A central feature of meiosis is the pairing and recombination of homologous chromosomes. Ustilago maydis, a biotrophic fungus that parasitizes maize, has long been utilized as an experimental system for studying recombination, but it has not been clear when in the life cycle meiotic recombination initiates. U. maydis forms dormant diploid teliospores as the end product of the infection process. Upon germination, teliospores complete meiosis to produce four haploid basidiospores. Here we asked whether the meiotic process begins when teliospores germinate or at an earlier stage in development. When teliospores homozygous for a cdc45 mutation temperature sensitive for DNA synthesis were germinated at the restrictive temperature, four nuclei became visible. This implies that teliospores have already undergone premeiotic DNA synthesis and suggests that meiotic recombination initiates at a stage of infection before teliospores mature. Determination of homologous recombination in plant tissue infected with U. maydis strains heteroallelic for the nar1 gene revealed that Nar + recombinants were produced at a stage before teliospore maturation. Teliospores obtained from a spo11D cross were still able to germinate but the process was highly disturbed and the meiotic products were imbalanced in chromosomal complement. These results show that in U. maydis, homologous recombination initiates during the infection process and that meiosis can proceed even in the absence of Spo11, but with loss of genomic integrity. M EIOSIS has long held fascination for evolutionary biologists, cytologists, and geneticists alike by the puzzle of its evolutionary origin, the chromosome choreography, and its role in generating genetic diversity (Wilkins and Holliday 2009). The distinctive events of meiosis include pairing of homologs along their entire lengths, extensive recombination between homologs, suppression of sisterchromatid separation in the first division, and absence of S phase at the start of the second division. Attention of geneticists has centered primarily on the second step-namely genetic recombination during pairing and its function in maintaining genomic integrity and in generating new linkage relationships among genes. As reciprocal recombination events are essential for proper chromosome segregation at the first meiotic division, this process has profound implications in human development and health (Nagaoka et al.
2012). The central question of how homologous chromosomes recognize each other and pair to enable recombination has provoked a great deal of research in a variety of experimental systems. Certain core principles underlie the process, but there is enormous variation in mechanical operations and regulatory mechanisms from one organism to the next and uncertainty in how DNA information is used for pairing and recombination (Pawlowski et al. 2007;Schvarzstein et al. 2010;Storlazzi et al. 2010;Tsai and Mckee 2011;Lake and Hawley 2012). Thus, to geneticists the study of meiosis is irresistibly interesting.Ustila...