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Processes and Stages of Development BIOSC 041 Reproduction & Development Reference: Lecture; Chapters 46 & 47 The Processes of Development v Development § A series of progressive changes in shape, form, function § Begins with fertilization of egg with sperm à zygote § Continues throughout an organism’s life cycle v Development consists of § Growth § Determination § Differentiation § Morphogenesis v The processes of development v Influence of cytoplasmic segregation and induction in cell determination v Stages of development 1. Cytoplasmic determinants (egg and sperm) 2. Fertilization 3. Cleavage: repackaging the cytoplasm 4. Gastrulation: producing the body plan 5. Neurulation: initiating the nervous system 6. Extraembryonic membranes The Processes of Development v Growth § Occurs by cell division and/or expansion § Starts with zygote § Repeated mitotic cell divisions increase cell # § May or may not increase actual size of embryo The Processes of Development The Processes of Development v The fertilized egg is totipotent § it can give rise to all other cell types of the organism v Morphogenesis § The shaping of the multicellular body and its organs § Results from pattern formation § Organization of differentiated tissues into specific structures (e.g., limbs, organs) § Influenced by signal transduction pathways that control: § Cell movement within developing embryo (animals) § Apoptosis (programmed cell death) § Ultimately under genetic control v Determination § Commitment of a cell to a particular fate § Loss of totipotency § Influenced by the extracellular environment and the cell itself acting on the its own genome v Differentiation § Follows determination § Results from differential gene expression § Cells become structurally and functionally specialized 1 Cell Determination: Cytoplasmic Segregation & Induction v Chemical signals control cell differentiation v Two signal mechanisms are known 1. Cytoplasmic segregation § A factor (signal) is unequally distributed in the zygote and ends up in some daughter cells but not others § These cytoplasmic determinants are often transcription factors (proteins that bind DNA to control gene expression) 2. Induction Cell Determination: Cytoplasmic Segregation v Polarity occurs early in development § Establishes major body axes § Anterior–posterior axis (head – tail) § Dorsal–ventral axis (back – belly) § The earliest step in differentiation v Materials are distributed unequally in egg and zygote § Dividing cells receive unequal amounts § Generates polarity in early embryo § A factor (signal) is produced and secreted by some cells to induce other cells to differentiate Cytoplasmic Segregation: Asymmetry in the Early Embryo (Part 1) Cytoplasmic Segregation: Asymmetry in the Early Embryo (Part 2) Cell Determination: Induction The Stages of Animal Development Example: development of the vertebrate eye 1. Developing forebrain bulges out at both sides & forms optic vesicles 2. Optic vesicles signal surface tissue à induce it to form lens placodes 3. Lens placodes differentiate into lenses 4. Developing lens induces surface tissue to develop into the cornea 1. Cytoplasmic determinants: Eggs and Sperm are not created equal 2. Fertilization: the onset of development 3. Cleavage: Repackaging the Cytoplasm 4. Gastrulation: Producing the Body Plan 5. Neurulation: Initiating the Nervous System 6. Extraembryonic Membranes 2 1. Sperm & Egg differ in size 2. Fertilization v Eggs contain cytoplasmic determinants § Regulate expression of genes affecting developmental fate of cells § Strong influence on early development § Cytoplasmic factors set anterior/posterior, and dorsal/ ventral axes v Union of gametes to form zygote (usually diploid) The acrosomal reaction (Fig 47.3) 2. Fertilization v Contact of (only 1!) sperm with egg’s surface § Initiates metabolic reactions within egg that trigger onset of embryonic development 1. Acrosomal reaction: 1-3 seconds after contact § Sperm releases hydrolytic enzymes that digest material surrounding the egg § Depolarization of egg cell membrane = fast block to polyspermy 2. Cortical reaction: rise in Ca2+ stimulates cortical granules 1 Contact. The sperm cell contacts the egg’s jelly coat, triggering exocytosis from the sperm’s acrosome. 2 Acrosomal reaction. Hydrolytic enzymes released from the acrosome make a hole in the jelly coat, while growing actin filaments form the acrosomal process. This structure protrudes from the sperm head and penetrates the jelly coat, binding to receptors in the egg cell membrane that extend through the vitelline layer. 3 Contact and fusion of sperm 4 Entry of and egg membranes. A hole sperm nucleus. is made in the vitelline layer, allowing contact and fusion of the gamete plasma membranes. The membrane becomes depolarized, resulting in the fast block to polyspermy. Sperm plasma membrane Sperm nucleus 5 Cortical reaction. Fusion of the gamete membranes triggers an increase of Ca2+ in the egg’s cytosol, causing cortical granules in the egg to fuse with the plasma membrane and discharge their contents. This leads to swelling of the perivitelline space, hardening of the vitelline layer, and clipping of sperm-binding receptors. The resulting fertilization envelope is the slow block to polyspermy. to release contents into perivitelline space Acrosomal process Basal body (centriole) Fertilization envelope Sperm head Actin Acrosome Jelly coat (Cumulus) Sperm-binding receptors Fused plasma Cortical membranes granule Perivitelline Hydrolytic enzymes space Vitelline layer (zona pelucida) Egg plasma membrane Cortical granule membrane EGG CYTOPLASM 2. Fertilization Types of Fertilization v These changes cause the formation of a fertilization envelope § Functions as a slow block to polyspermy 1. Internal fertilization § Male transfers sperm via a penis or other organ into female’s body 2. External fertilization § “Spawning”- fishes, amphibians, reptiles Salmon spawning video: http://www.youtube.com/watch?v=SOllUTBYZtE 3 3. Cleavage The Mammalian Zygote Becomes a Blastocyst (Part 1) v Rapid DNA replication and mitosis § Little actual growth § Little gene expression v Formation of blastula § Large zygote à morula à blastula with blastocoel § Blastomeres = Individual cells of blastula v Influenced by: § Yolk (impedes cleavage furrow- cleavage often incomplete/superficial) § Maternal cytoplasmic determinants § Mitotic spindle orientation determines cleavage plane § Radial pattern à frogs, sea urchins § Spiral pattern à molluscs (snail shells) § Rotational pattern à (mammals = us) Cleavage in the Mammalian Embryo Cleavage in Xenopus (frog): Fate Maps v Specific blastomeres generate specific tissues & organs § Determined by unequal complements of nutrients and cytoplasmic determinants in the egg Day 6-7 Day 1-2 Day 5 Day 2-3 Day 3-4 4. Gastrulation: Producing the Body Plan (Review) Gastrulation can look very different between groups v Gastrulation § Blastula transforms into embryo with three germ layers, and defined body axes v Three germ layers form: § Endoderm à inner germ layer § Gives rise to digestive, circulatory, and respiratory tracts § Mesoderm à middle germ layer § Bone, muscle, liver, heart, and blood vessels § Ectoderm à outer germ layer § Epidermis, nervous system 4 Gastrulation in Sea Urchins Gastrulation in the Frog Embryo *Primary embryonic organizer The dorsal lip is very important to development Gastrulation in the Chick Embryo 5. Organogenesis and Neurulation Neurulation in the Frog Embryo (Part 1) v Gastrulation produces an embryo with three germ layers v Organogenesis occurs next- formation of organs and organ systems v Neurulation occurs early in organogenesis- begins the formation of the nervous system in vertebrates 5 Neurulation in the Frog Embryo (Part 2) 6. Extraembryonic Membranes v Extraembryonic membranes § Originate from germ layers of embryo § Function in nutrition, gas exchange, waste removal v In the chicken, § Yolk sac forms first § Extension of the endodermal tissue of the hypoblast § Constricts at the top to create a tube continuous with embryo gut § Yolk is digested by the endodermal cells of the yolk sac § Nutrients transported through blood vessels lining outer surface of yolk sac Chick Embryo Extraembryonic Membranes Human Embryo Development The Mammalian Placenta Summary of Development/Reproduction 6 Useful video resources from Khan Academy Stages of development (video, ~10 min): https://www.khanacademy.org/partner-content/crash-course1/crashcourse-biology/v/crash-course-biology-115 Genes (such as Hox) that regulate development (video, ~10 min): https://www.khanacademy.org/partner-content/crash-course1/crashcourse-biology/v/crash-course-biology-116 7