Meiosis and mitosis Meiotic prophase (diakinesis) in a sporocyte of Ophioglossum reticulatum, showing about 630 bivalents. Meiotic chromosome dance The function of meiosis is to generate cells that contain exactly half of the genetic materials of the parental cells and that develop into germ cells. Chromosome rarrangements could occur during meiosis, get fixed in populations, and eventually can contribute to genetic differentiation and speciation. Mitosis vs. Meiosis I II Meiosis I and II Meiosis II (equational division) • prophase • metaphase • anaphase • telophase Meiosis I (reductional division) • prophase leptotene zygotene pachytene diplotene diakinesis • metaphase • anaphase • telophase Prophase I • leptotene • zygotene • pachytene • diplotene • diakinesis (a – f) prophase I (a) early zygotene (b - d) early to late pachytene (e) diplotene (f) diakinesis (g, h) metaphase I (i, j) anaphase I (k) telophase I prophaseI MEIOSISIMEIOSISII Meiotic divisions I and II in the rye (Secale cereale) (l) prophase II (m) metaphase II (n) anaphase II (o) telophase II (four haploid pollen mother cells) Meiotic divisions I and II in Arabidopsis thaliana MEIOSISIMEIOSISII (A - H) prophase I (A) leptotene (B) zygotene (C) pachytene (D) diplotene (E) diakinesis (F) metaphase I (G) anaphase I (H) telophase I (I) prophase II (J) metaphase II (K) anaphase II (L) telophase II (M) four newly formed nuclei IV a IV b IV a IV b Prophase I in Arabidopsis thaliana as revealed by chromosome painting NOR CEN leptotene zygotene pachytene 139 clones of a BAC tiling path covering Arabidopsis chromosome 4 were divided into 11 pools of 8-18 BACs. Individual pools were labelled either by biotin-dUTP (red) or digoxigenin-dUTP (green) for painting of either the long arm (113 BACs) or the entire chromosome (139 BACs). early prophase I CEN Key events of meiosis I Chromosome recognition and pairing - the mechanism is not known Synapsis - synaptonemal complex (SC) - the link between synapsis and recombination is not well understood Meiotic recombination - process of formation of doublestrand breaks (DSBs) and their subsequent repair - results in formation of crossover and non-crossover products Links between chromosome pairing, synapsis and recombination are not well undestood. Recombination plays a key role in unifying meiotic events in prophase I. Homologous chromosome recognition and pairing Schematic of centromere behavior in early meiotic prophase I in a hypothetical diploid organism with two pairs of chromosomes Homologous chromosome pairing facilitated by telomere and centromere clustering/coupling/pairing Homologous chromosome pairing facilitated by telomere and centromere clustering 2n = 28 CENs TELs leptotene zygotene telomere bouquet (centromere pairing is prior to telomere clustering) Arabidopsis: telomere clustering around nucleolus Armstrong et al. 2001, JCS Prophase I – pachytene (synapsis of homologous chromosomes) synaptonemal complex in Arabidopsis thaliana Synaptonemal complex (SC) - consists of two lateral elements (le) connected by a central element (ce) [the lateral elements formed as axial elements (AEs, also called the chromosome axis) in leptotene] - the central element assembles following chromosome pairing during zygotene - recombinational nodule (rn) – place of crossingover (genetic exchange between homologues Recombination (crossing over) and chiasmata Prophase I: crossing over • SPO11 protein introduces DSBs into chromosomal DNA • DNA ends adjacent to these breaks are later bound by RAD51 and DMC1 that catalyze singleend invasion of the broken DNA ends into the homologous chromosome Recombination: double-strand breaks (DSBs) and their repair dHJ: double-Holliday junction; SDSA: synthesis-dependent strand annealing; COs: crossovers; NCOs: noncrossovers D-loop Recombination (crossing over) and chiasmata chiasma formation Plants – male gametogenesis Plants – famale gametogenesis Plants – fertilization 3x endosperm