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Developmental Stages Ø Early Development • Fertilization • Cleavage • Gastrulation • Neurulation Ø Later Development • Organogenesis • Larval molts • Metamorphosis • Aging Ø Rapid cell divisions • Drosophila 8 , Xenopus 20 – no transcription – S and M phases only • mammals 12-24 hr, chick 6-12 hr – transcription occurs Ø Produces a hollow embryo • blastocoel Ø mesoderm induction • signals from endoderm induce ectodermal cells 1 Birds, Reptiles, & Fishes Eggs Telolecithal eggs that completely support embryogenesis; undergo meroblastic cleavage In contrast to holoblastic cleavage, where the entire zygote divides into blastomeres, meroblastic cleavage leaves a large portion of the zygote uncleaved. There are 2 types of meroblastic cleavage, discoidal & superficial. 2 Birds, Reptiles, & Fishes Eggs Discoidal: Birds & reptiles: 1st cleavage is meridional. Starts at the animal pole but does not progress far. The 2nd and 3rd cleavages are also m e r i d i o n a l . T h e 4th cleavage is equatorial, and it creates a layer of small cells on top of the huge uncleaved area below (yolk). 3 Birds, Reptiles, & Fishes Eggs Blastoderm: when cleavage progressed such that there are many blastomeres in the animal pole; blastoderm. Chicken eggs have a blastoderm of 60,000 cells when the egg is laid. 4 Birds, Reptiles, & Fishes Eggs The next step eggs is formation of the upper & lower blastoderm. Epiblast: (epi = upon) is the upper layer & forms the embryo proper. Hypoblast: (hypo = under) is the bottom layer that will form extra embryonic endoderm that surrounds the yolk. What is the counterpart in mammals? 5 Birds, Reptiles, & Fishes Eggs Blastocoel: lies between the 2 layers. Subgerminal space: lies between the hypoblast & yolk. 6 The mitotic spindle determines the orientation of the cleavage plane Blastomeres can cleave either equatorially or meridionally. Cytokinesis usually directly follows mitosis, except for superficial cleavage. How does a blastomere know to divide meridionally or equatorially? 7 The mitotic spindle determines the orientation of the cleavage plane Mitotic spindles work to keep the cell round in shape. Experiment: It is possible to control how tightly blastomeres adhere by changing the concentration of calcium. High calcium concentrations cause more cell – cell attachment. Low calcium causes minimal attachment. The effect is likely mediated by adhesion molecules such as cadherin. 8 The mitotic spindle determines the orientation of the cleavage plane When blastomeres adhere they have a longer axis, and the mitotic spindle is almost always oriented parallel to this axis. As the cell becomes more spherical in low calcium medium, the mitotic spindle orientation starts to become random. 9 Mammalian eggs have rotational cleavage that is holoblastic The mammalian egg is a little slow. It begins to cleave in the oviduct and continues until it implants in the wall of the uterus (1 cleavage / 24 hr). Asynchronous cleavage: mammalian embryos are unusual in that they have asynchronous cleavage. Not all blastomeres divide at the same time. 10 Mammalian eggs have rotational cleavage that is holoblastic The first cleavage is meridional, and the second cleavage is rotational. The 2 blastomeres divide in different planes (one is equatorial and one is meridional. 11 Mammalian embryos undergo compaction at the 8 cell stage At first, the blastomeres of mammalian embryos have a loose arrangement, and touch only at the basal surfaces. After compaction, blastomeres adhere tightly, maximizing the area of contact. During compaction, each blastomere undergoes polarization. Tight junctions develop on the outer surface, allowing proteins to specialize. Cells take up fluids from uterine environment & secrete into the blastocoel. 12 Mammalian embryos undergo compaction at the 8 cell stage Gap junctions form on the outer cells to aid in intercellular communication. 13 Blastula Morula; 16-cell stage, embryo enclosed in the zona pellucida. - 14 A blastocoel develops as cleavage proceeds to the 32-64 cell stage After compaction at the 8-16 cell stage, there are 2 types of blastomeres. Outside blastomeres are tightly joined and number about 9-14. They surround 2-7 inside blastomeres that are loosely joined. Cavitation: the outside blastomeres start to take up fluid from the uterus and pump it into the center, creating the blastocoel. The blastocyst is the hallmark of early embryonic development in mammals. 15 A blastocoel develops as cleavage proceeds to the 32-64 cell stage Trophoblast: a structure consisting of outside blastomeres, this contributes to forming the placenta. Inner cell mass: this gives rise to the embryo, & develops from the inside blastome res 16 Embryonic stem cells can be cultured from the inner cell mass • C e l l s i n t h e i n n e r c e l l m a s s a r e undifferentiated, they multiply indefinitely, & are known as embryonic stem cells. • Stem cells are totipotent = they have the potential to form any tissue. These cells are of great scientific and medical importance. • They can be removed from the embryo, genes can be introduced into the cells. 17 Embryonic stem cells can be cultured from the inner cell mass Interestingly, if you inject adult, differentiated cells back into the environment of the morula or blastula, they become undifferentiated, and they can re-differentiate to form many parts of the body. 18 Development of monozygotic or identical twins Monozygotic twins develop from one zygote by splitting at various stages of development (from the 2 cell to the blastocyst stage). The stage of splitting effects the overall structure of the embryo and extraembryonic membranes. 19 Development of monozygotic or identical twins What are conjoined twins and how do they arise? Where do fraternal twins come from? None identical twins 20 Implantation After fertilization, it takes 2-3 days for the zygote to travel to uterus. Blastulation Blastula à embryo arrives in uterus(4) hatches from zona pellucida and contact uterus. Blastocyst surrounded by ZP prevents premature implantation and ectopic pregnancy. 21 Implantation Hatching lyses of ZP • involves à a trypsin-like protease --Trophoblast cells secrete proteases which degrades the endometrial wall and blastocyst embeds About 7 days after fertilization, embryo implants (progesterone is the important hormone) in the uterus by releasing enzyme from trophoblasts . 22 Implantation • Enzymes degrade endometrium and implants blastocyst (stage of embryonic development) • Failure of implantation & placentation common cause of spontaneous abortion (70% of all pregnancies) 23 Implantation 24 Implantation 25 Implantation (implanta tion chamber) Invasive Implant. 26