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Developmental Zoology (ZOO 228.1.0) Gatrulation 1 Developmental Stages Ø Early Development • Fertilization • Cleavage • Gastrulation • Neurulation Ø Later Development • Organogenesis • Larval molts • Metamorphosis • Aging § § Cell movements occur in distinct ways in different genera Invertebrates § invagination of cells on the future ventral side § Vertebrates § fish & amphibians § spherical embryos § involution of mesoderm on future dorsal side at blastopore lip 2 Developmental Stages Ø Early Development • Fertilization • Cleavage • Gastrulation • Neurulation Ø Later Development • Organogenesis • Larval molts • Metamorphosis • Aging Ø Vertebrates • Reptiles, birds and some mammals (humans) – disc embryos – de-lamination of ectoderm to form endoderm & mesoderm – delaminating cells involute at Henson s node, primitive streak and groove form & progresses along dorsal midline 3 Developmental Stages Ø Early Development Ø Vertebrates • mammals (mouse) • Fertilization – cup embryos • Cleavage – ingression at the dorsal • Gastrulation midline (primitive streak) as in chick and • Neurulation human Ø Later Development • Organogenesis • Larval molts • Metamorphosis • Aging 4 Questions How does a single celled zygote become a gastrula? How are layers of tissue established? How do cells get to their proper positions for development? • In part, this is accomplished by various types of Morphogenetic Movements. • Induction Differentiation • Organogenesis is the formation of the organs (Organo = organs, genesis = creation) • Arises from the layering of cells that occurs during gastrula stage • The layers are germ layers; they have specific fates in the developing embryo: Differentiation – Endoderm • The innermost layer • Goes on to form the gut – Mesoderm • In the middle • Goes on to form the muscles, circulatory system, blood and many different organs – Ectoderm • The outermost • Goes on to form the skin and nervous system Differentiation of Primary Germ Layers (from the gastrula) Ectoderm Mesoderm Endoderm Nervous system Epidermis of skin Skeleton Digestive tract Muscles Respiratory system Liver, pancreas Circulatory system Gonads Bladder Morphogenetic Movements Epiboly Morphogenetic Movements Invagination, Evagination Morphogenetic Movements Involution Epiboly over a fold Involution! Morphogenetic Movements Delamination Morphogenetic Movements Ingression Morphogenetic Movements Cell Migration Cell Migration Morphogenetic Movements “Intercalation: rows of cells move between one another, creating an array of cells that is longer (in one or more dimensions) but thinner.” “Convergent Extension: rows of cells intercalate, but the intercalation is highly directional.” - involved in formation of the primitive streak in avian and mammalian embryos. Morphogenetic Movements Differential Growth .. .. .. .. time .... .. .. ... .. .. .. .. .. .. Fate Maps Mother cell Cell division Cell type A Cell type B Cell division places daughter cells in different environments, which can lead to different cell fate choices Fate Maps Induction: information from neighbors influences cell fate inducer responder Induction Competence: ability to respond to a certain inductive signal responder Cell not competent to respond inducer Induction Succesive inductions: can generate many cell types from just a few interactions inducer Cell not competent to respond responder Types of Signals Inducer Responder Types of Signals Signals can act globally throughout the body Induction An initial difference can be amplified into many cell types How is cell fate determined -- the dorsal-ventral axis in newts How is cell fate determined -- the dorsal-ventral axis in newts Spemann & Mangold: the organizer can influence neighboring cells to form a secondary body axis: Figure 7.17 Induction Induction of the organizer can influence neighboring cells to form a secondary body axis: Induction If one transplants a second inducer of the organizer the embryo forms two body axes It is not birth, marriage or death, but gastrulation, which is truly the most important time in your life. - Lewis Wolpert (1986) Gastrulation Cell Movements Relevant to Gastrulation Getting cells inside Spreading tissues out Moving cells around Making tissues longer Convergence/extension Gastrulation • Definition = migration and division of cells to set up the 3 primary germ layers.! • What positions do presumptive germ layers occupy in the blastula, before gastrulation? ! – Experimental answers to this question provide our basis for understanding cell movements that occur during gastrulation.! • FATE MAPS = diagram of blastula/blastodisc showing the fate of each part. ! Gastrulation • Fate mapping technique developed by Vogt in 1920s. Involves …! – Marking surface of blastula with vital dyes! – Dyes retained by cells for prolonged periods, but don t interfere with normal cellular processes! – Follow movements of marked cells during gastrulation to ultimate locations in later embryos! Sample Fate Map of Frogs