CHAPTER 13: MEIOSIS AND SEXUAL LIFE CYCLES Overview: Variations on a theme One of the characteristics of life is the

e ability of organisms to reproduce their own kind. o Another rules is that offspring resemble their parents more than they do unrelated individuals. The transmission of traits from one generation to the next is called inheritance, or heredity. Along with inherited similarity, there is also variation, differences between members of the same species. Genetics is the scientific study of heredity and hereditary variation. Concept 13.1 Offspring acquire genes from parents by inheriting chromosomes. Inheritance of Genes Parents endow their offspring with coded information in the form of hereditary units called genes. The genes we inherit from our mothers and fathers are our genetic link to our parents, and they account for family resemblances. The genetic program is written in the language of DNA, the polymer of four different nucleotides. Inherited info is passed on in the form of each gene's specific sequence of DNA nucleotides. Most genes program cells to synthesize specific enzymes and other proteins, whose cumulative action produces an organism's inherited traits. In animals and plants, reproductive cells called gametes are the vehicles that transmit genes from one generation to the next. Every species has a characteristic number of chromosomes. o For example, humans have 46 chromosomes in their somatic cells- all cells of the body expect the gametes and their precursors. o Each chromosome consists of a single long DNA molecule elaborately coiled in association with various proteins. A gene's specific location along the length of a chromosome is called the gene's locus. Comparison of Asexual and Sexual Reproduction Organisms that reproduce asexually have offspring that are their exact genetic copies of themselves. In asexual reproduction, a single individual is the sole parent and passes copies of all its genes to its offspring without the fusion of gametes. o reproduce by mitotic cell division in which DNA is copied and allocated equally to tow daughter cells. o The genomes of the offspring are virtually exact copies of the parent's genome.

 An individual that reproduces asexually gives rise to a clone, a group of genetically identical individuals. In sexual reproduction, two parents give parents give rise to offspring that have unique combinations of genes inherited from the two parents. Concept 13.2 Fertilization and meiosis alternate in sexual life cycles. A life cycle is the generation-to-generation sequence of stages in the reproductive history of an organisms, from conception to production of its own offspring. Sets of Chromosomes in Human Cells In humans, each somatic cell has 46 chromosomes. During mitosis, chromosomes become condensed. A display of the chromosome pairs of a cell arranged by size and shape is called a karyotype. The two chromosomes composing a pair have the same length, centromere position, and staining pattern: These are called homologous chromosomes, or homologs. The two distinct chromosomes referred to as X and Y are an important exception t the general pattern of homologous chromosomes in human somatic cells. o Because they determine an individual's sex, the X and Y chromosomes are called sex chromosomes. The other chromosomes are called autosomes. Any cell with two chromosome sets is called a diploid cell and has a diploid number of chromosomes, abbreviated 2n. Unlike somatic cells, gametes contain a single set of chromosomes. Such cells are called haploid cells, and each has a haploid number of chromosomes (n). Behavior of Chromosome Sets in the Hum life Cycle The human life cycle begins when a haploid sperm from the father fuses with a haploid egg from the mother. This union of gametes, culminating in fusion of their nuclei is called fertilization. o The resulting fertilized egg, or zygote, is diploid because it contains two haploid sets of chromosomes bearing genes representing the maternal and paternal family lines. The only cells of the human body not produces by mitosis are the gametes, which develop from specialized cells called germ cells in the gonads- ovaries in females and testes in males. In sexually reproducing organisms, gamete formation involves a type of cell division called meiosis. This type of cell division reduces the number of sets of chromosomes from two to one in the gametes, counterbalancing the doubling that occurs at fertilization. o In animals, meiosis occurs only in germ cells.

 The processes of fertilization and meiosis are the hallmarks of sexual reproduction in plans, fungi, and protists as well as in animals. The Variety of Sexual Life Cycles The timing of meiosis and fertilization in the life cycle varies, depending on the species. First type of life cycle is of humans and most animal cells. o Gametes are the only haploid cells. Meiosis occurs in germ cells during the production of gametes which undergo no further cell division prior to fertilization. After fertilization, the diploid zygote divides by mitosis, producing a multicellular organism that is diploid. Plants and some species of algae exhibit a second type of life cycle called alternation of generations. o This type includes both diploid and haploid stages that are multicellular. The diploid stage is called the sporophyte. Haploid stage is called gametophyte. The third type of life cycle occurs in most fungi and some protists, including some algae. o After gametes fuse and form a diploid zygote, meiosis occurs without a multicellular diploid offspring developing. Meiosis produces not gametes but haploid cells that then divide by mitosis and give rise to either unicellular descendants or a haploid organism carries out further mitoses, producing the cells that develop into gametes. The only diploid stage found in these species is the single-celled zygote. Only diploid cells can go through meiosis Concept 13.3 Meiosis reduces the number of chromosome sets from diploid to haploid Many steps of meiosis resemble corresponding steps in mitosis. Meiosis is preceded by the duplication of chromosomes. However, this single duplication is followed by not one but two consecutive cell divisions, called meiosis I and meiosis II. o These 2 divisions result in 4 daughter cells (rather than the two daughter cells of mitosis), each with only half as many chromosomes as the parent cell. The Stages of Meiosis For a single pair of homologous chromosomes in a diploid cell, both members of the pair are duplicated and the copies sorted into four haploid daughter cells. The sister chromatids are two copies of one chromosome, closely associated all along their length; this association is called sister chromatid cohesin. Together, the sister chromatids make up one duplicated chromosome. Meiosis I: separates homologous chromosomes o Prophase I: duplicated homologous chromosomes pair and exchange segments. Chromosomes begin to condense, and homologs loosely pail along their lengths, aligned gene by gene.

 Paired homologs become physically connected to each other along their lengths by a zipper like protein structure, the synaptonemal complex; this state is called synapsis. Crossing over, a genetic rearrangement between non-sister chromatids involving the exchange and corresponding segments of DNA molecules, begins during pairing and synaptonemal complex formation, and is completed while homologs are in synapsis. Each homologous pair ahs one or more X-shaped regions called chiasmata. It exists at the point where a crossover has occurred. o Metaphase I: Chromosomes line up by homologous pairs. Pairs of chromosomes are arranged at the metaphase plate with one chromosome facing each pole. Both chromatids of one homolog are attached to kinetochore microtubules form one pole; those of the other homolog are attached to microtubules from the opposite pole. o Anaphase I: Each pair of homologous chromosomes separates. Breakdown of proteins responsible for sister chromatid cohesion along chromatid arms allows homologs separate. The homologs move toward opposite poles, guided by the spindle apparatus. Sister chromatid cohesion persists at the centromere, causing chromatids to move as a unit toward the same pole. o Telophase I and Cytokinesis: Two haploid cells form; each chromosome still consists of two sister chromatids. Beginning of Telophase I, each half of the cell has complete haploid set of duplicated chromosomes. Cytokinesis usually occurs simultaneously with Telophase I, forming two haploid daughter cells. In animal cells, a cleavage furrow forms. In plant cells, a cell plate forms. In some species, chromosomes decondense and nuclear envelopes form. NO chromosome duplication occurs between meiosis I and meiosis II. Meiosis II: Separates sister chromatids. o Prophase II: A spindle apparatus forms. In late prophase II, chromosomes move toward the metaphase II plate. o Metaphase II: The chromosomes are at the metaphase plate. Because of crossing over, the two sister chromatids of each chromosome are not genetically identical. o Anaphase II: Breakdown of proteins holding sister chromatids together at the centromere allows the chromatids to separate. The chromatids move toward opposite poles as individual chromosomes.

o Telophase II and Cytokinesis: Nuclei form, the chromosomes begin decondensing, and Cytokinesis occurs. The meiotic division of one parent cell produces four daughter cells, each with a haploid set of chromosomes. The four daughter cells are genetically distinct from one another and from one another and from the parent cell. COMPARISON OF MITOSIS AND MEIOSIS Property Mitosis Meiosis DNA Replication Occurs during interphase Occurs during interphase before mitosis begins before meiosis I begins Number of divisions One, including prophase, Two, each including metaphase, anaphase, prophase, metaphase, and Telophase anaphase, and Telophase Synapsis homologous Does not occur Occurs during prophase I chromosomes along with crossing over between non-sister chromatids; resulting chiasmata hold pairs together due to sister chromatid cohesion. Number of Daughter cells Two, each diploid and Four, each haploid, and genetic composition genetically identical to containing half as many the parent cell chromosomes as the parent cell; genetically different from the parent cell and from each other. Role in the animal body Enables multicellular Produces gametes; adult to arise from reduces number of zygote; produces cells for chromosomes by half and growth, repair, and in introduces genetic some species, asexual variability among the reproduction gametes. A Comparison of Mitosis and Meiosis Basically, meiosis reduces the number of chromosome sets from two to one, whereas mitosis conserves the number of chromosome sets. Therefore, meiosis produces cells that differ genetically identical to their parent cell and to each other. Three events unique to meiosis occur during meiosis I: o Synapsis and crossing over: During prophase I Duplicated homologs pair up, and the formation of the synaptonemal complex between them holds them in synapsis. Crossing over also occurs during prophase I. Synapsis and crossing over normally do not occur during prophase of mitosis. o Homologous pairs ate the metaphase plate: AT metaphase I of meiosis

Chromosomes are positioned at the metaphase plate as pairs of homologs, rather than individual chromosomes, as in metaphase of mitosis. o Separation of homologs: At anaphase I of meiosis. The duplicated chromosomes of each homologous pair move toward opposite poles, but the sister chromatids of each duplicated chromosome remain attached. In anaphase of mitosis, by contrast, sister chromatids separate. How do sister chromatids stay together through meiosis I but separate from each other in meiosis II and mitosis? o Sister chromatids are attached along their lengths by protein complexes called cohesions. o In mitosis this attachment lasts until the end of metaphase, when enzymes cleave the cohesin, freeing the sister chromatids to move to opposite poles of the cell. o In meiosis, sister chromatid cohesion is released in two steps, one at the start of anaphase I and one at anaphase II. In metaphase I, homologs are held together by cohesion between sister chromatid arms in regions beyond points of crossing over, where stretches of sister chromatids now belong to different chromosomes. Sister chromatid cohesion and crossing over, acting together, play an essential role in the lining up of chromosomes by homologous pairs at metaphase I. Meiosis I is called the reductional division because it halves the number of chromosome sets per cell- a reduction from two sets (the diploid state) to one set (the haploid state). Meiosis II is called the equational division because the sister chromatids separate, producing haploid daughter cells.

Concept 13.4 Genetic variation produced in sexual life cycles contributes to evolution Mutations are the original source of genetic diversity. These changes in an organism's DNA create the different versions of genes known as alleles. Once these differences arise, reshuffling of the alleles during sexual reproduction produces the variation that results in each member of sexually reproducing population having a unique combination of traits. Origins of Genetic Variation Among Offspring In sexually reproducing species, the behavior of chromosomes during meiosis and fertilization is responsible for the variation that arises in each generation. Independent Assortment of Chromosomes At metaphase I, the homologous pairs, each consisting of one maternal and paternal chromosome, are situated at the metaphase plate. Because each pair of homologous chromosomes is positioned independently of the other pairs at metaphase I, the first meiotic division results in each pair sorting its maternal/paternal

homologs into daughter cells independently of every other pair. This is called independent assortment. The number of combinations possible for daughter cells formed by meiosis of a diploid cell with n=2 is four: two possible arrangements for the first pair times two possible arrangements for the second pair. Crossing Over As a consequence of independent assortment of chromosomes during meiosis, each of us produces a collection of gametes differing greatly in their combinations of the chromosomes we inherited from our two parents. Crossing over produces recombinant chromosomes, individual chromosomes that carry genes (DNA) derived from two different parents. o In meiosis in humans, an average of one to three crossover events occur per chromosome pair, depending on the size of the chromosomes and the position of their centromeres. Random Fertilization The random nature of fertilization adds to the genetic variation arising from meiosis. The fusion of a male gamete during fertilization will produce a zygote with any of about 70 trillion diploid combinations. The Evolutionary Significance of Genetic Variation Within Populations Genetic variation is the raw material for evolution by natural selection. Mutations are the original source of this variation; the production of new combinations of variant genes in sexual reproduction generates additional genetic diversity. Animals that reproduce only asexually are quite rare, underscoring the apparently great advantage of genetic diversity.