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F ro g E m b r yo l o gy 2 Sirwan M. Mohammed [email protected] Learning objectives By the end of this lecture students should be able to: Understand the concept of blastulation in frog embryo Know Frog’s early and late gastrulation. Describe neurilation & fate map of frog embryo frog egg frog sperm Gamete Formation Fertilization Cleavage Blastulation →Gastrulation midsectional views Organ Formation top view side view Growth, Tissue Specialization Stages of Embryonic Development The blastula leads to the gastrula through gastrulation which involves cell divisions, cell migrations and cell rearrangements Fertilized egg divides mitotically fluid filled cavity - blastocoel- begins to form Blastula - hollow ball of cells - has between 25 and 27 cells blastocoel The Blastula - Polarity Animal Pole Blastomeres Blastocoel Vegetal Pole Frog Blastula (section) ectoderm blastocoel mesoderm endoderm vegetal pole frog gastrulation mechanism Gastrulation begins when a small indented crease, 1 the dorsal lip of the blastopore, appears on one side of the blastula. The crease is formed by cells changing shape and pushing inward from the surface (invagination). Additional cells then roll inward over the dorsal lip (involution) and move into the interior, where they will form endoderm and mesoderm. Meanwhile, cells of the animal pole, the future ectoderm, change shape and begin spreading over the outer surface. SURFACE VIEW Animal pole CROSS SECTION Blastocoel Dorsal lip Vegetal pole of blastopore Blastula Blastocoel shrinking The blastopore lip grows on both sides of the 2 embryo, as more cells invaginate. When the sides of the lip meet, the blastopore forms a circle that becomes smaller as ectoderm spreads downward over the surface. Internally, continued involution expands the endoderm and mesoderm, and the archenteron begins to form; as a result, the blastocoel becomes smaller. 3 Late in gastrulation, the endoderm-lined archenteron Blastocoel remnant has completely replaced the blastocoel and the three germ layers are in place. The circular blastopore surrounds a plug of yolk-filled cells. Dorsal lip of blastopore Archenteron Ectoderm Mesoderm Endoderm Key Future ectoderm Future mesoderm Future endoderm Yolk plug Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Yolk plug Gastrula The developmental fate of cells depends on their history and on inductive signals Coupled with morphogenetic changes, development requires timely differentiation of cells at specific locations Two general principles cause differentiation: During early cleavage divisions, embryonic cells must become different from one another After cell asymmetries are set up, interactions among embryonic cells influence their fate, usually causing changes in gene expression Organ stages of development occur after tissue stages Newly formed mesoderm cells lie along main longitudinal axis of animal and combine to form a presumptive notochord Formation of the neural tube Thickening of cells, neural plate, appears along the dorsal surface of the embryo Neural folds develop on either side of neural groove Coelom appears and neural tube is complete At this point, the embryo is called a neurula Anterior end of the neural tube develops into the brain, and the rest becomes the spinal cord LE 47-14a Early in vertebrate organogenesis, the notochord forms from mesoderm, and the neural plate forms from ectoderm Neural folds LM 1 mm Neural fold Neural plate Notochord Ectoderm Mesoderm Endoderm Archenteron Neural plate formation Neural fold Neural plate Neural crest Outer layer of ectoderm Neural crest Neural tube Formation of the neural tube The neural plate soon curves inward, forming the neural tube Development of neural tube and coelom in a frog embryo Eye SEM Somites Neural tube Notochord Coelom Archenteron (digestive cavity) Somites Tail bud 1 mm Neural crest Somite Mesoderm lateral to the notochord forms blocks called somites Lateral to the somites, the mesoderm splits to form the coelom Classic studies using frogs – Gave indications that the lineage of cells making up the three germ layers created by gastrulation is traceable to cells in the blastula Epidermis Central nervous system Epidermis Notochord Mesoderm Endoderm Neural tube stage (transverse section) (a) Fate map of a frog embryo. The fates of groups of cells in a frog blastula (left) were determined in part by marking different regions of the blastula surface with nontoxic dyes of various colors. The embryos were sectioned at later stages of development, such as 47.23a the neural tube stage shown on the right, and the locations of the dyed cells determined. Blastula Figure Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Later studies developed techniques – That marked an individual blastomere during cleavage and then followed it through development (b) Cell lineage analysis in a tunicate. In lineage analysis, an individual cell is injected with a dye during cleavage, as indicated in the drawings of 64-cell embryos of a tunicate, an invertebrate chordate. The dark regions in the light micrographs of larvae correspond to the cells that developed from the two different blastomeres indicated in the drawings. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Derivatives of 3 Germ layers in adult vertebrates ECTODERM • Epidermis of skin and its derivatives (including sweat glands, hair follicles) • Epithelial lining of mouth and rectum • Sense receptors in epidermis • Cornea and lens of eye • Nervous system • Adrenal medulla • Tooth enamel • Epithelium or pineal and pituitary glands MESODERM • Notochord • Skeletal system • Muscular system • Muscular layer of stomach, intestine, etc. • Excretory system • Circulatory and lymphatic systems • Reproductive system (except germ cells) • Dermis of skin • Lining of body cavity • Adrenal cortex Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings ENDODERM • Epithelial lining of digestive tract • Epithelial lining of respiratory system • Lining of urethra, urinary bladder, and reproductive system • Liver • Pancreas • Thymus • Thyroid and parathyroid glands References Campbell, N.A. and Reece,J.B.(2005)Biology.7th edition.Pearson Education,Inc. Hickman C. P, L.S. Roberts, A.Larson ,H.I'Anson.(2004)Integrated principles of zoology,12th edition,McGraw-Hill Companies, Inc.Chapter7&8 Slack, J.M.W.(2006)Essential developmental biology.Blackwell Publishing Ltd,Second edition,Chap 7 Mader,S.S,(2009) . Concepts of Biology,McGraw-Hill Companies, Inc.,Chapter35