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During meiotic prophase the repair of self-inflicted DNA double-strand break (DSB) damage leads to meiotic recombination in yeast. We employed a genetic screen to specifically characterize cellular functions that become essential after this DSB formation. As a result a new allele of  MRE11,  termed  mre11S  (for S eparation of functions) was isolated that allows initiation but not processing and repair of meiotic DSBs similar to the well-characterized  rad50S  allele. In contrast, the mre11-1  allele blocks initiation of meiotic DSBs as reported previously by others. The  mre11S  allele, which is mutated in the 5′ part of the gene, can partially complement  mre11 alleles disrupted close to the 3′ end that cannot initiate DSBs when homozygous. This suggests homodimerization of the Mre11 protein and the presence of separate domains for DSB initiation and 5′ resection. The fact that two genes,  RAD50  and  MRE11, required for DSB processing are also essential for DSB initiation dictates a model in which a bifunctional initiation/repair complex is required to initiate meiotic recombination. A subset of  mre11S  nuclei was shown to perform extensive but partially nonhomologous synapsis. We propose that the unprocessed DSBs present in  mre11S  allow for synapsis, but that homologous synapsis is only ensured at a later stage of recombination.

mre11S—a yeast mutation that blocks double-strand-break processing and permits nonhomologous synapsis in meiosis