Urinary System • Kidney = Nephrons dorsal to coelom. • Nephron = Tubular structure that reabsorbs substances from a coelomic fluid-derived or plasma-derived filtrate Nephron Structure • Coelomic funnel = cilliated opening of nephron to coelom – where coelomic fluid enters the nephron • Bowman’s capsule = end of nephron that surrounds a mass of capillaries (Glomerulus) – where plasma enters the nephron • Concentrated fluid waste (urine) is formed that usually is rich in nitrogenous (e.g. urea, uric acid) waste. Human Kidney Development Nephron Development • Intermediate mesoderm forms nephric tubules in nephric ridges. • Some remain contiguous with the coelom of the lateral plate mesoderm. dorsal aorta int. mesoderm coelom Nephron Function Vertebrate Embryo (section) neural crest cells sclerotome dermatome myotome splanchnic mesoderm somatic mesoderm gut intermediate mesoderm lateral plate mesoderm Vertebrate Embryo (section) neural crest cells sclerotome dermatome myotome splanchnic mesoderm gut somatic mesoderm intermediate mesoderm lateral plate mesoderm Vertebrate Embryo (section) sclerotome dermatome myotome dorsal & ventral mesenteries gut intermediate mesoderm coelom lateral plate mesoderm (splanchnic mesoderm) Vertebrate Embryo (section) vertebra horizontal septum dermatome myotome dorsal & ventral mesenteries (splanchnic mesoderm) gut intermediate mesoderm coelom lateral plate mesoderm Vertebrate Embryo (section) vertebra horizontal septum dermatome intermediate mesoderm myotome dorsal mesentery (splanchnic mesoderm) gut coelom lateral plate mesoderm Nephron Development Kidney Regions • Nephric ridges • Pronephros = Segmental; develops 1st; coelomic funnel (Lost in adults, except Myxini). • Mesonephros = Segmental; posterior to pronephros; coelomic funnel & Bowman’s capsule (Lost in adult amniotes). • Metanephros = Develop segmentally but obscured by nephron proliferation (but segmental in hagfishes); posterior to mesonephros; Bowman’s capsule. Vertebrate Mesoderm Somitomeres Somites Lateral Plate Mesoderm Intermediate Mesoderm Kidney Development • Archinephric duct = Pronephros initiates formation; drains pro-, meso-, & sometimes meta-nephros. • Accessory duct = Develops longitudinally from the archinephric duct (in some elasmobrachs and salamanders); drains the meso- and meta-nephros. • Metanephric Duct (ureter) = Develops as bud from base of the archinephric duct; drains metanephros (Only in amniotes). Hagfish Kidney pronephros mesonephros metanephros archinephric duct Opisthonephric Kidney pronephros mesonephros metanephros accessory duct archinephric duct Metanephric Kidney pronephros mesonephros metanephros archinephric duct Adult Kidney Types • Holonephric Kidney = pro-, meso-, and segmental meta-nephric tubules • Opisthonephric Kidney = mesonephric and metanephric tubules drained by archinephric and possibly accessory ducts. • Metanephric Kidney = composed only of non-segmental metanephric tubules; drained by metanephric duct (ureter). Human Kidney Development Human Urinary Adult Kidney Group Myxini Petromyzontida Chondrichthys Actinopterygii Dipnoi Lissamphibia Mammalia Reptilia Type holonephric opisthonephric opisthonephric opisthonephric opisthonephric opisthonephric metanephric metanephric Tubules pro-, meso-, & meta- meso- & metameso- & metameso- & metameso- & metameso- & metametameta- Subphylum Vertebrata metanephric kidney; ureters opisthonephric kidney function of pronephric tubules reduced Nitrogenous Waste • Ammonia = NH4+, toxic, excreted in large amounts by many aquatic organisms (not all). hagfishes, lampreys, Actinopterygii, lungfishes, coelacanths, amphib., crocodylians, aquatic turtles. • Urea = CO(NH2 ), less toxic, gnathostomes (Actinopterygiians use it less, Chondrichthyians and Mammals most). • Uric Acid = C4O3N4H3, less toxic, slightly soluble in water, precipitates in cloaca. Requires least amount of water to excrete. Only reptiles produce in large amounts. Subphylum Vertebrata uric acid urea ancestrally ammonia function of pronephric tubules reduced Reproductive System • Gonad = Organ that produces gametes and hormones. • Testis = Male gonad • Seminiferous tubules = site of sperm formation in testis. • Ovary = Female gonad • Follicle = Specialized cells that surround a developing ovum in the ovary. • Gamete = 1n sex cell (sperm/ ovum). Gonadal Histology Gonad Development • Indifferent gonad (Genital ridges) = the early gonad; neither testis nor ovary. • Germ cells = will form gametes. Migrate from endoderm via dorsal mesentery. • Primary Sex Cords = coelomic mesothelium of ridge grows into mesenchyme of ridge. ♂ primary sex cords seminiferous tubules ♀ primary sex cords form and disappear • Secondary Sex Cords = mesothelium again grows into mesenchyme of genital ridge. ♀ secondary sex cords ovarian follicles Vertebrate Embryo (section) vertebra horizontal septum dermatome myotome genital ridge dorsal mesentery (splanchnic mesoderm) gut intermediate mesoderm coelom lateral plate mesoderm Gonad Development Gonad Development Male Urogenital Ducts • ALL Non-amniotes = archinephric ducts drain kidneys. • Amniotes = metanephric ducts drain kidneys. • Hagfishes & Lampreys = no testis ducts. • Gnathostomes = archinephric ducts drain the testes (also drain the kidneys in all nonamniote ganthostomes). EXCEPTION - Teleosts & Amia = new gonoduct drains testes (not the archinephric duct) Hagfish and Lamprey Male Ducts kidney testis archinephric duct abdominal pore Male Genital Ducts (Non-amniote) kidney testes abdominal pore archinephric duct Male Genital Ducts (Non-amniote) kidney testes archinephric duct abdominal pore Male Genital Ducts (Amniote) metanephric ducts kidney testes archinephric duct Male Genital Ducts (Teleost or Amia) kidney testes gonoduct archinephric duct Female Urogenital Ducts • ALL Non-amniotes = archinephric ducts drain kidneys. • Amniotes = metanephric ducts drain kidneys. • Hagfishes & lampreys = no ovary ducts. • Gnathostomes = Müllerian ducts open to the coelom capture ova. (archinephric ducts lost in female amniotes) In birds the right Müllerian duct is reduced and the right ovary is usually absent EXCEPTION - Teleosts & Amia = new gonoduct not open to coelom captures ova (not the Müllerian duct) Hagfish and Lamprey Female Ducts kidney ovary archinephric duct abdominal pore Female Genital Ducts (Non-amniote) ovaries kidney Müllerian duct archinephric duct abdominal pore Female Genital Ducts (Amniote) ovaries metanephric ducts kidney Müllerian duct Female Genital Ducts (Teleost or Amia) kidney ovaries archinephric duct gonoduct Female Genital Ducts Teleosts Male Genital Ducts Male Urogenital Ducts Group Petromyzontida from Kidney archinephric archinephric Chondrichthys accessory & arch. Myxini Polypter-, Ascipenser-, & Lepisosteiformes archinephric Teleostei & Amia archinephric archinephric Dipnoi Lissamphibia Mammalia Reptilia accessory & arch. metanephric metanephric from Testis none none archinephric archinephric male gonoduct archinephric archinephric archinephric archinephric Female Urogenital Ducts Group Myxini Petromyzontida Chondrichthys Polypter-, Ascipenser-, & Lepisosteiformes Teleostei & Amia Dipnoi Lissamphibia Mammalia Reptilia from Kidney archinephric archinephric archinephric archinephric from Ovary none none Müllerian duct Müllerian duct archinephric archinephric archinephric metanephric metanephric female gonoduct Müllerian duct Müllerian duct Müllerian duct Müllerian duct Subphylum Vertebrata archinephric duct lost in females; metanephric serves archinephric duct kidneys serves testes in males; Müllerian duct in females Class Actinopterygii Müllerian duct lost; new gonoducts for ovary and testis Amniota right ovary and oviduct (Müllerian) reduced or lost Human Genitalia • The human genitalia are sexually homologous developing from the “same” developmental precursors. Human Genital Development urinary Early Male reproductive tubercle glans penis folds scrotum Female glans clitoris labia Human Genitalia Development Human Genital Intersexuality Human Female Reproductive Human Male Reproductive Gonadal Descent • In both male and female humans the gonads move inferiorly, in males dramatically so into the scrotum. Cryptorchidism Comparative Vertebrate Genitalia • Genitalia = external or extruded reproductive structures • Intromittent organ = structure for transfer of sperm to facilitate internal fertilization (but not necessary for it); Intromittent organs evolved repeatedly. • Internal fertilization: Chondrichthyans, Amniotes, & Miscellaneous others Cloaca • Cloaca = (sewer in Latin) ectodermal, posterior chamber; intestine, urinary system, and reproductive system “empty” here. • Cloaca lost in Therian mammals, Actinopterygiians, and chimaeras. Separate, external anal and urogenital openings. Chondrichthyan Male Genitalia • Claspers = paired pelvic fin derived structures; grooved for sperm transfer. Amniote Genitalia • Amniotes ancestrally develop a genital tubercle. Homologous to glans penis and glans clitoris in mammals. • Genital tubercle often regresses in female non-mammalian amniotes. • Lepidosaurs and most birds have secondarily lost the genital tubercle. Amniote Genital Development urinary Early Male reproductive tubercle glans penis folds scrotum Female glans clitoris labia Male Genitalia • Median penis = in amniotes; derived from genital tubercle; erectile tissue and a sulcus spermaticus (groove). Secondarily lost in lepidosaurs and most birds (ratites, ducks & geese retain) Mammals = closed sulcus spermaticus that in therians also extends the urethra Many mammals have a baculum (bone in the penis) Testudinean Male Genitalia Crocodylian Genitalia Rhea Male Genitalia Monotreme Male Genitalia • Slightly forked penis (four-headed). • Testes remain abdominal (no scrotum). echidna penis Therian Testes • Scrotum sac containing the testes outside of the male’s body. • Adult testes usually are in a scrotum or to under the skin’s surface. Marsupial Genitalia • Forked penis in most male marsupials. (scrotum anterior to penis) • Paired vaginas in all female marsupials. sugar glider penis Kangaroo Male Genitalia Eutherian Genitalia • Simple penis in most male euterians. (scrotum posterior to penis) • Single vagina in all female eutherians. Hyaena Genitalia two female hyaenas Lepidosaur Male Genitalia • Hemipenes = paired cloacal structures; grooved for sperm transfer. cloaca lost claspers Subphylum Vertebrata genital tubercle; median penis & clitoris cloaca forked penis & paired vaginas sulcus spermaticus closed (tubular) vagina(s); scrotum; cloaca lost (or reduced) hemipenes; gen. tubercle lost Amniota genital tubercle lost in most genital tubercle - median penis & clitoris