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2010/10/1 Cell and Embryology Textbook: Wolpert L, Beddington R, Jessell T, Lawrence P, Meyerowitz E, Smith J. (2007) Principles of Development. 3th ed. London: Oxford university press. Gilbert SF. (2003) Development Biology. 7th ed. Sunderland: Sinaure Associates Inc. Schedule introduction Basic concept Exam I Fertilization Exam II Model systems Exam III Patterning the vertebrate body plan I: Axes and germ layer Exam VI Patterning the vertebrate body plan II: the mesoderm and early nervous Exam V Development of nematodes, fish, sea urchins ascidians and slime mold Exam VI Human embryology or development biology Final exam VII http://www2.nsysu.edu.tw/MR-embryology/index.htm 1 2010/10/1 • development Why Study Development? I am fearfully and wonderfully made. (Psalm 139) There are a Handful of Major Model Organisms Overview of basic embryonic development 2 2010/10/1 Zebrafish (Danio (Danio rerio) rerio) -- A Vertebrate Model Zebrafish as a High-throughput Model for biomedical Research and Therapeutic Development •It is 3 cm long •Short generation time •Large clutch size Forward Genetics: ENU mutagenesis Insertional mutagenesis •External fertilization Large number of offspring Optically clear embryos Short generation time S ll Si Small Size Reverse Genetics: Transgenic fish Tilling with ENU Morpholino injection •Transparent embryos •Rapid development Small Molecule Screens: Predictive of higher vertebrates Delivery by injection or soaking http://zfin.org/ and http://www.nih.gov/science/models/zebrafish/ Carcinogenesis: Aqueous delivery Similar to human tumors Model organism cleavage The organism chosen for understand broad biological principles is called a model organism. ARABIDOPSIS THAMAN MUS MUSCULUS DROSOPHILA MELANOGASTER (COMMON WALL CRES (MOUSE) (FRUIT FLY) CAENORHABDITIS ELEGANS (NEMATODE) Genomics: Sequenced Genome cDNA projects Microarrays DANIO RERIO (ZEBRAFISH) once fertilization is completed, a succession of rapid cleavage ensues the cells undergo the 1) S (DNA synthesis) 2) M (mitosis) phases of the cell cycle but often no G1 and G2 phases No human, why? 0.25 mm Figure 21.2 3 2010/10/1 cleavage in echinoderm embryo Cleavage to neurulation the embryo does not enlarge during this period but simply partitions the cytoplasm of the zygote into many smaller cells, called blastomeres and each with its own nucleus Gastrulation: External view Gastrulation: Internal view 4 2010/10/1 Caenorhabditis elegans Xenopus Drosophila 5 2010/10/1 Experimental Approaches • What causes cell differentiation: cytoplasm or nucleus? • • • • – Defect experiment: Defect experiments Isolation experiments Recombination experiments Transplantation experiments Spemann and Mangold’s Discovery of Induction (1924) Isolation experiment Conditional specification? 6 2010/10/1 after fertilization, embryonic development proceeds through cleavage, gastrulation, and organogenesis • Many different structures – Are derived from the three embryonic y germ g layers y during g organogenesis 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 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 Figure 47.16 fertilization in sea urchin model fertilization in sea urchin model fig 47.3 cortical reaction: sperm binding activate a signal transduction pathway involving 2 second messengers, IP3 and d DAG, that cause Ca2+ to be released from the egg’s endoplasmic reticulum (ER) into the cytosol a surge in Ca2+ levels in the cytoplasm fig 11.12 7 2010/10/1 fertilization in sea urchin model fertilization in sea urchin model cortical reaction: the Ca2+ release from the ER begins at the site of sperm entry and then propagates in a wave across the fertilized egg fig 47.5 fig 47.4 cleavage in echinoderm embryo cleavage once fertilization is completed, a succession of rapid cleavage ensues the cells undergo the 1) S (DNA synthesis) 2) M (mitosis) phases of the cell cycle but often no G1 and G2 phases fig 47.7 the embryo does not enlarge during this period but simply partitions the cytoplasm of the zygote into many smaller cells, called blastomeres and each with its own nucleus 8 2010/10/1 cleavage in frog embryo cleavage in frog embryo morula: the first 5 to 7 divisions form a cluster of embryonic cells blastocoel a fluid-filled cavity of the early embryobegins to form within the morula and is fully formed in the blastula blastula: a hollow ball of embryonic cells the body axes has been determined before fertilization and been i t intensively i l studies t di iin many ffrog spp polarity of the zygote except the egg of mammals, the eggs and zygotes of animals have a definite polarity is due to uneven distributions of mRNAs, proteins, and yolk cleavage in bird embryo cleavage in bird embryo fig 47.10 meroblastic cleavage: the incomplete division of a yolk-rich egg, i.e. cleavage of the fertilized egg is restricted to the small disk of yolk-free yolk free cytoplasm and cannot penetrate through the dense yolk the yolk remains uncleaved holoblastic cleavage: the complete division of eggs having little yolk (as in sea urchins) or a moderate amount of yolk (as in frogs) blastoderm the avian equivalent of the blastula blastomeres devidedinto 2 layers 1)) epiblast: upper 2) hypoblast: lower blastocoel the cavity between epiblast and hypoblast layers 9 2010/10/1 gastrulation in sea urchin embryo gastrulation gastrula: some of the cells at or near the surface of the blastula move to an interior location and the embryo becomes 3-germ-layer embryo allows cells to interact with each other in new ways ectoderm: forms the outer layer of the gastrula endoderm: the embryonic digestive tract mesoderm: partly fills the space between the ectoderm and the endoderm eventually, these 3 cell layers develop into all the tissues and organs of the adult animal gastrulation in frog embryo gastrulation in bird embryo • • involution: a process along the blastopore, future endoderm and mesoderm cells on the surface f rollll over th the edge d off th the lip into the interior of the embryo blastocoel collapses and displaced by archenteron which is formed by the tube of endoderm fig 47.12 47 12 10 2010/10/1 organogenesis in frog embryo organogenesis in frog embryo somites: condensations occur in strips of mesoderm lateral to the notochord, which separate into blocks off somites, being arranged serially on both sides along the notochord parts of the somites dissociate into individual mesenchymal cells, which migrate to new locations vertebrae: t b th notochord the t h d functions as a core around mesodermal cells fig 47.14 organizer region inductive signals and the cell fate fig 47.25 Spemann and Mangold g in 1920s dorsal lip of the blastopore functions as an organizer by initiating a chain of inductions 11 2010/10/1 formation of the limb in chick model formation of the limb in chick model fig 47.27 homeotic genes and pattern formation homeotic genes and pattern formation in the development of the body segments of Drosophila and Mus organogenesis adult derivatives of the three embryonic germ layers in vertebrates fig 47.16 12 2010/10/1 cytoskeleton in morphogenesis tab 6.1 13