Survey
* Your assessment is very important for improving the workof artificial intelligence, which forms the content of this project
* Your assessment is very important for improving the workof artificial intelligence, which forms the content of this project
Fertilization. Embryonic development Maria Kazakova, PhD Fertilization • Union of a spermatozoa and an ovum • Initiates human embryonic development • Determines the sex of the offspring – in human Fertilization - steps Spermatozoa are not fully capable of fertilization 1. Capacitation - a process that strips the coat of glycoprotein molecules of the spermatozoa 2. Acrosomal reaction – the cap of the acrosome breaks down and releases hydrolytic enzymes – hyaluronidase, acrosin 3. The corona radiata has been penetrated 4. Contact with zona pellucida – species specific 5. Fusion of membranes The completion of meiosis II 6. Fusion of pronuclei – 30 min., after the first step of fertilization 1. Capacitation - a process that strips the coat of glycoprotein molecules of the spermatozoa Sperms cannot fertilize oocytes when they are newly ejaculated. The process of capacitation takes 5-7 hours in humans Pro-Acrosin (inactive) is converted to acrosin (active) Capacitation occurs in the uterus and oviducts and is facilitated by substances of the female genital tract. o Capacitation alters two crucial aspects of sperm behavior: - It greatly increases the motility of the flagellum - It makes the sperm capable of undergoing the acrosome reaction 2. Acrosomal reaction – the cap of the acrosome breaks down and releases hydrolytic enzymes – hyaluronidase, acrosin - Occurs when sperms come into contact with the corona radiata of the oocyte Acrosome: - is a lysosomal-like compartment derived from the Golgi. It has a low pH and contains soluble hydrolases (serine protease acrosin) 3. The corona radiata has been penetrated 4. Contact with zona pellucida – species specific Contact with zona pellucida – species specific Zona pellucida is composed of three glycoproteins - ZP1, ZP2 and ZP3 Scanning electron micrograph of a human sperm contacting a hamster egg. The zona pellucida of the egg has been removed. The ability of an individual’s sperm to penetrate hamster eggs is used as an assay of male fertility. Penetration of more than 10-25% of the eggs is considered to be normal. Molecular Biology of the Cell (© Garland Science 2008) 5. Fusion of membranes The completion of meiosis II • The secondary oocyte – arrested in metaphase of the 2nd meiotic division forms the mature egg (ovum) and second polar body. Polyspermy barriers 1. A change in the egg plasma membrane Sea urchin by rapid depolarization In mammalian eggs, the mechanism is not known 2. Cortical reaction – releases various enzymes that change the structure of the zona pellucida so that the sperm cannot bind to or penetrate it. 6. Fusion of pronuclei – 30 min., after the first step of fertilization - sea urchin the chromosomes in each pronucleus condense the pronuclear envelopes break down without fusing together the male and female chromosomes intermix in the cytoplasm form the metaphase of the first mitotic spindle Release contents of the cortical granules Inactivation of ZP3 Cleave ZP2 Harding the zona pellucida Molecular Biology of the Cell (© Garland Science 2008) The coming together of the sperm and egg pronuclei after mammalian fertilization Molecular Biology of the Cell (© Garland Science 2008) Embryonic development Life cycles and the evolution of developmental patterns Descriptive embryology - the idea of a generalizable life cycle. Each animal, whether an earthworm, an eagle, or a beagle, passes through similar stages of development. When does embryonic development begin? The Stages of Animal Development Four essential processes by which a multicellular organism is made: 1. Cell proliferation – producing many cells from one 2. Cell specialization - creating cells with different characteristics at different positions 3. Cell interaction - coordinating the behavior of one cell with that of its neighbors 4. Cell movement – rearranging the cells to form structured tissues and organs The stages of development between fertilization and hatching are collectively called embryogenesis. 1. Initiated by the fusion of genetic material from the two gametes the sperm and the egg. 2. Fertilization, stimulates the egg to begin development. 3. Cleavage Cleavage is a series of extremely rapid mitotic divisions where in the enormous volume of zygote cytoplasm is divided into numerous smaller cells. These cells are called blastomeres, and by the end of cleavage, they generally form a sphere known as a blastula. A. The pattern of cleavage is influenced by the amount of yolk in the egg. In eggs with less yolk, cleavages are equal, and the resulting blastomeres are of similar size. B. If the yolk is localized, such as in frog eggs, then cleavages are unequal - the cells derived from the yolky region (the vegetal pole) are larger than those derived from the region without yolk (the animal pole). 4. Gastrulation – the series of extensive cell rearrangements. The embryo contains three germ layers: the ectoderm, the endoderm, and the mesoderm. 5. Organogenesis - the cells interact with one another and rearrange themselves to produce tissues and organs. Principles of experimental embryology I. Environmental Developmental Biology 1. Environmental sex determination In the marine worm Bonellia viridis - is thus determined by external, environmental factors (the presence or absence of bonellin). Larvae become: males if they make physical contact with a female and females if they end up on the bare sea floor 2. Adaptation of embryos and larvae to their environments Phenotypic variations caused by environmental differences are often called morphs. European map butterfly, Araschnia levana, which has two seasonal phenotypes The spring morph is bright orange with black spots The summer form is mostly black with a white band The change from spring to summer morph is controlled by changes in both day length and temperature during the larval period. 3. The developmental mechanics of cell specification 3.1. Specification - cells are capable to differentiate autonomously when placed in a neutral environment such as a petri dish or test tube. 3.2. Determination - cells are capable to autonomously even when placed into another region of the embryo. Three basic modes of commitment: Autonomous Conditional Syncytial Autonomous specification Characteristic of most invertebrates. Specification by differential acquisition of certain cytoplasmic molecules present in the egg. Blastomere fates are generally invariant. Cell type specification precedes any large-scale embryonic cell migration. Produces "mosaic" ("determinative") development: cells cannot change fate if a blastomere is lost. Conditional specification Characteristic of all vertebrates and few invertebrates. Specification by interactions between cells. Relative positions are important. Variable cleavages produce no invariant fate assignments to cells. Massive cell rearrangements and migrations precede or accompany specification. Capacity for "regulative" development: allows cells to acquire different functions Syncytial specification Syncytium - a cytoplasm that contains many nuclei. Characteristic of most insect classes. Specification of body regions by interactions between cytoplasmic regions prior to cellularization of the blastoderm. Variable cleavage produces no rigid cell fates for particular nuclei. After cellularization, conditional specification is most often seen. 4. Morphogenesis and Cell Adhesion There are two major types of cell arrangements in the embryo: - epithelial cells, which are tightly connected to one another in sheets or tubes - mesenchymal cells, which are unconnected to one another and which operate as independent units. Morphogenesis is brought about through a limited repertoire of variations in cellular processes: 1. the direction and number of cell divisions; 2. cell shape changes; 3. cell movement; 4. cell growth; 5. cell death; 6. changes in the composition of the cell membrane or secreted products. Morphogenesis and Cell Adhesion Cadherins: the major cell adhesion molecules critical for establishing and maintaining intercellular connections crucial to the spatial segregation of cell types the organization of animal form Cadherins + other cadherins on adjacent cells = Catenins In vertebrate embryos, several major cadherin classes have been identified: E-cadherin, P-cadherin, Ncadherin, EP-cadherin, Protocadherins