and dynein. Sperm dynein, which is the principal protein in the arms of the axonemal microtubules, has been shown, 0 be a divalent cation-activated ATPase, Sex Chromosomes: X and ¥ Spermatozoa ‘The process of sperm formation in most mammals results in two types of spermatozoa relative to sex chromatin. Mam. malian males are heterogemetic in that one-half ofthe sper- matoroa contain an X-chromosome and the other half Y-chromosome. Of the two types of gametes produced by ‘mammalian males, spermatozoa carrying the X-chromosome produce female embryos upon fertilization of an oocyte, whereas spermatozoa carrying the Y-chromosome produce male embryos. The males of avian species, however, are omogametic in that they produce spermatozoa with only ‘one Kind of sex chromosome. Sex determination in birds ‘occurs with the egg. Although the difference in DNA content bemeen X-chromosome-bearing and Y-chromosome-bearing sper- matozoa in domestic livestock is only about 3 to 4%, this ‘small difference can be resolved using fluorescent staining and flow cytometric analyses. Furthermore, flow cytometer: ‘have been modified so that they can sort viable mammalian spermatozoe into relatively pure X and Y sperm populations. When these sorted sperm were inseminated into females che sex ratio of the progeny was nearly identical to that ‘predicted by the ratio of X-spermatocoa to Y-spermatozoa ip the flow-sorted inseminate. Considerable effore is being ed on developing this appruch as a practical means ining the sex of domestic livestock. seminiferous epithelium, lining the seminiferous tu- is composed of two basic cell types: the Sertoli cells the developing germ cells. The germ cells undergo 2 tinuous ceries cf cellular divisions and developmental beginning at the periphery and progressing towards lumen of the tubule (Fig, 7-4). The stem cals, called togonia, divide several times before forming sperma- ‘The spermatocytes then utalergo meiosis, thereby ing the DNA content of the cells to one-haif that of tie cells. This series of cellular divisions, including the iferation of the spermatogonia and the meiotic divisions, 26 spermavocytogeness. The haploid ccils resulting this process are called spermatids. The spermatids then 8 progressive series of seruccual and developmental 8 to form spermatozoa. These metamorphic changes Known as :permiogenesis. 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The “ars preleptotene sperm eytesfeom stages I, the Golgi phase (steps 1 to 3 the maturation phage (steps 13 1974:140167-180) Tmatids into cells to be released into the lumen of the iniferous tubule. The reshaping of the nucleus and aero. ne of each spermatid, initiated during the previous phace ces spermatozoa characteristic foreach species" pusleus, the chromatin granules undergo pr tdeneetion as the transivioral proceins are rep e-—— [The varios seeps in spermatogenesis in the ull hpianing particular cellular assacation of the 12 cell typer are: 3, tei leptotene spermatocyte 2 Sry ef candy spenmaroye: through I are ep of spermicgensschowicg the cop phase (ters and 14). (Adapted 10 8.86 A spermazoponium, sages of the cyele of the 1, B, successive stages of spermatogonia, PEs gotene spermatocyte; P, pachytene spermato, 4 %0 7) the acrosome phase (steps 8 to 12} and fiom Bemdton WE, Desjardins C. Am } Anat procamines as they lunufoemly fils form a fine homogeneous material that the entire sperm nucleus (Fig. 7-5, steps F fibrous sheath and che coarse fibers are formed around the axo. theme. 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The analogy breaks dows, gre ft tharthestem cele ivide many mes during the spermaoge, and primary spermarocite sages resulting in mote than a honieed formed sperm or graduating sensors, (A ps PT, ed, Reproduction in Bamescic Animale th of Soa Academic Press, 1991.) (Bouom) Isolated spermatogonia cel Mvorw Laven. The basement me = that nds FEontractile my..id cells (F wf che ceil jancuams of this Cuarrer 7: Srenuatozon ann Semivat Plasma 103 rey E1GURE 7-7. Loops of the seminiferous nubules,the rete testis, and the excurrent duet yen of the ram. The pathway by which spermore, 8 are transported to the exterior it indicated by tows, (Mediéed itom Setchell BP, In: Cole HH, Coppe PT, eds. Repredhetion in Domestic Animals. New York: Academie Press 1977) apposition of the adjacent cell membranes. This barrier; however, is not well developed in the bull, ram, or boa ard may be a relatively unimportant permeability barriee in the testis of farm animals. Senrots Ceut Junctions. The principal permeability barrier between the blood and testis is thought to be the complexes at junctions between adjacent Sertoli cells, These Serroli-Sertali junctions, which are situated neat the cellular tase, contain multiple zones of adhesion (tight junctions) where the opposing membranes are fused. The occluding Junetions divide the seminiferous tubules into two distince compartments: (1) a basal comparement containing sper ‘matogonia and preleptotenc spermatocytes and (2) an als tninal compartment, containing the more advanced stages of spermatocytes and spermatids, shich freely communicates with the lumen of the tubule Th compartment is freely accessible t0 compo- ents that have previously penetrated the myoid layer. The second barrier composed of the occluding junctions be- ‘ween Sertoli cells, demonstrates a wide range of permeabil om complete exclusion of some substances to early of others. This differential permeabiticy ap tant in maintaining an environment suitable on of the tubuies. 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The endocrine control of erticuar function in mam: ‘The hypothalams secretes gonsdoteopin, a hormone releasing se (GaRH), thac stiles the secetion of LH and FSH fore scerior pituitary. The LH stimulates the intesttial cells of Lay ce androgens, mainly restosterone. The androgens ate secre he bloodstream mhere they cause the development of secondary sEaracteritics of the male and development and maintenance of snsle reproductive wee. The androgens suppress GnRH, LH, and escretion by negative feedback onthe pituiteryand hyp. chalarns erone is also secreted ints the seminiferous tubule where its for maintenance ! spermatogenesis. The FSH interacts with 1s on the Sertoli cells to cause proction of anirogen-hinding SABP), conversion of testosterone a cihydrotestosterone and n, stimulation of the spermatoertogeness, completion of sperm (spermition) and secretion of inhibin. The inhibin secreted she bloodstrcam hus @ nepative feedhack ctiect on FSH bot ne BBE secretion, (From Kalrenback CC, Dunn TO. Endvetinology ot fection. Ini Hafes ESE. Repeodaction in Facin Animals deh ed elphia: Lea & Febiger, is.) (Citarren 7: Speumaroz0a AND SeMINaL PLaswa 105 sis in hypophysectomized rats, provided treatiient begins immediately after removal of the pituitary. Ocher species, however, require FSH in addition to the steroid for mainte, nance of spermatogenesis. Other pituitary hormones (e-¢., hormone, and thytoid-stimulating hoe. mone) may have secondary roles in support of testicular fanction. The testes not only produce the major androgen, testos- terone, but also a series of related steroid hormones. The ‘major action of androgens is on the Sertoli cells rather than, directly on the germ cells, The mayoid cells also appear to bbe androgen dependent. This steroid dependency is met by pulssele production of androgens by the interstitial Leydig cells, which are adjacent to the seminiferous tubules (Fig. 7-10), The Leydig cells are stimulated to secrete androgens bby pulses of pituitary LH. The androgens produced by the Leydig cells not only diffuse into the adjacent Sertoli cells but are secreted into the blood where they feed back both ac the hypothalamus and the pituitary to block release of additional LH (Fig. 7-10). The other principal gonadotzo- phin, FSH, stimulates production of ABP and inhibin by the Sercoli cell. ABP forms a complex with androgen and is carried along with the spermatozoa into the epididymis, ‘The epithelial cells of the epididymis require relatively high levels of androgen for normal function. Inhibin has « nega- tive feedback effect on FSH secretion but not on LH. Al- though much of the testosterone sccreted into the serninifer- ous tubules is converted into ditydrotestosterone (DHT) by the enzyme, Sa-steroid reductase, some of the testosterone is converted vo estrogens by the enzyme aromatase. A relatively high level of testosterone is required for spermmatid manu. ration. Sertoli and germ cells reciprocally regulate each other's cyclic secretion of proteins along the length ofthe seminifer- ‘ous tubles. This process is amplified by the mysid cells through transforming growdh factors, humoral modulators, and extsacellular matrix modification. Activin and inhibin, which are secreted by the Sertoli cells, have remarkable characteristics to generate a diverse ‘ceries of signals, Activins, potent FSH-teleasing dimers (dimers of inhibin B-subunits), have paracrine (inhibiting growth hormone and adrenocorticotzopin secretion), and autocrine (stimu: lating FSH secretion) mechanisms. Activin and inhibin also act within the gonads as autocrine and paracrine ‘modulators of the production of steroids, other hormones, and growth factors. Transforming giowth factor (TGF), 2 multifunctional may be involved with regulation of testicular function. Tio groups of compounds are invoived with regulation of the hypothalamo-pituitary-gonadal axis. These include (2) newrotransmitters, dopamine, serotonin, and norepi- J other peptides. 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They Ss their fectlcing ability, then their motility, and Shey disintegrate, Ejaculates collected after prolonged 2 high percentage of degenerating icance of seminal plasma is question- f= that pregnancy can be induced in some species ination with epididymal spermatozoa. It, however, fo be an essential component in natural mating it serves as a carrier and protector ofthe spermato, importance of this role varies because spermatoron lated directly into the urerus in some species (eg, ‘sod mare). Seminal plasma appears to be more impor, 8 of the ewe and cow where the ejacu including the testes, epididymides, and accessory ‘of the male (Fig. 7-11). The only accessory gland ‘0 all mammals is the prostate. The epididjznis or ‘tonal analogue and the vas deferens are the only ‘organs present in male birds and reptiles. Constituents of Seminal Plasma »lasma contains unusually high levels of citric acid, eins fructose, glycerylphosphorylcholine, and sor, (Table 7-1). Appreciable quantities of ascorbic act, cids, peptides, proteins, lipids, fatty aces, and nu enymes are also present. Antimicrobial constituents ing immunoglobulins ofthe IgA class are constituents Sst plasma. In addition, a variety of hormonal sub. including androgens, estrogens, prostaglandins, LH, chorionic gonadotrophin-like material, growth, relaxin, thyroid re- aad enkephalins have been detected in “Seoxile character of spermatozoa provides an terns of assessing their physiologic status. But by itself, is nor an accurate predictor of potentis! ‘aracity. The energy required gor motility isappar- Serived trom intracellular scores bf ATP (adenosine CHAnTER 7: Sronmatozoa AND Semintat PLasma 107 Es a5 | eh GLAND [7] TF GLAND ies SEMEN Fiours 7-11. Ejaculaed semen of most of in addition oa smal! amount of testicular uid, contibacions ‘gveral accesory organs including the epididymis (CAPUT CORPS EPID and CAUDA EPID), ampulary glands (AMP), vesicular land. prostate gland, and bulbourthral glands. The relative contribision of the glans vary noe only among species, but also among invades ‘ich @ species and among ejacuates from the sane mieal farm animals is compored axeunang, equssadso810) ssulsognay amare “008 -oxpod jenusiod areuruny2 “Caymnou FRe2129q ‘vonwiedaid uoneuruzsm susie] ‘sovpogqiue wiodsaue’areumint ‘=n a10)99 passayy pe pauses Joypour uupds ao] “quadsooce84Q aus ‘mpets toon Jatpo 20 "uONN| Tues ponsardoG) 08 es spony “upoU OT SEH wonenaela apenloney Paugpoyy “@>HIOST MHA “wIfOrg men uareds (so) sensseus wads oeusepdo ‘pros aun my susie yo aomeynsnueworsyy wonenipsgjennyy onmirede> ant uy tanop mss pue dn ins dai 20) vonendnew 830!9g agen’ wsods 2se2:0u) uumop wig Soygonb unas 3563294] ‘wads Jo woR=P>g din ways ead ‘wou syuauoduno> ayog yea Jo sso] tsasard sosoooud 2100 Tsuon aanoe ar], soueaquiows yeoxonEuLsas jo sesso niodsuen anne ap jo fay axp ures 1 pass atos “foypiow jo sovosd Burunsuoo-ABouo ofp 205 Pon st LV oxp 30 ot Winowp|y “yLV OTe ABuoue axp jo 2800 2A sooo eororeunnds ap “ossaooxd aqoqene9 54H BUSES ‘oom texp ASsous Jo uoranpoxd ay uF aeIayP 2008 Age “roprstoo st etpuoyonn atp wm paaeooy # Yor ‘Seawped aanopixo sm Tate pure appxcip vox ptt 0 soroMI 30 Tunopezeang ap io1y Amynsor azeanakd 20 sxe] ap Busn jo sueaw atp sepisomd Garanse drovesdsex stay “uaHAxo jo souasaid axp Ut sovensgns Jo Sate © osn EozowEULIOdS, uopmadsoy ‘yoneuwesyy JE}OyAA UI 9sn 10} voxOLUIIEdS Jo esos Suuunp weeuod} s| sRsueDeIey> sty] “suOMpUCD Digorseue 2tp r2pun aajazns or eoroxeweds smoyye ‘resins purse yediouud yp st asorony asnevaq fatanoe onAjoronA Apoasiog ayow 10 “ayanse omsjoosiS sty] ‘poe INDE] OF “yget 2abgag 9 21 RENN Pe 9 “TEAREY wey UI voNDRPOHI [pe 35a SVEN 1 pH wR “OR6I "ABTA 9 OT ANSPPUINA FO PF HUEY wry oF woponpoNsy pe aS HAH “MOM MOY PRAY se-so go-¥0 sone ut aaaed zo-s00 s0-¥o oso st aid ening 5-90 wt sues zI-01 Best 0-100 se 6 06-06 €0-0ro sous ad s1-10 oor-ve cos w-08 ort oreo sea #9 rI-80 oe cos sta zo-t0 ost-oor ost ws (re w3401) (qu) suvinoveg Gy mE SG Spun NOULYEINIONOD s0-9T0A, ews {ar onaamug 40 ONINNION, NoLLneoway Jo ,OTCISAHE HT LIVd BOTthe sperm cel. Inthe absence of exogenous substrates, sper ‘smatozoa use their intracellular stores of plasmalogen to pro- vide energy on a short term basis. IMMUNOLOGIC ASPECTS OF SPERMATOZOA Antigenicity of Spermatozoa ‘An important function of the blood-testis barcier is the immunologic isolation of developing gametes. Its impor- tance is thar the spermatocytes, spermatids, and spermatozoa are readily recognized as forcign cells by the immune system ‘ofthe adule male. Thus, sequestering of the developing germ cells behind an immunologic barrier prevents an adult male fiom developing antibodies against his own sperm, Autoimmunity Immunization of a male against spermatozoa or isolated spermatozoal antigens results in varying degrees of autoim- ‘mune orchitis (inflammation of the testis) and, in some cases, cesation of spermatogenesis. As.aresult of autoimmu- nity, the concentration of antispermatozoal antibodies ean be high in the seminal fluid. Such spermatozoal specific antibodies can interfere with spermatozoal_ functioning thereby reducing male fertility. In more severe cases, damage to the blood-testis barrier or the epididymal portion of the excurrent duct system through traumatic injury or infection often results in autoimmune orchitis affecting both testes. Vasectomy, the transectioning and ligation of the vas deferens, also resulis in varying degrees of ‘esticular inflammation. The severity of the orchitis de- Pends on the species and the postoperative time. The autoimmune response of vasectomized males is thought x occur because intermittent distention and rupture of ligated duct results in release of spermatozoa into thé In Vitro EVALUATION OF SEMEN Males should be evaiuaced for seminal quality before use as breeding animals (Table 7-2). Once the semen has been collected from the male, several approaches may be used to ases the quality of the sample. Classic semen evaluation sechniques, as well as new automated semen assessment fechnologies, yield oniy estimates of potential fertilizing ‘capacity of spermatozoa, CHAPTER 7: SPERMATOZOA AND StMINAL PLasma 109. ASSISTED REPRODUCTIVE cs TECHNOLOGIES Various techniques for micromanipulation of spermatozo (Table 7-3) ean be used to enhance the reproductive poten’ tal of genetically superior sires. In vitro fertilization (IVE) procedures can be used if sperm numbers are insufficient for fercilization to take place within the female tract. Another technique isto physically insert a single spermatozoon into an appropriately matured oocyte using micromanipulation.. This procedure, which is called intracytoplasmic sperm in- ection (ICSI), can overcome a variety of male infertility problems. The application of this method remains contro- versial in that the actual fertilizing sperm is selected by- ‘Passing the screening systems inherent to the female repro- ductive tact that prevent defective spermatozoa from fertilizing oocytes. Considerable laboratory skill is required when manipulating the gametes in vito, SuGGESTED READING ‘What is semen? How does semen analysis asist in understanding the reproductive stas ofthe male? In: Robaire B, Pryor Jl ‘Trasler JM. eds. Handbook of Andeology. Am Soc Andrology, ‘Amman RP, Hatemersted¢ RH. nitro evaluation ofspenm quality: ‘An opinion. J Androl 1993;14:397-406, Dados: JP, Demoulin A. Structure and functions of the zestis. In: ‘ThitaultC, Levasseur MC, Hunter RHF, eds. Reproduction in Mammals and Man. Part: Ellipses, 1993. Gamer DL. Ancillary tests of bull sperm semen quality. In: Van Camp SD, ed. Veterinary Clinics of North America: Food ‘Animal Practice. 1997513:313-330, Carer DL. Arefcial Insemination. In: Cupps PT, ed. Reproduc- ton in D.smestie Animals 4th ed. San Diego, CA: Academic Press, 1991. Johnson L. Spermatogeness. In: Cupps PT, ed. Reproduction in Domestic Animals. 4th ed. San Diego, CA: Academie Press, 1991, Laraming GE. Marshall's Physiology of Reproduction: Vol. 2, Re- production in the Male. New York: Churchill Livingstone, 1990. ‘Mann T, Lurwak-Mann C. Male Reproductive Function and Se- men. New York: Springer-Verlag, 1981 (Otc RS. Breeding soundness of bulls. fn: Current Therapy in Theri ceenology: Diagnosis, Treatment and Prevention of Repro. ductive Diseases in Small and Large Animels. Philadelp hi: WB Saunders, 1986:125-136, ‘Senger FL. Pathways te Pregnancy and Parturition, Pullman, WA, ‘Current Conceptions, 1997,