12.1. Halving the Chromosome Number

    A. Sexual Reproduction

        1. Meiosis is nuclear division reducing chromosome number from diploid (2n) to haploid (n) number.
        2. Haploid (n) number is half the diploid number of chromosomes.
        3. Requires gamete formation and then fusion of gametes to form a zygote.
        4. A zygote always has a full or diploid (2n) number of chromosomes.
        5. If gametes contained same number of chromosomes as body cells, doubling would soon fill cells.
        6. Belgian cytologist found Ascaris egg and sperm contain two chromosomes; body cells would contain four.

    B. Homologous Pairs of Chromosomes

        1. In a diploid body cell, chromosomes occur as pairs.
            a. Each set of chromosomes is a homologous pair; each member is a homologous chromosome or homologue.
            b. Homologues look alike; they have same length and centromere position; have similar banding pattern
                when stained.
            c. A location on one homologue contains the same types of gene which occurs at the same locus on other
                homologue.
        2. Chromosomes duplicate just before nuclear division.
            a. Duplication produces two identical parts called sister chromatids, held together at centromere.
            b. Non-sister chromatids do not share the same centromere.
        3. One member of each homologous pair is inherited from either male or female parent; one member of
            each homologous pair is placed in each sperm or egg.

    C. Overview of Meiosis

        1. Meiosis involves two nuclear divisions and produces four haploid daughter cells.
        2. Each daughter cell has half the number of chromosomes in the diploid parent nucleus.
        3. Meiosis I is the nuclear division at the first meiotic division.
            a. Prior to meiosis I, DNA replication occurs and each chromosome has two sister chromatids.
            b. During meiosis I, homologous chromosomes come together and line up (cause unknown) in synapsis.
            c. During synapsis, the two sets of paired chromosomes lay alongside each other as bivalents or a tetrad.
            d. Crossing over is an exchange of homologous segments between non-sister chromatids of bivalent during
                meiosis I; results in genetic recombination.
            e. After crossing over occurs, sister chromatids of a chromosome are no longer identical.
        4. Meiosis II
            a. No replication of DNA is needed between meiosis I and II because chromosomes were already doubled.
            b. During meiosis II, centromeres divide; daughter chromosomes derived as sister chromatids separate.
            c. Chromosomes in the four daughter cells have only one chromatid.
            d. Counting the number of centromeres verifies that parent cells were diploid, each daughter cell is haploid.
            e. In the animal life cycle, daughter cells become gametes that fuse during fertilization; this restores the
                diploid number in body cells.

12.2. Genetic Recombination

    A. Genetic Recombination

        1. Due to genetic recombination, offspring have a different combination of genes than their parents.
        2. Without recombination, asexual organisms must rely on mutations to generate variation among offspring;
             this is sufficient because they have great numbers of offspring.

    B. Crossing-Over Introduces Variation

        1. Crossing-over results in exchange of genetic material between non-sister chromatids.
        2. At synapsis, homologous chromosomes are held in position by a nucleoprotein lattice (the synaptonemal complex).
        3. As the lattice of the synaptonemal complex breaks down at beginning of anaphase I, homologues are temporarily
            held together by chiasmata, regions were the non-sister chromatids are attached due to crossing-over.
        4. The homologues separate and are distributed to separate cells.
        5. Due to crossing-over, daughter chromosomes derived from sister chromatids are no longer identical.

    C. Independent Assortment of Homologous Chromosomes

        1. Independent assortment in a cell with only three pairs of chromosomes is eight possible combinations.
        2. In humans with 23 pairs of chromosomes, the combinations possible are 8,388,608 possible combinations.

    D. Fertilization

        1. Meiosis increases variation.
        2. When gametes fuse at fertilization, chromosomes donated by parents combine.
        3. Chromosomally different zygotes from same parents are 70,368,744,000,000 combinations
            possible without crossing over.
        4. If crossing over occurs once, then  4,951,760,200,000,000,000,000,000,000 combinations of
            genetically different zygotes are possible for one couple.

12.3. Meiosis Has Phases

    A. Number of Phases

        1. Both meiosis I and meiosis II have four phases: prophase, metaphase, anaphase and telophase.

    B. Prophase I

        1. Nuclear division is about to occur: nucleolus disappears; nuclear envelope fragments; centrosomes
            migrate away from each other; and spindle fibers assemble.
        2. Homologous chromosomes undergo synapsis forming bivalents; crossing over may occur at this time
            in which case sister chromatids are no longer identical.
        3. Chromatin condenses and chromosomes become microscopically visible.

    C. Metaphase I

        1. During prometaphase I, bivalents held together by chiasmata have moved toward the metaphase plate.
        2. In metaphase I, there is a fully formed spindle and alignment of the bivalents at the metaphase plate.
        3. Kinetochoares are regions just outside centromeres; they attach to spindle fibers call kinetochore spindle fibers.
        4. Bivalents independently align themselves at the metaphase plate of the spindle.
        5. Maternal and paternal homologues of each bivalent may be oriented toward either pole.

    D. Anaphase I

        1. The homologues of each bivalent separate and move toward opposite poles.
        2. Each chromosome still has two chromatids.

    E. Telophase I

        1. Only occurs in some species.
        2. When it occurs, the nuclear envelope reforms and nucleoli reappear.

    F. Interkinesis

        1. This period between meiosis I and meiosis II is similar to interphase of mitosis.
        2. However, no DNA replication occurs.

    G. Meiosis II

        1. During metaphase II, the haploid number of chromosomes align at metaphase plate.
        2. During anaphase II, centromeres divide and daughter chromosomes move toward the poles.
        3. At the end of telophase II and cytokinesis, there are four haploid cells.
        4. Due to crossing-over, each gamete can contain chromosomes with different types of genes.
        5. In animals, the haploid cells mature and develop into gametes.
        6. In plants, the daughter cells become spores and divide to produce a haploid adult generation.
        7. In some fungi and alge, a zygote results from gamete fusion and immediately undergoes meiosis;
            therefore, the adult is always haploid.

12.4. Comparison of Meiosis with Mitosis

    A. How Meiosis I Differs from Mitosis

        1. DNA is replicated only once before both mitosis and meiosis; in mitosis there is only one nuclear
            division; in meiosis there are two nuclear divisions.
        2. During prophase I of meiosis, homologous chromosomes pair and undergo crossing-over; this does
            not occur during mitosis.
        3. During metaphase I of meiosis, paired homologous chromosomes align at the metaphase plate; in
            mitosis individual chromosomes align.
        4. During anaphase I in meiosis, homologous chromosomes with centromeres intact separate and move
            to opposite poles; in mitosis at this stage, sister chromatids separate and move to poles.

    B. How Meiosis II Differs from Mitosis

        1. Events of meiosis II are same stages as in mitosis.
        2. However, the nuclei contain the haploid number of chromosomes in meiosis.
        3. Mitosis produces two daughter cells; meiosis produces four daughter cells.

12.5. Viewing the Human Life Cycle

    A. Life Cycle has Both Meiosis and Mitosis

        1. Life cycle refers to all reproductive events between one generation and next.
        2. In animals, the adult is always diploid [Instructors note: some bees, etc. have haploid male adults].
        3. In animals, it occurs during production of gametes; adult is diploid and gametes are haploid.
        4. Mosses are haploid most of their cycle; oak trees are diploid most of their cycle.
        5. In fungi and some algae, organisms you see is haploid and produces haploid gametes.
        6. In males, meiosis is part of spermatogenesis, the production of sperm, and occurs in the testes.
        7. In females, meiosis is part of oogenesis, the production of eggs cells, and occurs in the ovaries.
        8. After birth, mitotic cell division is involved in growth and tissue regeneration.

    B. Spermatogenesis and Oogenesis in Humans

        1. Spermatogenesis
            a. In the testes of males, primary spermatocytes with 46 chromosomes divide meiotically to form two
                secondary spermatocyes, each with 23 duplicated chromosomes.
            b. Secondary spermatocyes divide to produce four spermatids, also with 23 daughter chromosomes.
            c. Spermatids then differentiate into sperm (spermatozoa).
            d. Meiotic cell division in males always results in four cells that become sperm.
        2. Oogenesis
            a. In the ovaries of human females, primary oocytes with 46 chromosomes divide meiotically to form two
                cells, each with 23 duplicated chromosomes.
            b. One of the cells, a secondary oocyte, receives most cytoplasm; the other cell, a polar body, disintegrates
                or divides again.
            c. A secondary oocyte begins meiosis II and then stops at metaphase II.
            d. At ovulation, the secondary oocyte leaves the ovary and enters an oviduct where it may meet a sperm.
            e. If a sperm enters secondary oocyte, oocyte is activated to continue meiosis II to completion; result is
                a mature egg and another polar body, each with 23 daughter chromosomes.
            f. Polar bodies serve to discard unnecessary chromosomes and retain most of the cytoplasm in the egg.
            g. The cytoplasm serve as a source of nutrients for the developing embryo.

 

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