Você está na página 1de 70

Fisiologia Endcrina

The Ages of Man

http://www.get-back-on-track.com/en/professionals/00_meta/07_praesentationen/p_con_0007_03_00.php

Endocrinologia da reproduo masculina


REVISO DAS ESTRUTURAS DO APARELHO REPRODUTOR MASCULINO ADULTO
Profa. Dra. Cristina Maria Henrique Pinto
CFS/CCB/UFSC Junho de 2006

O APARELHO REPRODUTOR MASCULINO ADULTO


Aes da Testosterona: aptido para a penetrao e fecundao
Urinary bladder

Pubic symphysis

Saladin, Anatomy and Physiology, cap. 27, 2002

As glndulas acessrias

e a composio do smen

GLNDULAS ACESSRIAS E A COMPOSIO DO SMEN O smen composto aproximadamente por 10% de esperma e fluido testicular, 30% de secrees da prstata e 60% de secrees das vesculas seminais.

Saladin, Anatomy and Physiology, cap. 27, 2002

GLNDULAS ACESSRIAS E A COMPOSIO DO SMEN O smen composto aproximadamente por 10% de esperma e fluido testicular, 30% de secrees da prstata e 60% de secrees das vesculas seminais.

Secrees das vesculas seminais incluem frutose, enzima coagulante e prostaglandinas, dentre outras. A frutose o substrato energtico para os sptz. A enzima coagulante favorece que o smen se torne uma secreo agregada, o que ajuda sua propulso pela vagina. As prostaglandinas diminuem a viscosidade do muco cervical e estimulam a peristalse reversa do tero.

Saladin, Anatomy and Physiology, cap. 27, 2002

GLNDULAS ACESSRIAS E A COMPOSIO DO SMEN O smen composto aproximadamente por 10% de esperma e fluido testicular, 30% de secrees da prstata e 60% de secrees das vesculas seminais.

Secrees da prstata: incluem citrato, fibrinolisina, Ccio, Zinco, fosfatase cida, dentre outros. O citrato fonte energtica. A fibrinolisina age como um anticoagulante do smen, o que ajuda na mobilidade do esperma. pH alcalino (neutralizao do pH
cido do lquido tub. seminferos)

Saladin, Anatomy and Physiology, cap. 27, 2002

GLNDULAS ACESSRIAS E A COMPOSIO DO SMEN O smen composto aproximadamente por 10% de esperma e fluido testicular, 30% de secrees da prstata e 60% de secrees das vesculas seminais.

Secrees das glndulas bulbouretrais


secretam muco lubrificante que contm galactose, dentre outros. So emitidas antes da ejaculao. Saladin, Anatomy and Physiology, cap. 27, 2002

GLNDULAS ACESSRIAS E A COMPOSIO DO SMEN O smen composto aproximadamente por 10% de esperma e fluido testicular, 30% de secrees da prstata e 60% de secrees das vesculas seminais.

Tipicamente , 2-5 mL de smen so expelidos por ejaculao. Cada mL de smen pode conter 50-130 milhes de sptz

Saladin, Anatomy and Physiology, cap. 27, 2002

Espermatognese
Reviso dos tipos celulares dos tbulos seminferos e do espao intersticial

VISO MICROSCPICA DO TESTCULO

http://academic.pgcc.edu/~aimholtz/AandP/206_ONLINE/Repro/malerepro1.html

Tipos celulares nos tbulos seminferos

clulas reprodutoras
(espermatognias, espermatcitos, espermtides e espermatozides)

e clulas sustentaculares (Sertoli)

http://academic.pgcc.edu/~aimholtz/AandP/206_ONLINE/Repro/malerepro1.html

TBULO SEMINFERO E TIPOS CELULARES

membrana basal

clulas sustentaculares (Sertoli)

espermatognias

espermatcitos primrios

espermatcitos secundrios

espermtides

Seminiferous tubules form the mass of the testes and are the sites of spermatogenesis. Seminiferous tubules are composed of a thick layer of spermatogenic cells (most numerous) and sustentacular (Sertoli) cells which rest on a basement membrane. The spermatogenic cells-spermatogonia, primary spermatocytes, secondary spermatocytes, spermatids and spermatozoa--represent different cell stages in spermatogenesis (setas amarelas). The outlines of sustentacular cells are not distinct. Maturing spermatozoa are found embedded, head first, in the sustentacular cells, which provide mechanical support, protection and possibly nutrition for the developing spermatozoa.

http://trc.ucdavis.edu/mjguinan/apc100/modules/Reproductive/mammal/testis3/test is6.html

ESPERMATOGNESE

Espermatognese
o processo pelo qual as clulas-tronco se desenvolvem em espermatozides maduros. Existem 3 fases: (1) Espermatocitognese, (2) Meiose, and (3) Espermiognese.

1. Espermatocitognese (tambm chamada mitose): clulas-tronco (espermatognia do Tipo A) dividemse mitoticamente e produzem as clulas que ir se diferenciar (espermatognia do Tipo B).

compartimento basal

http://www.cvm.okstate.edu/instruction/mm_curr/histology/MR/HiMRP4.htm

Espermatognese
o processo pelo qual as clulas-tronco se desenvolvem em espermatozides maduros. Existem 3 fases: (1) Espermatocitognese, (2) Meiose, and (3) Espermiognese. 2. Meiose: Clulas em prfase da 1 diviso meitica so os espermatcitos primrios. Estes, ao completarem a diviso meitica so chamados espermatcitos secundrios. Rapidamente ocorre a 2 diviso meitica, originando as espermtides.

compartimento basal

compartimento apical ou adluminal

http://www.cvm.okstate.edu/instruction/mm_curr/histology/MR/HiMRP4.htm

Espermatognese
o processo pelo qual as clulas-tronco se desenvolvem em espermatozides maduros. Existem 3 fases: (1) Espermatocitognese, (2) Meiose, and (3) Espermiognese. 3. Espermiognese: a metamorfose das espermtides esfricas a espermatozides alongados. Durante a espermiognese formado o acrossoma e o flagelo.

compartimento basal

compartimento apical ou adluminal

http://www.cvm.okstate.edu/instruction/mm_curr/histology/MR/HiMRP4.htm

Espermiognese
As mudanas durante a espermiognese envolvem transformaes da espermtide esfrica a espermatozide maduro: (1) formao do acrossoma, (2) mudanas nucleares, (3) desenvolvimento do flagelo, (4) reorganizao do citoplasma e organelas celulares e (5) o processo de liberao da cl. de Sertoli (espermiao).

http://www.endotext.org/male/male1/maleframe1.htm

Espermiognese
As mudanas durante a espermiognese envolvem transformaes da espermtide esfrica a espermatozide maduro: (1) formao do acrossoma, (2) mudanas nucleares, (3) desenvolvimento do flagelo, (4) reorganizao do citoplasma e organelas celulares e (5) o processo de liberao da cl. de Sertoli (espermiao).

http://rbp.fmrp.usp.br/didatico/Embriologia-Propedeutico/embrio__propedeutico%202004_tema1.PDF

Veja mais sobre Gametognese

Online course in embryology for medicine students developed by the universities of Fribourg, Lausanne and Bern (Switzerland) with the support of the Swiss Virtual Campus

veja tambm: dicas de WEBsites de Embriologia

Espermatognese

http://academic.pgcc.edu/~aimholtz/AandP/206_ONLINE/Repro/malerepro1.html

EPIDDIMO

http://fisiologia.med.up.pt/teoricoprtc.html

Epiddimo

EPIDDIMO

http://microanatomy.net/Male_Reproductive/Lecture_40_Male_Reproductive_Childs_4_slides_per_page.pdf

CAPACITAO DO ESPERMATOZIDE
ALTERAO FUNCIONAL DO ESPERMATOZIDE QUE OCORRE NA TROMPA UTERINA. REQUER UM TEMPO PARA QUE OCORRA (2 OU AT >6 HORAS)

PELO MENOS DOIS FNOMENOS SO IMPORTANTES: -O AUMENTO DA TAXA DE BATIMENTO DO FLAGELO E A ACELERAO DO MOVIMENTO DO ESPERMATOZIDE

-REAO ACROSSMICA NO SPTZ QUE PERMITE A FUSO COM O OVO: FRAGMENTAO E PERDA DO ACROSSOMA COM A LIBERAO DE ENZIMAS E PROTEASES QUE PERMITIRO AO SPTZ PENETRAR E SE FUNDIR AO OVO.

O ENTENDIMENTO DESTES MECANISMOS IMPORTANTE PARA A FERTILIZAO IN VITRO

A fisiologia das clulas testiculares envolvidas na espermatognese

As clulas testiculares envolvidas na espermatognese

parcrina

Clulas de Leydig

Clulas de Sertoli

parcrina

Clulas germinativas

parcrina

T
Clulas miides

testculo

T: testosterona

Clulas de Sertoli e as clulas espermatognicas


As clulas germinativas no possuem receptores para T e FSH. Quem os possui so as clulas de Sertoli que, sob influncias desses hormnios, propiciam a espermatognese.

http://www.cvm.okstate.edu/instruction/mm_curr/histology/MR/HiMRP4.htm

ntima relao entre as clulas de Sertoli e as clulas espermatognicas

Cross-section of a seminiferous tubule from an adult Sprague-Dawley rat showing the organization of testicular cells and the intimate relationships between Sertoli and germ cells. S, Sertoli cell nucleus; SG, spermatogonium; PS, pachytene spermatocyte; RS, round spermatid; ES, elongated spermatid. Cheng and Mruk 2002, Physiological Reviews

Clulas de Sertoli e as clulas espermatognicas Em todos os estgios de diferenciao, as clulas espermatognicas esto em ntimo contato com as clulas de Sertoli que provem a estas: -suporte estrutural (microtbulos) -suporte metablico (fornecem lactato para spct 1) -regulao do meio interno dos tbulos seminferos (formam a barreira hemato-testicular,
fagocitose)

-secretam protenas: transportadoras de nutrientes (Fe, Cu e Vit. A) ABP (ptn ligante de andrgenos), dentre outras. As cls. de Sertoli so tambm endcrinas: AMH (diferenciao sexual) Inibina B e Ativina (regulao da secreo de FSH).
http://www.cvm.okstate.edu/instruction/mm_curr/histology/MR/HiMRP4.htm

Clulas de Sertoli e as clulas espermatognicas BARREIRA HEMATO-TESTICULAR (junes firmes ou do tipo tight)

-comunicao entre vrias clulas de Sertoli -proteo para as clulas em desenvolvimento contra a resposta auto-imune. -isola as clulas germinativas do resto do organismo

traador eletrondenso no atravessa alm do compartimento basal

http://microanatomy.net/Male_Reproductive/Lecture_40_Male_Reproductive_Childs_4_slides_per_page.pdf

Clulas de Sertoli e a migrao das clulas espermatognicas O nmero de clulas de Sertoli determina a taxa mxima de produo de espermatozides

A schematic drawing that illustrates extensive changes in tight junction (TJ) and cell-cell actin-based adherens junction (AJ) dynamics during spermatogenesis and spermiogenesis in the mouse. This figure was prepared based on reviews and reports cited in sections III and V. Among the AJs in the testis, four functional complexes are known to exist to date, which include cadherin/catenin complex, nectin/afadin complex, tubulobulbar complex, and ES (see sect. V). The ES is composed of basal and apical ES constituted possibly by 6 4- and 6 1-integrins, respectively (); however, their binding partner(s), if any, is not known. It is possible that laminin 1 1 1-chains and 3-chains constitute the binding partners for the basal and apical integrins in the ES, respectively () (see sect. VC1). While it is certain that 6 1-integrins are found between Sertoli cells and developing spermatids in the apical ES (, it remains to be determined if 6 4-integrins can be found between Sertoli cells and developing spermatocytes and spermatogonia (type B) in the basal ES, or it is restricted only to the interface of Sertoli cells and the basement membrane. ES, ectoplasmic specialization, a modified testis-specific AJ. Physiological Reviews, Cheng and Mruk 2002

Sertoli secreta protenas ligantes de hormnios sexuais esterides


ABP Ptn ligante de testosterona/andrgenos biodisponibilidade parcrina (100x+)

Outra protena ligante importante (secretada pelo fgado): SSBG Globulina ligante de esterides sexuais biodisponibilidade perifrica Crystal structure of the N-terminal
domain of human SSBG in complex with dihydrotestosterone.

por estrgenos e hormnios tireoideanos


por andrgenos, glicocorticides, GH e insulina
http://www.scielo.br/scielo.php?script=sci_arttext&pid=S0004-27302002000500002&lng=en&nrm=iso http://www.mdc-berlin.de/~crystal/Structures/images/sex_2.html

As clulas testiculares envolvidas na espermatognese Clulas intersticiais ou de Leydig secretoras de Testosterona

parcrina

Clulas de Leydig

Clulas de Sertoli

parcrina

Clulas germinativas

parcrina

T
Clulas miides

testculo

As clulas testiculares envolvidas na espermatognese Clulas intersticiais ou de Leydig secretoras de Testosterona

http://www.cvm.okstate.edu/instruction/mm_curr/histology/MR/HiMRP4.htm

Regulao endcrina e parcrina da espermatognese

Fig. 46-16 Interactions among the various cells of the testis in the hormonal regulation of spermatogenesis. Berne et al., 2004

Regulao endcrina e parcrina da espermatognese


NVEIS BAIXOS DE FSH NVEIS ELEVADOS DE FSH

INIBINA B
por influncias parcrinas dos espermatcitos sobre as clulas de Sertoli

INIBINA B

Testosterona

clulas de Leydig

LH

Endocrine and paracrine regulation of spermatogenesis. The preprogrammed development of sperm from spermatogonia is facilitated and maintained at a basal rate by T (testosterone) effects on Sertoli cells that are transmitted to and elicited by germ cells at various stages. The basal rate of production is augmented by FSH (follicle-stimulating hormone). When the rate is too low, signals from various stages of sperm development (B, spermatogonium B; P, primary spermatocyte; S, spermatid; Spz, spermatozoa) decrease output of inhibin B from the Sertoli cell. This results in increased FSH secretion from the pituitary gland. Likewise, when spermatogenesis is too brisk, Sertoli cells release more inhibin B, which suppresses FSH secretion and turns down the rate of spermatogenesis. Berne et al., 2004

Berne et al., 2004

Terapia com Testosterona pode diminuir a produo de espermatozides


Injees de testosterona podem suprimir a secreo de FSH e LH, afetando a produo de esperma. Notar a recuperao da contagem de esperma quando hCG e hFSH so utilizados em uma nica dose.
http://www.endotext.org/male/index.htm

Os hormnios sexuais masculinos

Os hormnios sexuais masculinos: Testosterona e derivados

LH
20,22-desmolase 17-hidroxylase

VIAS DE SNTESE DE ESTERIDES NAS CLULAS DE LEYDIG

Pathways of synthesis of gonadal steroid hormones. Testosterone is the major secretory product of the testis. Estradiol and progesterone are the major secretory products of the ovary. Enzymes are
(1) (2) (3) (4) (5) (6) (7) (8) 20,22-desmolase 17-hydroxylase 17,20-desmolase 17-OH-steroid dehydrogenase 3-ol-dehydrogenase and 4,5-isomerase 5-reductase 3-reductase aromatase

17,20-desmolase

VIAS DE SNTESE DE aromatase ESTERIDES NAS CLULAS-ALVO


17-OHesteride desidrogenase 3-OHSD aromatase

5-reductase 2

3-reductase

P-450 SCC: protena reguladora aguda esteroidognica 3-OHSD: 3-OH-steroid dehydrogenase

Berne et al., 2004

http://www.endotext.org/male/index.htm

Hormnio luteinizante (LH)

estrona

Secretion
17 estradiol testosterona

testosterona

veja legenda no prximo slide

diidrotestosteron a
diidrotestosterona

http://www.endotext.org/male/male2/male2.htm

Legenda da Figure anterior Pathways of testosterone biosynthesis and action. In men, testosterone biosynthesis occurs almost exclusively in mature Leydig cells by the enzymatic sequences illustrated. Cholesterol originates predominantly by de novo synthesis pathway from acetyl-CoA with luteinizing hormone regulating the rate-limiting step, the conversion of cholesterol to pregnenolone within mitochondria, while remaining enzymatic steps occur in smooth endoplasmic reticulum. The D5 and D4 steroidal pathways are on the left and right, respectively. Testosterone and its androgenic metabolite, dihydrotestosterone, exert biological effects directly through binding to the androgen receptor and indirectly through aromatization of testosterone to estradiol, which allows action via binding to the ER. The androgen and ERs are members of the steroid nuclear receptor superfamily with highly homologous structure differing mostly in the C-terminal ligand binding domain. The LH receptor has the structure of a G-protein linked receptor with its characteristic seven transmembrane spanning helical regions and a large extracellular domain which binds the LH molecule which is a dimeric glycoprotein hormone consisting of an a subunit common to other pituitary glycoprotein hormones and a b subunit specific to LH. Most sex steroids bind to sex hormone binding globulin (SHBG) which binds tightly and carries the majority of testosterone in the bloodstream.
http://www.endotext.org/male/male2/male2.htm

Aes diretas e indiretas da Testosterona


regulao das gonadotrofinas espermatognese

LH

T
Testculos

T
DHT

diferenciao sexual: estimulao ducto de Wolff virilizao externa des. prstata maturao sexual na puberdade

T
5-Reductase

DHT

AR: receptor de andrognios

Clula-alvo

Manuteno funcional dos rgos sexuais e dos caracteres sexuais 2rios

T: testosterona; DHT: diidrotestosterona

rgos acessrios, pele da regio genital e folculos pilosos Wilson, 2003

Aes diretas e indiretas da Testosterona


testculos, ossos e SNC regulao das gonadotrofinas
aromatase

LH E2 E2

espermatognese diferenciao sexual: estimulao ducto de Wolff virilizao externa des. prstata maturao sexual na puberdade

T
Testculos

T
DHT

T
5-Reductase

DHT

ER: receptor de estrognio AR: receptor de andrognios

Clula-alvo

Manuteno funcional dos rgos sexuais e dos caracteres sexuais 2rios

T: testosterona; DHT: diidrotestosterona; E2: estradiol

rgos acessrios, pele da regio genital e folculos pilosos Wilson, 2003

DIRECT AND INDIRECT EFFECTS OF TESTOSTERONE

http://www.get-back-on-track.com/en/professionals/00_meta/07_praesentationen/p_con_0007_01_01.php

REGULAO HORMONAL DA ESPERMATOGNESE Clulas de Leydig LH Testosterona

Espermatognese Diferenciao e amadurecimento dos gametas masculinos

Clulas de Sertoli FSH

Metabolism of plasma testosterone in extraglandular tissues

Metabolism of plasma testosterone in extraglandular tissues. Testosterone can be metabolized to either active or excretory metabolites. Active metabolites such as dihydrotestosterone may be further metabolized to excretory metabolites. HSD, hydroxysteroid dehydrogenase. Willians, 2003 Willians, 2002

A regulao da funo testicular pelas gonadotrofinas e hormnios testiculares na vida adulta

http://acpmedicine.com/acpmedicine/chapters/CH0305.htm

The anterior pituitary and the hypothalamus are connected by the hypophyseal portal vasculature. Releasing or inhibiting hormones secreted by hypothalamic neurons enter the primary plexus of the hypophyseal portal vasculature. They flow down the long portal veins in the pituitary stalk to the secondary plexus, a capillary network that enmeshes the cells of the anterior pituitary. The anterior pituitary cells secrete their hormones in response to the releasing hormones. Because neither the hypothalamus nor the anterior pituitary is isolated by the blood-brain barrier, feedback signals have direct access to both sites of regulation. The posterior pituitary is made up of the terminal portions of neurons whose origin is the hypothalamus. (ACTH adrenocorticotropic hormone; ADH antidiuretic hormone; FSHfolliclestimulating hormone; GHgrowth hormone; LHluteinizing hormone; PRLprolactin; TSHthyroidstimulating hormone)

A adenohipfise e suas relaes com o SNC


corpos celulares dos nn. hipotalmicos secretores de GnRH/LHRH (ncleo arqueado)

Hormnio liberador de gonadotrofinas GnRH/LHRH

LH e FSH

http://www.biosbcc.net/barron/physiology/endo/hypopit.htm

Hormonal relationships in the braintesticular axis.


(Green arrows and plus signs indicate stimulatory effects; red arrows and bars indicate inhibitory effects.)

GnRH from the hypothalamus stimulates the anterior pituitary to secrete FSH and LH. FSH stimulates Sertoli (sustentacular) cells to secrete androgen-binding protein (ABP). LH stimulates interstitial cells to secrete testosterone. In the presence of ABP, testosterone stimulates spermatogenesis. Testosterone also stimulates development of secondary sex organs, and secondary sex characteristics, and stimulates libido. Testosterone has a negative effect on the hypothalamus and pituitary, reducing GnRH secretion and pituitary sensitivity to GnRH. Sertoli cells secrete inhibin which selectively inhibits FSH without reducing testosterone secretion.
http://www.mhhe.com/biosci/esp/2002_general/Esp/default.htm

http://www.get-back-on-track.com/en/professionals/01_androgenbildung/p_pop_01_02_00_01.php?flash=1

A Adenohipfise e os gonadotrfos

gonadotrfos

AS AES DAS GONADOTROFINAS E DA TESTOSTERONA NAS CLULAS TESTICULARES ENVOLVIDAS NA ESPERMATOGNESE


ADENOHIPFISE

LH
estimula a sntese das enzimas da esteroidognese

FSH
Inibina
proliferao e diferenciao das clulas de Sertoli

Clulas de Leydig

Clulas de Sertoli

parcrina

Clulas germinativas

parcrina

T
Clulas miides

testculo

LH: hormnio luteinizante; FSH: hormnio foliculoestimulante

Fluctuation of peripheral vein plasma LH levels and portal vein plasma GnRH levels in unanesthetized, ovariectomized female sheep. Each pulse of LH is coordinated with a pulse of GnRH. This supports the view that pulsatility of LH release is dependent on pulsatile stimulation of the pituitary by GnRH.

Berne et al., 2004

Flutuaes pulsteis dos nveis de LH plasmticos so refletidos nas flutuaes pulsteis e seu hormnio-alvo, a testosterona. Frequncia de pulsos do FSH muito menor.

Berne et al., 2004

Diminui a frequncia de pulso de secreo de GnRH


testosterona 17-Estradiol
(ncleo arqueado)

testosterona

Testosterona

Inibina

Efeitos do FSH e LH (separadamente) na produo de esperma humano

Fig. 46-15 Individual effects of FSH and LH on human sperm production. Normal men were given sufficient testosterone to suppress endogenous FSH and LH secretion by negative feedback. As a result, sperm density declined to very low but detectable levels. Selective restoration of either FSH or LH individually raised sperm levels. However, neither gonadotropin alone could return sperm production to normal. Berne et al., 2004

Hormones of the Male Reproductive System Now, let's take a look at the hormonal control of spermatogenesis. The hormonal pathway is sometimes referred to as the brain-testicular axis. The hypothalamus releases gonadotropinreleasing hormone (GnRH). GnRH travels to the anterior pituitary gland and causes it to release follicle-stimulating hormone (FSH) and luteinizing hormone (LH). LH binds to the interstitial cells of the testes and causes them to secrete testosterone. For this reason, LH can be referred to as interstitial cellstimulating hormone (ICSH). Testosterone will promote spermatogenesis. However, it must be concentrated within the seminiferous tubules. FSH acts on the sertoli cells, causing them to release androgenbinding protein (ABP). ABP binds to and concentrates testosterone within the seminiferous tubules. Testosterone also acts on other body tissues to create secondary sexual characteristics, including pubic, axillary and facial hair, enlargement of the larynx, and increased bone and muscle mass. If testosterone levels rise too high, it begins to inhibit the release of GnRH from the hypothalamus and FSH and LH from the anterior pituitary. If sperm count rises too high, the sertoli cells release the hormone inhibin, which inhibits GnRH and FSH release. These negative feedback processes prevent testosterone levels from rising too high and spermatogenesis from going too fast.

http://academic.pgcc.edu/~aimholtz/AandP/206_ONLINE/Repro/malerepro1.html

Produo de testosterona ao longo da vida do homem

Schematic diagram of the different phases of male sexual function during life as indicated by mean plasma testosterone level and sperm production at different ages. Willians, 2003

Rato macho adulto

orquiectomia

veremos na aula prtica

http://salmon.psy.plym.ac.uk

Diferentes abordagens na busca por contraceptivos masculinos (?)

Current approaches for male contraception. This figure depicts the current approaches in the field utilizing different routes to perturb male fertility at the level of the hypothalamus, pituitary gland, testis and epididymis, and spermatozoa per se. Physiological Reviews, Cheng and Mruk, 2002

Diferentes abordagens na busca por contraceptivos masculinos (?)

http://www.healcentral.org Disponibiliza vdeos/animaes de contedo muito bom


(buscando pela palavra-chave physiology, foram encontrados 84 vdeos).

Aqui esto alguns exemplos:

DIFERENCIAO GONADAL

EIXO HIPOTLAMOHIPOFISRIO GONADAL

DIFERENCIAO SEXUAL DO SNC

veja tambm: Endocrinologia da reproduo feminina:

foliculognese
Profa. Dra. Cristina Maria Henrique Pinto
CFS/CCB/UFSC Junho de 2006