Distributing Meiotic Crossovers for Optimal Fertility and Evolution

During meiosis, homologous chromosomes of a diploid cell are replicated and, without a second replication, are segregated during two nuclear divisions to produce four haploid cells (including discarded polar bodies in females of many species). Proper segregation of chromosomes at the first division requires in most species that homologous chromosomes be physically connected. Tension generated by connected chromosomes moving to opposite sides of the cell signals proper segregation. In the absence of the required connections, called crossovers, chromosomes often segregate randomly and produce aneuploid gametes and, thus, dead or disabled progeny. To be effective, crossovers must be properly distributed along chromosomes. Crossovers within or too near the centromere interfere with proper segregation; crossovers too near each other can ablate the required tension; and crossovers too concentrated in only one or a few regions would not re-assort most genetic characters important for evolution. Here, we discuss current knowledge of how the optimal distribution of crossovers is achieved in the fission yeast Schizosaccharomyces pombe, with reference to other well-studied species for comparison and illustration of the diversity of biology.