8. SEXUAL DIFFERENTIATION. Germinal cells, gonocytes. Indifferent stage INDIFFERENT STAGE
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1 8. SEXUAL DIFFERENTIATION INDIFFERENT STAGE Early in pregnancy, (within 10-15% f the pregnancy s expected length) a genital ridge is frmed in the sides f the embrynic tissue, ventral t the mesnephrs r temprary kidneys. Initially, the genital ridge is made f celmic epithelium with mesenchyme tissue. Subsequently, this area is invaded by germinal cells riginating in the ylk sac (Figs. 8-1, 8-2). These clusters f cells are the indifferent gnads (Fig. 8-3). Germinal cells, gncytes Future gametes precursrs Originate in ylk sac Migrates t gnadal regin rgans Accumulates due t: Attractin by chemtactic substances Failure t survive elsewhere Figure 8-1. Generalities abut gncytes Nrmally the indifferent gnads will evlve int testis if the gentype f the cnceptus is XY and varies if the cnceptus is XX. The name indifferent gnads r biptential tissue is given because at this develpmental stage, regardless f the genetic makeup f the cnceptus, the gnads have the ptential t be either varies r testis. Up until recently, the understanding was that unless ther substances were present, these indifferent gnads were destined t becme female rgans. Recent discveries, hwever, reveal that the indifferent gnads have t be induced t becme either male r female rgans. Fllwing the establishment f the indifferent gnads, the embry develps tw pairs f ducts. One pair f mesnephric ducts, als knwn as the Wlfian ducts, which have the ptential t further develp int the epididymis, ductus deferens, and ureter f the male reprductive tract. Their riginal rle is t drain fluids frm the mesnephrs t the urgenital sinus. The ther pair is the paramesnephric ducts, als knwn as Müllerian ducts, which have the ptential t develp int the viducts, uterus, cervix and a prtin f the vagina (Fig. 8-3). Gnads begin Indifferent stage Pair f ducts capable f becming male rgans Pair f ducts capable f becming female rgans Underdevelped ptential external genitalia Nervus system t be primed Figure 8-3. Cnditins existing at the indifferent stage f develpment Figure 8-2. Origin f germinal cells prir t migrating t the area f the future indifferent gnads At this stage, the embry has a urgenital sinus, als knwn as the claca, with the ptential t develp int female r male external genitalia, as well as an incipient central nervus system, which culd be primed t functin in the pattern f a male r female animal. At this develpmental pint the embry is cnsidered t be in an indifferent stage (Fig. 8-4). V BS 122 Physilgy II 70 Class f 2016
2 The pre-sertli cells synthesize Anti-Müllerian Hrmne (AMH), als knwn as Müllerian Inhibiting Substance. AMH belngs t a superfamily f plypeptide hrmnes called Transfrming Grwth Factr beta (TGFbeta). AMH triggers the migratin f the epithelial cells t the central part f the mesnephrs. The mesnephric cells becme pre-sertli cells which surrund the germinal cells r gncytes. AMH als triggers the regressin f the Müllerian r paramesnephric ducts thrugh induced cellular apptsis (Figs. 8-5, 8-6, 8-7). Figure 8-4. Schematic representatin f the structures visible at the indifferent stage Male develpment The sex-determining regin Y gene (SRY), lcated in the Y chrmsme appears t be respnsible fr the initial rganizatin f the indifferent gnad int the male structures. The SRY gene is activated, by still unknwn mechanisms, during a very precise windw f time in the early develpmental perid. As a result f its activatin, a prtein named testis-determining factr is prduced and acts upn sme f the mesnephric cells in the indifferent gnadal regin. These cells then develp int pre-sertli cells (Fig. 8-5). Figure 8-6. Schematic representatin f the structures visible during early develpment in the male and the female Figure 8-5. Pathway leading t the establishment f male sex rgans and develpment f male like behaviur Figure 8-7. In late develpment the new metanephrs develps the accessry sex glands and the bladder V BS 122 Physilgy II 71 Class f 2016
3 Gncytes will be eventually cnverted t nn-mtile spermatgnia, which will divide by mitsis until stpped by the Sertli cells. The final utcme is the frmatin f the primitive sex crds which will then remain quiescent until puberty withut any further frmatin f spermatgnia. Simultaneusly, sme mesenchymal cells will differentiate int interstitial r Leydig cells. These cells will start t prduce teststerne (T), which in turn will be cnverted t dihydrteststerne (DHT) in the urgenital sinus. DHT is respnsible fr the virilisatin f the external genitalia and fr the frmatin f accessry secretry glands in males (Fig. 8-7). Female develpment If the indifferent tissue is nt expsed t AMH, the Müllerian r paramesnephric duct is nt reabsrbed as a cnsequence f induced apptsis. In the female anther gene, the DAX1, (dsage-sensitive sex reversal, adrenal hypplasia critical regin, n chrmsme X, gene 1) is activated at this time. This gene is lcated in the X chrmsme. The activatin f the DAX1 gene results in the develpment f the indifferent gnad int varies, the grwth f the paramesnephric r Müllerian duct int viduct and uterine structures (Figs. 8-6, 8-8). Female develpment Later than male's Germinal cells 6 t 7 X 10 6 at six mnths 1 t 4 X 10 5 at puberty 400 ptential va Figure 8-9. Generalities abut female develpment This time lapse permits a clear divisin between males and females and appears t allw fr the ptential manifestatin f the male characteristics. In the develping female there is a rapid prliferatin f germinal cells in the incipient vary. The number f germinal cells then decreases twards parturitin with a further decrease at the time f puberty (Fig. 8-9). Figure 8-8. Sequence f events leading t the develpment f females At the same time, the Wlffian r mesnephric ducts start atrphying. The metanephrs starts develping int the future kidneys and starts functining by 30% f the pregnancy, while the mesnephrs becmes the vary (Fig. 8-10). The changes frm the indifferent stage twards the establishment f female rgans start apprximately tw weeks after the change twards males (Figs. 8-8, 8-9). Figure Pictrial descriptin f the rgans in female late develpment Pseud hermaphrditism Pseud hermaphrditism refers t the cnditin whereby an animal has chrmsmes and gnads f ne sex and the external genitalia f the ppsite sex, r with cmpnents f bth sexes (Figs. 8-11, 8-12). V BS 122 Physilgy II 72 Class f 2016
4 Pseud hermaphrditism Result f a develpmental abnrmality where the genetic makeup f the gnads are f ne gender while the external and / r internal genitalia is f the ther gender, r an incmpletely differentiated rgan Figure Pseudhermaphrditism This abnrmality may be bserved in genetic females which have been expsed t elevated levels f andrgens during gestatin (Fig. 8-13). The surce f andrgens can be an abnrmality f the fetal adrenal r maternal secretins. In the case f males, pseudhermaphrditism may be triggered by a lack f sufficient teststerne prductin by the fetal testes. It can als be induced by a resistance r a lack f respnse f the urgenital sinus t andrgens. This resistance can be the cnsequence f a lack f andrgen receptrs in this tissue. The phenmenn can als be triggered by the failure in the cnversin f teststerne t DHT by the enzyme 5α- reductase (Fig. 8-14). Pseudhermaphrditism in a genetically male animal Male testicular abnrmality N MIS = female internal genitalia N T = female genitalia Andrgen resistance N DHT N receptrs Figure Pseud hermaphrditism in a pig. Tw descended testicles and a clear vulva are present Pseud hermaphrditism in a genetically female animal Female expsed t andrgens Adrenal abnrmality Maternal andrgens Figure Pssible causes f a female develping pseudhermaphrditism Figure Pssible causes f a male develping pseudhermaphrditism A cmmn ccurrence in cattle is the prductin f freemartins. A freemartin ccurs as a cnsequence f a translcatin f teststerne prduced by the bull calf int the circulatin f the heifer calf at the site f the ctyledns. This takes place because the chrin f bth cnceptuses anastmses is frming a cmmn bld supply (Fig. 8-15). The teststerne acts upn the reprductive tract f the female inducing the frmatin f male-like internal rgans, at the same time that the AMH causes partial regressin f the paramesnephric ducts f the female cnceptus. Usually, the result is an vtestis and a fragmented uterus lacking sme f its parts. The final utcme is a phentypic female that is sterile in 92% f the cases. Furthermre, the freemartin displays bullish-like V BS 122 Physilgy II 73 Class f 2016
5 behaviur because the expsure f the CNS t teststerne during fetal life has primed her t functin in a male-like fashin. A sund management practice is t treat any female brn as a twin t a bull, as a male and d nt keep it fr breeding. Freemartinism Sexual mdificatin f a female twin, by the in uter transfer f bld frm a male fetus Vascular anastmses between the placentas permits the transfer f bld frming and germ cells amng them 92% f female twin calves brn with a bull calf are freemartin A single brn female may be freemartin if the twin calf dies early in gestatin Figure Characteristics f a freemartin V BS 122 Physilgy II 74 Class f 2016
11. SEXUAL DIFFERENTIATION. Germinal cells, gonocytes. Indifferent stage INDIFFERENT STAGE
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