Iately regulate the Rad51 nucleoprotein filament dynamics in the course of meiosis.RECOMBINATION in between homologous sequences promotes genomic stability. As such, a wide selection of DNAassociated proteins and protein complexes regulate the recombination process. Within chromatin, the chromosome structure in all probability also directly impacts the assembly and disassembly of protein complexes involved in recombination. To know the molecular mechanisms that govern assembly and disassembly of those complexes, homologous recombination throughout meiosis is often utilised as a model system. Meiotic recombination is initiated when Spo11, a meiosisspecific topoisomerase-like protein, generates double-strand breaks (DSBs) (Keeney 2001). The 59 ends of those DSBs are speedily resected to create a 39-overhanging stretch of single-stranded DNA (ssDNA). The ssDNA is used to search for homologous double-stranded DNAs (dsDNAs). Once homology among the ss- and dsDNAs are matched, the ssDNA is invaded in to the dsDNA. Strand invasion leads to DNA syn-Copyright 2013 by the Genetics Society of America doi: 10.1534/genetics.113.150615 Manuscript received February 20, 2013; accepted for publication June five, 2013 Supporting information is offered online at http://www.genetics.org/lookup/suppl/ doi:10.1534/genetics.113.150615/-/DC1. 1 Present address: Division of Radiation Genetics, Kyoto University, School of Medicine, Yoshida Konoe, Sakyo-ku Kyoto 606-8501 Japan. 2 Corresponding author: Institute for Protein Study, Osaka University, 3-2 Yamadaoka, Suita, Osaka 565-0871, Japan. E-mail: [email protected] together with the 39 finish in the invading strand as the primer. The resulting intermediate is then converted into the singleinvasion intermediate (SEI) and then a DNA structure with double Holiday junctions (dHJ) (Schwacha and Kleckner 1994; Hunter and Kleckner 2001). Resolution of dHJ structures generally benefits in chromosomal crossover (CO) (Allers and Lichten 2001; Hunter and Kleckner 2001). Alternatively, newly synthesized DNA from the invading strand right after the displacement can anneal with ssDNA from the other finish on the DSB, which results in the formation of a noncrossover (NCO) product (through the synthesis-dependent strand-annealing pathway). In meiosis, the recombination occurs preferentially amongst homologous chromosomes as opposed to amongst sister chromatids. Immediately after the resection of DSBs, Replication Protein A (RPA) binds to the ssDNA, followed by the binding of Rad51 (a homolog of bacterial RecA). Rad51 types nucleoprotein filaments on ssDNA, thereby catalyzing the invasion on the ssDNA into a homologous DNA duplex (Ogawa et al.Atorvastatin 1993; Sung 1994).CNTF Protein, Human Rad51 mediates the strand invasion for the duration of each mitosis and meiosis (Shinohara 1992), whereas Dmc1, an additional RecA homolog that also forms filaments on ssDNA, participates only in meiotic recombination (Bishop et al.PMID:23910527 1992). The Rad51- and Dmc1-type filaments are structurally related (Ogawa et al. 1993; Sheridan et al. 2008), but have veryGenetics, Vol. 194, 85972 Augustdifferent in vivo functions. Although, in mitosis, Rad51 is essential for homology search and strand invasion, in meiosis, Rad51 plays an accessary part for Dmc1 (Cloud et al. 2012). The collaborative action of each Rad51 and Dmc1 ensures interhomolog bias for the meiotic recombination. In vivo assembly of a Rad51-ssDNA filament is a highly dynamic approach and is extensively regulated. Many auxiliary proteins [e.g., Rad52 and also the Rad55 ad57 complicated, the.