Abstract

Replicated sister chromatids are held in close association from the time of their synthesis until their separation during the next mitosis. This association is mediated by the ring-shaped cohesin complex that appears to embrace the sister chromatids. Upon proteolytic cleavage of the α-kleisin cohesin subunit at the metaphase-to-anaphase transition by separase, sister chromatids are separated and segregated onto the daughter nuclei. The more complex segregation of chromosomes during meiosis is thought to depend on the replacement of the mitotic α-kleisin cohesin subunit Rad21/Scc1/Mcd1 by the meiotic paralog Rec8. In Drosophila, however, no clear Rec8 homolog has been identified so far. Therefore, we have analyzed the role of the mitotic Drosophila α-kleisin Rad21 during female meiosis. Inactivation of an engineered Rad21 variant by premature, ectopic cleavage during oogenesis results not only in loss of cohesin from meiotic chromatin, but also in precocious disassembly of the synaptonemal complex (SC). We demonstrate that the lateral SC component C(2)M can interact directly with Rad21, potentially explaining why Rad21 is required for SC maintenance. Intriguingly, the experimentally induced premature Rad21 elimination, as well as the expression of a Rad21 variant with destroyed separase consensus cleavage sites, do not interfere with chromosome segregation during meiosis, while successful mitotic divisions are completely prevented. Thus, chromatid cohesion during female meiosis does not depend on Rad21-containing cohesin.

Highlights

  • During meiosis, haploid germ cells are generated from diploid parental cells by two consecutive cell divisions without intervening DNA replication

  • synaptonemal complex (SC) formation commences with the establishment of the axial elements (AE) which represent a scaffold running alongside the paired sister chromatids within each homolog

  • Meiosis is a specialized form of cell division that ensures production of germ cells with the right number of chromosomes, so that at fertilization the embryo receives complete sets of paternal and maternal chromosomes

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Summary

Introduction

Haploid germ cells are generated from diploid parental cells by two consecutive cell divisions without intervening DNA replication. Before the first meiotic division, homologous chromosomes are paired into bivalents and the two sister centromeres in each homolog are constrained to behave as a functional unit. The two homologous centromeres of each bivalent are bi-oriented in the spindle and segregated apart during the first meiotic division. Crossovers are generated by meiotic recombination between non-sister chromatids of homologous chromosomes. Fungi and plants double strand breaks (DSBs) mark the first event of meiotic recombination, and DSBs are required for the intimate pairing (synapsis) of homologous chromosomes during the extended prophase of meiosis I. SC formation commences with the establishment of the axial elements (AE) which represent a scaffold running alongside the paired sister chromatids within each homolog. Proper C(3)G localization requires C(2)M but not vice versa [3]

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