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Life History and Embryology of Vertebrates Hazel Anne L. Tabo, MS Embryology Study of development from fertilization to embryo. The nature of relationship from: ancestor to offspring can be studied. It reveals ancestry. Haeckel: “Ontogeny recapitulates phylogeny.” (development repeats evolution). De Beers: Recapitulation is not the only relationship between embryos and ancestors. Embryo (De Beers, 1930) Embryonic structures may or may not be present from their ancestors or descendants. Embryonic structures - vestigial (lost); or retained to adulthood. Developmental stages may be repeated in earlier stage but not in later stages. Developmental sequence may or may not be altered in the descendant. Gametes Gametes (sex cells, haploid chromosomes) union to form diploid organism. Gametes perform meiotic division haploidy Sperms (males) have various head appearance: spherical, spatulate, hooked, lancet-shaped, spiraled, capped (acrosome). Sperm: Head (nucleus); Neck (mitochondria); Flagella (movement). Egg cells: Microlecithal (small yolk); Mesolecithal (moderate yolk); Macrolecithal (abundant yolk). Gametogenesis – formation of gametes OOCYTE (egg cell; ovum) Developed and released by ovarian follicle; vitelline membrane (nourishes the egg) In therian mammals: egg is enclosed in thick zona pellucida, corona radiata, cells of ovarian follicle. Other vertebrates: egg is enclosed after ovulation; Jelly layers (amphibians); albumen (birds); horny, membranous (fishes) or calcareous shells (reptiles, birds). Microlecithal: Amphioxus; Eutherian mammals (placental nourishment) Mesolecithal: Lampreys; some fishes; Amphibians Macrolecithal: Most fishes; Reptiles; Birds; Monotremes (egg-laying mammals) Fertilization In therian mammals: Penetration of sperm from corona radiata to vitelline membrane of the egg. Fertilization involves enzymatic and physical interactions between sperm acrosome and egg cortex. Union of gametes: restores the diploid # of chromosomes. Activation of egg to refract additional sperm entry. Zygote (fertilized egg) Embryo (multicellular) FERTILIZATION Cleavage Blastula – structure from series of cell division into multicellular (embryo). Blastomeres – daughter cells Blastocoel - coelom / cavity Yolk – nutrition of developing embryo; process of cleaving & blastula is dependent on the yolk present. Morula (blastula) – multiple numbered cells ready for uterine implantation. Cleavage: 2-celled; 4-celled; 8-celled; 16-celled; 32-celled stages of blastula Vertebrate blastula Composed of single tissue layer with hundred of cells. The cells exhibit cellular polarity related to the axis of the adult body. Polarity: 1) animal pole - embryo 2) vegetal pole – yolk (gut) Blastocoel – body cavity (animal pole) BLASTULA (blastocoel) (vegetal pole) Blastula: Animal and vegetal poles Animal pole – EPIBLAST or EPIMERE; developing embryo nourished by the vegetal pole (dorsal cells--ECTODERM) Vegetal pole – HYPOBLAST or HYPOMERE or developing yolk; nourishes the embryo GUT (ventral cells--ENDODERM) Microlecithal (Amphioxus) – “holoblastic” (total cleavage furrows penetrate the entire yolk); equal-sized blastomeres Animal pole Blastocoel or Gut Vegetal pole Cleavage in Amphibians Mesolecithal - holoblastic; unequal-sized blastomeres Vegetal pole – larger blastomeres; nourishes the embryo; slower development Animal pole – smaller blastomeres; developing embryo Blastocoel – towards animal pole Amphibians Cleavage in Birds (Aves) Macrolecithal – “meroblastic” (partial cleavage); unequal-sized blastomeres Vegetal pole – large size yolk mass; too great to be penetrated by cleavage furrow Animal pole – is relatively small (blastoderm) Narrow blastocoel Birds: Yolk > Embryo Cleavage in Mammals Microlecithal - holoblastic; unequal-sized blastomeres Blastula Blastocyst (inner cell mass formation) Trophoblast – superficial layer of cells Blastocoele – towards vegetal pole Embryoblast – animal pole; epimeric cells Endometrium – inner lining of the uterus (mother) Cleavage patterns followed by holoblastic (total cleavage) and meroblastic (partial cleavage) eggs Holoblastic Radial (sea urchin, amphioxus) Bilateral (tunicates, amphibians) Spiral (annelids, mollusks) Rotational (mammals) Meroblastic Discoidal (fish, birds, reptiles) Superficial (insects) Gastrulation: formation of three germ layers Blastula Gastrula Animal pole – EPIBLAST or EPIMERE; developing embryo (dorsal cells--ECTODERM) Vegetal pole – HYPOBLAST or HYPOMERE or Yolk; nourishes the embryo GUT (ventral cells-ENDODERM) Germ layers: – 1) Ectoderm – from epiblast (animal pole); outermost layer – 2) Mesoderm – middle layer; MESENCHYME – 3) Endoderm – from hypoblast (vegetal pole); innermost layer (animal pole) BLASTULA (blastocoel) (vegetal pole) Ectoderm Mesoderm Endoderm Gastrula Gastrulation Cells migrate to the interior of the blastula, consequently forming two (in diploblastic animals) into three (triploblastic) germ layers. The embryo during this process is called a gastrula. Among the different animals, different combinations of the following processes occur to place the cells in the interior of the embryo: – Epiboly - expansion of one cell sheet over other cells – Ingression - cells move with pseudopods – Invagination - forming the mouth, anus, and archenteron – Delamination - the external cells divide, leaving the daughter cells in the cavity – Polar proliferation In most animals a blastopore is formed at the point where cells are entering the embryo. Two major groups of animals can be distinguished according to the blastopore's fate. In deuterostomes the anus forms from the blastopore (formerly the mouth), while in protostomes it develops into the mouth. Blastopore Blastopore – an opening that resulted from invagination of cells towards embryo A. Protostomes B. Deuterostomes Holoblastic Gastrula: mesoderm formation Vegetal (yolk) pole – folds inward towards animal pole double-walled cup Cup opening – blastopore continues to another end Gastrocoel – body cavity (coelom) Ectoderm – uppermost layer of cells Archenteron – primitive gut (yolk) lined by endoderm Mesoderm – middle layer formed from dorsolateral outpocketing of archenteron Notochord – anlagen of nervous system (primitive), formed from dorsal wall of archenteron Ectoderm Mesoderm Endoderm Archenteron (primitive gut) Gastrula Holoblastic gastrula Notochord – anlagen of nervous system (primitive), formed from dorsal wall of archenteron Chordamesoderm – roof of the archenteron, forms the notochord in the midline and somites (series of paired mesodermal tissue blocks) No outpocketing from the gut tube - schizocoel Coelom – formed by splitting of the hypoblast in the somites (schizocoel – pouch form) Mammalian embryo Blastocoel – indirectly filled with maternal fluid and enlarges Blastocyst Delamination of endoderm (hypoblast) from inner cell mass and cover the trophoblast Blastoderm – (or embryoblast) flattened inner cell mass forms primitive streak (notochord CNS) Mesoderm – forms extraembryonic membranes (fetal membranes) and body of embryo (musculoskeletal and circulatory system); Mesenchymal tissue Coelom – forms by schizocoel (pouch form) Embryoblast or Blastoderm inner cell mass + primitive streak Primitive streak – notochord (anlagen of CNS); promotes neurulation Germ layers Ectoderm – Central nervous system, retina and lens, cranial and sensory, ganglia and nerves, pigment cells (melanocytes), head connective tissue, epidermis of skin, hair, mammary glands. Mesoderm – musculoskeletal system, circulatory system, dermis of skin, connective tissue, urogenital system, heart, blood (lymph cells), and spleen. Endoderm – gastrointestinal system (gut); stomach, colon, liver, pancreas, urinary bladder; lining of urethra, epithelial parts of trachea, lungs, pharynx, thyroid, parathyroid, intestine. Neurulation Overlaps with gastrulation establishing the central nervous system (CNS) Gastrula Neurula Neurocoel – coelom Chordamesoderm – induces thickening of the ectoderm into a neural plate Neural crest cells – (Ectomesenchyme) considered as the 4th germ layer which arised from ECTODERM forms cartilage & bones of the head, pharyngeal cartilages; peripheral nerve ganglia, some glandular tissues; melanocytes Neural plate – ectodermal cells Neural folds – formed from neural plate; fold of cells that arches and meet at the mid-dorsal line and forms the neural tube. Neural tube – encloses the neurocoel (cavity) Neurocoel – anlagen of the dorsal hollow nerve cord (+) chordates Organogenesis Migration of primordial germ cells Holoblastic embryo: Establishes the head and tail Meroblastic embryo: 3 germ layers spread faced down on the uncleaved yolk Embryo increase in length while the archenteron becomes part of the gut tube Schizocoely Enterocoelom (blastopore becomes anus while the mouth opens anteriorly. Organogenesis: Endoderm Endoderm form the lining of the gut. Archenteron: Foregut, Midgut & Hindgut Gut Major Structures Foregut Epithelium: oral cavity, nasal cavity, pharynx, gill, esophagus, lung Midgut Visceral organs: stomach, liver, gall bladder, pancreas, intestines, germ cells of gonads (ovary & testis), part of the yolk sac membrane, part of the allantois Hindgut Urinary bladder; cloaca / anus Organogenesis: Mesoderm Mesoderm layer Major structure Outer Epimere – Dermatome Skin Dermis Middle Epimere – Myotome Muscles: axial, appendicular, branchiomeric, hypobranchial Inner Epimere – Sclerotome Vertebral column Chordamesoderm Notochord Intermediate mesoderm Kidney; Urogenital ducts Somatic hypomere Bones: ribs, sternum (breast plate), appendicular skeleton, some appendicular muscle, parietal peritoneum (outer abdominal wall); genital ridge; part of amnion and chorion Splanchnic hypomere Blood, heart, gut, smooth muscle, visceral peritoneum (inner abdominal wall); yolk sac and allantois Organogenesis: Ectoderm Ectoderm layer Major structure Somatic ectoderm Skin epidermis; Teeth enamel; stomodeum (mouth); proctodeum (anus/cloaca); gill epithelium; part of amnion and chorion Neural plate ectoderm Brain; Spinal cord Epidermal placodes Capsules: olfactory (nasal); optic (eye); otic (ear); epibranchial capsules; neuromast (electroreceptors organs); ganglia of some cranial nerves Ectomesenchyme (Neural crest) Spinal ganglia; splanchnocranium; neurocranium (part); dermatocranium (part); teeth dentine; eye cornea; chromatophores (pigment cells); branchiomeric muscles (part); aortic arches; heart septum (divides the right and left side of heart) Extraembryonic membranes (fetal membranes) and Placenta Amphibians – yolk cleaves and directly incorporated into somatic cells (body cells) Yolk supply – limited Early hatching Larva No fetal membranes required to sustain larva Fishes Yolk remains uncleaved, eggs are laid in water; yolk sac is the only fetal membrane formed. Yolk sac – formed from 3 germ layers to absorb the yolk into the body Respiration and excretion are direct contact with the environment Reptiles and Birds (Avians) Primitive streak extraembryonic mesoderm splits forms extraembryonic coelom SPLANCHNOPLEURE Splanchnopleure + adjacent Yolk = Yolk sac Allantois (vascular) – under the shell; near the chorioallantoic membrane. Allantois (respiration, receives excretory wastes; absorbs albumen; takes some minerals from the shell) Somatopleure + head fold of amnion = Chorion (outer) and Amnion (inner) Amnion – “water bag” contains amniotic fluid that bathes the embryo; cushions the embryo (shelter for growth) Eutherian mammals Placental mammals – nourish their young in the uterus, exchange and nutrition of fetus occurs between fetal and maternal bloodstream by PLACENTA Yolk sac and Allantois – vascularized (blood vessels) supplies nutrition to placenta Chorion and Amnion – avascular (no blood vessels) cannot support placenta Allantoic circulation (birds & reptiles) – homologous to umbilical circulation (mammals) Amnion – sac filled with amniotic fluid bathes the fetus; allows the early fetus to move freely and protects the fetus from pressure of the maternal abdomen Amniotes – collective terms for reptiles, birds and mammals because the possess amnion. PLACENTAL FORMATION Maternal contribution: Chorion (lies between allantois & uterus) incorporated into the placenta CHORIOALLANTOIC Membrane Fetal contribution: Allantois (in most mammals) as fetus grows, allantois decreases in size Prototherians – egg-laying mammals; deposits egg in a pouch (oviparous) Metatherians – marsupials (pouched animals) – no typical placenta Eutherians – placental mammals (viviparous)