32. Johnson-Schlitz DM, Flores C, Engels WR (2007) Multiple-pathway analysis of double-strand break repair mutations in Drosophila. PLoS Genet 3: e50. doi:10.1371/journal.pgen.0030050. 33. McKim KS, Hayashi-Hagihara A (1998) mei-W68 in Drosophila mela- nogaster encodes a Spo11 homolog: evidence that the mechanism for initiating meiotic recombination is conserved. Genes Dev 12: 2932–2942. 34. Neuman-Silberberg FS, Schupbach T (1993) The Drosophila dorsoventral patterning gene gurken produces a dorsally localized RNA and encodes a TGF alpha-like protein. Cell 75: 165–174. 35. Abdu U, Brodsky M, Schupbach T (2002) Activation of a meiotic checkpoint during Drosophila oogenesis regulates the translation of Gurken through Chk2/Mnk. Curr Biol 12: 1645–1651. 36. Abdu U, Klovstad M, Butin-Israeli V, Bakhrat A, Schupbach T (2007) An essential role for Drosophila hus1 in somatic and meiotic DNA damage responses. J Cell Sci 120: 1042–1049. 37. Liu H, Jang JK, Kato N, McKim KS (2002) mei-P22 encodes a chromosome- associated protein required for the initiation of meiotic recombination in Drosophila melanogaster. Genetics 162: 245–258. 38. Silva E, Tiong S, Pedersen M, Homola E, Royou A, et al. (2004) ATM is required for telomere maintenance and chromosome stability during Drosophila development. Curr Biol 14: 1341–1347. 39. Abdu U, Gonzalez-Reyes A, Ghabrial A, Schupbach T (2003) The Drosophila spn-D gene encodes a RAD51C-like protein that is required exclusively during meiosis. Genetics 165: 197–204. 40. Ghabrial A, Ray RP, Schupbach T (1998) okra and spindle-B encode components of the RAD52 DNA repair pathway and affect meiosis and patterning in Drosophila oogenesis. Genes Dev 12: 2711–2723. 41. McCaffrey R, St Johnston D, Gonzalez-Reyes A (2006) Drosophila mus301/ spindle-C encodes a helicase with an essential role in double-strand DNA break repair and meiotic progression. Genetics 174: 1273–1285. 42. Mehrotra S, McKim KS (2006) Temporal analysis of meiotic DNA double- strand break formation and repair in Drosophila females. PLoS Genet 2: e200. doi:10.1371/journal.pgen.0020200. 43. Harrison JC, Haber JE (2006) Surviving the breakup: the DNA damage checkpoint. Annu Rev Genet 40: 209–235. 44. Brodsky MH, Sekelsky JJ, Tsang G, Hawley RS, Rubin GM (2000) mus304 encodes a novel DNA damage checkpoint protein required during Drosophila development. Genes Dev 14: 666–678. 45. Brodsky MH, Nordstrom W, Tsang G, Kwan E, Rubin GM, et al. (2000) Drosophila p53 binds a damage response element at the reaper locus. Cell 101: 103–113. 46. Brodsky MH, Weinert BT, Tsang G, Rong YS, McGinnis NM, et al. (2004) Drosophila melanogaster MNK/Chk2 and p53 regulate multiple DNA repair and apoptotic pathways following DNA damage. Mol Cell Biol 24: 1219–1231. Dm brca2 in DNA Repair and Checkpoints 47. Bi X, Gong M, Srikanta D, Rong YS (2005) Drosophila ATM and Mre11 are essential for the G2/M checkpoint induced by low-dose irradiation. Genetics 171: 845–847. 48. Jaklevic B, Uyetake L, Lemstra W, Chang J, Leary W, et al. (2006) Contribution of growth and cell cycle checkpoints to radiation survival in Drosophila. Genetics 174: 1963–1972. 49. Weiss RS, Leder P, Vaziri C (2003) Critical role for mouse Hus1 in an S- phase DNA damage cell cycle checkpoint. Mol Cell Biol 23: 791–803. 50. Ludwig T, Chapman DL, Papaioannou VE, Efstratiadis A (1997) Targeted mutations of breast cancer susceptibility gene homologs in mice: lethal phenotypes of Brca1, Brca2, Brca1/Brca2, Brca1/p53, and Brca2/p53 nullizygous embryos. Genes Dev 11: 1226–1241. 51. Brough R, Wei D, Leulier S, Lord CJ, Rong YS, et al. (2008) Functional analysis of Drosophila melanogaster BRCA2 in DNA repair. DNA Repair (Amst) 7: 10–19. 52. Brenneman MA, Wagener BM, Miller CA, Allen C, Nickoloff JA (2002) XRCC3 controls the ﬁdelity of homologous recombination: roles for XRCC3 in late stages of recombination. Mol Cell 10: 387–395. 53. Chen Y, Pane A, Schupbach T (2007) Cutoff and aubergine mutations result in retrotransposon upregulation and checkpoint activation in Drosophila. Curr Biol 17: 637–642. 54. Ellison V, Stillman B (2003) Biochemical characterization of DNA damage checkpoint complexes: clamp loader and clamp complexes with speciﬁcity for 59 recessed DNA. PLoS Biol 1: e33. doi:10.1371/journal.pbio.0030033 55. Bermudez VP, Lindsey-Boltz LA, Cesare AJ, Maniwa Y, Grifﬁth JD, et al. (2003) Loading of the human 9-1-1 checkpoint complex onto DNA by the checkpoint clamp loader hRad17-replication factor C complex in vitro. Proc Natl Acad Sci U S A 100: 1633–1638. 56. Delacroix S, Wagner JM, Kobayashi M, Yamamoto K, Karnitz LM (2007) The Rad9-Hus1-Rad1 (9-1-1) clamp activates checkpoint signaling via TopBP1. Genes Dev 21: 1472–1477. 57. Lee J, Kumagai A, Dunphy WG (2007) The Rad9-Hus1-Rad1 Checkpoint Clamp Regulates Interaction of TopBP1 with ATR. J Biol Chem 282: 28036– 28044. 58. Weiss RS, Matsuoka S, Elledge SJ, Leder P (2002) Hus1 acts upstream of chk1 in a mammalian DNA damage response pathway. Curr Biol 12: 73– 77. 59. Zou L, Cortez D, Elledge SJ (2002) Regulation of ATR substrate selection by Rad17-dependent loading of Rad9 complexes onto chromatin. Genes Dev 16: 198–208. 60. Rubin GM, Spradling AC (1982) Genetic transformation of Drosophila with transposable element vectors. Science 218: 348–353. 61. Van Doren M, Broihier HT, Moore LA, Lehmann R (1998) HMG-CoA reductase guides migrating primordial germ cells. Nature 396: 466–469. PLoS Genetics | www.plosgenetics.org 0012 February 2008 | Volume 4 | Issue 2 | e31