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    1.1. Introduction

    This chapter will deal with the reproductiveanatomy of the mare, detailing the indi-vidual structures along with their function.Further accounts may be found in othertexts, such as Ashdown and Done (1987),Frandson and Spurgen (1992), Ginther(1992), Kainer (1993), Dyce et al. (1996), Bone(1998), Senger (1999), Bergfelt (2000) andHafez and Hafez (2000). The reproductivetract of the mare may be considered as aY-shaped tubular organ with a series of con-strictions along its length. The perineum,vulva, vagina and cervix can be consideredas the outer protective structures, providingprotection for the inner, more delicate struc-tures, the uterus, Fallopian tubes and

    ovaries, responsible for fertilization andembryo development. Figure 1.1, taken afterslaughter, shows the reproductive structuresof the mare, and Figs 1.2 and 1.3 illustratethese diagrammatically. Each of these struc-tures will be dealt with in turn in the follow-ing account.

    1.2. The Vulva

    The vulva (Fig. 1.4) is the external area ofthe mares reproductive system, protectingthe entrance to the vagina. The outer area ispigmented skin with the normal sebaceousand sweat glands, along with the nerve andblood supply normally associated with theskin of the mare. The inner area is lined by

    1

    The Reproductive Anatomy of the Mare

    1.1. Introduction 11.2. The Vulva 11.3. The Perineum 3

    1.3.1. Protection of the genital tract 41.4. The Vagina 61.5. The Cervix 81.6. The Uterus 91.7. The Utero-tubular Junction 10

    1.8. The Fallopian Tubes 111.9. The Ovaries 11

    1.9.1. Follicular development and ovulation 121.10. Conclusion 15

    CAB International2003. Equine Reproductive Physiology, Breeding and Stud Management2nd edn (M.C.G. Davies Morel) 1

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    mucous membrane and is continuous with

    the vagina. The upper limit (the dorsal com-missure) is situated approximately 7 cmbelow the anus. Below the entrance to thevagina, in the lower part of the vulva (theventral commissure), lie the clitoris and thethree sinuses (ventral, medial and lateral)associated with it. These sinuses are ofimportance in the mare as they provide anideal environment for the harbouring ofmany venereal disease (VD) bacteria, such asTaylorella equigenitalis (causal agent for conta-gious equine metritis (CEM)), Klebsiella pneu-

    moniae and Pseudomonas aeruginosa. Hence,

    this area is regularly swabbed in mares priorto covering and, indeed, in theThoroughbred industry, such swabbing iscompulsory (McAllister and Sack, 1990;Ginther, 1992; Horse Race Betting LevyBoard, 2001). Within the walls of the vulvalies the vulva constrictor muscle, runningeither side of the length of the vulval lips.This muscle acts to maintain the vulval sealand to invert and expose the clitoral areaduring oestrus, known as winking(Ashdown and Done, 1987; Kainer, 1993).

    2 Chapter 1

    Fig. 1.1. The mares reproductive tract after slaughter and dissection (see also Fig. 1.3).

    kidney

    broad ligaments

    uterine bodyInfundibulum

    ovaryFallopian tube

    uterine horn

    sacral vertebra

    rectumanuscervixvaginavulva

    clitoris

    Fig. 1.2. A lateral view (from the side) of the mares reproductive tract.

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    1.3. The Perineum

    The perineum is a rather loosely definedarea in the mare, but includes the outer

    vulva and adjacent area, along with theanus and the surrounding area. In the marethe conformation of this area is of clinicalimportance, due to its role in the protectionof the genital tract from the entrance of air.Malconformation in this area predisposesthe mare to a condition termed pneu-movagina, or vaginal wind-sucking, inwhich air is sucked in and out of the vaginathrough the open vulva. Along with thispassage of air also go bacteria, which bom-bard the cervix, exposing it to unacceptablyhigh levels of contamination, which it is

    often unable to cope with, especially dur-ing oestrus when it is less competent.Passage of bacteria into the higher, moresusceptible parts of the mares tract mayresult in bacterial infections, such as CEM,and other venereal diseases, leading toendometritis. Chapter 19 gives furtherdetails on the causes of VD infections in themare, all of which adversely affect fertiliza-tion rates (Ginther, 1992; Easley, 1993;Kainer, 1993).

    Reproductive Anatomy of the Mare 3

    Fallopian tube

    uterine horn

    ovary

    broad ligaments

    uterine body

    cervix

    bladder

    vagina

    urethra

    vulva

    Fig. 1.3. The mares reproductive tract: a diagrammatic representation of Fig. 1.1.

    Fig. 1.4. The vulval area of the mare: in this

    instance the conformation of the perineal area is

    poor with the anus sunken cranially, opening the

    vulva up to faecal contamination.

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    1.3.1. Protection of the genital tract

    As mentioned previously, adequate protec-tion of the genital tract is essential to preventthe adverse affects of pneumovagina. Thereare three seals within the tract: the vulvalseal, the vestibular or vaginal seal and thecervix. These are illustrated in Fig. 1.5.

    The perineal area plus the vulva con-strictor muscle in the walls of the vulvaform the vulval seal. The vestibular seal isformed by the natural apposition of thewalls of the posterior vagina, where it sitsabove the floor of the pelvic girdle and thehymen, if still present. The tight musclering within the cervix forms the cervicalseal. This series of seals is affected by theconformation of an individual and also bythe stage of the oestrous cycle (Figs1.61.8).

    The ideal conformation is achieved if80% of the vulva lies below the pelvic floor.A simple test can be used to assess this. If asterile plastic tube is inserted through thevulva into the vagina and allowed to reston the vagina floor, the amount of vulva

    lying below this tube should be approxi-mately 80% in well-conformed mares. Thistechnique is illustrated diagrammatically inFig. 11.3.

    If the ischium of the pelvis is too low, thenthe vulva tends to fall towards the horizontalplane, as seen in Fig. 1.6. This opens up thevulva to contamination by faeces, increasingthe risk of uterine infection. Pascoe (1979a)suggested that mares should be allocated aCaslick index derived by multiplying theangle of inclination of the vulva with the dis-tance from the ischium to the dorsal com-misure. This index can then be used toclassify mares into three types and so predictthe likely occurrence of endometritis (Fig. 1.7).

    The effect of poor conformation of theperineum area may be alleviated by aCaslicks vulvoplasty operation, developedby Dr Caslick in 1937 (Caslick, 1937). Thelips on either side of the upper vulva are cut,and the two sides are then sutured together.The two raw edges heal together, as in thehealing of an open wound, and hence sealthe upper part of the vulva. The hole left atthe ventral commisure is adequate for urina-tion but prevents the passage of faeces intothe vagina (Fig. 1.9).

    The chance that a mare requiring aCaslicks operation will pass on the trait to

    her offspring is reasonably high. This, cou-pled with the fact that the operation site hasto be cut to allow mating and foaling, castsdoubt on whether such mares should be

    4 Chapter 1

    Fig. 1.5. The seals of the mares reproductive tract during dioestrus.

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    bred. Mares that have been repeatedly cutand resutured become increasingly hard toperform a Caslicks operation on, as the lipsof the vulva become progressively morefibrous and therefore difficult to suture. Insuch cases, a procedure termed a Pouret maybe carried out (Pouret, 1982). This is a moremajor operation and involves the realign-

    ment of the anus as well as the vulva.This perineal malformation is particularlyprevalent in Thoroughbred mares. It iscausal to both pneumovagina (collectionof air within the vagina) and urinovagina(collection of urine within the vagina), bothconditions being precursors for endometritisand hence infertility. The condition tends tobe exacerbated in mares that lack conditionand also in multiparous, aged mares. Its con-tinued existence is largely due to the selec-tion of horses for athletic performance rather

    than reproductive competence (Caslick, 1937;Pascoe, 1979a; Pouret, 1982; Le Blanc, 1991;Easley, 1993).

    The oestrous cycle also has an effect on thecompetence of the three seals. Further detailsof the effect of the oestrous cycle on thereproductive tract are detailed in Chapter 3.However, in summary, oestrus results in the

    slackening of all three seals, due to a relax-ation of the muscles associated with thereproductive tract, especially the cervix(Fig. 1.8). This allows intromission at cover-ing but also decreases the competence of thereproductive-tract seals and so increases thechance of bacterial invasion. In part this iscompensated for by elevated oestradiol levelscharacteristic of oestrus, which enhance themares immunological response, thus reduc-ing the chance of uterine infection, despitethe increased chance of bacterial invasion.

    Reproductive Anatomy of the Mare 5

    Fig. 1.6. The effect of conformation on the competence of the vulval, vestibular and cervical seals in themare. (a) A low ischium (pelvic floor) results in an incompetent vestibular seal. In this case the vulval seal

    is still competent; therefore, infection risk is limited. (b) A low ischium results in an incompetent vestibularseal. In this case, the vulval seal is also incompetent; therefore, infection risk is increased. (c) An incom-petent vestibular and vulval seal plus a sloping perineal area results in a significant infection danger,especially from faecal contamination.

    (a) (b)

    (c)

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    6 Chapter 1

    Fig. 1.7. (Left-hand column) A lateral view of the

    relationship between the anus, vulva and ischium,

    indicating: (a) Type I mare with good conformation,

    Caslick index < 50 (b = 23 cm, a < 10) no

    Caslick required. (b) Type II mare with poor confor-

    mation, predisposing to type III in later life, Caslick

    index 50100 (b = 67 cm, a = 1020) no imme-

    diate need for a Caslick but likely in later life. (c)

    Type III mare with very poor conformation, including

    vulva lips in a horizontal plane, Caslick index > 150

    (b= 59 cm, a 30) Caslick required immedi-

    ately, significant chance of endometritis and a reduc-

    tion in reproductive success.

    1.4. The Vagina

    The vagina of the mare is on average1823 cm long and 1015 cm in diameter. Inthe well-conformed mare, the floor of thevagina should rest upon the ischium of thepelvis and the walls are normally collapsedand apposed, forming the vestibular seal.The hymen, if present, is also associated with

    this seal and divides the vagina into anterior(cranial) and posterior (caudal) sections. Insome texts the posterior vagina is referred toas the vestibule. The urethra, from the blad-der, opens just caudal to the hymen. Withinthe body cavity, the vagina is mainly coveredby the peritoneum and is surrounded byloose connective tissue, fat and bloodvessels. The walls of the vagina are muscularwith a mucous lining; the elasticity conferredby the muscle layer allows the major stretch-ing required at parturition (Fig. 1.10).

    Fig. 1.8. The effect of oestrus on the competence

    of the vulval, vestibular and cervical seals in the

    mare: oestrus causes a relaxation of the seals and

    therefore an increase in infection danger.

    dorsalcommissure

    ischium

    vulva

    ventralcommissure

    anus

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    The vagina acts as the first protector andcleaner of the system, containing acidic to neu-tral secretions, originating from the cervix,which are bactericidal. The vagina itself isaglandular, except just cranial to the vulval

    lips, where secretory glands are located. Thesesecretions, however, have the disadvantage ofattacking the epithelial cell lining of thevagina, necessitating the protective mucouslayer, and of being spermicidal. Thus, at ejacu-

    Reproductive Anatomy of the Mare 7

    Fig. 1.9. A Caslick operation in the mare showing (a) the cutting of the vulval lips, (b) suturing and (c)

    the finished job.

    (a) (b)

    (c)

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    lation, sperm are deposited into the top of thecervix and/or bottom of the uterus, to avoidthe detrimental effect of the relatively high pHwithin the vagina. The exact composition ofvaginal secretion is controlled by the cyclicalhormonal changes of the mares reproductivecycle (Ginther, 1992; Kainer, 1993).

    1.5. The Cervix

    The cervix lies at the entrance to the uterus.It is a tight, thick-walled, sphincter muscle,acting as the final protector of the system.

    In the sexually inactive, dioestrous state, itis tightly contracted, white in colour andmeasures on average 68 cm long by45 cm in diameter; cervical secretion isminimal and thick in consistency. The mus-cle tone and therefore cervix size, alongwith its secretion, are again governed bycyclic hormonal changes. Muscle tonerelaxes during oestrus and there is anincrease in secretion, easing the passage ofthe penis into the entrance of the cervix.The oestrous cervix appears pink in colourand may be seen protruding or floweringinto the vagina (Fig. 1.11; Lieux, 1970).

    8 Chapter 1

    Fig. 1.10. The internal surface of the mares vagina (the coin measures 21 mm in diameter).

    Fig. 1.11. The oestrus cervix protrudes (or flowers) into the vagina of the mare.

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    The lining of the cervix consists of a seriesof folds or crypts, as shown in Fig. 1.12. Thesecrypts are continuous with the smaller foldsin the uterine endometrium and enable thesignificant expansion of the cervix required atparturition (Ginther, 1992; Kainer, 1993).

    1.6. The Uterus

    The uterus of the mare is a hollow muscularorgan joining the cervix and the Fallopiantubes (Fig. 1.12). This upper part of the tractincluding the uterus is attached to the lumbarregion of the mare by two broad ligaments,outfoldings of the peritoneum, on either sideof the vertebral column. The broad ligamentsprovide the major support for the reproduc-tive tract (Fig. 1.13) and can be divided into

    three areas: the mesometrium, attached to theuterus; the mesosalpinx, attached to theFallopian tubes; and the mesovarium,attached to the ovaries (Ginther, 1992).

    The uterus is divided into two areas, thebody and the horns. The body of the uterusnormally measures 1820 cm long and812 cm in diameter and divides into twouterine horns, which are approximately 25 cmlong and which reduce in diameter from 46cm to 12 cm as they approach the Fallopiantubes (Fig. 1.14). The size of the uterus isaffected by age and parity, older multiparousmares tending to have larger uteri. The uterusof the mare is termed a simplex uterus bipar-titus, due to the relatively large size of theuterine body compared with the uterinehorns (60: 40 split). This differs from that inother farm livestock, where the uterine hornsare the more predominant feature. The lack ofa septum dividing the uterine body is alsonotable (Hafez and Hafez, 2000). In situ theuterine walls are flaccid and intermingle withthe intestine, the only lumina present beingthose formed between the endometrial folds.

    The uterine wall (Fig. 1.13) consists of three

    layers: an outer serous layer (perimetrium)continuous with the broad ligaments, a cen-tral muscular layer (myometrium) and aninner mucous-membrane lining (endo-metrium). The central myometrial layer con-sists of external longitudinal muscle fibres, acentral vascular layer and internal circular mus-cle fibres. It is this central myometrial layer thatallows the considerable expansion of the uterusduring pregnancy and provides the force forparturition. The inner endometrium is arrangedin 1215 longitudinal (Fig. 1.12) folds and

    Reproductive Anatomy of the Mare 9

    Fig. 1.12. The internal surface of the uterus, illus-

    trating the endometrial folds of the uterine lining.

    lumbar vertebra

    broad ligaments

    serous layermyometrial layerendometrium

    Fig. 1.13. Cross-section through the abdomen of the mare illustrating the reproductive-tract support

    provided by the broad ligaments.

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    comprises luminal epithelial cells, stroma ofconnective tissue or lamina propria andassociated epithelial glands and ducts.Collagenous connective-tissue cores supportthese folds. The activity and therefore appear-ance of these endometrial glands are depen-dent on the cyclical hormonal changes. It isthis endometrial layer that is largely responsi-ble for supporting the developing conceptusand for placental attachment and develop-ment (Ashdown and Done, 1987; Ginther,1992, 1995; Kainer, 1993; Sertich, 1998).

    1.7. The Utero-tubular Junction

    The utero-tubular junction is a constrictionor sphincter formed by a high concentrationof muscle cells from the circularmyometrium of the Fallopian tube. The junc-

    tion, which appears as a papilla in theendometrium, separates the end of the uter-ine horns from the beginning of the Fallopiantubes (Fig. 1.15). Fertilization takes place inthe Fallopian tubes, and only fertilized ovacan pass through this junction and on to the

    10 Chapter 1

    Fig. 1.14. The convoluted Fallopian tube running through the mesovarian section of the broad ligaments,

    from the uterine horn on the right to the ovary on the left.

    Fig. 1.15. The utero-tubular junction in the mare, as seen from the uterine horn side (the dark colour of

    the uterine endometrium is not natural but allows easier identification of the utero-tubular junction).

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    uterus for implantation and further develop-ment. Fertilized ova appear to actively con-trol their own passage, possibly via alocalized secretion of prostaglandin E (PGE)(Ball and Brinsko, 1992), leaving the unfertil-ized ova on the Fallopian-tube side of thejunction. These then gradually degenerate(Ginther, 1992; Kainer, 1993; Fig. 1.14).

    1.8. The Fallopian Tubes

    The mare has two Fallopian tubes or oviductsof 2530 cm in length, which are continuouswith the uterine horns (Fig. 1.14). The diame-ter of these tubes varies slightly along theirlength, being 25 mm at the isthmus end,nearest the uterine horn, and increasing to510 mm at the ampulla, nearest the ovary.The division of the Fallopian tube between theisthmus and ampulla is approximately equal.The Fallopian tubes lie within peritoneal folds,which form the mesosalphinx part of thebroad ligaments. They have walls very similarin structure to the uterus, but thinner, com-posed of three layers: the outer, fibrous,

    serous layer, continuous with the mesos-alpinx; a central myometrial layer of circularand longitudinal muscles fibres; and an innermucous membrane. Fertilization takes placein the ampulla, a region lined with fimbriae(hair-like projections), which act to waftunfertilized ova into the ampulla to await thesperm and to waft fertilized ova out of theampulla and on towards the utero-tubularjunction. The ampulla of each Fallopian tubeends in the infundibulum, a funnel-like open-ing close to the ovary (Sisson, 1975).

    The infundibulum in the mare is closelyassociated with a specific part of the ovarytermed the ovulation fossa, which is unique tothe mare and is the only site of ova release; inother mammals ovulation may occur over thewhole surface of the ovary. The infundibulumis, therefore, relatively hard to distinguish inthe mare, not being so evident as a funnel-shaped structure surrounding the whole ovary.The infundibulum is lined, like the ampulla, byfimbriae, which attract and catch the ova, guid-ing them towards the entrance of the Fallopiantubes (Ginther, 1992; Kainer, 1993).

    1.9. The Ovaries

    The ovaries of the mare are both cytogenicand endocrine in function, producinggametes (ova) and hormones. They are evi-dent as two bean-shaped structures situatedventrally to (below) the fourth and fifth lum-bar vertebrae and supported by the mesovar-ium part of the broad ligaments. They makethe total length of the reproductive tract in themare in the region of 5060 cm. In the sexu-ally inactive stage, i.e. during the non-breed-ing season, the mares ovaries measure 24cm in length and 23 cm in width and arehard to the touch, due to the absence of devel-oping follicles. During the sexually activestage when the mare is in season, theyincrease in size to 68 cm in length and 34cm in width; they are also softer to the touch,due to the development of fluid-filled follicles(Fig. 1.16). Older, multiparous mares tend toshow larger ovaries up to 10 cm in length.

    The convex outer surface or border of theovary is attached to the mesovarian section ofthe broad ligaments (Fig. 1.16) and is theentry point for blood and nerve supply; the

    concave inner surface is free from attachmentand is the location of the ova fossa. The wholeovary is contained within a thick protectivelayer, the tunica albuginea, except for the ovafossa. The tissue of the ovary in the mare isarranged as the inner cortex (active gamete-producing tissue) and the outer medulla (sup-porting tissue). Ova release at ovulationoccurs only through the ova fossa, and all fol-licular and corpora lutea development occursinternally, within the cortex of the ovary(Witherspoon, 1975). The mare differs in theseaspects from other mammals, in which themedulla and cortex are reversed, ovulationoccurring over the surface of the ovary and allfollicular and corpora lutea developmentoccurring on the outer borders. Rectal palpa-tion, as a clinical aid to assess reproductivefunction in the mare, is not, therefore, as easyto perform as it is in other farm livestock, forexample, the cow. However, with the adventof ultrasound, reproductive assessment ofovarian characteristics in the mare is nowquite accurate (Ginther, 1992, 1995; Kainer,1993; Sertich, 1998; Hafez and Hafez, 2000).

    Reproductive Anatomy of the Mare 11

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    1.9.1. Follicular development and

    ovulation

    The ovary is made of two basic cell types:interstitial cells (stroma), which provide

    support; and germinal cells, which producethe ova. The number of potential ova con-tained within the female ovary is dictatedprior to birth; subsequently no addition tothe pool of ova can be made. These veryimmature ova are termed oogonia, andthere are many more than an individualwill use within her reproductive lifetime.These oogonia, with their full complementof chromosomes (64) and surrounded by asingle layer of epithelial cells, are termedprimordial follicles. At birth, the ovary con-tains many thousands of these primordialfollicles. After birth and prior to puberty,some oogonia start development to pri-mary oocytes, and these, surrounded bytheir epithelial, or granulosa, cells, undergothe first stages of meiosis. They then awaitpuberty, when hormone secretion from theanterior pituitary drives their furtherdevelopment.

    From puberty onwards, primary oocytesdevelop and complete the final stages ofmeiosis at varying rates, designed to ensurethat a regular supply of developed follicles

    is available for ovulation every 21 days dur-ing the breeding season. Not all primaryfollicles are destined to ovulate, as many arewasted along the way, degenerating andbecoming atretic; normally (in 75% of cases)

    just one reaches the stage ready for ovula-tion (Davies Morel and OSullivan, 2001).

    As the primary follicle develops, its sur-rounding epithelial cells differentiate intofollicular epithelial cells, which secrete follic-ular fluid, filling the cavity surrounding theoocyte. The follicle grows in size as fluidaccumulation increases. The primary oocytenow also increases in size and develops athick, outer, jelly-like layer, the zona pellu-cida; it is now termed a secondary oocyteand has a haploid number of chromosomes(32). The secondary oocyte now becomesassociated with one inner edge of the follicleand lies on a mound of follicular cells calledcumulus oophorus. The stroma immediatelysurrounding the follicle becomes organizedinto a double-lined membrane, the thecamembrane, the inner layer of which is vascu-larized, the outer layer not. The folliclescontinue to develop (normally to in excess of3 cm in diameter) and are termed Graafianfollicles (Fig. 1.17).

    This seems to be a critical stage inequine follicular development, for now a

    12 Chapter 1

    Fig. 1.16. The ovaries of the mare. Note the difference in size between the ovary on the left, which is

    inactive, and the one on the right, which is active. The concave surface (position of the ova fossa) and

    the convex surface (entry point for blood and nerve supply) of the ovary are clearly seen.

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    decision is made as to which of the devel-oping follicles is destined to develop fur-ther for ovulation and which ones willdegenerate. In 75% of cases only one follicleis destined to ovulate and hence develop

    further. In 23% of cases two follicles maydevelop further and ovulate, the remainderbeing triplets or more (Davies Morel andOSullivan, 2001). This decision seems to begoverned by the follicles ability torespond to elevated levels of circulatingoestrogen, follicle-stimulating hormone(FSH) and luteinizing hormone (LH),though the exact mechanisms are un-clear (Fay and Douglas, 1987; Roy andGreenwald, 1987).

    In those follicles destined to ovulate, fol-licular diameter increases and at the sametime they appear to move within the stromaof the ovary and orientate themselves toawait ovulation through the ova fossa.Ovulation of the mature follicle occurs intwo stages, which normally (99% of occa-sions) occur concurrently. The two stagesare follicular collapse and ova release. Thewhole process may take from a matter ofseconds up to a few hours, with ova releaseoccurring at the later stages of ovulation (J.Newcombe, Wales, 2001, personal commu-nication). The ova and follicular fluid are

    released through the ova fossa to be caughtby the infundibulum and passed down theFallopian tube for potential fertilization.Ovulation of follicles of diameter less thanor greater than 3 cm does occur but this is

    the exception (Sirosis et al., 1989; Gintherand Bergfeldt, 1993).

    After the release of the ova and follicularfluid, the old follicle collapses and the thecamembrane and remaining follicular epithe-lial cells become folded into the old follicu-lar cavity. Bleeding, from the theca interna,occurs into the centre of this cavity, forminga clot. This clot, the theca cells and anyremaining follicular epithelial cells make upthe corpus luteum (CL) or yellow body.Blood capillaries and fibroblasts theninvade the CL. It is initially a reddish-pur-ple colour. As the CL ages, it becomesbrowner in colour and, if the mare is notpregnant, regresses to yellow and thenwhite (corpus albicans) as it becomes non-functional. The luteal tissue is then gradu-ally replaced with scar tissue (Fig. 1.18;Vogelsang et al., 1987; Del Campo et al.,1990; Kainer, 1993; Pierson, 1993; Ginther,1995; Sertich, 1998; Hafez and Hafez, 2000).Figures 1.191.21 show sections through amares ovary, illustrating the presence ofdeveloping follicles and CL.

    Reproductive Anatomy of the Mare 13

    follicular fluid

    theca externa

    theca interna

    granulosa cells

    corona radiatazona pellucida

    vitelline membraneoocyte nucleus

    epithelial cells

    cumulus oophorus

    Fig. 1.17. The equine Graafian follicle.

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    14 Chapter 1

    stroma

    primordialfollicle

    fibrouscapsule

    oogonium

    corpus luteum

    ova fossa epithelial cells

    primary oocytes

    follicular fluid

    cumulus oophorus

    granulosa cellstheca membrane

    Fig. 1.18. Diagrammatic representation of follicular development and ovulation within the ovary.

    Fig. 1.19. Cross section taken through the two ovaries pictured in Fig. 1.16. Note in the active ovary the

    corpora lutea (dark mass at the top left) and the large follicle (hollow or space at the top right).

    Fig. 1.20. A cross section taken through an active ovary illustrating the presence of a large follicle (3 cm

    in diameter) at the bottom of the ovary.

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    1.10. Conclusion

    It can be concluded that the reproductive tractof the mare is a remarkable system, designed

    not only to maximize the chance of fertilizationand subsequent maintenance of the resultingconceptus in a sterile environment, but alsoto expel that conceptus successfully at term.

    Reproductive Anatomy of the Mare 15

    Fig. 1.21. A cross section taken through an active ovary illustrating a large corpus luteum at the top of

    the ovary.