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22/05/12 Scholarships 2012- 2013 The Ed Fund is currently accepting applications for next year’s scholarships. Please let returning students know that it’s a simple one-page application for both meritand need-based awards, and direct them to https://northseattle.edu/edfund-scholarships to learn more! The deadline is May 31. Commit to Complete Signing Ceremony. • Phi Theta Kappa Alpha Epsilon Omega Chapter and North Seattle Community College • http://www.cccompletioncorps.org/ • Today, Weds. 23 May at 1:15 • in the hallway on the way to the cafeteria • President Mark Mitsui will lead this ceremony. • Students are encouraged to attend! Animal Development Common themes Differences INTERCONNECTEDNESS REDUCE – REUSE – RECYCLE 1 22/05/12 Figure 47.1 1 mm Fertilization • Molecules and events at the egg surface play a crucial role in each step of fertilization – Sperm penetrate the protective layer around the egg – Receptors on the egg surface bind to molecules on the sperm surface – Changes at the egg surface prevent polyspermy, the entry of multiple sperm nuclei into the egg The Acrosomal Reaction • The acrosomal reaction is triggered when the sperm meets the egg • The acrosome at the tip of the sperm releases hydrolytic enzymes that digest material surrounding the egg © 2011 Pearson Education, Inc. 2 22/05/12 Figure 47.3-5 Sperm plasma membrane Sperm nucleus Basal body (centriole) Sperm head Acrosome Jelly coat Sperm-binding receptors Fertilization envelope Acrosomal process Actin filament Cortical Fused granule plasma membranes Hydrolytic enzymes Perivitelline space Vitelline layer Egg plasma membrane EGG CYTOPLASM • Gamete contact and/or fusion depolarizes the egg cell membrane and sets up a fast block to polyspermy • The Cortical Reaction • Fusion of egg and sperm also initiates the cortical reaction • Seconds after the sperm binds to the egg, vesicles just beneath the egg plasma membrane release their contents and form a fertilization envelope • The fertilization envelope acts as the slow block to polyspermy • The cortical reaction requires a high concentration of Ca2+ ions in the egg © 2011 Pearson Education, Inc. • Development occurs at many points in the life cycle of an animal • This includes metamorphosis and gamete producEon, as well as embryonic development © 2011 Pearson Education, Inc. 3 22/05/12 Egg Ac&va&on • The rise in Ca2+ in the cytosol increases the rates of cellular respiraEon and protein synthesis by the egg cell • With these rapid changes in metabolism, the egg is said to be acEvated • The proteins and mRNAs needed for acEvaEon are already present in the egg • The sperm nucleus merges with the egg nucleus and cell division begins © 2011 Pearson Education, Inc. Fer&liza&on in Mammals • FerElizaEon in mammals and other terrestrial animals is internal • SecreEons in the mammalian female reproducEve tract alter sperm moElity and structure • This is called capacitaEon, and must occur before sperm are able to ferElize an egg © 2011 Pearson Education, Inc. • Sperm travel through an outer layer of cells to reach the zona pellucida, the extracellular matrix of the egg • When the sperm binds a receptor in the zona pellucida, it triggers a slow block to polyspermy • No fast block to polyspermy has been idenEfied in mammals • In mammals the first cell division occurs 12-‐36 hours aUer sperm binding • The diploid nucleus forms aUer this first division of the zygote © 2011 Pearson Education, Inc. 4 22/05/12 Figure 47.5 Zona pellucida Follicle cell Sperm nucleus Sperm basal body Cortical granules Cleavage • FerElizaEon is followed by cleavage, a period of rapid cell division without growth • Cleavage parEEons the cytoplasm of one large cell into many smaller cells called blastomeres • The blastula is a ball of cells with a fluid-‐filled cavity called a blastocoel © 2011 Pearson Education, Inc. Figure 47.6 50 µm (a) Fertilized egg (b) Four-cell stage (c) Early blastula (d) Later blastula 5 22/05/12 Cleavage Pa6erns • In frogs and many other animals, the distribuEon of yolk (stored nutrients) is a key factor influencing the paYern of cleavage • The vegetal pole has more yolk; the animal pole has less yolk • The difference in yolk distribuEon results in animal and vegetal hemispheres that differ in appearance © 2011 Pearson Education, Inc. • The first two cleavage furrows in the frog form four equally sized blastomeres • The third cleavage is asymmetric, forming unequally sized blastomeres • Holoblastic cleavage, complete division of the egg, occurs in species whose eggs have little or moderate amounts of yolk, such as sea urchins and frogs • Meroblastic cleavage, incomplete division of the egg, occurs in species with yolk-rich eggs, such as reptiles and birds © 2011 Pearson Education, Inc. Figure 47.7a-5 Animal pole Gray crescent Zygote 2-cell stage forming Blastocoel Vegetal pole 4-cell stage forming 8-cell stage Blastula (cross section) 6 22/05/12 Figure 47.7 Zygote 2-cell stage forming Gray crescent 0.25 mm 8-cell stage (viewed from the animal pole) 4-cell stage forming Animal pole 8-cell stage 0.25 mm Blastula (at least 128 cells) Vegetal pole Blastocoel Blastula (cross section) Regula&on of Cleavage • Animal embryos complete cleavage when the raEo of material in the nucleus relaEve to the cytoplasm is sufficiently large © 2011 Pearson Education, Inc. Concept 47.2: Morphogenesis in animals involves specific changes in cell shape, posiEon, and survival • AUer cleavage, the rate of cell division slows and the normal cell cycle is restored • Morphogenesis, the process by which cells occupy their appropriate locaEons, involves – Gastrula:on, the movement of cells from the blastula surface to the interior of the embryo – Organogenesis, the formaEon of organs © 2011 Pearson Education, Inc. 7 22/05/12 GastrulaEon • Gastrulation rearranges the cells of a blastula into a three-layered embryo, called a gastrula • The three layers produced by gastrulation are called embryonic germ layers – The ectoderm forms the outer layer – The endoderm lines the digestive tract – The mesoderm partly fills the space between the endoderm and ectoderm • Each germ layer contributes to specific structures in the adult animal Video: Sea Urchin Embryonic Development © 2011 Pearson Education, Inc. Figure 47.8 ECTODERM (outer layer of embryo) • Epidermis of skin and its derivatives (including sweat glands, hair follicles) • Nervous and sensory systems • Pituitary gland, adrenal medulla • Jaws and teeth • Germ cells MESODERM (middle layer of embryo) • Skeletal and muscular systems • Circulatory and lymphatic systems • Excretory and reproductive systems (except germ cells) • Dermis of skin • Adrenal cortex ENDODERM (inner layer of embryo) • Epithelial lining of digestive tract and associated organs (liver, pancreas) • Epithelial lining of respiratory, excretory, and reproductive tracts and ducts • Thymus, thyroid, and parathyroid glands Gastrula&on in Sea Urchins • Gastrulation begins at the vegetal pole of the blastula • Mesenchyme cells migrate into the blastocoel • The vegetal plate forms from the remaining cells of the vegetal pole and buckles inward through invagination • The newly formed cavity is called the archenteron • This opens through the blastopore, which will become the anus © 2011 Pearson Education, Inc. 8 22/05/12 Figure 47.9a-5 Blastocoel Animal pole Mesenchyme cells Filopodia Vegetal Vegetal pole plate Archenteron Blastocoel Archenteron Digestive tube (endoderm) Ectoderm Blastopore Key Mouth Mesenchyme (mesoderm forms future skeleton) Future ectoderm Future mesoderm Anus (from blastopore) Future endoderm Figure 47.9b Blastocoel Filopodia Mesenchyme cells Blastopore Archenteron 50 µm Variations • Frogs • Chicks • Humans 9 22/05/12 Gastrula&on in Frogs • Frog gastrulation begins when a group of cells on the dorsal side of the blastula begins to invaginate • This forms a crease along the region where the gray crescent formed • The part above the crease is called the dorsal lip of the blastopore • Cells continue to move from the embryo surface into the embryo by involution • These cells become the endoderm and mesoderm • Cells on the embryo surface will form the ectoderm © 2011 Pearson Education, Inc. Gastrula&on in Chicks • Prior to gastrulation, the embryo is composed of an upper and lower layer, the epiblast and hypoblast, respectively • During gastrulation, epiblast cells move toward the midline of the blastoderm and then into the embryo toward the yolk • The midline thickens and is called the primitive streak • The hypoblast cells contribute to the sac that surrounds the yolk and a connection between the yolk and the embryo, but do not contribute to the embryo itself © 2011 Pearson Education, Inc. Figure 47.11 Fertilized egg Primitive streak Embryo Yolk Primitive streak Epiblast Future ectoderm Blastocoel Migrating cells (mesoderm) Endoderm Hypoblast YOLK 10 22/05/12 Gastrula&on in Humans • Human eggs have very liYle yolk • A blastocyst is the human equivalent of the blastula • The inner cell mass is a cluster of cells at one end of the blastocyst • The trophoblast is the outer epithelial layer of the blastocyst and does not contribute to the embryo, but instead iniEates implantaEon © 2011 Pearson Education, Inc. Figure 47.12 Endometrial epithelium (uterine lining) 1 Blastocyst reaches uterus. Uterus Inner cell mass Trophoblast Blastocoel 2 Blastocyst implants (7 days after fertilization). Expanding region of trophoblast Maternal blood vessel Epiblast Hypoblast Trophoblast 3 Extraembryonic membranes start to form (10–11 days), and gastrulation begins (13 days). Expanding region of trophoblast Amniotic cavity Epiblast Hypoblast Yolk sac (from hypoblast) Extraembryonic mesoderm cells (from epiblast) Chorion (from trophoblast) 4 Gastrulation has produced a three-layered embryo with four extraembryonic membranes. Amnion Chorion Ectoderm Mesoderm Endoderm Yolk sac Extraembryonic mesoderm Allantois Developmental AdaptaEons of Amniotes • The colonizaEon of land by vertebrates was made possible only aUer the evoluEon of – The shelled egg of birds and other repEles as well as monotremes (egg-‐laying mammals) – The uterus of marsupial and eutherian mammals © 2011 Pearson Education, Inc. 11 22/05/12 • In both adaptaEons, embryos are surrounded by fluid in a sac called the amnion • This protects the embryo from desiccaEon and allows reproducEon on dry land • Mammals and repEles including birds are called amniotes for this reason © 2011 Pearson Education, Inc. • The four extraembryonic membranes that form around the embryo – The chorion funcEons in gas exchange – The amnion encloses the amnioEc fluid – The yolk sac encloses the yolk – The allantois disposes of waste products and contributes to gas exchange © 2011 Pearson Education, Inc. Organogenesis • During organogenesis, various regions of the germ layers develop into rudimentary organs • Early in vertebrate organogenesis, the notochord forms from mesoderm, and the neural plate forms from ectoderm © 2011 Pearson Education, Inc. 12 22/05/12 Figure 47.13a Neural folds Neural fold 1 mm Neural plate Notochord Ectoderm Mesoderm Endoderm Archenteron (a) Neural plate formation • The neural plate soon curves inward, forming the neural tube • The neural tube will become the central nervous system (brain and spinal cord) Video: Frog Embryo Development © 2011 Pearson Education, Inc. Figure 47.13b-3 Neural fold Neural plate Neural crest cells Neural crest cells (b) Neural tube formation Outer layer of ectoderm Neural tube 13 22/05/12 • Neural crest cells develop along the neural tube of vertebrates and form various parts of the embryo (nerves, parts of teeth, skull bones, and so on) • Mesoderm lateral to the notochord forms blocks called somites • Lateral to the somites, the mesoderm splits to form the coelom (body cavity) © 2011 Pearson Education, Inc. Figure 47.13c Eye SEM Neural tube Notochord Coelom Somites Tail bud 1 mm Neural crest cells Somite (c) Somites Archenteron (digestive cavity) 14