Relationship between incomplete synapsis and chiasma localization

One of the subjects within the meiotic field that has been actively investigated in the recent years is the temporal and functional relationships between meiotic recombination, cohesin loading and synaptonemal complex (SC) assembly. Although the study of meiotic mutants has shed some light, many questions remain to be answered. Here, we have studied this topic in the orthopteran Paratettix meridionalis, a species with telocentric chromosomes, which shows two unusual cytological features: pairing and synapsis of homologues during prophase I are restricted to the non-centromeric distal regions and extremely distal chiasma localization in metaphase I bivalents. In order to determine whether there is a relationship between both phenomena, we have used: (1) a spreading technique for following the ultrastructure of SC assembly and (2) immunofluorescence for SMC3 and SMC1alpha cohesin subunits, which mark the development of the axial element (a SC component); the histone gamma-H2AX, which mostly labels the sites of double-strand breaks; and the recombinase RAD51. Spermatocytes showed conspicuous polarization of both the maturation of cohesin axes and the initiation of meiotic recombination events. Consequently, it is proposed that maturation of cohesin axes, which begins in very distal regions, could drive the latter loading of recombinases to such regions. This restricted distribution of recombination events along homologues would finally be responsible for the incomplete pairing and synapsis observed in all autosomes of the complement and hence for chiasma localization.