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  AP BIOLOGY:
Chapter Twelve Outline

INTRODUCTION TO MEIOSIS

                Different Chromosome Numbers Are Found in One Organism
                        Gametes contain half as many chromosomes as somatic cells
                        Zygotes are produced by fusion of gametes
                                Each gamete contains a single complement of genetic material
                                The zygote contains two copies of each chromosome
                        Fusion of gametes is called fertilization or syngamy
                        Fusion of body cells would sequentially increase chromosome number

                Meiosis Is an Important Event   fig 12.1
                        Serves to stabilize the chromosomes number
                        Is a reduction division process

THE SEXUAL LIFE CYCLE

                Fertilization and Meiosis Constitute a Cycle of Sexual Reproduction     fig 12.2
                        Body cells of adult are diploid and possess two sets of chromosomes
                        Gametes are haploid and possess a single set of genetic material
                        An individual inherits genes from its father and its mother     fig 12.3
                                In humans, 23 chromosomes from the mother's egg 
                                In humans, 23 chromosomes from the father's sperm 

                Life Cycles Show Pattern of Alternating Chromosome Numbers
                        Alternate between diploid and haploid number    fig 12.4
                        After syngamy the zygote divides by mitosis
                        All adult somatic cells are genetically identical to the zygote

                Sexual Reproduction Varies by Kingdom
                        Unicellular organisms
                                Individual cells function directly as gametes
                                Zygote may divide mitotically or meiotically
                        Plants
                                Haploid cells are produced through meiosis
                                Cells divide mitotically to form multicellular haploid phase
                                Special haploid cells differentiate into eggs or sperm
                        Animals 
                                Gamete-producing cells differentiate from somatic cells early
                                Referred to as germ line cells
                                Somatic cells are diploid and reproduce by mitosis
                                Diploid gamete-producing cells, produce haploid gametes by meiosis

THE STAGES OF MEIOSIS

                Meiosis Consists of Two Rounds of Nuclear Division
                        Process has much in common with mitosis
                        Different due to two unique features
                                Homologous chromosomes (homologues) pair along their length
                                        Crossing over:  genetic exchange occurs between pairs
                                        Chromosomes come together along equatorial plate
                                        Homologues drawn to opposite poles
                                        Clusters at poles are haploid 
                                        Each chromosome still composed of two chromatids
                                Chromosomes do not replicate between divisions
                                        Second division is nearly identical to mitosis
                                        Sister chromatids dissimilar because of crossing over
                        Process is continual, arbitrarily divided into stages
                        Two stages called meiosis I and meiosis II
                        Each stage divided into prophase, metaphase, anaphase and telophase     fig 12.5
                        Meiosis prophase I more complex than mitosis prophase

                The First Meiotic Division
                        Prophase I
                                Individual chromosomes become visible under light microscopy
                                Each chromosome replicated thus present as two sister chromatids
                                Ends of chromatids attach to nuclear envelope at specific sites
                                        Membrane sites of homologues are adjacent
                                        Members of homologous pairs brought close together
                                Homologues line up side by side in synapsis     fig 12.6
                                Forms resultant synaptonemal complex
                                        Held together by protein latticework
                                        Each gene held in precise register with its corresponding gene
                                        DNA duplexes unwind
                                        Single strands of DNA pair with complementary strand from other homologue
                                Synapsis initiates a complex event called crossing over
                                        DNA segments are exchanged between sister chromatids
                                        When process is complete synaptonemal complex breaks down
                                Homologous chromosomes released from the nuclear membrane
                                Homologues do not separate completely
                                        Sister chromatids held together by their common centromere
                                        Paired homologues held together at points of crossing over
                                Points of crossing over may be visible as X-shaped chiasma      fig 12.7
                                        Chiasma indicates that two chromatids have exchanged parts
                                        Chiasma move to ends of arms as chromosomes separate
                Metaphase I
                                Nuclear envelope disperses, microtubules form spindle as in mitosis
                                Different from mitosis as homologues are paired
                                        Terminal chiasmata:  when chiasma reaches ends of chromosome
                                        One side of centromere faces outward
                                        One side of centromere faces other homologue    fig 12.8
                                        Spindle microtubules can only attach to outer face of centromere
                                        Centromere of each homologue attached to only one pole
                                Joined homologues line up on metaphase plate
                                Attachment of homologue to a pole is random     fig 12.9
                        Anaphase I
                                With completion of spindle attachment, microtubules shorten
                                Chiasma broken, centromeres pulled toward each pole
                                        Chromosomes dragged along
                                        Individual centromeres not pulled apart 
                                        Sister chromatids do not separate
                                Each pole has a complete set of haploid chromosomes
                                        Each set contains one member of each homologous pair
                                        Poles receive homologues randomly 
                                Genes on different chromosomes assort independently
                        Telophase I
                                Each pole has full complement of chromosomes
                                Each chromosome exists as sister chromatids joined by centromere
                                Chromatids are not identical because of crossing over
                                Cytokinesis may or may not occur at this point

                The Second Meiotic Division     fig 12.10
                        Is a simple mitotic division using the products of meiosis I
                        Two clusters at either pole divide mitotically
                                Spindle apparatus binds to sides of centromeres
                                Centromeres divide
                                Chromatids drawn to opposite poles
                        End result is four haploid complements of chromosomes
                        Nuclear envelopes reform
                        In animals, cells develop into gametes
                        In plants, fungi, protists cells may proliferate via mitotic divisions

WHY SEX?

                Not All Reproduction Is Sexual
                        Asexual reproduction
                                Individual inherits all chromosomes from one parent
                                Individual is genetically identical to parent
                        Bacterial cells reproduce by binary fission
                        Protists divide asexually unless under stress
                        Multicellular organisms
                                May reproduce by budding off localized masses of cells
                                Sponges reproduce asexually by fragmentation
                        Development from an unfertilized egg via parthenogenesis
                                Example:  bees 
                                        Fertilized eggs become diploid females
                                        Unfertilized eggs become haploid males
                                Example:  vertebrates   fig 12.11

                Evolutionary Rationale for Sexual Reproduction
                        Problems associated with sexual reproduction
                                Advantage to species which benefit from genetic variability
                                Evolution occurs because of changes at level of the individual
                        Recombination is evolutionarily both constructive and destructive
                                Segregation of chromosomes disrupts beneficial gene combinations
                                Diverse progeny will be less well-adapted than parents
                                Complex adaptations are less likely to benefit from recombination
                        Trends for asexually-reproducing organisms 
                                Live in harsh habitats
                                Premium on well-adapted, genetically uniform individuals
                        Benefit to sexual reproduction is yet unknown
                                Meiotic recombination among protists is often absent
                                Sex may only occur under stressful conditions
                                In some protists diploid is transient or only haploid phase exists
                                        With stress haploids fuse forming diploid zygote
                                        Resulting diploid may not persist
                        Sex may have evolved in protists to repair DNA damage
                                Particularly double-stranded breaks in DNA
                                Breaks induced by radiation or chemicals
                                Repair of such damage is necessary in longer-lived organisms
                                DNA repair through mechanism of synaptonemal complex
                                Transient diploid stage allows for such repair
                                Special yeast mutations
                                        Repair system inactivated for double-strand breaks
                                        Crossing over also prevented

THE EVOLUTIONARY CONSEQUENCES OF SEX

                Principal Factors in the Evolution of the Eukaryotes
                        Reassortment of genetic material occurs during meiosis
                                Represents an enormous factor in initiation of genetic variability
                                        In humans 23 chromosomes are from each parent
                                        Each chromosome segregates independently of all others
                                        Gamete possibilities equals 223 (over eight million)
                                Fertilization squares the number of possibilities (70 trillion)
                        Crossing over further adds to the variability

                Evolutionary Consequences of Sex Are Profound
                        Genetic diversity is raw material of evolution
                        Pace of evolution increased with greater genetic diversity

                Evolutionary Process Is Revolutionary and Conservative
                        Revolutionary as the pace is quickened by genetic variability
                        Conservative as variation is not always favored by selection
                                Acts to preserve existing combinations of genes
                                Greater in asexual organisms that are not highly mobile
                                Live in extremely demanding habitats
                        In vertebrates, the evolutionary premium is on versatility, thus sex



 

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