Genital - Male Development

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Human Y chromosome SRY region.jpg
Urogenital male structures cartoon
Urogenital Male

The male and female reproductive systems develop initially "indifferently", it is the product of the Y chromosome SRY gene that makes the "difference".

The paired mesonephric ducts (Wolffian ducts) contributes the majority of male internal genital tract.

Embryonic gonad development leads to the mesonephric/paramesonephric duct changes, while the external genitaila remain indeterminate in appearance through to the fetal period.

Importantly its sex chromosome dependence, late embryonic/fetal differential development, complex morphogenic changes, long time-course, hormonal sensitivity and hormonal influences make it a system prone to many different abnormalities.

There are also separate pages describing: Y Chromosome | spermatozoa | testis | epididymis | ductus deferens | prostate | penis | Category:Male

Historic Embryology
Caspar Friedrich Wolff (1734-1794)
Theoria Generationis 1774.jpg

Caspar Friedrich Wolff (1734-1794) was a German embryologist and anatomist best known today for identifying the Wolffian duct (mesonephric duct; ductus deferens, epididymis), Wolffian body (mesonephros) and Wolffian cyst (mesonephric origin uterine broad ligament cyst) that bear his name. Thought also to be a founder of the germ layer theory. His doctorate dissertation Theoria generationis (1774) discarded the developmental theory of preformation. Later in his career, his teaching in Berlin was opposed by the professors of the Medical-Surgical College, who had guild privileges to teach medicine.

Genital Links: genital | Lecture - Medicine | Lecture - Science | Lecture Movie | Medicine - Practical | primordial germ cell | meiosis | endocrine gonad‎ | Genital Movies | genital abnormalities | Assisted Reproductive Technology | puberty | Category:Genital
Female | X | X inactivation | ovary | corpus luteum | oocyte | uterus | vagina | reproductive cycles | menstrual cycle | Category:Female
Male | Y | SRY | testis | spermatozoa | ductus deferens | penis | prostate | Category:Male
Historic Embryology - Genital 
General: 1901 Urinogenital Tract | 1902 The Uro-Genital System | 1904 Ovary and Testis | 1912 Urinogenital Organ Development | 1914 External Genitalia | 1921 Urogenital Development | 1921 External Genital | 1942 Sex Cords | 1953 Germ Cells | Historic Embryology Papers | Historic Disclaimer
Female: 1904 Ovary and Testis | 1904 Hymen | 1912 Urinogenital Organ Development | 1914 External Genitalia | 1914 Female | 1921 External Genital | 1927 Female Foetus 15 cm | 1927 Vagina | 1932 Postnatal Ovary
Male: 1887-88 Testis | 1904 Ovary and Testis | 1904 Leydig Cells | 1906 Testis vascular | 1909 Prostate | 1912 Prostate | 1914 External Genitalia | 1915 Cowper’s and Bartholin’s Glands | 1920 Wolffian tubules | 1935 Prepuce | 1935 Wolffian Duct | 1942 Sex Cords | 1943 Testes Descent | Historic Embryology Papers | Historic Disclaimer

Some Recent Findings

Model male second trimester androsterone synthesis
Model male second trimester androsterone synthesis[1]
Male urogenital histology
Male urogenital development (week 8, stage 22)
  • Alternative (backdoor) androgen production and masculinization in the human fetus[1] "Masculinization of the external genitalia in humans is dependent on formation of 5α-dihydrotestosterone (DHT) through both the canonical androgenic pathway and an alternative (backdoor) pathway. The fetal testes are essential for canonical androgen production, but little is known about the synthesis of backdoor androgens, despite their known critical role in masculinization. ...Results show that androsterone is the principal backdoor androgen in the male fetal circulation and that DHT is undetectable (<1 ng/mL), while in female fetuses, there are significantly lower levels of androsterone and testosterone. In the male, intermediates in the backdoor pathway are found primarily in the placenta and fetal liver, with significant androsterone levels also in the fetal adrenal. Backdoor intermediates, including androsterone, are only present at very low levels in the fetal testes. This is consistent with transcript levels of enzymes involved in the alternate pathway (steroid 5α-reductase type 1 [SRD5A1], aldo-keto reductase type 1C2 [AKR1C2], aldo-keto reductase type 1C4 [AKR1C4], cytochrome P450 17A1 [CYP17A1]), as measured by quantitative PCR (qPCR). These data identify androsterone as the predominant backdoor androgen in the human fetus and show that circulating levels are sex dependent, but also that there is little de novo synthesis in the testis. Instead, the data indicate that placental progesterone acts as substrate for synthesis of backdoor androgens, which occurs across several tissues. Masculinization of the human fetus depends, therefore, on testosterone and androsterone synthesis by both the fetal testes and nongonadal tissues, leading to DHT formation at the genital tubercle. Our findings also provide a solid basis to explain why placental insufficiency is associated with disorders of sex development in humans."
  • Alterations of sex determination pathway in the genital ridges of males with limited Y chromosome genes[2] "We previously demonstrated that in the mouse only two Y chromosome genes are required for a male to produce an offspring with the help of assisted reproduction technologies (ART): testis determinant Sry and spermatogonial proliferation factor Eif2s3y. Subsequently, we have shown that the function of these genes can be replaced by transgenic overexpression of their homologues, autosomally encoded Sox9 and X-chromosome encoded Eif2s3x. Males with Y chromosome contribution limited to two (XEif2s3yOSry), one (XEif2s3yOSox9 and XOSry, Eif2s3x) and no genes (XOSox9, Eif2s3x) produced haploid germ cells and sired offspring after ART. However, despite successful assisted reproductive outcome, they had smaller testes and displayed abnormal development of the seminiferous epithelium and testicular interstitium. Here we explored whether these testicular defects originated from altered pro-testis and pro-ovary factor signaling in genital ridges at the time of sex determination."

More recent papers  
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Search term: Male Embryology | Male Development | Male Genital Development

Testis Development
Older papers  
These papers originally appeared in the Some Recent Findings table, but as that list grew in length have now been shuffled down to this collapsible table.

See also the Discussion Page for other references listed by year and References on this current page.

  • Tissue-specific roles of Fgfr2 in development of the external genitalia[3] "Congenital anomalies frequently occur in organs that undergo tubulogenesis. Hypospadias is a urethral tube defect defined by mislocalized, oversized, or multiple openings of the penile urethra. Deletion of Fgfr2 or its ligand Fgf10 results in severe hypospadias in mice, in which the entire urethral plate is open along the ventral side of the penis. In the genital tubercle, the embryonic precursor of the penis and clitoris, Fgfr2 is expressed in two epithelial populations: the endodermally derived urethral epithelium and the ectodermally derived surface epithelium. Here, we investigate the tissue-specific roles of Fgfr2 in external genital development by generating conditional deletions of Fgfr2 in each of these cell types. These results demonstrate that urethral tubulogenesis, prepuce morphogenesis, and sexually dimorphic patterning of the lower urethra are controlled by discrete regions of Fgfr2 activity." Fibroblast Growth Factor
  • Penile biometry on prenatal MR imaging [4]"Mean length values, including 95% confidence intervals and percentiles, were defined. Penile length as a function of gestational age was expressed by the regression equation: outer mean length= -5.514 + 0.622 *, and total mean length= -8.865 + 1.312 * (*= gestational weeks). The correlation coefficients were statistically significant (p < .001). The comparison between outer length on MRI and US data showed no significant differences, whereas total length on MRI and US data demonstrated significant differences (p< .001)." penis
  • Male reproductive tract abnormalities: More common after assisted reproduction?[5] "IVF and ICSI, by increasing the risks of prematurity, low birthweight, and multiple gestation, are indirect risk factors for developing male genital malformations. In infants with normal birhtweight or from singleton pregnancies, ICSI is a specific risk factor for hypospadias."
  • Temporal and spatial dissection of Shh signaling in genital tubercle development.[6] "Genital tubercle (GT) initiation and outgrowth involve coordinated morphogenesis of surface ectoderm, cloacal mesoderm and hindgut endoderm. GT development appears to mirror that of the limb. Although Shh is essential for the development of both appendages, its role in GT development is much less clear than in the limb. Here, by removing Shh at different stages during GT development in mice, we demonstrate a continuous requirement for Shh in GT initiation and subsequent androgen-independent GT growth."
  • Bmp7 expression and null phenotype in the urogenital system suggest a role in re-organization of the urethral epithelium. [7] "Signaling by Bone morphogenetic proteins (Bmps) has multiple and diverse roles in patterning and morphogenesis of the kidney, eye, limbs and the neural tube. ...Together, our analysis of Bmp7 expression and the null phenotype, indicates that Bmp7 may play an important role in re-organization of the epithelium during cloacal septation and morphogenesis of the genital tubercle."


Historic drawing of the testis
Historic drawing of the testis
  • Human Embryology (2nd ed.) Larson Chapter 10 p261-306
  • The Developing Human: Clinically Oriented Embryology (6th ed.) Moore and Persaud Chapter 13 p303-346
  • Before We Are Born (5th ed.) Moore and Persaud Chapter 14 p289-326
  • Essentials of Human Embryology, Larson Chapter 10 p173-205
  • Human Embryology, Fitzgerald and Fitzgerald Chapter 21-22 p134-152
  • Developmental Biology (6th ed.) Gilbert Chapter 14 Intermediate Mesoderm


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 ‎‎Urogenital Septum
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 ‎‎Male External
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 ‎‎Testis Descent
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 ‎‎Gonad Vascular
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Mouse Primordial Germ Cell Migration
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 ‎‎Germ Cell E9.0
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Development Overview

Three main stages during development, mesonephric/paramesonephric duct changes are one of the first male/female differences that occur in development, while external genitaila remain indeterminate in appearance for quite a while.

  1. Differentiation of gonad (Sex determination)
  2. Differentiation of internal genital organs
  3. Differentiation of external genital organs

The 2nd and 3rd stages dependent on endocrine gonad. Reproductive development has a long maturation timecourse, begining in the embryo and finishing in puberty. (More? Puberty Development)

Historic Images of Genital Changes

Urogenital Indifferent Urogenital Male Urogenital Female
Urogenital indifferent Urogenital male Urogenital female


Sry (sex-determining region on the Y chromosome) gene was found in 1990 on the Y and the first SOX gene identified, the sry gene encodes a "testis-determining factor" a 204aa protein (Mr 23884 Da).

Sry acts as a transcriptional activator (HMG type-high mobility group) binding to DNA and initiating male sex determination then regulating male development. The protein sequence is shown on this current page and the full genebank entry can also be seen. The sry protein has a HMG box that binds DNA by intercalating in the minor groove. Read about the mapping of the testis determining factor which is SRY.

The actual gene targets of SRY are still being determined but at least one downstream gene Sox9 has been identified. Another gene Dax1 (nuclear hormone-receptor superfamily member) when expressed as a transgene will antagonize Sry and also force dosage-sensitive sex reversal.

Mouse sex determination genes 01.jpg

Mouse sex determination genes[8]

Links: SRY | Y chromosome | OMIM - Sry | 3d Structure Of The Human Sry-Dna Complex

Human SOX Family  
Table - Human Sox Family
Approved Name Previous Symbols Synonyms Chromosome
SOX1 SRY-box 1 13q34
SOX2 SRY-box 2 3q26.33
SOX3 SRY-box 3 PHP Xq27.1
SOX4 SRY-box 4 6p22.3
SOX5 SRY-box 5 "L-SOX5, MGC35153" 12p12.1
SOX6 SRY-box 6 11p15.3
SOX7 SRY-box 7 8p23.1
SOX8 SRY-box 8 16p13.3
SOX9 SRY-box 9 "CMD1, CMPD1" SRA1 17q24.3
SOX10 SRY-box 10 "DOM, WS4, WS2E" 22q13.1
SOX11 SRY-box 11 2p25.2
SOX12 SRY-box 12 SOX22 20p13
SOX13 SRY-box 13 "Sox-13, ICA12, MGC117216" 1q32.1
SOX14 SRY-box 14 SOX28 3q22.3
SOX15 SRY-box 15 SOX20 "SOX27, SOX26" 17p13.1
SOX17 SRY-box 17 8q11.23
SOX18 SRY-box 18 20q13.33
SOX21 SRY-box 21 SOX25 13q32.1
SOX30 SRY-box 30 5q33.3
SRY sex determining region Y TDF Yp11.2
    Links: Developmental Signals - Sox | OMIM | HGNC | Tbx Family | Bmp Family | Fgf Family | Pax Family | R-spondin Family | Sox Family | Tbx Family

SRY Nuclear Import

SRY nuclear import model.jpg

A model for nuclear import of SRY from normal males and XY females.[9]

The distinct nuclear localization signals (NLSs) of SRY use different import pathways.

  • cNLS - recognized by IMPβ, docks the transport complex at the nuclear pore complex (NPC) and is then translocation through. After nuclear entry of the complex, RanGTP binds to IMPβ to trigger release of SRY; DNA binding by SRY may also facilitate the release process.
  • nNLS - mediates nuclear import through a novel pathway not utilizing conventional nuclear import factors such as IMPs but an unidentified "transport factor" (TF) suggested to be calcium-calmodulin.

Sry Target Genes

Cerebellin 4 precursor (Cbln4) - encodes a secreted protein expressed in Sertoli cells in the developing gonad.[10]

Androgen receptor - SRY interacts with and negatively regulates this receptor transcriptional activity.[11]

Gonad - Testis

See the detailed notes on testis development.

Links: Testis Development

Internal Genital

Mesonephric duct or Wolffian duct differentiates to form the male internal genital tract the vas deferens (ductus deferens, vas deferens or simply vas). Associated with the duct are the male prostate and accessory glands.

Human Mesonephric Duct position (week 6 to 11)
Mesonephric duct position week 6-11.jpg Schematic representations of the descent of the mesonephric duct (Wolffian duct, WD) or vas deferens. Anterior view.[12]

  • 6 weeks - (A) the WD opens to the urogenital sinus at a site adjacent to the ureteral orifice (UR).
  • 7–8 weeks - (B, C) rather than descent, there are individual variations in the WD position along the mediolateral axis as well as in left/right difference in morphology of the urogenital sinus (URS). The future bladder and urethra are not discriminated in the sinus.
  • 8–9 weeks - the bilateral upper angles (arrows) of the URS start upward growth toward the umbilicus.

  • 9 weeks - (D) depending on development of smooth muscles in the bladder as well as rhapdosphincter muscles of the urethra (RS), the descent of the vas deferens becomes evident. However, the epithelium is still same (arrowheads) between the future bladder and urethra.
  • 10–11 weeks - (E) a drastic upward growth of bladder smooth muscles as well as a developing prostate (PR) accelerates the descent of the vas.
Adult Ductus deferens
Testis histology 003.jpg
Adult Prostate
Prostate histology 01.jpg Prostate histology 02.jpg Prostate histology 03.jpg
Human prostate histology Corpora Amylacea Submucosal gland
(adult, low power overview) (adult, detail) (adult, high power detail)

External Genital

Historic diagram of external development (male left)
  • external genitalia are initially identical and undergo male and female differentiation under the influence or absence of steroidal sex hormones.
  • Indifferent stage ‐ cloaca divided by proliferating mesenchyme forming the urorectal septum which separates the ventral urogenital sinus from the dorsal rectum.
  • Difference stage ‐ locally in this region the presence or absence of dihydrotestosterone (DHT), generated from testosterone, determines male/female development.


Anti-Müllerian Hormone

TGF-beta signaling pathway[13]

Anti-Müllerian Hormone (AMH, Müllerian Inhibiting Substance, MIS, Müllerian Inhibiting Factor, MIF) is a secreted glycoprotein factor of the transforming growth factor-beta, TGF-beta superfamily, that regulates gonadal and genital tract development. (Gene locus 19p13.3)

In the male embryo, the Sertoli cell secrete AMH and inhibit paramesonephric (Mullerian) duct development. This secreted hormone also acts to differentiate the Leydig cells (interstitial cells).

Ligand of TGF-beta (transforming growth factor-beta) superfamily and receptor binds to the anti-Mullerian hormone receptor type 2. Signalling pathway activate SMAD family transcription factors that regulate gene expression.

In postnatal males, AMH increases during the first month, reaching peak level at 6 months of age, and then slowly declines during childhood falling to low levels in puberty.

In reproductive age women, AMH is produced in the ovary by the granulosa cells surrounding preantral and small antral follicles and serum levels may reflect the remaining follicle cohort and decrease with age.

Sertoli cells release mainly a prohormone (proAMH), that is cleaved by subtilisin/kexin-type proprotein convertases or serine proteinases. The cleaved protein forms a stable complex (AMHN,C). Therefore the circulating AMH is a mixture of proAMH and AMHN,C. It has been suggested that proAMH may be activated within the gonads and also by its endocrine target-cells.

Preproprotein proteolytically processed to generate N- and C-terminal cleavage products, that homodimerize and associate to form a biologically active noncovalent complex. (see Protein Atlas)

Male testosterone and AMH level graph.jpg

Male testosterone and AMH levels

Ovary AMH

During ovary follicle development, the granulosa cells secrete AMH and it may have a role in follicular recruitment and development.[14] and may also function in postnatal elevation of FSH secretion in females.[15]

Other AMH Tissues

  • The placenta has also been shown to both synthesise AMH and express its receptors.[16]
  • AMH receptors have been identified in both the pituitary and brain.[15]

Links: TGF-beta | OMIM - AMH

Dihydrotestosterone (DHT)

Male presence of Dihydrotestosterone (DHT, 5α-dihydrotestosterone, androstanolone, 5α-androstan-17β-ol-3-one).
  • locally in this region leads to genital tubercle growth, form
  • genital folds (urethral) initial maintenance and then fusion, forming perineal and penile raphe.
  • labioscrotal swellings become the scrotum.
Testosterone metabolism

Links: Endocrinology - Diagram of the development of the external genitalia | image


Androgen and Digit ratio (2D:4D)

Androgen and Digit ratio (2D:4D

The ratio of 2nd and 4th finger (D, digit) length. This ratio has been suggested to relate to high fetal testosterone concentration (males have lower 2D:4D than females) and has been shown for several species.[17] Although a study in mice has not shown the same correlation.[18] There have been some suggestions that the ratio may also be an indicator of various neurological abnormalities.

To measure (2D:4D) - using your right hand palm up, measure the index finger (2) and ring finger (4) length from palm to tip. Dividing the index finger by the ring finger gives the 2D:4D ratio, average women ratio is 1, average men is 0.98.

Additional Images


  1. 1.0 1.1 O'Shaughnessy PJ, Antignac JP, Le Bizec B, Morvan ML, Svechnikov K, Söder O, Savchuk I, Monteiro A, Soffientini U, Johnston ZC, Bellingham M, Hough D, Walker N, Filis P & Fowler PA. (2019). Alternative (backdoor) androgen production and masculinization in the human fetus. PLoS Biol. , 17, e3000002. PMID: 30763313 DOI.
  2. Ortega EA, Salvador Q, Fernandez M & Ward MA. (2018). Alterations of sex determination pathway in the genital ridges of males with limited Y chromosome genes. Biol. Reprod. , , . PMID: 30285093 DOI.
  3. Gredler ML, Seifert AW & Cohn MJ. (2015). Tissue-specific roles of Fgfr2 in development of the external genitalia. Development , 142, 2203-12. PMID: 26081573 DOI.
  4. Ishiguro T, Tamagawa S & Ogawa H. (1992). [Changes of pupil size in brain death patients]. Seishin Shinkeigaku Zasshi , 94, 864-73. PMID: 1484906
  5. Funke S, Flach E, Kiss I, Sándor J, Vida G, Bódis J & Ertl T. (2010). Male reproductive tract abnormalities: more common after assisted reproduction?. Early Hum. Dev. , 86, 547-50. PMID: 20674196 DOI.
  6. Lin C, Yin Y, Veith GM, Fisher AV, Long F & Ma L. (2009). Temporal and spatial dissection of Shh signaling in genital tubercle development. Development , 136, 3959-67. PMID: 19906863 DOI.
  7. Wu X, Ferrara C, Shapiro E & Grishina I. (2009). Bmp7 expression and null phenotype in the urogenital system suggest a role in re-organization of the urethral epithelium. Gene Expr. Patterns , 9, 224-30. PMID: 19159697 DOI.
  8. de Lau WB, Snel B & Clevers HC. (2012). The R-spondin protein family. Genome Biol. , 13, 242. PMID: 22439850 DOI.
  9. Harley VR, Layfield S, Mitchell CL, Forwood JK, John AP, Briggs LJ, McDowall SG & Jans DA. (2003). Defective importin beta recognition and nuclear import of the sex-determining factor SRY are associated with XY sex-reversing mutations. Proc. Natl. Acad. Sci. U.S.A. , 100, 7045-50. PMID: 12764225 DOI.
  10. Bradford ST, Hiramatsu R, Maddugoda MP, Bernard P, Chaboissier MC, Sinclair A, Schedl A, Harley V, Kanai Y, Koopman P & Wilhelm D. (2009). The cerebellin 4 precursor gene is a direct target of SRY and SOX9 in mice. Biol. Reprod. , 80, 1178-88. PMID: 19211811 DOI.
  11. Yuan X, Lu ML, Li T & Balk SP. (2001). SRY interacts with and negatively regulates androgen receptor transcriptional activity. J. Biol. Chem. , 276, 46647-54. PMID: 11585838 DOI.
  12. Jin ZW, Abe H, Hinata N, Li XW, Murakami G & Rodríguez-Vázquez JF. (2016). Descent of mesonephric duct to the final position of the vas deferens in human embryo and fetus. Anat Cell Biol , 49, 231-240. PMID: 28127497 DOI.
  13. Pang K, Ryan JF, Baxevanis AD & Martindale MQ. (2011). Evolution of the TGF-β signaling pathway and its potential role in the ctenophore, Mnemiopsis leidyi. PLoS ONE , 6, e24152. PMID: 21931657 DOI.
  14. McLennan IS & Pankhurst MW. (2015). Anti-Müllerian hormone is a gonadal cytokine with two circulating forms and cryptic actions. J. Endocrinol. , 226, R45-57. PMID: 26163524 DOI.
  15. 15.0 15.1 Garrel G, Racine C, L'Hôte D, Denoyelle C, Guigon CJ, di Clemente N & Cohen-Tannoudji J. (2016). Anti-Müllerian hormone: a new actor of sexual dimorphism in pituitary gonadotrope activity before puberty. Sci Rep , 6, 23790. PMID: 27030385 DOI.
  16. Novembri R, Funghi L, Voltolini C, Belmonte G, Vannuccini S, Torricelli M & Petraglia F. (2015). Placenta expresses anti-Müllerian hormone and its receptor: Sex-related difference in fetal membranes. Placenta , 36, 731-7. PMID: 25972076 DOI.
  17. McIntyre MH. (2006). The use of digit ratios as markers for perinatal androgen action. Reprod. Biol. Endocrinol. , 4, 10. PMID: 16504142 DOI.
  18. Yan RH, Bunning M, Wahlsten D & Hurd PL. (2009). Digit ratio (2Dratio4D) differences between 20 strains of inbred mice. PLoS ONE , 4, e5801. PMID: 19495421 DOI.


Cohn MJ. (2011). Development of the external genitalia: conserved and divergent mechanisms of appendage patterning. Dev. Dyn. , 240, 1108-15. PMID: 21465625 DOI.

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Rey RA & Grinspon RP. (2011). Normal male sexual differentiation and aetiology of disorders of sex development. Best Pract. Res. Clin. Endocrinol. Metab. , 25, 221-38. PMID: 21397195 DOI.

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Chawengsaksophak K, Svingen T, Ng ET, Epp T, Spiller CM, Clark C, Cooper H & Koopman P. (2012). Loss of Wnt5a disrupts primordial germ cell migration and male sexual development in mice. Biol. Reprod. , 86, 1-12. PMID: 21900680 DOI.

Cools M, Wolffenbuttel KP, Drop SL, Oosterhuis JW & Looijenga LH. (2011). Gonadal development and tumor formation at the crossroads of male and female sex determination. Sex Dev , 5, 167-80. PMID: 21791949 DOI.

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Search Pubmed: Male Genital System Development | mesonephric duct


  • mesonephric duct - (Wollfian duct) An early developing urogenital paired duct system that initially runs the length of the embryo, that will differentiate and form the male reproductive duct system (ductus deferens). In females, this duct degenerates occasionally some remnants may remain associated in broad ligament.
  • Wolffian duct - (mesonephric duct, preferred terminology), A developmental duct that runs from the mesonephros to cloaca. The duct in male differentiates to form the ductus deferens and in female the same structure regresses. Historically named after Caspar Friedrich Wolff (1733-1794), a German scientist and early embryology researcher and is said to have established the doctrine of germ layers. (More? Caspar Friedrich Wolff)

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Cite this page: Hill, M.A. (2024, May 25) Embryology Genital - Male Development. Retrieved from

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