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Development of Multicellular Organisms Shu-Ping Lin, Ph.D. Institute of Biomedical Engineering E-mail: [email protected] Website: http://web.nchu.edu.tw/pweb/users/splin/ Date: 01.03.2011 Chromosome Numbers All are even numbers – diploid (2n) sets of homologous chromosomes! -ploid = number of copies of each chromosome. Haploid Diploid In humans … 23 chromosomes donated by each parent (total = 46 or 23 pairs). Sex cells divide to produce gametes (sperm or egg), occurs only in gonads (testes or ovaries). Male: spermatogenesis; Female: oogenesis: The form of cell division by which gametes, with half the number of chromosomes, are produced. Contain 22 autosomes and 1 sex chromosome. Are haploid (haploid number “n” = 23 in humans). Fertilization results in zygote with 2 haploid sets of chromosomes now diploid. Diploid cell; 2n = 46. (n=23 in humans) Meiosis is similar to mitosis with some chromosomal differences (Most cells in the body produced by mitosis). Only gametes are produced by meiosis, is sexual reproduction. Two divisions (meiosis I and meiosis II). Meiosis Leading to Sperm and Oocytes From Unfertilized Egg to Zygote # Meiotic cell division produces gametes 1. During interphase parent cell has 2n chromosomes. 2. Chromosomes duplicate and form sister chromatids. 3. Homolog pairs of sister chromatids then synapse and exchange genetic material. 4. Sister chromatids remain together, but homologs separate to opposing poles. 5. Nuclear membrane forms around the chromosomes, and equatorial plane contracts and separates the newly formed cells. 6. Next cell division occurs without DNA synthesis. 7. Chromatid pairs align on the equatorial plane. 8. Chromatids of each pair separate and move to opposing poles, and from then on the division proceeds as described for mitotic division. Meiosis I First division of meiosis Prophase 1: Each chromosome dupicates and remains closely associated. These are called sister chromatids. Crossing-over can occur during the latter part of this stage. Metaphase 1: Homologous chromosomes align at the equatorial plate. Anaphase 1: Homologous pairs separate with sister chromatids remaining together. Telophase 1: Two daughter cells are formed with each daughter containing only one chromosome of the homologous pair. Meiosis II Second division of meiosis: Gamete formation Prophase 2: DNA does not replicate. Metaphase 2: Chromosomes align at the equatorial plate. Anaphase 2: Centromeres divide and sister chromatids migrate separately to each pole. Telophase 2: Cell division is complete. Four haploid daughter cells are obtained. Meiosis – Key Differences from Mitosis Meiosis reduces the number of chromosomes by half. Daughter cells differ from parent, and each other. Meiosis involves two divisions, Mitosis only one. Meiosis I involves: Synapsis – homologous chromosomes pair up. Chiasmata form (crossing over of non-sister chromatids). In Metaphase I, homologous pairs line up at metaphase plate. In Anaphase I, sister chromatids do NOT separate. Overall, separation of homologous pairs of chromosomes, rather than sister chromatids of individual chromosome. Animation DIFFERENCES BETWEEN MITOSIS AND MEIOSIS EVENTS Occurrence Definition Number of daughter cells Prophase MITOSIS In all the body cells including germ cells. Only in the germ (reproductive) cells. It is an equational division. It is a reductional division. Only two Four Involves relatively few changes. Chromomeres Not visible in prophase. Synapsis Does not occur. Crossing over Does not occur. Metaphase Centromeres in Anaphase Centromeres in Metaphase Telophase Cytokinesis MEIOSIS Chromosomes arrange along the equator. Each centromere splits into two. Orient towards the equator while chromatids orient towards poles. Involves a series of changes in chromosomes distinguished into 5 substages. Visible in the leptotene stage of prophase -I Occurs in zygotene of prophase-I. Occurs in pachytene stage of prophase-I. Chromosomes arrange equally on either side of the equator in metaphase-I. Centromeres do not split in metaphase-l. Orient towards poles while chromatids orient towards the equator in metaphaseI Results in the formation of two daughter Telophase-II results in the formation of nuclei having the same no. of two daughter nuclei, each having half the chromosomes as that of parent cell. no. of chromosomes as that of parent cell. Follows immediately after karyokinesis. May or may not occur at the end of first karyokinesis Mitosis vs. Meiosis Polarization of Fertilized Frog Egg (a) Unfertilized egg has radial symmetry around an axis that passes through cell centrosome and nucleus. Hemisphere that contains centrosome and nucleus is called animal hemisphere, whereas lower hemisphere contains large numbers of yolk platelets full of nutrients. Point of contact between egg and second polar body coincides with animal pole of egg. Entry of sperm into egg induces rotation of cell cortex around cell center(b). This movement is initiated by molecular motors such as myosin and involves relative sliding movement of cell cortex on underlying cytoskeleton. Cortical rotation is resisted by viscous forces exerted by surrounding fluid(c). Resultant moment exerted by fluid is proportional to the rate of rotation of cortex as well as cell radius squared. This moment has opposite sense of direction to angular velocity of cortex. Since this is the only external moment acting on the cell, Newton’s laws of motion dictate that inner region of cell must move in opposite direction of cortex (c). As a result of this complex movement, radial symmetry is destroyed. Bisection of Eight-cell Sea Urchin Embryos in Two Different Midplanes Division of asymmetric fertilized egg results in cells with differing cell contents represented by different shading. Horizontal plane in the figure separates each cell into chemically distinct upper hemisphere (animal hemisphere) and lower hemisphere (vegetal hemisphere). When embryo is bisected vertically, resulting 4-cell clusters develop into small, but normal, larva (a). Bisection along middle horizontal plane leads to different result (b). Cells from animal pole remain embryonic, whereas cells from vegetal plane develop into small, but abnormal, embryo. Three cycles of divisions of a fertilized egg give rise to 3 different cell types. DNA-binding proteins can block parts of DNA from action of transcription machinery. Idealized DNA molecules shown in figure belong to stem cells (top row) and cells that differentiated into cell types 1 and 2 (bottom row). Cleavage refers to the first stage of embryonic development during which cells divide rapidly without significantly increasing overall mass volume of zygote. Amphibian and mammary embryos exhibit different courses of development. Frog zygote develops into a spherical shell of cells called blastula at the late stage of cleavage (a), whereas human zygote develops into what is called blastocyst (b). Outer layer of cells is called trophoblast. Embryonic cells massed at top of fluid-filled sack (blastocoel) give rise to human embryo. Gastrulation phase of embryonic development, during which cells change their shape and move within the embryo. Cells move as individual cells or as a sheet of cells. Figure shows some of modes of cell movement observed during gastrulation. Invagination involves a layer of cells moving inward toward the center of blastula. Inward migration of single cells from outer layer of embryo is known as ingression. Term of delamination is used to characterize inward movement of newly formed daughter cells while keeping shell of blastula intact. Involution refers to folding of cell layers, leading to new cell-cell contacts and opportunities for induction. Compact domain of embryo extending to cover large surface area through cell movements is characterized as convergent extension. Induction by direct cell-cell contact (a) and by diffusion of a morphogen through the embryo (b). 3 germ layers developed during gastrulation: ectoderm, mesoderm, and endoderm. Through a series of cell movements and inductive interactions, these 3 layers of cells give rise to all the tissues in the body. Generation of different cell types through induction. 4 Essential Processes Cell proliferation: producing many cells from one Cell specialization: creating cells with different characteristics at different positions Cell interaction: coordinating the behavior of one cell with that of its neighbors Cell movement: rearranging the cells to form structured tissues and organs Fruit Fly Embryo Nuclear division is not accompanied by cell division until about 20003000 nuclei form. Maternal mRNA of bicoid gene is localized at anterior end, and mRNA of nanos gene is concentrated at posterior end Products of these genes play roles in the activation or repression of gene transcription involved in development. About 6 hours after fertilization, midsection of fruit fly embryo exhibits 14 stripes, each a few cells thick, with alternative expression of even-skipped (Eve) and fushitarazu (Ftz) proteins, both of which are involved in the generation of development of smallerscale patterns. Activator and Repressor Sites Transcription factors that bind to activator sites are shown as half circles, whereas those that bind to repressors are shown as triangles. High-affinity regulator sites are darker than low-affinity sites. Capital letters B and H stand for bicoid and hunchback proteins, respectively. Head-to-tail Pattern Formation in Fruit Fly Figure shows spatial distribution of transcription factors bicoid, hunchback, giant, and krüpple along axis of fruit fly larva between segments 1 and 3. Expression of even-skipped (Eve) protein in stripe 2 is regulated by these transcription factors. Hypothetical Homeobox Gene Language Hox genes can be either on (1) or off (0). Expression of Hox d2 and d3 in the context of no expression of other Hox genes specifies that those cells will go on to form tissue in the tail of mouse. If other Hox genes are inappropriately expressed or timing of expression is wrong, then ability of Hox d2 and Hox d3 to specify a tail fate is lost. Body pattern of mouse and spatial range of activation of hox genes that mediate mouse development. Stem Cells and Tissue Engineering Tissue engineering is a field of biotechnology that aims to generate living tissue from isolated cells, scaffolds, matrix proteins, and growth factors.