Outline for today’s lecture (Ch. 13) • Sexual and asexual life cycles • Meiosis • Origins of Genetic Variation – Independent assortment – Crossing over (“recombination”) Heredity • Transmission of traits between generations • Molecular basis of heredity is DNA replication • Gene is a specific segment of DNA • Physical location on the chromosome is called a genetic LOCUS (plural = “loci”) – e.g., the “eye-color locus”, Adh locus Asexual Life Cycles • Single (diploid) individual is the parent • Parent passes copies of ALL its genes to its offspring (reproduces “clonally”) • Various mechanisms – Mitotic cell division in unicellular Eukaryotes – Vegetative reproduction, e.g., plant cuttings, hydra budding – Parthenogenesis Sexual Life Cycles • Two (diploid) parents give rise to offspring • Offspring differ genetically from their parents and their siblings • GAMETES are haploid reproductive cells that transmit genes across generations Sexual Life Cycles • Key Point: Sexual reproduction → Genetic variation • MOST eukaryotes reproduce sexually at least sometimes • Most prokaryotes (e.g., bacteria) exchange genes at least occasionally Sexual Life Cycles – Human Example • 46 Chromosomes • 22 Homologous pairs, called “autosomes” – Same length – Same centromere position – Same sequence (+/-) – SAME GENES!! Sexual Life Cycles – Human Example • One pair of “sex chromosomes” – i.e., “sex-determining gene(s)” reside on these chromosomes • Females are XX • Males are XY • Only small region of homology (= same genes) between X, Y X Y Schematic drawing of a chromosome Diploid cell, n=3 BEFORE DNA replication • 3 Homologous Pairs – 2 autosomes – 1 sex chromosome (XX) • One homologous chromosome from each parent 2 2 X • DNA content = 2C 1 X • Ploidy = 2n 1 Diploid cell, n=3, AFTER DNA replication • 3 Homologous Pairs • One homologous chromosome from each parent = TWO SISTER CHROMATIDS 22 • DNA content = 4C 22 XX XX • Ploidy = 2n 11 11 Sexual Life Cycles - animals • Free-living stage is diploid • Gametes formed by meiosis • Haploid gametes merge genomes to form diploid zygote (“syngamy”) Sexual Life Cycles - Plants • Diploid sporophyte forms haploid spores by meiosis • Spores form gametophyte by mitosis • Gametophyte forms gametes by mitosis • Gametes merge to form diploid zygote Sexual Life Cycles - Fungi • Free-living, multicellular organism is haploid • Gametes formed by mitosis • Gametes merge to form diploid zygote • Zygote undergoes meiosis to form haploid cells Meiosis • RECALL: Function of MITOSIS is to faithfully replicate the parental genome in each daughter cell with no change in information content • Function of MEIOSIS is to produce haploid cells from diploid cells • Necessary for the formation of gametes • Necessary for sexual reproduction Meiosis – an overview • Interphase 1 – – Begin with two homologous chromosomes, – DNA content = 2C – Ploidy = 2n (diploid) Meiosis – an overview • Interphase 1 – – Chromosomes replicate – DNA content = 4C – Ploidy = 2n Meiosis – an overview • “Meiosis I” – Homologous chromosomes separate – Cell Division #1 – Result is TWO haploid (ploidy = n) cells with TWO SISTER CHROMATIDS of one of the two homologs Meiosis – an overview • “Meiosis II” – Sister chromatids separate – Cell Division # 2 – Result is FOUR haploid daughter cells, each with an unreplicated chromosome (= 1C) – Half as many chromosomes as the parent cell Meiosis I – early Prophase I Chiasmata • Homologous chromosomes pair • Synaptonemal complex (proteins) attaches homologs – “synapsis” • Homologs form tetrad Tetrad Meiosis I – late Prophase I Chiasmata • Chromosomes cross over, form “chiasmata” Spindle fiber • Exchange of DNA between homologs occurs at chiasma • Spindles form and attach to kinetochores as in mitosis Tetrad Meiosis I – Metaphase I • Chromosomes lined up on metaphase plate in homologous pairs • Spindles from one pole attach to one chromosome of each pair • Spindles from the other pole attach to the other chromosome of the pair Kinetochore Meiosis I – Anaphase I • Homologous chromosomes separate and move along spindle fibers toward pole • Sister chromatids remain attached at centromeres • Note that recombination has occurred! Meiosis I – Telophase and cytokinesis • Homologous chromosomes reach (opposite) poles • Each pole has complete haploid complement of chromosomes • Each chromosome consists of two sister chromatids Meiosis II – Prophase II • Spindle forms • Chromosomes move toward metaphase plate Meiosis II – Metaphase II • Chromosomes reach metaphase plate, as in mitosis • Kinetochores of sister chromatids attach to spindle fibers from opposite poles Meiosis II – Anaphase II • Centromeres of sister chromatids separate • Sister chromatids move toward opposite poles Meiosis II – Telophase and cytokinesis • Mechanism as before • Note that now FOUR HAPLOID DAUGHTER CELLS formed from each parent cell • Note that some chromosomes are recombinant, some are not Meiosis I - Summary Chiasma (site of crossing-over) Tetrad formed by synapsis of homologs Meiosis I - Summary Tetrads align at metaphase plate Meiosis I - Summary Homologous chromosomes separate Sister chromatids remain paired Meiosis II - Summary Sister chromatids separate Haploid daughter cells result Origins of Genetic Variation 1. Independent Assortment of Chromosomes • 2. Crossing over • 3. Recombination among chromosomes Recombination within chromosomes Random fertilization Independent Assortment of Chromosomes Independent Assortment of Chromosomes • Number of possible combinations of chromosomes within a gamete – Two homologs A, B: Mom = A1B1, Dad = A2B2 • Four combinations: A1B1, A1B2, A2B1, A2B2 – Three homologs: Mom = A1B1C1, Dad = A2B2C2 • Eight combinations: A1B1C1, A1B1C2, A1B2C1, A1B2C2, A2B1C1, A2B2C1, A2B1C2, A2B2C2 – n homologs: 2n combinations Crossing-over – Recombination within chromosomes • Averages ≥ 2 per chromosome per meiosis in humans, flies • If no crossing-over, genes on same chromosomes would always be inherited together Crossing-over – Recombination within chromosomes Human genome has ~20K genes. Suppose each gene assorts independently. How many combinations? Review: Mitosis vs. Meiosis Event DNA Replication # Cell Divisions # Daughter cells “Ploidy” of daughters Synapsis of homologs? Crossing-over (recombination) Biological Purpose Mitosis Interphase 1 2 2n (diploid) No No Duplicate cells faithfully Meiosis Interphase I 2 4 n (haploid) Yes Yes Generate gametes Meiosis, Genetic variation, and Evolution • Role of segregation • Role of crossing-over • What about LIMITS to evolution? – E.g., body size For Thursday: Introduction to Mendelian Genetics • Read Chapter 14 through p. 260