Meiosis: The Cellular Basis of Sexual Reproduction The Mechanisms of Meiosis Meiosis is based on the interactions and distribution of homologous chromosome pairs Meiosis produces four genetically different cells with half the parental chromosomes Sexual Reproduction Sexual reproduction produces offspring by union of male and female gametes (sperm and egg) Meiosis produces gametes with half chromosome number Evolutionary advantage: Genetic shuffling of sex Fertilization Fertilization fuses nuclei of egg and sperm Zygote cell produced by fertilization Restores parental chromosome number Homologous Chromosome Pairs Paternal chromosomes from male parent, maternal from female parent Homologous chromosome pairs Alleles may be different within homologous pairs Meiosis separates homologous pairs Before meiosis, diploid (2n) After meiosis, haploid (n) Diploid Haploid Meiotic Cell Cycle Mitosis and meiosis compared Both: Similar cell divisions, meiosis divides twice Mitosis: Two identical daughter cells, diploid Meiosis: Four genetically different cells, haploid Premeiotic interphases similar to mitotic interphase Chromosomes copied into sister chromatids Meiosis I Meiosis I: First meiotic division Recombination exchanges segments between homologues Produces two haploid cells with chromatids attached Meiosis II Meiosis II: Second meiotic division Sister chromatids separate into separate cells Produces 4 recombined haploid cells Meiotic Divisions Homologous chromosome pair A diploid cell showing the two chromosomes of a homologous pair. Replication during premeiotic interphase Sister chromatids As a result of replication, each chromosome consists of two sister chromatids. Chromosome pairing during prophase I of meiosis Fig. 11-2a, p. 223 While the homologous chromosomes are paired, they may undergo recombination by exchanging segments. First meiotic division Second meiotic division The first meiotic division separates the homologs, placing one in each of the two cells resulting from the division. These products have the haploid number of chromosomes, but each chromosome still consists of two chromatids. The second meiotic division separates the sister chromatids, placing one in each cell resulting from the division. Fig. 11-2b, p. 223 Meiosis animation Meiotic Cell Cycle Prophase I Sister chromatids condense to chromosomes Synapsis (pairing of homologs) Tetrads (Fully paired homologs) Recombination mixes alleles across tetrads Prometaphase I Nuclear envelope breaks down Kinetochores attach to polar spindles Meiotic Divisions Meiotic Cell Cycle Metaphase Tetrads align on metaphase plate Homologs segregate, move to poles (sister chromatids attached) Nondisjunction creates abnormal chromosome numbers Telophase I and Anaphase I and Interkinesis No change in chromosomes Spindle disassembles Meiotic Divisions Meiotic Cell Cycle Prophase II, Prometaphase II, and Metaphase II Chromosomes condense, spindles form Nuclear envelope breaks, kinetochores form Chromosomes align on metaphase plate Anaphase II and Telophase II Spindles separate chromatids Spindles disassemble New nuclear envelopes form Meiotic Divisions Meiotic Divisions Comparison of Meiosis and Mitosis Comparison of Meiosis and Mitosis Sex Chromosomes in Meiosis Sex chromosomes Different in males and females Human females XX, males XY XX fully homologous, XY homologous in short region Sex Chromosomes in Meiosis Meiosis and sex chromosome inheritance Gametes produced by females may receive either X chromosome Gametes produce by males may receive either X or Y chromosome Sex determination Sex – sexual phenotype Male and female determination – mechanism by which sex is established Sex Mechanisms That Generate Variability Recombination depends on physical exchanges between homologous chromatids Segregation of maternal and paternal chromosomes is random Random joining of male and female gametes in fertilization adds additional variability Recombination of Chromatids Recombination (crossing over) Key genetic shuffle of prophase I Tetrads held together at synaptonemal complex Two of four chromatids exchange alleles Chiasmata or crossovers are points of exchange Crossing Over Prophase I Long and complex Pachytene = Synaptonemal complex Elaborate protein structure Crossing over occurs – homologous chromosomes exchange genetic information Synaptonemal Complex Random Segregation Random segregation Key genetic shuffle of metaphase I Each chromosome of a homologous pair may randomly end up at either spindle pole Any combination of maternal and paternal chromosomes can be segregated to gametes 2X number of possible combinations Genetic Variation Independent segregation of homologous pairs of chromosomes 23 pairs of chromosomes 223 or >8X106 different combinations Random Spindle Connections Random Fertilization Random chance of male and female gamete forming zygote Meiosis allows randomness necessary for Mendelian laws of inheritance Recombination, random segregation, and random fertilization are mechanisms of randomness Gamete Diversity animation Meiosis in humans Difference between male and female are remarkably different Meiosis in humans Spermatogenesis Production of sperm from primordial germ cells Begins at puberty Each day more than 100 million sperm are made in each human testicle Over a lifetime a male produces 1012 -1013 sperm (Stem cells capable of self renewal) Developmental Biology, 6th Ed., Gilbert Developmental Biology, 6th Ed., Gilbert Motile nucleus Meiosis in humans Oogenesis Production of eggs from primordial germ cells Human embryo by 7 months gestation, 7 million germ cells After 7 months gestation, most germ cells die, the remaining oogonia enter the first meiotic division Meiosis in humans Primary oocytes, progress through the first meiotic prophase until the diplotene stage, at which point they are maintained until puberty (~12 years later) Of the millions of primary oocytes present at birth only about 400 mature during a women’s lifetime With the onset of adolescence, groups of oocytes periodically resume meiosis Meiosis in humans Oogenic meiosis also differs from spermatogenic meiosis in its placement of the metaphase plate. At telophase, one of the two daughter cells contains hardly any cytoplasm, whereas the other cell has nearly the entire volume of cellular constituents The smaller cell is called the first polar body, and the larger cell is referred to as the secondary oocyte. Meiosis in humans During the second division of meiosis, a similar unequal cytokinesis takes place Most of the cytoplasm is retained by the mature egg (ovum), and a second polar body receives little more than a haploid nucleus. Thus, oogenic meiosis conserves the volume of oocyte cytoplasm in a single cell rather than splitting it equally among four progeny Developmental Biology, 6th Ed., Gilbert New Information Research suggests women can make new eggs throughout their lives What defects might occur if there are errors in meiosis? Question 80-90% of abnormalities in chromosome number result from nondisjunction at maternal Meiosis I Why might this be more common in female gametogenesis than male gametogenesis? Mistakes in Meiosis animation Explain to your neighbor how you will tell the difference between these phases Metaphase of mitosis and metaphase I of meiosis Metaphase of mitosis and metaphase II of meiosis Metaphase I and metaphase II of meiosis Telophase of mitosis and telophase of meiosis