what is synapsis in meiosis
Nucleic Acids Res. doi: 10.1371/journal.pgen.1000702, Woltering, D., Baumgartner, B., Bagchi, S., Larkin, B., Loidl, J., de los Santos, T., et al. Remote homology search had previously identified Rec102 as the Topo VIB-like subunit in S. cerevisiae (Robert et al., 2016). Genetics 170, 7185. Bookshelf doi: 10.1016/S0092-8674(00)80609-1, Kugou, K., Fukuda, T., Yamada, S., Ito, M., Sasanuma, H., Mori, S., et al. Spo11 evolved from the catalytic subunits of a type IIB topoisomerase, Topo VI (Bergerat et al., 1997; Keeney et al., 1997). Here, we have developed single, double and triple zip4 TILLING mutants and a CRISPR Ttzip4-B2 mutant, to determine the effect of ZIP4 genes on synapsis and CO formation in the tetraploid wheat cultivar 'Kronos'. Curr. Fertility Sterility (2015) 103(4):9069. 2:e12. Synapsis can increase the variability because synapsis and crossing-over (between two different chromatids) occur during meiosis phase 1. Meiosis is preceded by an interphase consisting of the G 1, S, and G 2 phases, . doi: 10.1101/gad.1099003, Nandabalan, K., and Roeder, G. S. (1995). (C) SC assembly leads to the removal of Hop1 and DSB proteins from the axis, thereby shutting down further DSB formation. Semin. Turner JM. The Meiotic Checkpoint Network: Step-by-Step Through Meiotic Prophase. Science 263, 515518. In mutants with recombination or synapsis defects, checkpoint activation via Mec1 activates Mek1, which inhibits Ndt80 activity and leads to the extension of prophase (Figure 9A, circuit 4; Acosta et al., 2011; Gray et al., 2013; Prugar et al., 2017). Thus, many details about meiotic DSB-repair and crossover formation during in vitro meiosis remain largely unknown. Cell 13, 389401. The structure of Ski8p, a protein regulating mRNA degradation: implications for WD protein structure. The tight pairing of the homologous chromosomes is called synapsis (Figure 2). Finally, the core complex binds with high affinity to the ends of DNA duplexes in vitro (Claeys Bouuaert et al., 2021; Figure 4C, iv). In contrast to mitosis, molecular mechanisms and regulation of meiosis are much less understood. However, in the current culture system, 3 to 4 clear pachytene cells per microscope slide, from day 8 to day 14 (Figure2A), were observed to reach full synapsis of the autosomes, marked by SYCP1 (Figure2B), while clear XY bodies were marked by MDC1 (Figure2B) or H2AX and MDC1 (Figure2C), which is consistent with pachytene spermatocytes in vivo (Figures2B, C). At day 6, the in vitro-induced meiotic cells were dissociated by accutase (Thermo Fisher Scientific), and were subsequently seeded on the additional 12-well plates containing undifferentiated mGSCs and SK49 cells. The presence of a strong hotspot suppresses the DSB activity of an adjacent hotspot (Wu and Lichten, 1994; Xu et al., 1995; Keeney et al., 2014). Recombinational DNA Double-Strand Breaks in Mice Precede Synapsis. A duplicated copy of the meiotic gene ZIP4 preserves up to 50% pollen viability and grain number in polyploid wheat. doi: 10.1074/jbc.M114.625293, Lichten, M., and Goldman, A. S. H. (1995). This work was supported by the European Research Council under the European Unions Horizon 2020 Research and Innovation Program (ERC Grant Agreement 802525) and by the Fonds National de la Recherche Scientifique (FNRS MIS-Ulysse Grant F.6002.20). In many organisms, synapsis and homologous recombination are interdependent: recombination promotes SC formation and SCs are required for crossing-over. Indeed, yeast-two-hybrid (Y2H) experiments showed strong interactions between Spo11 and Ski8, and between Rec102 and Rec104 (Arora et al., 2004; Kee et al., 2004; Maleki et al., 2007). DNA-driven condensation assembles the meiotic DNA break machinery. Holoenzyme assembly and ATP-mediated conformational dynamics of topoisomerase VI. 31, 11641173. In S. pombe, blocking DNA replication also abolishes meiotic DSB formation (Ogino and Masai, 2006). For instance, retinoic acid does not only induce spermatogonial differentiation and meiotic initiation; once germ cells have entered meiosis, pachytene spermatocytes also produce RA to coordinate spermatogenesis (41). doi: 10.1038/nsmb1264, Corbett, K. D., and Berger, J. M. (2003). What is the role of MRX? In contrast to the other core complex proteins, Ski8 is not meiosis-specific. The C-terminal coiled-coil domain of Red1 forms a tetrameric parallel-antiparallel -helical bundle (West et al., 2019). In spermatocytes in vivo, the first meiotic prophase is tightly regulated and prolonged. doi: 10.1038/s41418-017-0015-1, 25. 31, 18801893. *Correspondence: Corentin Claeys Bouuaert, corentin.claeys@uclouvain.be, Meiosis: from Molecular Basis to Medicine, View all The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. Genetics 167, 593605. Mol. Nature 479, 237240. From day 4 to day 6, the same amount of additional 12-well plates was prepared and pre-coated with laminin. Endo T, Freinkman E, de Rooij DG, Page DC. doi: 10.1038/nsmb928, Wojtasz, L., Daniel, K., Roig, I., Bolcun-Filas, E., Xu, H., Boonsanay, V., et al. Ichijima Y, Ichijima M, Lou Z, Nussenzweig A, Camerini-Otero RD, Chen J, et al. Genes Cells 11, 615622. doi: 10.1111/gtc.12138. (B) The tethered loop-axis model for DSB formation. doi: 10.1016/j.yexcr.2014.08.024, Hunter, N. (2015). 276, 4190641912. Temporospatial coordination of meiotic dna replication and recombination via DDK recruitment to replisomes. doi: 10.1038/nsmb.2116, Hollingsworth, N. M., Goetsch, L., and Byers, B. End-to-end polymerization of the coiled-coil is thought to underlie axis assembly (West et al., 2019). A., Hyppa, R. W., and Smith, G. R. (2004). As the nuclear envelope begins to break down, the proteins associated with homologous chromosomes bring the pair close to each other. Cell. doi: 10.1126/science.290.5492.806, Borde, V., Lin, W., Novikov, E., Petrini, J. H., Lichten, M., and Nicolas, A. 311, 303326. Biomolecular condensates: organizers of cellular biochemistry. Xrs2 is essential for the nuclear localization of Mre11 (Tsukamoto et al., 2005). Rev. BLM helicase ortholog Sgs1 Is a central regulator of meiotic recombination intermediate metabolism. doi: 10.1007/s10577-007-1150-1, Madrona, A. Y., and Wilson, D. K. (2004). Biol. Meiosis is a process where germ cells divide to produce gametes, such as sperm and egg cells. Mol. (A) Cartoon illustrating the arrangement of the different subunits in the core complex. The catalytic tyrosine is located in the 5Y-CAP domain and the Toprim domain coordinates Mg2+ ions important for catalysis. doi: 10.1101/cshperspect.a016618, Hunter, N., and Kleckner, N. (2001). doi: 10.1146/annurev.genet.33.1.603, Zickler, D., and Kleckner, N. (2015). Nat. (B) Meiotic recombination is initiated by Spo11-mediated DSB formation and leads to the formation of crossovers via a ZMM-dependent double Holliday Junction (dHJ) resolution pathway or non-crossovers by synthesis-dependent strand annealing. Long-Term Proliferation in Culture and Germline Transmission of Mouse Male Germline Stem Cells. Iwasaki, D., Hayashihara, K., Shima, H., Higashide, M., Terasawa, M., Gasser, S. M., et al. Mer2 binds directly to both nucleosomes and axial proteins as the keystone of meiotic recombination. Proc. EMBO J. In addition, break formation displays sequence bias within and around the footprint of Spo11 and at the cleavage site, with a preference for cleavage 3 of a C (Murakami and Nicolas, 2009; Pan et al., 2011; Figure 2A). doi: 10.1016/j.tcb.2015.07.009, Chen, C., Jomaa, A., Ortega, J., and Alani, E. E. (2014). doi: 10.1093/emboj/17.5.1497, Arora, C., Kee, K., Maleki, S., and Keeney, S. (2004). In comparison to previous culture system, the in vitro-generated spermatocytes were transferred after meiotic initiation to a second culture dish. Sci. In accordance with the very inefficient formation of meiotic crossovers, these M-phase cells only displayed pairs of sister chromatids (univalents) with CREST-stained centromeres located at the ends of the paired sister chromatids (Figure4A). (2014). In humans, the haploid cells made in meiosis are sperm and eggs. Geijsen N, Horoschak M, Kim K, Gribnau J, Eggan K, Daley GQ. Cell 69, 439456. The G 1 phase, which is also called the first gap phase, is the first phase of the interphase and is focused on cell growth. The COMPASS subunit Spp1 links histone methylation to initiation of meiotic recombination. Cell. Nat Rev Genet (2001) 2(4):28091. This may have implications regarding the first steps of DSB processing, since Spo11 could cap the DNA ends during resection and perhaps after strand invasion has initiated. In addition, scar-less repair by non-homologous end joining of meiotic DSBs that have undergone resection in a Drosophila strain with homolog pairing defects (Mcm5A7) provided further support for end-capping by Spo11-oligonucleotides after resection had initiated (Hatkevich et al., 2020). (2019). To our knowledge, only one study described the formation of chiasmata and bivalent chromosome pairs in in vitro-generated MI-spermatocytes, however, the meiotic recombination process and subsequent crossover formation was not described (2). ATM controls meiotic double-strand-break formation. Dev. Rec114Mei4 forms a complex with a 2:1 stoichiometry where the C-terminus of Rec114 homodimerizes and interacts with the N-terminus of Mei4 (Figures 6A,B). PLoS Genet. The reason may be that some DSB sites were not repaired timely and remained present at synapsed chromosomes. doi: 10.1007/s00412-006-0055-7, Klein, F., Mahr, P., Galova, M., Buonomo, S. B. C., Michaelis, C., Nairz, K., et al. PMC Cell Sci. Recombinationless meiosis in Saccharomyces cerevisiae. Figure 10. National Library of Medicine If the complex traps two duplexes like Topo VI, what is the physiological relevance of this activity, since Spo11 activity presumably does not require strand passage? doi: 10.1128/mcb.1.10.891, Martini, E., Borde, V., Legendre, M., Audic, S., Regnault, B., Soubigou, G., et al. The red dotted lines connecting two proteins represent their respective interaction domains. DDK is bound to the replisome via interactions with the fork protection complex (FPC). J. Biol. Cell 16, 597608. (2011). synapses occurs during prophse 1 in meiosis is the pairing of two homologous chromosomes that occurs during meiosis. (A) Domain structure of Rec114, Mei4, and Mer2 with regions involved in protein-protein and protein-DNA interactions (Claeys Bouuaert et al., 2021). Prophase I Early in prophase I, the chromosomes can be seen clearly microscopically. Asy2/Mer2: an evolutionarily conserved mediator of meiotic recombination, pairing, and global chromosome compaction. Royo H, Prosser H, Ruzankina Y, Mahadevaiah SK, Cloutier JM, Baumann M, et al. doi: 10.1016/j.molcel.2010.11.032, Zakharyevich, K., Tang, S., Ma, Y., and Hunter, N. (2012). However, during in vitro meiosis, we observed that, although chromosome synapsis was complete, still very few crossovers were detected. QL is recipient of a China Scholarship Council grant (201706300107). doi: 10.1016/0092-8674(90)90216-2, Hong, E. L., Shinohara, A., and Bishop, D. K. (2001). Recombination is a prominent feature of meiosis in which it plays an important role in increasing genetic diversity during inheritance. Hum Reprod (2018) 33(1):8190. Synapsis of Homologous Chromosomes In human somatic cells, there are 46.
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