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Development of the Penis and Scrotum > > Embryology

MALE EXTERNAL GENITALIA Includes the penis, penile , and scrotum (which houses the testes). During gestation, development is heavily influenced by produced by the Leydig (aka, interstitial) cells of the testes. Weeks 3-4

• Period of undifferentiated primordial genitalia

• The , which is the precursor to the penis, emerges.

is bordered by the primorida of the genital tubercle, and is surrounded by mesenchymal cells.

• Anterior fusion of the primordia forms the definitive genital tubercle.

— Posteriorly, the unfused portion gives rise to the urogenital (aka, genital) folds, which enclose the cloacal membrane. — Lateral to the folds the genital swellings (aka, labioscrotal swellings) arise. Weeks 7-12 The has reached the cloacal membrane and now forms the perineal body; male differentiation begins.

• The perineal body divides the cloacal membrane into urogenital and anal membranes, which then rupture, forming the and anus.

• The genital tubercle enlarges and is now referred to as the phallus.

• The urogenital folds are now the urethral folds, as they line the urethral groove.

— The roof of the groove comprises an endodermal expansion.

• Distally is the , and the glans plate, which is a solid endodermal remnant of the cloacal membrane.

• The genital swellings are now the scrotal swellings.

• Posteriorly, the perineum separates the urethral folds and groove from the anus.

Week 12 The urethral folds and scrotal swellings fuse at the midline.

• Fusion of the urethral folds forms the floor of the spongy urethra within the shaft of the penis.

• Midline raphes reflect fusion of the tissues.

• The surrounding mesenchyme will give rise to the erectile tissues and vasculature of the penis.

• Formation of the glans portion of the urethra occurs separately, via canalization of the glans plate (the exact mechanism is uncertain).

• At the midline, the raphe reflects the fusion of the urethral folds.

1 / 2 9 Months Folds are fully fused, and urethral opening is at the tip of the penis. The scrotal sac houses the testes. Clinical Correlation

• Hypospadia

— Incomplete fusion of the urethra leads to hypospadias, in which the urethral opening is displaced ventrally. Summary: Genital tubercle -> Shaft and glans of the penis Genital folds -> Floor of the spongy urethra Genital swellings -> Scrotum

TESTICULAR DESCENT:&NBSP;

Week 7 The testes are originally located within the abdominal cavity, deep to the (aka, retroperitoneal).

• The caudal genital ligament attaches to the superior pole of the testes.

• The anchors the inferior pole to the scrotal swelling, inferiorly.

Week 12 The peritoneum begins to evaginates, forming the as it grows towards and into the scrotal swelling. A* s it migrates, the vaginal process creates a cavity as it pushes through the abdominal wall layers, ultimately giving rise to the inguinal canals. 9 Months The testes have traveled through the inguinal canals to reach the scrotal sac; thus, in the mature form, the testes are external to the pelvic cavity.

• Though the caudal genital ligament regresses, the gubernaculum persists to anchor the testis.

- — The gubernaculum directs the testis during its migration through the to the scrotal sac; although the exact mechanisms are uncertain, shortening of the gubernaculum paired with growth of the abdominal organs (and, consequently, increased intra-abdominal pressure) likely facilitate testes descent.

Clinical Correlation

• Cryptorchidism

— Failure of one or both testes to descend into the scrotal sac; if the testes remain undescended too long after birth, surgery may be required. Additional resources:

- Li, Y., Sinclair, A., Cao, et al. Canalization of the Urethral Plate Precedes Fusion of the Urethral Folds during Male Penile Urethral Development: The Double Zipper Hypothesis. Journal of Urology: 193(4): 1353-1360 (2015).

- Hadidi, A., Roessler, J., Coerdt, W. Development of the Human Male Urethra: A Histochemical Study on Human . Journal of Pediatric Surgery. 49: 1146-1152. (2014).

- Cunha, G.R., Sinclair, A., Risbridger, G., Hutson, J. & Baskin, L.S. Current Understanding of Hypospadias: Relevance of Animal Models. Nature Reviews in Urology.(2015) doi: 10.1038/nrurol.2015.57. UC San Francisco: 898757. Retrieved from: http://escholarship.org/uc/item/60s7k7n0 February 15, 2017.

- Singh, V. Textbook of Clinical Embryology. Elsevier Health Sciences. (2012).

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