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Overview • DNA replication is prerequisite to cell division. • DNA is replicated by using each strand as template for synthesis of the complementary strand. – one strand is synthesized continuously, the other discontinuously – all nucleotides are added at the 3’ end of the growing strand • Replicated chromosomes are partitioned to nuclei of daughter cells in mitosis. • Meiosis distributes replicated chromosomes to haploid daughter cells in two nuclear and cell divisions. Cell division in eukaryotes • Mitosis – – – – an asexual division equal division of replicated chromosomes occurs with any number of chromosomes part of cell cycle • Cell cycle – – – – G1: gap (growth) S: DNA replication, forms sister chromatids G2: gap (growth) M: mitosis (nuclear division) and cytokinesis 1 Mitosis • Nuclear division, usually with cell division • Divided into four continuous stages – prophase • chromosomes condense and attach to spindle via kinetochore • nucleolus and nuclear membrane disappear – metaphase: chromosomes align on equatorial plane of cell – anaphase: sister chromatids separate and migrate to opposite poles – telophase: cell division and reformation of nucleus and nucleolus 2 Mitotic machinery • Sister chromatids (replicated chromosomes) attached by centromere – identical in DNA sequence by rules of DNA replication (base pairing) – centromere acts as binding site for kinetochore, a site for attachment of microtubules • Spindle apparatus – microtubules assembled from tubulin – motor proteins – provides force for separation of chromatids Meiosis • Specialized cell division to halve chromosome number • Compensates for doubling of chromosome number by fertilization • Occurs in meiocyte at fixed point in life cycle – premeiotic S phase to replicate chromosomes – meiocyte divides twice to yield tetrad of four haploid cells – meiocyte effectively 4n prior to division, yielding four 1n cells Meiotic divisions • Replicated homologous chromosomes pair along their length (synapsis) – form tetrad of four chromatids – paired nonsister chromatids may undergo crossing-over • At first meiotic division, chromosomes segregate to opposite poles – spindle attaches only one side of centromere • At second meiotic division, sister chromatids segregate to opposite poles Consequences of meiosis • Formation of four haploid cells, each with one complete copy of genome • Recombination – crossing-over in prophase I – independent assortment of nonhomologous chromosomes – results in genetic diversity • Meiosis occurs at some point in life cycle of all sexually reproducing organisms 3 Chapter 5 The Inheritance of SingleGene Differences Alleles at single locus Overview • In matings, precise phenotypic ratios are produced in descendants as a result of chromosome segregation. • In heterozygotes, alleles segregate equally into meiotic products. • Progeny ratios can be predicted from known genotypes of parents. • Parental genotypes can be inferred from phenotypes of progeny. • In many organisms, sex chromosomes determine sex. • X-linked genes can show different phenotypic ratios in male and female progeny. • In humans, single-gene traits can be studied in pedigrees. • Organelle genes are inherited maternally. Meiotic chromosome segregation • In meiosis, each of the four haploid products receives one of each kind of chromosome – A/A homozygotes → all get A chromosomes – A/a heterozygotes → half get A chromosomes half get a chromosomes • As a consequence of chromosome segregation, alleles of heterozygotes segregate equally 4 Equal segregation • First observed by Mendel in crosses with peas • Readily observed using some fungi and protists in which all four haploid products of meiocyte (tetrad) can be analyzed A a A a Aa Crosses • Controlled mating of two individuals – obtain desired genotype – deduce genotypes of parents • Selfing: individual with both reproductive organs reproduces (crosses) to itself • × symbol is used to indicate a cross – a haploid mating: A × a – a diploid mating: A/a × A/a • A/– : – represents either A or a Diploid crosses (1) • Three possible diploid genotypes A/A a/a A/a • Six possible diploid crosses Cross A/A × A/A a/a × a/a A/A × a/a A/a × A/A A/a × a/a A/a × A/a Genotypic ratio A/A a/a A/a 1A/A:1A/a 1A/a:1a/a 1A/A:2A/a:1a/a Phenotypic ratio all A all a all A all A 1A:1a 3A:1a 5 Diploid crosses (2) • Crosses between individuals heterozygous for the same single gene are also called monohybrid crosses • A heterozygote for unexpressed recessive allele is sometimes called a carrier, particularly in humans • A cross between an unknown genotype (e.g., A/–) and the homozygous recessive genotype (a/a) is called a testcross 6