Male Reproductive System TESTES - The testes have both exocrine and endocrine functions Exocrine – seminiferous tubules o Compound, coiled, tubular glands that “secrete” cells (i.e. spermatozoa) o Lined by stratified epithelium made of developing spermatozoa and supportive sertoli cells Endocrine – interstitial (Leydig) cells o Secrete hormones, testosterone, for development and maintenance of accessory organs - Structure of the testes Carries the peritoneum during descent to form the tunica vaginalis Testis proper is surrounded by CT and vascular layers o Tunica albuginea (CT) o Tunica vasculosa (vessels) Tunica vasculosa projects a conical mass of CT – mediastinum testis – into the testis Mediastinum parenchyma subdivides testes into ~300 lobuli testis (lobules) Each lobule contains 1-4 convoluted seminiferous tubules - Enclosed by basal lamina - Surrounded by 3-4 layers of smooth muscle cells Myoid (peritubular) cells - Each tubule is ~150-300 µm in diameter and 30-80 cm long - Each tubule is U-shaped with ends opening into the rete testis - Tubules lined with seminiferous epithelium for spermatogenesis Developing sperm cells Sertoli (Sustentacular) cells Each seminiferous tubule continues near the mediastinum and transitions into a straight tubule – tubulus rectus Tubulus rectus continues into the rete testis Rete testes continues into excurrent ducts SPERMATOGENESIS - The process by which stem cells develop into mature spermatozoa - Three phases Spermatocytogenesis (mitosis) Meiosis Spermiogenesis - Spermatogenic cells Spermatogonia o “Mitotic” Cells o Stem cells that replenish and populate o Lie along basement membrane Type A spermatogonia o Stem cells that give rise to both type A (themselves) and Type B spermatogonia o Have a rounded nuclei with very fine chromatin grains and one or two nucleoli Type B spermatogonia o Are not stem cells o Divide repeatedly o Have rounded nuclei with chromatin granules of variable size and one nucleolus Chromatin granules often attach to the nuclear membrane - Meiotic cells Primary spermatocytes o Luminal and larger than spermatogonia Always visible in cross-sections o Immediately enter prophase of the first meiotic division Extremely prolonged phase (22 days) o Completion of the first meiotic division results in the formation of secondary spermatocytes Secondary spermatocytes o Smaller than primary spermatocytes o Rapidly enter and complete the second meiotic division Seldom seen in histological preparations Spermatids o Closest to luminal part of the seminiferous epithelium o Small (diameter ~10 µm) with a very light nucleus - - - - Spermiogenesis Terminal phase of spermatogenesis Consists of differentiation and maturation of the newly-formed spermatids into spermatozoa Golgi phase o Formation of the acrosome vesicle from hydrolytic granules formed in the Golgi Cap phase o Acrosomal vesicle forms the head cap over the nucleus o Centrioles (2) assemble at the caudal pole of the nucleus and help organize spindle fibers Distal centriole gives rise to an outgrowing flagellum Acrosomal phase o Nucleus elongates and chromatin condenses (stains darker) o Spermatid rotates with head towards the basal lamina and tail towards the lumen o Acrosome stains brightly with PAS stain due to its content of hydrolytic enzymes Maturation phase o Completion of nuclear condensation o Arrangement of mitochondria around the axoneme of the flagellum o Release of the spermatid into the tubular lumen o Spermatid reduces in size by extruding cytoplasm and organelles – forms a “residual body” o Residual bodies are phagocytosed by Sertoli cells, lost in tubular lumen, and partly autolysed Spermatogenic cycle Defined as the time it takes for the reappearance of the same stage within a given segment of the tubule o ~ 48-60 days from meiosis to mature spermatozoa In testes cross-section, sperm cells can be seen to differentiate in distinct associations with other cells o Each spermatogenic association is classified as a “stage” of the seminiferous epithelial cycle Number of stages within a spermatogenic cycle varies between species Number of cycles required for the completion of spermatogenesis varies between species Sertoli cells Hormone responsive support cells Pleomorphic - two or more structural forms during a life cycle Provide nutrient support Secrete ABP, inhibin, activin Connected by tight junctions through which germ cells move Tight junctions form the blood-testis barrier Interstital tissue Located between seminiferous tubules Composed of CT, blood vessels, lymphatics and Leydig (interstitial) cells o Leydig cells secrete testosterone and estrogens Sertoli Cells Leydig Cells DUCTS - Excurrent ducts Tubulus rectus (straight tubule) – lined by low columnar cells Rete testes – lined by flattened to cuboidal epithelium Ductus efferens (efferent duct) o Lined by pseudostratified columnar epithelium with cilia o Carry spermatozoa from the testes to the epididymis o Absorptive cells Ductus epididymis o Spermatozoa-carrying duct that forms the epididymis o Organized into head (caput), body (corpus), and tail (cauda) o Lined by tall, pseudostratified stereociliated columnar epithelium and basal epithelium o Spermatozoa are stored within the epididymis while they mature to become sperm Epididymis Epididymis Vas deferens (ductus deferens) o Carry sperm from the epididymis to the ejaculatory duct in anticipation of ejaculation o Transverses the inguinal canal toward urethra o Lined by pseudostratified columnar epithelium with sterocilia o Lamina propria and thick muscularis with inner and outer layers o Terminates at the ampulla (enlargement of the duct) near the prostate Ejaculatory duct o Short duct formed via joining of distal ductus deferens and a duct from the seminal vesicle o Carries sperm and semen through the prostate gland, emptying into the urethra o Lined by simple or pseudostratified epithelium o Wall is composed of fibrous tissue ACCESSORY GLANDS - Functions of accessory gland products Nourish and activate the spermatozoa Clear urethral tract prior to ejaculation Transport vehicle for spermatozoa in the female tract Plug female tract after placement of spermatozoa to help ensure fertilization - Gland structure varies highly in organization and distribution among species Usually described as branched tubular or branched tubuloalveolar - Seminal vesicles (vesicular gland) Develop from vas deferens Elongated sacs (~4 cm long, ~2 cm wide) that taper where they unite with the vas deferens Each seminal vesicle consists of one coiling tube (~15cm long) Mucosa is composed of thin-branched columnar or pseudostratified epithelium Muscularis layer on inner and outer walls Seminal vesicles - Prostate gland Largest accessory sex gland Equivalent to the mammary gland in females Empty into excretory ducts of urethra Promotes movement of spermatozoa Helps form a vaginal plug that ensures fertilization Secretions help neutralize vaginal secretions o More serous (dogs) o More mucous, contains high levels of fructose and citric acid (bull) Stromal cells have 5-alpha reductase to produce dihydrotestosterone Structure o Compound tubuloalveolar gland lined by low cuboidal to low columnar epithelium Epithelium contains secretory and basal cells o Grossly divided into two parts (development varies with species) – body, disseminate part o Secretory ducts lined by simple columnar to transitional epithelium Secretions empty directly into urethra o Concretions (solid masses) may be present in secretory end pieces and parts of the duct system - Bulbourethral glands All domestic species (not dog) have these glands Small, paired tubular or tubuloalveloar glands o Composed of simple columnar and secretory epithelium Capsule of dense CT contains smooth and skeletal m. of the bulbocavernous and urethral mm. Have separate ducts that enter the urethra Functions of mucous secretions o Clears the urethra of urine o Lubricates urethra and vagina o May be energy source for spermatozoa Urethra Connects bladder and passage for spermatozoa Structure o Has prostatic, membranous (pelvic), and spongiose (penile) portions o Mostly lined with transitional epithelium Tip is squamous epithelium - Bulbourethral gland (boar) o o o o o - Penis Propria submucosa contains erectile tissue of cavernous sinuses Also has branched tubular mucous glands and the urethral glands (species variable) Colliculus Seminalis Openings for ductus deferens, prostate and vesicular glands Tunica muscularis near bladder consists of striated urethral muscle Tunica muscularis of membranous region consists of smooth muscle Cavernous erectile tissue – corpus cavernosum – is found in CT beneath epithelium Erectile tissue – corpus spongiosum – is more pronounced in the penile urethra Shared outlet for urine and spermatozoa Copulatory organ Mostly consists of erectile tissue Structure o Paired corpora cavernosa Dorsal columns of erectile tissue surrounded by dense CT (tunica albuginea) Erectile tissue consists of trabeculae of irregular cavernous blood spaces lined by endothelial cells and surrounded by smooth muscle cells, nerves and dense CT o Corpus spongiosum o Glans penis – erectile tissue, bone, fibrocartilage