Premeiotic events and meiotic chromosome pairing

There is practical difficulty in identifying when meiosis begins. Moreover, because of contradictory definitions there is ambiguity and some confusion as to when, in terms of the cell cycle, premeiosis ends and meiosis begins. Nevertheless, results for several organisms show clearly that meiotic chromosome behaviour is affected by premeiotic events and especially by events during the final premeiotic mitosis and/or premeiotic interphase. This review considers only premeiotic events which do (or might) affect meiotic chromosome pairing by their effect on genomic characters, such as: chromosome number, homology, condition and position, with particular emphasis on the last. Interpreted in its widest sense 'premeiotic events affecting meiotic chromosome pairing' must include karyogamy. Moreover, while karyogamy is the normal means of achieving the diploid chromosome number and pairs of homologues essential for normal chromosome pairing, it is not the only way, as illustrated by the remarkable premeiotic adaptations seen in the apogamous ferns and the frog Rana esculenta. Little is known about the condition (including the molecular organization) of chromosomes during their approach and switch to meiosis. However, completion during premeiosis of some DNA synthesis may be essential for normal meiotic chromosome pairing. Various results (including different effects of colchicine given first at different premeiotic stages) have been claimed as evidence of one or other type of premeiotic spatial ordering of chromosomes which might favour, or be essential for, meiotic chromosome pairing. Chromosome placement has been studied recently using the electron microscope, serial thin-section, reconstruction technique. This has revealed clear evidence of non-random spatial placement of chromosomes in non-meiotic and premeiotic cells. For example, in root-tip cells of barley, Hordeum vulgare L. cv. Tuleen 346 (2n = 2x = 14), it showed: a significant spatial separation of two haploid chromosome sets; no somatic association of homologues; but a significant expression of a predicted mean order of heterologues (namely, linkage groups -4-1-2-3-5-6-7-) within haploid sets. This spatial arrangement of chromosomes might have important implications for the switch from premeiotic to meiotic development, if it occurred in germ line cells. Consequently, metaphase cells of Tuleen 346 at the final premeiotic mitosis were reconstructed next. Analysis of centromere positions showed a significantly ordered arrangement of haploid sets, and of heterologues within haploid sets, essentially the same as that previously noted for root-tip cells.(ABSTRACT TRUNCATED AT 400 WORDS)