Recombination at work for meiosis

Current Opinion in Genetics and Development

Volumn 8, Issue 2, 1998, Pages 200-211

  Smith, Kathleen N ^a   Nicolas, Alain ^a  

^a INSTITUT CURIE   (France)

Author keywords

[No Author keywords available]

Indexed keywords

ARTICLE; BIODIVERSITY; CELL CYCLE; CHROMOSOME SEGREGATION; GENETIC RECOMBINATION; MEIOSIS; NONHUMAN; PRIORITY JOURNAL; REPRODUCTION;

EID: 0032055005     PISSN: 0959437X     EISSN: None     Source Type: Journal    
DOI: 10.1016/S0959-437X(98)80142-1     Document Type: Article

References (105)

1. Kleckner N. Meiosis: how could it work? Proc Natl Acad Sci USA. 93:1996;8167-8174.
2. Roeder GS. Meiotic chromosomes: it takes two to tango. Genes Dev. 11:1997;2600-2621.
3. Heyting C. Synaptonemal complex: structure and function. Curr Opin Cell Biol. 8:1996;389-396.
4. Bascom-Slack CA, Ross LO, Dawson DS. Chiasmata, crossovers, and meiotic chromosome segregation. Adv Genet. 35:1997;253-284.
5. Szostak JW, Orr-Weaver TL, Rothstein RJ, Stahl FW. The double-strand-break repair model for recombination. Cell. 33:1983;25-35.
6. Stahl F. Meiotic recombination in yeast: coronation of the double-strand break repair model. Cell. 87:1996;965-968.
7. Lichten M, Goldman ASH. Meiotic recombination hotspots. Annu Rev Genet. 29:1995;423-444.
8. Baudat F, Nicolas A. Clustering of meiotic double-strand breaks on yeast chromosome III. of outstanding interest Proc Natl Acad Sci USA. 94:1997;5213-5218 This work provides a map of sites of DSB formation across the 340kb of Chromosome III and demonstrates that they occur almost exclusively in promoter regions. The cumulative frequency of DSBs for Chromosome III suffices for at least one reciprocal exchange per meiosis, thereby ensuring disjunction.
9. Bullard SA, Kim S, Galbraith AM, Malone Re. double-strand breaks at the HIS2 recombination hotspot in Saccharomyces cerevisiae. of special interest Proc Natl Acad Sci USA. 93:1996;13054-13059 Analysis of DSB formation at HIS2 shows that strong breaks occur downstream of HIS2 (in the promoter region of the next open reading frame) and, more unexpectedly, within the HIS2 coding region itself. Surprisingly, a deletion that increases DSB formation can act in trans to raise the level of DSBs on an otherwise recombinationally 'colder' allele.
10. de Massy B, Rocco V, Nicolas A. The nucleotide mapping of DNA double-strand breaks at the CYS3 initiation site of meiotic recombination in Saccharomyces cerevisiae. EMBO J. 14:1995;4589-4598.
11. Liu J, Wu TC, Lichten M. The location and structure of double-strand breaks during yeast meiosis: evidence for a covalently linked DNA-protein intermediate. EMBO J. 14:1995;4599-4608.
12. Xu L, Kieckner N. Sequence non-specific double-strand breaks and interhomolog interactions prior to double-strand break formation at a meiotic recombination hot spot in yeast. EMBO J. 14:1995;5115-5128.
13. Xu F, Petes TD. Fine structure mapping of meiosis-specific double-strand DNA breaks at a recombination hotspot associated with an insertion of telomeric sequences upstream of the HIS4 locus in yeast. Genetics. 143:1996;1115-1125.
14. Klein S, Zenvirth D, Dror V, Barton AB, Kaback DB, Simchen G. Patterns of meiotic double-strand breaks on native and artificial yeast chromosomes. Chromosoma. 105:1996;276-284.
15. Klein S, Zenvirth D, Sherman A, Ried K, Rappold G, Simchen G. Double-strand breaks on YACs during yeast meiosis may reflect meiotic recombination in the human genome. of special interest Nat Genet. 13:1996;481-484 Yeast artificial chromosomes carrying human sequences from various loci were examined for DSB formation in yeast meiotic cells. The extent of breakage correlates well with the degree of meiotic recombination measured at these loci in humans, suggesting that DNA with a given sequence content adopts a similar chromatin configuration in both yeast and humans.
16. Nicolas A. Relationship between transcription and initiation of meiotic recombination: toward chromatin accessibility. Proc Natl Acad Sci USA. 95:1998;87-89.
17. Fan Q-Q, Petes TD. Relationship between nuclease-hypersensitive sites and meiotic recombination hot spot activity at the HIS4 locus of Saccharomyces cerevisiae. Mol Cell Biol. 16:1996;2037-2043.
18. Keeney S, Kleckner N. Communication between homologous chromosomes: genetic alterations at a nuclease-hypersensitive site can alter mitotic chromatin structure at that site both in cis and in trans. Genes Cells. 1:1996;475-489.
19. Fox ME, Smith GR. Control of meiotic recombination in Schizosaccharomyces pombe. Proc Nucleic Acid Res Mol Biol. 1998;. in press.
20. Mizuno K, Emura Y, Baur M, Kohli J, Ohta K, Shibata T. The meiotic recombination hot spot created by the single-base substitution ade6-M26 results in remodeling of chromatin structure in fission yeast. of outstanding interest Genes Dev. 11:1997;876-886 Meiotic chromatin at the wild-type ADE6 locus of S. pombe was compared to chromatin at this locus from strains homozygous for the M26 hot spot mutation. In M26 strains, a new MNase-hypersensitive site was detected at the position of the M26 mutation and meiotic induction of MNase hypersensitivity in the ade6 promoter was enhanced.
21. Kon N, Krawchuk MD, Warren BG, Smith GR, Wayne WP. Transcription factor Mts1/Mts2 (Atf1/Pcr1, Gad7/Pcr1) activates the M26 meiotic recombination hotspot in Schizosaccharomyces pombe. of outstanding interest Proc Natl Acad Sci USA. 94:1997;13765-13770 The subunits of the heterodimeric protein Mts1/Mts2, which binds to M26 are identical to the transcription factors Atf1 (Gad7) and Pcr1, respectively. Mutations in the MTS1 or MTS2 genes abolish hotspot activity at M26. Hypotheses concerning the mechanism of initiation or stimulation of recombination by the M26 heptamer at ADE6 and elsewhere in the S. pombe genome are presented.
22. Ohta KA, Nicolas M, Furuse A, Nabetani H, Ogawa H, Shibata T. Mutations in the MRE11, RAD50, XRS2 and MRE2 genes alter chromatin configuration at meiotic DNA double-stranded break sites in premeiotic and meiotic cells. Proc Natl Acad Sci USA. 95:1998;646-651.
23. Wu T-C, Lichten M. Factors that affect the location and frequency of meiosis-induced double-strand breaks in Saccharomyces cerevisiae. Genetics. 140:1995;55-66.
24. Fan Q-Q, Xu F, White MA, Petes TD. Competition between adjacent meiotic recombination hotspots in the yeast Saccharomyces cerevisiae. Genetics. 145:1997;661-670.
25. Rocco V, Nicolas A. Sensing of DNA non-homology lowers the initiation of meiotic recombination in yeast. Genes Cells. 1:1996;645-661.
26. Bascom-Slack Ca, Dawson DS. The yeast motor protein, Kar3p, is essential for meiosis I. of outstanding interest J Cell Biol. 139:1997;459-467 KAR3, which encodes a motor protein, is essential for meiosis, kar3 mutants block in prophase of meiosis I with incomplete SC and exhibit a defect in DSB formation and heteroallelic recombination. KAR3 may function in meiotic prophase to mediate homologous chromosome interactions either before or during DSB formation.
27. Nag DK, Kurst A. A 140-bp-long palindromic sequence induces double-strand breaks during meiosis in the yeast Saccharomyces cerevisiae. of special interest Genetics. 146:1997;835-847 A long palindromic sequence inserted in the yeast genome acts as a recombinational hot spot and significantly biases the direction of gene conversion. Palindromes of sufficient size may form cruciforms that are recognized by structure-specific endonucleases during meiosis, thereby generating the strong DSBs that are observed at sites of insertion.
28. Johzuka K, Ogawa H. Interaction of Mre11 and Rad50: two proteins required for DNA repair and meiosis-specific double-strand break formation in Saccharomyces cerevisiae. Genetics. 139:1995;1521-1532.
29. Nakagawa T, Ogawa H. Involvement of the MRE2 gene of yeast in the formation of meiosis-specific double-strand breaks and crossover recombination through RNA splicing. of outstanding interest Genes Cells. 2:1997;65-79 The MER2 gene product is required for meiotic DSB formation. The meiosis-specific splicing of MER2 pre-mRNA is impaired in a mre2 deletion mutant but DSB induction can be restored by overexpressing MER2 cDNA. The Mre2 protein has two RNA recognition motifs and may be required for the processing of other RNAs, as MER2 cDNA does not fully restore crossover frequencies in the mre2 background.
30. Shinohara A, Ogawa T. Homologous recombination and the roles of double-strand breaks. Trends Biochem Sci. 20:1995;387-391.
31. Keeney S, Kleckner N. Covalent protein-DNA complexes at the 5′ termini of meiosis-specific double-strand breaks in yeast. Proc Natl Acad Sci USA. 92:1995;11274-11278.
32. Keeney S, Giroux C, Kleckner N. Meiosis-specific DNA double-strand breaks are catalysed by Spo11, a member of a widely conserved protein family. of outstanding interest Cell. 88:1997;375-384 Large-scale isolation of a protein-DNA complex from meiotic rad50S diploids cells followed by protein microsequence identified the peptide covalently attached to the ends of DSBs as a fragment of the Spo11 protein.
33. Bergerat A, de Massy B, Gadelle D, Varoutas P-C, Nicolas A, Forterre P. An atypical topoisomerase II from archaea with implications for meiotic recombination. of outstanding interest Nature. 386:1997;414-417 A report of the cloning and sequencing of the genes encoding the A and B subunits of a topoisomerase II from the archaeon Sulfobolus shibatae. The A subunit shares similarities with three other eukaryotic proteins, including the Spo11 protein from S. cerevisiae. Mutagenesis of a single conserved tyrosine (Tyr135) in Spo11 produces a null phenotype for DSB formation and meiotic recombination, suggesting a mechanism of action involving transesterification.
34. Gardiner JM, Bullard SA, Chrome C, Malone RE. Molecular analysis of REC103, an early meiotic recombination gene in yeast. of special interest Genetics. 146:1997;1265-1274 Mutation of REC103, a newly early recombination gene, rescues the spore inviability of rad52 spo13 diploids. REC103 is identical to SK18 - a gene that inhibits the expression of double-stranded RNA viruses in yeast - and blocks translation of non-polyadenylated mRNA's in mitotic cells. In meiosis, the REC103/SK18 gene product might suppress the translation of a non-polyadenylated mRNA(s) that encodes a meiotic inhibitor.
35. Evans DH, Li YF, Fox ME, Smith GR. A WD repeat protein, Rec14, essential for meiotic recombination in Schizosaccharomyces pombe. Genetics. 146:1997;1253-1264.
36. Tavassoli M, Shayeghi M, Nasim A, Watts FZ. Cloning and characterisation of the Schizosaccharomyces pombe rad32 gene: a gene required for repair of double-strand breaks and recombination. Nucleic Acids Res. 23:1995;383-388.
37. Dolganov GM, Maser A, Novikov A, Tosto L, Chong S, Bressan DA, Petrini JHJ. Human Rad50 is physically associated with human Mre11: identification of a conserved multiprotein complex implicated in recombinational DNA repair. Mol Cell Biol. 16:1996;4832-4841.
38. Nairz K, Klein F. mre11S-a yeast mutation that blocks double-strand break processing and permits nonhomologous synapsis in meiosis. of outstanding interest Genes Dev. 11:1997;2272-2290 The authors of this paper describe a new allele of MRE11, termed mre11S (for separation of function), that accumulates unresected DSBs. Partial complementation of the mre11S allele - which has a mutation in the 5′ region of the gene - by mre11 alleles with mutations near the 3′ end (and which cannot initiate DSB formation) indicates that the Mre11 protein has separate domains controlling DSB initiation and resection.
39. Tsubouchi H, Ogawa H. A novel mre11 mutation impairs processing of double-strand breaks of DNA during both mitosis and meiosis. of outstanding interest Mol Cell Biol. 18:1998;260-268 rad58-4 is a new allele of MRE11. The rad58-4 mutation (also termed mre11-58S), which affects a phosphodiesterase motif, allows meiotic DSB formation but not processing. This phenotype is similar to that conferred by the rad50S allele, except that mre11-58S mutants are highly sensitive to methyl methane sulfonate (MMS), indicating that the phosphodiesterase consensus sequence is important for nucleolytic processing of DSB ends during both mitosis and meiosis.
40. Sharples GJ, Leach DRF. Structural and functional similarities between the SbcCD proteins of Escherichia coli and the Rad50 and Mre11 (Rad32) recombination and repair proteins of yeast. Mol Microbiol. 17:1995;1215-1220.
41. Malkova A, Ross L, Dawson D, Hoekstra MF, Haber JE. Meiotic recombination initiated by a double-strand break in rad50Δ yeast cells otherwise unable to initiate meiotic recombination. of outstanding interest Genetics. 143:1996;741-754 In this study, the HO endonuclease was expressed under the control of the meiotic-specific SPO13 promoter to create a DSB at a single site in the yeast genome. In the absence of the Rad50 or Xrs2 proteins, not all DSBs are repaired, and some recombination events occur, indicating that these proteins are important but not essential for the repair of HO-induced DSBs.
42. Prinz S, Amon A, Klein F. Isolation of COM1, a new gene required to complete meiotic double-strand break-induced recombination in Saccharomyces cerevisiae. of outstanding interest Genetics. 146:1997;781-795 A screen to isolate mutants that fail to yield viable spores but can be rescued by the spo11 and spo13 mutations pinpointed a new gene, COM1 (completion of meiotic recombination). The com1 deletion mutant confers a phenotype similar to that of rad50S and mre11S point mutants, the accumulation of unresected DSBs. COM1 and SAE2 [43] are the same gene.
43. McKee AHZ, Kleckner N. A general method for identifying recessive diploid-specific mutations in Saccharomyces cerevisiae, its application to the isolation of mutants blocked at intermediate stages of meiotic prophase and characterization of a new gene SAE2. of outstanding interest Genetics. 146:1997;797-815 A screen to isolate mutants that confer a defect in spore formation which is dependent upon the initiation of recombination yielded three new genes: SAE1, SAE2 and SAE3 (sporulation in the absence of Spo eleven). The phenotypes of the sae2 (same as com1) null mutation are described.
44. Schwacha A, Kleckner N. Identification of double Holliday junctions as intermediates in meiotic recombination. Cell. 83:1995;783-791.
45. Schwacha A, Kleckner N. Interhomolog bias during meiotic recombination: meiotic functions promote a highly differentiated interhomolog-only pathway. of outstanding interest Cell. 90:1997;1123-1135 The authors of this paper address the genetic requirements for inter-sister and inter-homolog JM formation. RED1, DMC1, and RAD51, RAD55, and RAD57 (which act coordinately) all promote inter-homolog interactions. JMs do not form at all in dmc1 strains, which accumulate hyper-resected DSBs and arrest in meiosis but upon return to mitotic growth conditions, inter-sister JMs appear and their formation is dependent on RAD51.
46. Roca AI, Cox MM. RecA protein: structure, function, and role in recombinational DNA repair. Prog Nucleic Acid Res Mol Biol. 56:1997;129-223.
47. Sung P. Yeast Rad55 and Rad57 proteins form a heterodimer that functions with replication protein A to promote DNA strand exchange by Rad51 recombinase. of outstanding interest Genes Dev. 11:1997;1111-1121 RPA is necessary for strand exchange, perhaps by removing secondary structure from single-stranded DNA but it competes with Rad51 for binding DNA. Rad55 and Rad57 have limited RecA homology and are required in vivo for recombination. The two proteins form a heterodimer in yeast cells. In vitro, inclusion of this complex with Rad51 and RPA stimulates the strand exchange reaction, suggesting that the Rad55-Rad57 heterodimer functions as a cofactor to overcome the inhibitory effect of RPA.
48. Sung P. Function of yeast Rad52 protein as a mediator between replication protein A and the Rad51 recombinase. of outstanding interest J Biol Chem. 272:1997;28194-28197 See annotation to [50].
49. Shinohara A, Ogawa T. Stimulation by Rad52 of yeast Rad51-mediated recombination. of outstanding interest Nature. 391:1998;404-407 See annotation to [50].
50. New JH, Sugiyama T, Zaitseva E, Kowalczykowski SC. Rad52 protein stimulates DNA strand exchange by Rad51 and replication protein A. of outstanding interest Nature. 391:1998;407-410 These three studies [48-50] demonstrate that inclusion of the Rad52 protein in the Rad51-promoted strand exchange reaction alleviates inhibition by RPA, suggesting that Rad52 functions as a cofactor for Rad51 activity. They constitute an important milestone in the development of in vitro reconstructions of homologous recombination in eukaryotes.
51. Benson FE, Baumann P, West SC. Synergistic actions of Rad51 and Rad52 in recombination and DNA repair. of outstanding interest Nature. 391:1998;401-404 Here, enzymatic activities comparable to those described for yeast Rad52 are demonstrated for the human Rad52 protein, which stimulates human Rad51-mediated strand exchange. This study underscores the evolutionary conservation of the strand exchange reaction from yeast to man.
52. Stassen NY, Logsdon JM, Vora GJ, Offenberg HH, Palmer JD, Zolan ME. Isolation and characterization of rad51 orthologs from Coprinus cinereus and Lycopersicon esculentum, and phylogenetic analysis of eukaryotic recA homologs. Curr Genet. 31:1997;144-157.
53. Shinohara A, Gasior S, Ogawa T, Kleckner N, Bishop DK. Saccharomyces cerevisiae recA homologues RAD51 and DMC1 have both distinct and overlapping roles in meiotic recombination. of special interest Genes Cells. 2:1997;615-629 Side-by-side comparison of rad51, dmc1, and double mutant phenotypes with respect to DSB processing, formation of recombinants, and progression through meiosis emphasizes that Rad51 and Dmc1 have complementary but not identical functions.
54. Muris DFR, Vreeken K, Carr AM, Murray JM, Smit C, Lohman PH, Pastink A. Isolation of the Schizosaccharomyces pombe RAD54 homolog, rhp54+, a gene involved in the repair of radiation damage and replication fidelity. J Cell Sci. 109:1996;73-81.
55. Kanaar R, Troelstra C, Swagemakers SM, Essers J, Smit B, Franssen JH, Pastink A, Bezzubova OY, Buerstedde JM, Clever B, et al. Human and mouse homologs of the Saccharomyces cerevisiae RAD54 DNA repair gene: evidence for functional conservation. Curr Biol. 6:1996;828-838.
56. Jiang H, Xie Y, Houston P, Stemke-Hale K, Mortensen UH, Rothstein R, Kodadek T. Direct association between the yeast Rad51 and Rad54 recombination proteins. J Biol Chem. 271:1996;33181-33186.
57. Clever B, Interthal H, Schmuckli-Maurer J, King J, Sigrist M, Heyer W-D. Recombinational repair in yeast: functional interactions between Rad51 and Rad54 proteins. EMBO J. 16:1997;2535-2544.
58. Klein H. RDH54, a RAD54 homolog in Saccharomyces cerevisiae, is required for mitotic diploid-specific recombination and repair and meiosis. of outstanding interest Genetics. 147:1997;1533-1543 RDH54, identified as a RAD54 homolog in the course of the yeast genome sequencing project, has a less severe mutant phenotype in mitosis than does the rad54 mutant. This paper demonstrates that RAD54 has important roles in mitotic DNA repair whereas RDH54 specializes in the meiotic repair of DSBs.
59. Shinohara M, Shita-Yamaguchi E, Buerstedde J-M, Shinegawa H, Ogawa H, Shinohara A. Characterization of the roles of the Saccharomyces cerevisiae RAD54 gene and a homologue of RAD54, RDH54/TID1, in mitosis and meiosis. of outstanding interest Genetics. 147:1997;1545-1556 This detailed investigation of the meiotic phenotypes of the rdh54 mutant shows that in the absence of RAD54, RDH54 is essential for the repair of DSBs after they have been processed, and for the formation of recombinants. The synergistic effects exhibited by a double mutant suggest that the two genes may function in different recombinational pathways during meiosis, or that each performs a different function in the same pathway.
60. Dresser ME, Ewing DJ, Conrad MN, Dominguez AM, Barstead R, Jiang H, Kodadek T. DMC1 functions in a Saccharomyces cerevisiae meiotic pathway that is largely independent of the RAD51 pathway. of outstanding interest Genetics. 147:1997;533-544 Analysis of new alleles of DMC1 indicates that nucleotide binding and Dmc1-Dmc1 homotypic interactions are important for function. The failure of Dmc1 to interact directly with Rad51 despite their similar mutant phenotypes and patterns of localization on meiotic chromosomes is attributed to sequences in the amino-terminal region of the protein. Specific interactions of Dmc1 with several other proteins, notably Tid1/Rdh54, are demonstrated.
61. Anderson LK, Offenberg HH, Verkuijlen WMHC, Heyting C. RecA-like proteins are components of early meiotic nodules in lily. of special interest Proc Natl Acad Sci USA. 94:1997;6868-6873 Lily offers many cytological advantages for the study of meiosis. Here, electron microscopic immunogold localization of spreads of zygotene and early pachytene synaptonemal complexes indicate that the Rad51 and/or Lim15 (a Dmc1 homolog) proteins are components of early recombination nodules, which are proposed to mark all sites at which strand exchange occurs.
62. Terasawa M, Shinohara A, Hotta Y, Ogawa H, Ogawa T. Localization of RecA-like recombination proteins on chromosomes of the lily at various meiotic stages. Genes Dev. 9:1995;925-934.
63. Page AW, Orr-Weaver TL. Stopping and starting the meiotic cell cycle. Curr Opin Genet Dev. 7:1997;23-31.
64. Rockmill B, Sym M, Scherthan H, Roeder GS. Roles for two RecA homologs in promoting meiotic chromosome synapsis. Genes Dev. 9:1995;2684-2695.
65. Li Z, Golub EI, Gupta R, Radding CM. Recombination activities of HsDmc1 protein, the meiotic human homolog of RecA protein. Proc Natl Acad Sci USA. 94:1997;11221-11226.
66. McKee AHZ, Kleckner N. Mutations in Saccharomyces cerevisiae that block meiotic prophase chromosome metabolism and confer cell cycle arrest at pachytene identify two new meiosis-specific genes SAE1 and SAE3. of special interest Genetics. 146:1997;817-834 Two new genes required for progression through meiosis are described. Mutants of SAE3 are phenotypically similar to dmc1 mutants, in that they accumulate hyper-resected DSBs, are deficient in the formation of recombinants, and arrest permanently in prophase. Double mutant analysis indicates that DMC1 and SAE3 are required for the same meiotic function. The gene contains an unusually small ORF of 50 codons.
67. Plug AW, Xu J, Reddy G, Golub EI, Ashley T. Presynaptic association of Rad51 protein with selected sites in meiotic chromatin. Proc Natl Acad Sci USA. 93:1996;5920-5924.
68. Keegan KS, Holtzman DA, Plug AW, Christenson Er, Brainerd EE, Flaggs G, Bentley NJ, Taylor EM, Meyn MS, Moss SB, et al. The Atr and Atm protein kinases associate with different sites along meiotically pairing chromosomes. Genes Dev. 10:1996;2423-2437.
69. Plug AW, Peters AFHM, Xu Y, Keegan KS, Hoekstra MF, Baltimore D, de Boer P, Ashley T. ATM and RPA in meiotic chromosome synapsis and recombination. Nat Genet. 17:1997;457-461.
70. Scully R, Chen J, Plug A, Xiao Y, Weaver D, Feunteun J, Ashley T, Livingston DM. Association of BRCA1 with Rad51 in mitotic and meiotic cells. of special interest Cell. 88:1997;265-275 BRCA1 mutations are responsible for most instances of inherited susceptibility to breast and ovarian cancer. The protein encoded by BRCA1 interacts physically with Rad51 and the two colocalize on both mitotic and meiotic chromosomes. These observations highlight novel roles for Rad51 that may have evolved in higher eukaryotes. S. cerevisiae lacks a BRCA1 homolog.
71. Petes TD, Pukkila PJ. Meiotic sister chromatic recombination. Adv Genet. 33:1995;41-62.
72. Mao-Draayer Y, Galbraith AM, Pittman DL, Cool M, Malone RE. Analysis of meiotic recombination pathways in the yeast Saccharomyces cerevisiae. of special interest Genetics. 144:1996;71-86 This work addresses the issue of inter-sister (intrachromosomal) versus interhomolog recombination by a genetic analysis of RED1 and HOP1 function in different strain backgrounds. red1 and hop1 mutations rescue the spore inviability of rad52 spo13 strains - as has been observed for mutations that prevent DSB formation - but not of rad50S spo13 strains. The activity that is defined by the rad50S mutation may be required only in an interchromosomal recombination pathway.
73. Smith AV, Roeder GS. The yeast Red1 protein localizes to the cores of meiotic chromosomes. J Cell Biol. 136:1997;957-967.
74. Hollingsworth NM, Ponte L. Genetic interactions between HOP1, RED1 and MEK1 suggest that MEK1 regulates assembly of axial element components during meiosis in the yeast Saccharomyces cerevisiae. of outstanding interest Genetics. 147:1997;33-42 These three genes, HOP, RED1 and HEK1, promote interchromosomal recombination and crossing over but not intrachromosomal recombination. A genetic analysis of interactions among Hop1, Red1, and Mek1 suggests that the stoichiometry of Hop1 and Red1, which localize to meiotic chromosomes, is important in controlling the early phases of SC development.
75. Zenvirth D, Loidl J, Klein S, Arbel A, Shemesh R, Simchen G. Switching yeast from meiosis to mitosis: double-strand break repair, recombination and synaptonemal complex. of special interest Genes Cells. 2:1997;487-498 After yeast cells enter the meiotic program, they can be induced to resume vegetative growth conditions before they complete sporulation. Recombination can be measured genetically in such cells; this paper presents experiments showing that the repair of meiotically induced DSBs does not require meiosis-specific functions and that DSB formation precedes the commitment to recombination mediated by DMC1.
76. Kolodner RD. Biochemistry and genetics of eukaryotic mismatch repair. Genes Dev. 10:1996;1433-1442.
77. Nicolas A, Petes T. Polarity of meiotic gene conversion in fungi: contrasting views. Experientia. 50:1994;242-252.
78. Alani E, Lee S, Kane MF, Griffith J, Kolodner RD. Saccharomyces cerevisiae MSH2, a mispaired base recognition protein, also recognizes Holliday junctions in DNA. of special interest J Mol Biol. 265:1997;289-301 The Msh2 protein is shown by filter-binding assays and electron microscopy to bind synthetic Holliday junctions, providing strong evidence for mechanistic links between mismatch repair and genetic recombination. The mismatch repair system may regulate heteroduplex tract length by recognition of mispaired bases in recombinational intermediates.
79. Kirkpatrick DT, Petes TD. Repair of DNA loops involves DNA-mismatch and nucleotide-excision repair proteins. of outstanding interest Nature. 387:1997;929-931 In addition to correcting mismatches and small insertions and deletions, Msh2 is required for the processing of a 26 bp insertion loop at HIS4. During meiotic recombination, msh2 mutants exhibit increased frequencies of post-meiotic segregation at HIS4. The unexpected involvement of the nucleotide excision repair system in the repair of large DNA loops is also suggested by aberrant segregation of this allele in the rad1 background.
80. Hunter N, Chambers SR, Louis EJ, Borts RH. The mismatch repair system contributes to meiotic sterility in an interspecific yeast. of outstanding interest EMBO J. 15:1996;1726-1733 Hybrids derived from crosses between S. cerevisiae and S. paradoxus are defective in meiosis. The rare viable spores they produce are highly aneuploid and have undergone low levels of recombination. Mutations in MSH2 or PMS1 alleviate this hybrid sterility. This result extends the observation made in bacteria that the mismatch repair system constitutes part of the genetic barrier between species.
81. West SC. Processing of recombination intermediates by the RuvABC proteins. Annu Rev Genet. 31:1997;213-244.
82. Gilbertson LA, Stahl FW. A test of the double-strand break repair model for meiotic recombination in Saccharomyces cerevisiae. Genetics. 144:1996;27-41.
83. Storlazzi A, Xu L, Cao L, Kleckner N. Crossover and noncrossover recombination during meiosis: timing and pathway relationships. Proc Natl Acad Sci USA. 92:1995;8512-8516.
84. Kaback DB. Chromosome-size dependent control of meiotic recombination in humans. Nat Genet. 13:1996;20-21.
85. Storlazzi A, Xu L, Schwacha A, Kleckner N. Synaptonemal complex (SC) component Zip1 plays a role in meiotic recombination independent of SC polymerization along the chromosomes. Proc Natl Acad Sci USA. 93:1996;9043-9048.
86. Chua PR, Roeder GS. Tam1, a telomere-associated meiotic protein, functions in chromosome synapsis and crossover interference. of special interest Genes Dev. 11:1997;1786-1800 S. cerevisiae can segregate heterologous chromosomes during meiosis by an alternative system termed distributive disjunction. The Tam1/Ndj1 protein is required for segregation of both homologous and heterologous chromosomes, and localizes to telomeres in meiotic cells. Mutants are recombinationally competent but exhibit reduced interference. The tam1/ndj1 mutation is the first to separate crossover frequency from crossover distribution.
87. Conrad MN, Dominguez AM, Dresser ME. Ndj1, a meiotic telomere protein required for normal chromosome synapsis and segregation in yeast. of special interest Science. 276:1997;1252-1255 Here, the Tam1/Ndj1 protein is shown to accumulate at telomeres and to promote distributive disjunction of linear but not circular minichromosomes, as well as to function in the segregation of homologous chromosomes. Tam1/Ndj1 may stabilize homology-dependent interactions and, in its absence, telomeres may interfere with the distributive pathway.
88. Shimanuki M, Miki F, Ding D-Q, Chikashige Y, Hiraoka Y, Horio T, Niwa O. A novel fission yeast gene, kms1+, is required for the formation of meiotic prophase-specific nuclear architecture. of special interest Mol Gen Genet. 254:1997;238-249 In fission yeast, meiotic cells undergo nuclear reorganization, which includes the clustering of telomeres near the spindle pole bodies. The kms1 mutant interferes with telomere localization, meiotic recombination, and chromosome segregation - indicating that the spatial organization of chromosomes in the nucleus is necessary for efficient pairing and recombination between homologs.
89. Hollingsworth NM, Ponte L, Halsey C. MSH5, a novel MutS homolog, facilitates meiotic reciprocal recombination between homologs in Saccharomyces cerevisiae but not mismatch repair. Genes Dev. 9:1995;1728-1739.
90. Pochart P, Woltering D, Hollingsworth NM. Conserved properties between functionally distinct MutS homologs in yeast. of outstanding interest J Biol Chem. 272:1997;30345-30349 Physical interactions between the Msh4 and Msh5 proteins are demonstrated here. Msh4 and Msh5 are not required for mismatch repair in mitotic and meiotic cells but instead influence the degree of homologous exchange during recombination, perhaps by binding to Holliday junctions.
91. Hunter N, Borts RH. Mlh1 is unique among mismatch repair proteins in its ability to promote crossing-over during meiosis. of special interest Genes Dev. 11:1997;1573-1582 Mutants in the MutL homolog MLH1 are defective in crossing-over as well as in repairing mismatches generated during recombination. Double mutant studies indicate that MLH1 acts in the same pathway as MSH4 and MSH5 - two MutS homologs which have no role in mismatch repair, with respect to promoting crossovers.
92. Hassold TJ. Mismatch repair goes meiotic. Nat Genet. 13:1996;261-262.
93. Baker SM, Bronner CE, Zhang L, Plug AW, Robatzek M, Warren G, Elliott EA, Yu J, Ashley T, Arnheim N, et al. Male mice defective in the DNA mismatch repair gene PMS2 exhibit abnormal chromosome synapsis in meiosis. Cell. 82:1995;309-319.
94. Baker SM, Plug AW, Prolla TA, Bronner CE, Harris AC, Yao X, Christie DM, Monell C, Arnheim N, Bradley A, et al. Involvement of mouse Mlh1 in DNA mismatch repair and meiotic crossing over. of outstanding interest Nat Genet. 13:1996;336-342 Mice deficient in the mismatch repair protein Mlh1 exhibit both somatic (microsatellite instability) and meiotic defects, including a reduction in chiasmata. These defects may explain the sterility of male and female mlh1 mice. The protein localizes to sites of crossing over on meiotic chromosomes.
95. Edelmann W, Cohen PE, Kane M, Lau K, Morrow B, Bennett S, Umar A, Kunkel T, Cattoretti G, Chaganti R, et al. Meiotic pachytene arrest in MLH1-deficient mice. of outstanding interest Cell. 85:1996;1125-1134 This paper also describes germline defects in male and female MLH1-deficient mice (see [94]). Comparison of the phenotypes of mice lacking the Mlh1 protein with mice deficient in other mismatch repair-related proteins, Msh2 and Pms2, demonstrates that the three proteins have independent functions in meiosis, despite their apparently related roles in mitotic DNA repair.
96. Hassold T, Abruzzo M, Adkins K, Griffin D, Merrill M, Millie E, Saker D, Shen J, Zaragoza M. Human aneuploidy: incidence, origin, and etiology. Environ Mol Mutagen. 28:1996;167-175.
97. Lamb NE, Freeman SB, Savage-Austin A, Pettay D, Taft L, Hersey J, Gu Y, Shen J, Saker D, May KM, et al. Susceptible chiasmate configurations of chromosome 21 predispose to non-disjunction in both maternal meiosis I and meiosis II. of outstanding interest Nat Genet. 14:1996;400-405 Analysis of 133 cases of trisomy 21 reveals that maternal Meiosis II nondisjunction is associated with increased proximal recombination, leading to an expansion of the genetic map for chromosome 21. This result is interpreted to indicate that all maternal non-disjunction, whether Meiosis I or (apparent) Meiosis II nondisjunction, results from events occurring in Meiosis I.
98. Ross LO, Maxfield R, Dawson D. Exchanges are not equally able to enhance meiotic chromosome segregation in yeast. of special interest Proc Natl Acad Sci USA. 93:1996;4979-4983 Using linear yeast artificial chromosomes to study the relationship between position of a crossover and its ability to promote disjunction, the authors find that exchanges in telomeric regions are correlated with higher levels of Meiosis I nondisjunction.
99. Orr-Weaver TL. Meiosis in Drosophila: seeing is believing. Proc Natl Acad Sci USA. 92:1995;10443-10449.
100. Koehler KE, Boulton CL, Collins HE, French RL, Herman KC, Lacefield SM, Madden LD, Schuetz CD, Hawley RS. Spontaneous X chromosome MI and MII nondisjunction events in Drosophila melanogaster oocytes have different recombinational histories. of outstanding interest Nat Genet. 14:1996;406-414 103 cases of spontaneous X chromosomal non-disjunction in Drosophila oocytes were analyzed for recombination events. Of these, 97 resulted from Meiosis I non-disjunction without exchange, or more rarely, with distal exchanges insufficient for proper chromosome segregation. In contrast, all six of the Meiosis II non-disjunction events were associated with proximal exchanges.
101. Orr-Weaver TL. Meiotic nondisjunction does the two-step. Nat Genet. 14:1996;374-375.
102. Lydall D, Nikosky Y, Bishop DK, Weinert T. A meiotic recombination checkpoint controlled by mitotic checkpoint genes. of outstanding interest Nature. 383:1996;840-843 A important study in defining how recombinational events are coordinated with the cell cycle. The checkpoint genes MEC1, RAD17, and RAD24 - which halt the mitotic cell cycle in response to DNA damage - are shown to mediate the meiotic arrest of cells with a dmc1 mutation. However, these mutations do not relieve meiotic lethality, or allow for the repair of the persistent DSBs that are seen in dmc1 single mutants. The three genes may function in meiosis by signalling the presence of unrepaired intermediates of recombination.
103. Xu L, Weiner BM, Kleckner N. Meiotic cells monitor the status of the interhomolog recombination complex. of outstanding interest Genes Dev. 11:1997;106-118 Here, mutations in the RED1 and MEK1/MRE4 genes are shown to alleviate meiotic arrest in dmc1, rad51, and zip1 cells. red1 and mek1/mre4 mutations confer similar effects with respect to DSB induction and processing, SC formation, and production of recombinants. The authors propose that the recombination complex actively inhibits the progression of the meiotic cell cycle until certain phases have been completed.
104. Galbraith AM, Bullard SA, Jiao K, Nau JJ, Malone RE. Recombination and the progression of meiosis in the yeast Saccharomyces cerevisiae. of special interest Genetics. 146:1997;481-489 The authors of this paper examine the kinetics of the first meiotic division in four mutants which are defective in DSB formation. In strains mutant for REC102, REC104 or REC114 but not MEI4, the reductional division occurs earlier than in wild-type diploids. These early genes are likely to help coordinate recombination with progression through meiosis.
105. Xu L, Ajimura M, Padmore R, Klein C, Kleckner N. NDT80, a meiosis-specific gene required for exit from pachytene in Saccharomyces cerevisiae. Mol Cell Biol. 15:1995;6572-6581.