction, Princeton, NJ, 1983, Veterinary Learning Systems Co)

constant at 14 to 15 days and is less affected by season presence of low circulating progesterone concentra-
than is the length of estrus. The length of estrus is more tion. Inhibin and estrogen produced by growing fol-
variable (range, 2 to 12 days, or more), typically with a licles have a negative feedback effect on release of FSH
longer duration early in the breeding season, perhaps (i.e., they inhibit FSH release). Progesterone produced
as the result of a less prominent luteinizing hormone by the CL has a negative feedback effect on release of
(LH) surge during this time period. The diameter of the LH from the anterior pituitary.
largest follicle at the time of luteolysis affects the inter-
val from onset of estrus to ovulation; larger follicles PHOTOPERIOD
present at corpus luteum (CL) regression typically
ovulate sooner, thus shortening the associated estrous
period. Increase Decrease
The regular pattern of the estrous cycle relies on the
delicate balance among hormones produced by the
pineal gland, hypothalamus, pituitary gland, ovaries, Retina
and endometrium (Figures 2-2 and 2-3). The neurosecre-
tory cells in the hypothalamus produce gonadotropin-
releasing hormone (GnRH). Axons of these cells proj- Pineal gland
ect into the perivascular space in the median eminence
at the origin of the pituitary stalk and episodically Melatonin
release GnRH into the hypothalamic-hypophyseal
Hypothalamus
(hypothalamic-pituitary) portal system, which trans-
ports the hormone to the anterior pituitary. GnRH GnRH
stimulates the synthesis and release of the gonadotro-
pin follicle-stimulating hormone (FSH) and LH from Pituitary gland
the anterior pituitary gland. These hormones enter the FSH
systemic circulation, and at the level of the ovaries, LH
FSH is responsible for follicular recruitment whereas Ovaries
LH is responsible for follicular maturation and pro-
duction of estrogen, ovulation, and luteinization. The Figure 2-2 A simplified version of hormonal regulation of the
estrous cycle of the mare. The regular pattern of the estrous
estrogen produced by maturing follicles (particularly cycle of the mare is controlled by the interplay among the
the dominant follicle) has a positive feedback on pineal gland, hypothalamus, pituitary, ovaries, and endome-
LH release (i.e., it promotes further LH release) in the trium. The role of the endometrium is depicted in Figure 2-3.
 12 CHAPTER 2 ■
Reproductive Physiology of the Nonpregnant Mare

of a group of follicles until one or perhaps two, or rarely

more, follicles begin preferential growth (i.e., become
dominant) to the remaining follicles. The selected follicle
continues to grow (⬎30 mm diameter) (Figure 2-5) until
GnRH it either ovulates or regresses. For mares with one fol-
licular wave during the estrous cycle, the wave emerges
at midcycle (approximately day 10 after ovulation). This

Pro LH
primary follicular wave results in an ultrasonographi-
ibin

ges
Est
cally identifiable dominant follicle approximately 7 days

FS
LH
H
en
Inh

tero
rog H
FS

H
before ovulation. For mares with two follicular waves
ro g

FS

en

ne
L
during the estrous cycle, the dominant follicle selected
Est

H
LH
FS

FS

LH

during the first follicular wave that begins during late

estrus or early diestrus sometimes ovulates (diestrous
ovulation). The incidence of secondary follicular waves
Follicle CL and diestrous ovulations is currently thought to be
higher in Thoroughbreds than in Quarter Horses and
ponies and may contribute (when ovulation occurs) to
longer diestrous intervals within a breed when com-
pared with mares of the same breed that do not have
diestrous ovulations.
At the onset of luteolysis, the largest follicle is typi-
PGF2 cally the one to enlarge and ovulate. Ovulation is a
rapid process, with most of the follicular fluid being
released within 2 minutes (50% to 90% evacuation
Estrus Diestrus
within 60 seconds) and with complete evacuation typi-
Figure 2-3 Schematic depiction of hormonal control of cally requiring 2 to 7 minutes. The remainder of the
ovarian activity in the mare. follicles that have already become atretic eventually
regress. In light breed horse mares, follicular diameter
at ovulation normally ranges from 30 to 50 mm (sel-
The follicular phase of the estrous cycle is character- dom ⬍35 mm) and usually approximates 40 to 45 mm,
ized by follicular growth with estrogen production, although smaller or larger follicles sometime ovulate.
which results in behavioral estrus (Figure 2-4). Many Smaller breeds (e.g., ponies and miniatures) tend to
follicles start the maturation process, but usually only ovulate smaller (ⱕ30 mm) follicles, and larger breeds
one follicle becomes dominant and ovulates. Follicular such as draft mares, especially Friesians, tend to ovu-
development typically occurs in one or two major waves late larger (⬎45-mm follicles). Ovulatory follicles are
during the estrous cycle. The term follicular wave has often larger early in the breeding season (March to
been used to describe the initially synchronous growth May) compared with those that ovulate in the peak of

Figure 2-4 Mare shows typical signs of estrus: squatting Figure 2-5 Cross-sectional view of mare ovary with large
with tail raised, urinating, and everting clitoris. preovulatory follicle.
 Reproductive Physiology of the Nonpregnant Mare ■
CHAPTER 2 13

the season (June and July). Most mares ovulate within

48 hours before the end of estrus; occasionally mares
ovulate after the end of estrus (usually on the day
when intensity of estrous signs is decreasing). The inci-
dence of double (multiple) ovulations averages 16%,
with Thoroughbred, Warmblood, and draft mares hav-
ing the highest incidence of multiple ovulations and
Quarter Horse, Appaloosa, and pony mares having the
lowest incidence.
The luteal phase is initiated at ovulation by the
formation of a progesterone-secreting CL (Figure 2-6),
which causes the mare to cease showing signs of
behavioral estrus (Figure 2-8). Maximum circulating
progesterone concentrations are reached by 6 days after
ovulation. A mare rarely shows behavioral signs of
estrus when plasma progesterone concentrations exceed
1 to 2 ng/mL, even when large follicles are present
on the ovaries. The life span of the corpus luteum is
dependent on the endogenous release of prostaglan-
din-F2 (PGF2) from the endometrium, which occurs
in bursts between days 13 and 16 after ovulation. The Figure 2-8 Mare shows typical signs of nonestrus: laid-back
PGF2␣ is absorbed into the uterine venous drainage ears, switching tail, and kicking.
and, unlike in the cow, enters the circulation and

reaches the ovaries primarily via systemic circulation.

Rapid luteolysis is caused by the PGF2␣, resulting in
a decline in circulating progesterone concentration
(a detectable decline occurs within 4 hours, with concen-
trations less than 1 ng/mL occurring within 40 hours of
the initial PGF2␣ release), which, in turn, removes the
block to LH secretion. Follicular maturation and behav-
ioral signs characteristic of the follicular phase of the
estrous cycle then ensue. Deviations (of several days) in
the length of the luteal phase are usually the result of
uterine disorders that lead to spurious secretion of
PGF2␣, such as acute endometritis (which shortens dies-
trus) or persistent luteal function (which prolongs dies-
Figure 2-6 Cross-sectional view of mare ovary. Corpus trus) from failure of the luteolytic mechanism (perhaps
luteum protrudes into ovulation fossa. (Photo courtesy of from failure of the endometrium to release sufficient
Dr. John Edwards.) amounts of PGF2␣ in late diestrus).

STAGING THE ESTROUS CYCLE VIA

Regressing
EXAMINATION
CL
Determination of the stage of the estrous cycle can be
Ovulation
fossa made via examination of the reproductive tract. Crite-
ria that are used for determining the stage of the cycle
Corpus include the size and softness of ovarian follicles and
hemorrhagicum
relaxation of the uterus and cervix, via palpation
per rectum. With the exception of determination of a
recent ovulation (i.e., typically within 0 to 3 days) (see
Figure 2-7), palpation for presence of a CL is not pos-
sible in the mare (e.g., the follicle ovulates into
the ovulation fossa and the CL does not protrude from
Figure 2-7 Cross section of mare ovary depicting recent the surface of the ovary). Use of transrectal ultrasonog-
ovulation (corpus hemorrhagicum). raphy is also helpful for staging the estrous cycle