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Gene Review

RAD51  -  recombinase RAD51

Saccharomyces cerevisiae S288c

Synonyms: DNA repair protein RAD51, YER095W
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Disease relevance of RAD51

  • Dmc1 and Rad51, yeast homologs of the E. coli RecA protein, are shown by immunostaining to localize to as many as 64 sites within spread meiotic nuclei [1].
  • Treatment of mouse 203G glioma cells with 100 nM of RAD51 antisense ODNs significantly enhanced the radiation-induced cell kill compared to control cells, and cells treated with sense or scrambled ODNs [2].
  • Chromosomal integration of LTR-flanked DNA in yeast expressing HIV-1 integrase: down regulation by RAD51 [3].
  • Our data allowed the identification of RAD51 as a novel in vitro IN cofactor able to down regulate the activity of this retroviral enzyme, thereby acting as a potential cellular restriction factor to HIV infection [3].
  • This review focuses on the biochemical and physical characteristics of DNA strand exchange proteins from three diverse organisms: RecA protein from E. coli, UvsX protein from Bacteriophage T4, and RAD51 protein from Saccharomyces cerevisiae [4].

High impact information on RAD51

  • In budding yeasts, there are two BIR pathways, one dependent on the Rad51 recombinase protein and one Rad51 independent; these two repair processes lead to different types of survivors in cells lacking the telomerase enzyme that is required for normal telomere maintenance [5].
  • We observed chromosome fusions only in mutant strains expressing Rad51 and Rad55 or when Tel1 was inactivated [6].
  • Meiotic recombination requires the meiosis-specific RecA homolog Dmc1 as well as the mitotic RecA homolog Rad51 [7].
  • Saccharomyces cerevisiae Mer3 helicase stimulates 3'-5' heteroduplex extension by Rad51; implications for crossover control in meiotic recombination [8].
  • Overexpressing SRS2 nearly eliminates crossovers, whereas overexpression of RAD51 in srs2Delta cells almost completely eliminates the noncrossover recombination pathway [9].

Chemical compound and disease context of RAD51


Biological context of RAD51


Anatomical context of RAD51


Associations of RAD51 with chemical compounds

  • Transcription of uvsC is induced by methyl-methane sulphonate (MMS), as is transcription of RAD51 of yeast [20].
  • The mus-25 mutant is epistatic to the mei-3 mutant for MMS sensitivity. mei-3, which is a homololog of the Saccharomyces cerevisiae gene RAD51, is a member of the uvs-6 epistasis group which contains several genes that are homologous to recombination repair genes in other organisms [21].
  • We show here that the Rad54 protein interacts with the Rad51 protein in vivo and in vitro and that the NH2-terminal 115 residues of the Rad54 protein are necessary for this interaction [22].
  • Saccharomyces cerevisiae lacking Snm1, Rev3 or Rad51 have a normal S-phase but arrest permanently in G2 after cisplatin treatment [23].
  • The pairing and strand exchange reaction requires adenosine triphosphate, a result consistent with the presence of a DNA-dependent adenosine triphosphatase activity in RAD51 protein [24].

Physical interactions of RAD51

  • Dmc1 and, by inference, Rad51 form complexes before synapsis as monitored by immunostaining for Zip1 protein [1].
  • Yeast Rad52 protein interacts with Rad51 protein, binds single-stranded DNA and stimulates annealing of complementary single-stranded DNA [25].
  • Rad55 was shown to interact with Rad51 and Rad57 but not with itself [26].
  • These results suggest that the presence of the Rad51 homologous recombination complex in a top3 background facilitates creation of detrimental intermediates by Sgs1 [27].
  • Accordingly, N-terminal truncation mutants of Rdh54 that fail to bind Rad51 are also impaired for functional interactions with the latter [28].
  • Hed1 binds Rad51 with high affinity and specificity [29].

Enzymatic interactions of RAD51


Regulatory relationships of RAD51

  • Rad52 protein stimulates DNA strand exchange by Rad51 and replication protein A [25].
  • Yeast Rad55 and Rad57 proteins form a heterodimer that functions with replication protein A to promote DNA strand exchange by Rad51 recombinase [32].
  • A class of rad51 alleles was isolated that suppresses the requirement for RAD55 and RAD57 in DNA repair, but not the other accessory factors [33].
  • Furthermore, an increased dosage of SWI6 enhanced the transcript level of the RAD51 gene and also the recombination frequency in meiosis [34].
  • Rad54 protein stimulated Rad51/Rpa-mediated DNA strand exchange by specifically increasing the kinetics of joint molecule formation [35].

Other interactions of RAD51

  • Dmc1 and Rad51 colocalize and are therefore likely to act together during recombination [1].
  • The DNA-binding properties of hRad52 indicate that Rad52 is involved in an early stage of Rad51-mediated recombination [12].
  • One mutation identified, rad59, reduced recombination 1200-fold in the presence of a rad51 mutation, but only 4- to 5-fold in a wild-type background [36].
  • In yeast, mutations of this type include rad50S, dmc1, rad51, and zip1 [13].
  • Lack of Rad54p does not significantly impair Rad51p recruitment to MAT or its initial association with HML alpha; however, Rad54p is required at or before the initiation of DNA synthesis after synapsis has occurred at the 3' end of the invading strand [30].

Analytical, diagnostic and therapeutic context of RAD51

  • RAD51 relieved the negative dominance of each of these alleles either by competitive titration or functional activation of mutant or heterologous Rad52 proteins [37].
  • First, flow cytometric measurements of DNA content and immunofluorescence were used to determine the phase-specific levels of RAD51 and RAD52 protein expression in irradiated and control populations [38].
  • To further substantiate this, we measured the induction of the DNA repair gene RAD51 in RAD51-LACZ fusion strains using the dsb repair and recombination deficient mutant rad52 and the corresponding wild type, and we determined the formation of dsb by pulsed-field gel electrophoresis [39].
  • Molecular dissection of interactions between Rad51 and members of the recombination-repair group [40].
  • Using site-directed mutagenesis of highly conserved residues of human Rad51 (hRad51) and gene targeting of the RAD51 locus in chicken DT40 cells, we examined the importance of Rad51's highly conserved ATP-binding domain [41].


  1. RecA homologs Dmc1 and Rad51 interact to form multiple nuclear complexes prior to meiotic chromosome synapsis. Bishop, D.K. Cell (1994) [Pubmed]
  2. In vitro and in vivo potentiation of radiosensitivity of malignant gliomas by antisense inhibition of the RAD51 gene. Ohnishi, T., Taki, T., Hiraga, S., Arita, N., Morita, T. Biochem. Biophys. Res. Commun. (1998) [Pubmed]
  3. Chromosomal integration of LTR-flanked DNA in yeast expressing HIV-1 integrase: down regulation by RAD51. Desfarges, S., San Filippo, J., Fournier, M., Calmels, C., Caumont-Sarcos, A., Litvak, S., Sung, P., Parissi, V. Nucleic Acids Res. (2006) [Pubmed]
  4. DNA strand exchange proteins: a biochemical and physical comparison. Bianco, P.R., Tracy, R.B., Kowalczykowski, S.C. Front. Biosci. (1998) [Pubmed]
  5. Break-induced replication and recombinational telomere elongation in yeast. McEachern, M.J., Haber, J.E. Annu. Rev. Biochem. (2006) [Pubmed]
  6. Recombination and the Tel1 and Mec1 checkpoints differentially effect genome rearrangements driven by telomere dysfunction in yeast. Pennaneach, V., Kolodner, R.D. Nat. Genet. (2004) [Pubmed]
  7. A protein complex containing Mei5 and Sae3 promotes the assembly of the meiosis-specific RecA homolog Dmc1. Hayase, A., Takagi, M., Miyazaki, T., Oshiumi, H., Shinohara, M., Shinohara, A. Cell (2004) [Pubmed]
  8. Saccharomyces cerevisiae Mer3 helicase stimulates 3'-5' heteroduplex extension by Rad51; implications for crossover control in meiotic recombination. Mazina, O.M., Mazin, A.V., Nakagawa, T., Kolodner, R.D., Kowalczykowski, S.C. Cell (2004) [Pubmed]
  9. Srs2 and Sgs1-Top3 suppress crossovers during double-strand break repair in yeast. Ira, G., Malkova, A., Liberi, G., Foiani, M., Haber, J.E. Cell (2003) [Pubmed]
  10. Chromosome instability and defective recombinational repair in knockout mutants of the five Rad51 paralogs. Takata, M., Sasaki, M.S., Tachiiri, S., Fukushima, T., Sonoda, E., Schild, D., Thompson, L.H., Takeda, S. Mol. Cell. Biol. (2001) [Pubmed]
  11. Rad51 protein involved in repair and recombination in S. cerevisiae is a RecA-like protein. Shinohara, A., Ogawa, H., Ogawa, T. Cell (1992) [Pubmed]
  12. Synergistic actions of Rad51 and Rad52 in recombination and DNA repair. Benson, F.E., Baumann, P., West, S.C. Nature (1998) [Pubmed]
  13. Meiotic cells monitor the status of the interhomolog recombination complex. Xu, L., Weiner, B.M., Kleckner, N. Genes Dev. (1997) [Pubmed]
  14. Brca2 is involved in meiosis in Arabidopsis thaliana as suggested by its interaction with Dmc1. Siaud, N., Dray, E., Gy, I., Gérard, E., Takvorian, N., Doutriaux, M.P. EMBO J. (2004) [Pubmed]
  15. The human and mouse homologs of the yeast RAD52 gene: cDNA cloning, sequence analysis, assignment to human chromosome 12p12.2-p13, and mRNA expression in mouse tissues. Shen, Z., Denison, K., Lobb, R., Gatewood, J.M., Chen, D.J. Genomics (1995) [Pubmed]
  16. Yeast cell-free system that catalyses joint-molecule formation in a Rad51p- and Rad52p-dependent fashion. Nagaraj, V., Norris, D. Biochem. J. (2000) [Pubmed]
  17. RAD51 and DMC1 form mixed complexes associated with mouse meiotic chromosome cores and synaptonemal complexes. Tarsounas, M., Morita, T., Pearlman, R.E., Moens, P.B. J. Cell Biol. (1999) [Pubmed]
  18. Rad51-deficient vertebrate cells accumulate chromosomal breaks prior to cell death. Sonoda, E., Sasaki, M.S., Buerstedde, J.M., Bezzubova, O., Shinohara, A., Ogawa, H., Takata, M., Yamaguchi-Iwai, Y., Takeda, S. EMBO J. (1998) [Pubmed]
  19. RAD51 homologues in Xenopus laevis: two distinct genes are highly expressed in ovary and testis. Maeshima, K., Morimatsu, K., Shinohara, A., Horii, T. Gene (1995) [Pubmed]
  20. Cloning, sequencing, disruption and phenotypic analysis of uvsC, an Aspergillus nidulans homologue of yeast RAD51. van Heemst, D., Swart, K., Holub, E.F., van Dijk, R., Offenberg, H.H., Goosen, T., van den Broek, H.W., Heyting, C. Mol. Gen. Genet. (1997) [Pubmed]
  21. Characterization of the Neurospora crassa mus-25 mutant: the gene encodes a protein which is homologous to the Saccharomyces cerevisiae Rad54 protein. Handa, N., Noguchi, Y., Sakuraba, Y., Ballario, P., Macino, G., Fujimoto, N., Ishii, C., Inoue, H. Mol. Gen. Genet. (2000) [Pubmed]
  22. Direct association between the yeast Rad51 and Rad54 recombination proteins. Jiang, H., Xie, Y., Houston, P., Stemke-Hale, K., Mortensen, U.H., Rothstein, R., Kodadek, T. J. Biol. Chem. (1996) [Pubmed]
  23. Saccharomyces cerevisiae lacking Snm1, Rev3 or Rad51 have a normal S-phase but arrest permanently in G2 after cisplatin treatment. Grossmann, K.F., Ward, A.M., Moses, R.E. Mutat. Res. (2000) [Pubmed]
  24. Catalysis of ATP-dependent homologous DNA pairing and strand exchange by yeast RAD51 protein. Sung, P. Science (1994) [Pubmed]
  25. Rad52 protein stimulates DNA strand exchange by Rad51 and replication protein A. New, J.H., Sugiyama, T., Zaitseva, E., Kowalczykowski, S.C. Nature (1998) [Pubmed]
  26. Functional differences and interactions among the putative RecA homologs Rad51, Rad55, and Rad57. Johnson, R.D., Symington, L.S. Mol. Cell. Biol. (1995) [Pubmed]
  27. Mutations in homologous recombination genes rescue top3 slow growth in Saccharomyces cerevisiae. Shor, E., Gangloff, S., Wagner, M., Weinstein, J., Price, G., Rothstein, R. Genetics (2002) [Pubmed]
  28. Yeast recombination factor Rdh54 functionally interacts with the Rad51 recombinase and catalyzes Rad51 removal from DNA. Chi, P., Kwon, Y., Seong, C., Epshtein, A., Lam, I., Sung, P., Klein, H.L. J. Biol. Chem. (2006) [Pubmed]
  29. Hed1 regulates Rad51-mediated recombination via a novel mechanism. Busygina, V., Sehorn, M.G., Shi, I.Y., Tsubouchi, H., Roeder, G.S., Sung, P. Genes Dev. (2008) [Pubmed]
  30. In vivo roles of Rad52, Rad54, and Rad55 proteins in Rad51-mediated recombination. Sugawara, N., Wang, X., Haber, J.E. Mol. Cell (2003) [Pubmed]
  31. Spontaneous and double-strand break-induced recombination, and gene conversion tract lengths, are differentially affected by overexpression of wild-type or ATPase-defective yeast Rad54. Kim, P.M., Paffett, K.S., Solinger, J.A., Heyer, W.D., Nickoloff, J.A. Nucleic Acids Res. (2002) [Pubmed]
  32. Yeast Rad55 and Rad57 proteins form a heterodimer that functions with replication protein A to promote DNA strand exchange by Rad51 recombinase. Sung, P. Genes Dev. (1997) [Pubmed]
  33. Mutations in yeast Rad51 that partially bypass the requirement for Rad55 and Rad57 in DNA repair by increasing the stability of Rad51-DNA complexes. Fortin, G.S., Symington, L.S. EMBO J. (2002) [Pubmed]
  34. Meiotic role of SWI6 in Saccharomyces cerevisiae. Leem, S.H., Chung, C.N., Sunwoo, Y., Araki, H. Nucleic Acids Res. (1998) [Pubmed]
  35. Rad54 protein stimulates heteroduplex DNA formation in the synaptic phase of DNA strand exchange via specific interactions with the presynaptic Rad51 nucleoprotein filament. Solinger, J.A., Lutz, G., Sugiyama, T., Kowalczykowski, S.C., Heyer, W.D. J. Mol. Biol. (2001) [Pubmed]
  36. A Rad52 homolog is required for RAD51-independent mitotic recombination in Saccharomyces cerevisiae. Bai, Y., Symington, L.S. Genes Dev. (1996) [Pubmed]
  37. Dominant negative alleles of RAD52 reveal a DNA repair/recombination complex including Rad51 and Rad52. Milne, G.T., Weaver, D.T. Genes Dev. (1993) [Pubmed]
  38. Cell cycle-dependent protein expression of mammalian homologs of yeast DNA double-strand break repair genes Rad51 and Rad52. Chen, F., Nastasi, A., Shen, Z., Brenneman, M., Crissman, H., Chen, D.J. Mutat. Res. (1997) [Pubmed]
  39. DNA photodamage, repair, gene induction and genotoxicity following exposures to 254 nm UV and 8-methoxypsoralen plus UVA in a eukaryotic cell system. Averbeck, D., Averbeck, S. Photochem. Photobiol. (1998) [Pubmed]
  40. Molecular dissection of interactions between Rad51 and members of the recombination-repair group. Krejci, L., Damborsky, J., Thomsen, B., Duno, M., Bendixen, C. Mol. Cell. Biol. (2001) [Pubmed]
  41. The essential functions of human Rad51 are independent of ATP hydrolysis. Morrison, C., Shinohara, A., Sonoda, E., Yamaguchi-Iwai, Y., Takata, M., Weichselbaum, R.R., Takeda, S. Mol. Cell. Biol. (1999) [Pubmed]
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