Maize, with its excellent forward genetics and male sterility screens, was used to identify .50 meiotic mutants representing at least 35 genes that affect key prophase processes such as pairing, synapsis, and homologous recombination. Most of these mutants were found by Inna Golubovskaya during the course of her remarkable career as a cytogeneticist. In addition to undertaking general cytological surveys to classify mutant phenotypes, Golubovskaya focused her efforts on characterizing several key regulatory mutants: ameiotic1 (am1), required to establish the meiotic cell cycle in maize; absence of first division (afd1), required for proper prophase chromosome morphology and for meiotic sister-chromatid cohesion leading to a reductive chromosome segregation at the first meiotic division; and plural abnormalities of meiosis (pam1), required for the clustering of telomeres on the nuclear envelope needed for pairing and synapsis. Her dramatic childhood in Leningrad during its siege in World War II, her fortuitous education in genetics at Leningrad State University, her continued research at the forward-looking Institute of Cytology and Genetics of the USSR Academy of Science Siberian branch, her plight at the fall of the Soviet Union, and her work in America helped engender a unique and valuable plant geneticist. Inna Golubovskaya related this personal history to the authors in conversation.
MEIOSIS is the specialized cell division required in all eukaryotes with a sexual life cycle to produce gametes with a haploid content of chromosomes. During meiosis one round of DNA replication is associated with two rounds of chromosome segregation. The general progression of meiosis is conserved evolutionarily, and hence meiotic prophase and chromosome segregation are similar in plants, animals, and fungi. Following S phase, at leptotene, chromosomes condense and the two sister chromatids are held together along their length by the sister-chromatid cohesin complexes that help to form the axial element that runs the length of the leptotene chromosome. The double-strand breaks that initiate homologous recombination usually occur at this stage. At the leptotene-zygotene transition there is a transient remodeling of chromosome architecture that can include the attachment and clustering of telomeres on the nuclear envelope. These events facilitate the pairing of homologous chromosomes. Coincident with pairing, the homologs zip up-i.e., synapse-as a tripartite synaptonemal complex (SC) forms along the length of the two chromosomes between their axial elements. At pachytene, the homologs are completely synapsed and homologous recombination is completed, leading to crossing over between homologs and chiasmata formation. During diplotene and diakinesis, the SCs fall apart as chromosomes condense further. The homologs are held together by the chiasmata until they segregate away from each other at the first meiotic division (MI). During the subsequent second meiotic division (MII), sister chromatids separate from each other to ...