HL Meiosis Syllabus Statements 10.1 HL Understandings of Meiosis 1. Chromosomes replicate in interphase before meiosis. 2. Crossing over is the exchange of DNA material between nonsister homologous chromatids. 3. Crossing over produces new combinations of alleles on the chromosomes of haploid cells. 4. Chiasmata formation between non-sister chromatids can result in an exchange of alleles. 5. Homologous chromosomes separate in meiosis 1. 6. Sister chromatids separate in meiosis II. 7. Independent assortment of genes is due to the random orientation of pairs of homologous chromosomes in meiosis 1 Applications and Skills • 1. Draw diagrams to show chiasmata formed by crossing over. Diagrams of chiasmata should show sister chromatids still closely aligned, except at the point where crossing over occurred and a chiasma was formed. Syllabus statements 10.1.1 Describe the behavior of the chromosomes in the phases of meiosis. 10.1.2 Outline the formation of chiasmata in the process of crossing over. Use diagrams 10.1.3 Explain how meiosis results in an effectively infinite genetic variety in gametes through crossing over in prophase I and random orientation in metaphase I. 10.1.4 Distinguish between autosomes and sex chromosomes. 10.1.5 Explain how crossing over between non-sister chromatids of a homologous pair in prophase 1 can result in an exchange of alleles. Use diagrams 10.1.6 State Mendel’s law of independent assortment. 10.1.7 Define linkage group. Retro: • 1. Compare diploid chromosome numbers of Homo sapiens, Pan troglodytes, Canis familiaris, Oryza sativa, Parascaris equorum. • 2. Define genome size (total length of DNA in an organism). • 3. Compare genome size of T2 phage, Escherichia coli, Drosophila melanogaster, Homo sapiens and Paris japonica. • 4. Describe Cairns’ technique for measuring the length of DNA molecules by autoradiography. • 5. Description of methods used to obtain cells for karyotype analysis e.g. chorionic villus sampling and amniocentesis and the associated risks. • 6. Draw diagrams to show the stages of meiosis resulting in the formation of 4 haploid cells. 1. Compare diploid chromosome numbers of Homo sapiens, Pan troglodytes, Canis familiaris, Oryza sativa, Parascaris equorum. The number of chromosomes is a characteristic feature of members of a species. The diploid number is the number of chromosomes present in normal body cells (not gametes). The diploid number varies with some species having fewer large chromosomes and others having a greater number of small chromosomes. Five examples of chromosome number Homo sapiens (humans) 46 Pan troglodytes (chimpanzee) 48 Canis familiaris (dog) 78 Oryza sativa (rice) 24 Parascaris equorum 4 (Horse threadworm) 1. Explain why none of the species has an odd number of chromosomes. 2. Discuss, using these data, the hypothesis that the more complex an organism is, the more chromosomes it has. 3. Explain why the size of the genome of a species cannot be deduced from the number of chromosomes. 4. Suggest, using this table, a change in chromosome structure that may have occurred during human evolution. 3. Compare genome size of T2 phage, Escherichia coli, Drosophila melanogaster, Homo sapiens and Paris japonica. • Genome size (total length of DNA in an organism). This varies by a huge amount with the smallest genomes those of viruses. Given in million base pairs. • T2 phage (virus that attacks E. coli): 0.18 • Escherichia coli (gut bacterium): 5 • Drosophila melanogaster (fruit fly): 140 • Homo sapiens (humans): 3000 • Paris japonica (woodland plant) : 150,000 Trends • The prokaryote has the smallest genome. • Genome size of eukaryotes depends on the size and number of chromosomes. • It is correlated with the complexity of the organism, but is not directly proportional. • The reason for this is that the proportion of DNA that acts as functional genes is very variable and also the amount of gene duplication varies. 4. Describe Cairns’ technique for measuring the length of DNA molecules by autoradiography. Stolen from Allott and Mindorf (2014) • John Cairns produced images of DNA molecules from E. coli using this technique: • Cells were grown for 2 generations in a culture medium containing tritiated thymidine (thymine linked to deoxyribose). Tritium is a radioactive isotope of hydrogen…radioactively labelled DNA produced by replication. • Cells placed in dialysis membrane and their cell walls using lysozyme. Cells gently burst with DNA released onto surface of dialysis membrane. • Thin film of photographic emulsion applied to the surface of the membrane and left in darkness for 2 months. Some of the atoms of tritium in the DNA decayed and emitted high energy electrons, which react with the film. • At end of 2 months, film was developed and examined under a microscope. Dark grains appeared every place tritium atom decayed indicating position of DNA. • Images of E coli showed that chromosome was single, circular molecule with a length of 1,100 um. (length of E. coli is 2 um). Autoradiography John Cairns 1963 Chromosome of E. coli 5. Description of methods used to obtain cells for karyotype analysis e.g. chorionic villus sampling and amniocentesis and the associated risks. Preparation of Human Karyogram (Karyotype is a property of a cell – the number and type of chromosomes present in the nucleus) The Human Life Cycle Meiosis Overview Review of Definitions 1. Somatic cells: non sex cells. 2. Sex cells (gametes): egg (ova), sperm 3. Autosome: chromosome that is not a sex chromosome (does not determine gender). 4. Sex chromosomes: dissimilar chromosomes that determine an individual’s sex. In humans: X, Y 5. Homologous chromosomes: pairs of chromosomes that have the same size, centromere position, staining pattern, location of genes. Definitions continued 6. Fertilization: the union of two gametes to form a zygote. 7. Zygote: a diploid cell that results from the union of 2 haploid gametes. 8. Meiosis: special type of cell division that produces haploid cells and compensates for the doubling of chromosome number that occurs at fertilization. Key Differences with Mitosis 1. Meiosis is a reduction division: gametes have ½ number of chromosomes as parent cell. 2. Meiosis creates genetic variation: 4 daughter cells genetically different from parent cell and from each other. 3. Meiosis is 2 successive nuclear divisions. Meiosis I Meiosis II Stages of Meiotic Cell Division a) Interphase I: precedes meiosis i. Chromosomes replicate ii. Each duplicated chromosome consists of 2 identical sister chromatids attached at their centromere. iii. Centriole pairs in animal cells also replicate into two pairs. Meiosis I: Segregates the 2 chromosomes of each homologous pair and reduces the chromosome number by one-half. b)Prophase I: 90% of the time required for meiosis. i. Longer and more complex than prophase of mitosis. ii. Chromosomes condense – start to coil up and become shorter and thicker. iii. Synapsis occurs: homologous chromosomes come together as pairs. Uses the synaptenemal complex to accomplish this. Chromosome coiling (nucleosome) and pairing of bivalents with synaptonemal complex Prophase I continued iv. Each chromosome has 4 chromatids, so that each homologous pair in synapsis appears as a complex of 4 chromatids of a tetrad. v. In each tetrad, sister chromatids of the same chromosome are attached at their centromeres. vi. Nonsister chromatids are linked by X-linked chiasmata, sites where homologous strand exchange or crossing over occurs. vii. Chromosomes thicken further and detach from the nuclear envelope. viii. Centriole pairs move apart, toward poles, and spindle microtubules form between them. ix. Nuclear envelope and nucleoli disperse. Chromosome Tetrad Meiosis I continued c) Metaphase I: tetrads (bivalents) are aligned on the metaphase plate, having moved there during this stage. i. Chromosomes continue to shorten and thicken. ii. Spindle microtubules attach to the kinetochore region of the centromeres. iii. Bivalents line up on the equator so that centromeres of homologues point towards opposite poles. iv. Chiasmata slide towards the ends of the chromosomes causing the shapes of the bivalents to change. v. At the end, chromosomes start to move. Metaphase 1: 2n = 4; 2n = 6 Chiasmata terminalizing Meiosis I continued d) Anaphase I: homologues separate and are moved towards the poles by the spindle apparatus. (Bivalents separate). This halves the chromosome number. i. Each chromosome consists of 2 chromatids (sister chromatids remain attached at the centromeres). ii. Because of crossing over, the 2 chromatids are not identical. iii. At the end, chromosomes reach the poles. Meiosis I continued e) Telophase I and cytokinesis: spindle apparatus continues to separate homologous chromosome pairs until the chromosomes reach the poles. i. Each pole has a haploid set of chromosomes composed of 2 sister chromatids. ii. Nuclear membranes form around the groups of chromosomes at each pole. iii. Chromosomes uncoil partially. iv. Cytokinesis occurs simultaneously forming 2 daughter cells. Cleavage furrows form in animal cells and cell plates form in plant cells. Meiosis I: reduction division – separation of homologous chromosomes Meiosis II: Separation of sister chromatids Interphase II: May not really exist per se f) Two cells either enter a brief period of interphase or immediately proceed to the second division of meiosis. i) DNA is not replicated. Meiosis II: separates sister chromatids of each chromosome g) Prophase II: chromosomes become shorter and thicken again by coiling. i. Centrioles move to poles (animal cells). ii. Nuclear membrane breaks down. iii. Spindle apparatus forms and chromosomes move towards the metaphase II plate. Meiosis II continued h) Metaphase II: chromosomes align singly on the metaphase plate. i. Spindle microtubules attach to the centromeres. ii. Chromosomes line up on equator. iii. Centromeres divide iv. Kinetochores of sister chromatids point towards opposite poles. Metaphase 2: 2n = ? Meiosis II continued i) Anaphase II: sister chromatids separate and move toward opposite poles of the cell. j) Telophase II and Cytokinesis: nuclei form at opposite poles of the cell. i) nuclear membranes reform around the groups of chromatids at each pole. ii) Cytokinesis produces 4 cells total. iii) Chromosomes uncoil, nucleoli appear iv) In most organisms, develop into gametes Comparison of mitosis and meiosis Summary Comparison Meiosis and fertilization Primary sources of genetic variation in sexually reproducing organisms. Sexual reproduction provides genetic variation by: 1. Independent Assortment 2. Crossing over during Prophase I 3. Random fertilization by gametes Independent Assortment of chromosomes 1. Orientation of the homologous pair of chromosomes (one maternal and one paternal) is RANDOM. 50-50 chance that each gamete receives maternal or paternal derived chromosome. 2. Each homologous pair of chromosomes orients independently of the other pairs at metaphase I. Ist meiotic division results in independent assortment of maternal and paternal chromosomes. Alternative arrangements of 2 homologous chromosome pairs Definition: Independent Assortment Independent Assortment: The random distribution of maternal and paternal homologues to the gametes. OR Independent Assortment: The presence of an allele of one of two genes in a gamete has no influence over which allele of another gene is present in the same gamete. Independent Assortment leads to Genetic Variation Process produces 2n possible combinations of maternal and paternal chromosomes in gametes. If n = 2 then 22 = 4. If n = 23 (human) then 223 or 8,000,000 different possibilities before crossing over Crossing Over Definition: the exchange of genetic material between homologues (homologous portion of 2 non-sister chromatids trade places). X-shaped chiasmata are the visible evidence of this process. Produces chromosomes that contain genes from both parents. In humans, an average of 3 crossovers/chromosome pair. Crossing Over continued Most of the time, crossing over can occur without loss of genetic material because of the precise base to base pairing of homologues involving the formation of the synaptonemal complex, a protein structure that brings the chromosomes into close association. The results of crossing over during meiosis Process of Crossing Over: 10.1.2 Benefits of Crossing Over: 10.1.5 1. Chiasmata hold together homologues during Pro. & Met. I. 2. Allows recombination of linked genes. Breaks up linkage groups/parental combinations. Crossing over results in an exchange of alleles • Parental combinations of linked genes cannot be broken up without crossing over. • Crossing over occurs between nonsister chromatids of a homologous pair in prophase 1 between the loci of the 2 linked genes . • Parentals: abc, ABC • Crossing over occurs between B and C • Position of chiasma formed by crossing over • The 4 chromatids separate into 4 nuclei produced by meiosis. • Recombinants produced = abC, ABc Linkage groups • Definition: All the genes that have their loci on the same chromosome type form a linkage group. Example of Gene Linkage Random Fertilization: Result of sexual reproduction In humans an egg cell with 1 of 8,000,000 different possibilities will be fertilized by a sperm cell that is also 1 of 8,000,000 possibilities, resulting in a zygote that can have 1/64,000,000,000,000 possible diploid combinations. Evolutionary Advantages of Genetic Variation in Populations Basis of the theory of natural selection is inheritable variation. Darwinian mechanism for evolutionary change is natural selection: Most fit organism leaves behind the most young a) Increases the frequency of inheritable variations that favor the reproductive success of some individuals over others. Adaptation continued b) Results in adaptation, the accumulation of inheritable variations that are favored by the environment. c) Genetic variation allows populations to cope with new environmental conditions. In the Red Queen, Matt Ridley, suggests that the major reason that sex has been selected for is that genetic variation allows species (including humans) to adapt to evolving parasites and pathogens. d) Two sources of genetic variation: sexual reproduction and mutation (rarer).