Abstract

Normal females developing at 25 degrees C produce their first population of oocytes at 132 +/- 2 hr post oviposition. Entrance of the oocytes into premeiotic interphase signals initiation of DNA replication which continues for 30 hr. Coincidentally, extensive SCs appear, averaging 50 microns (132 hr), peaking at 75 microns (144 hr), and continuing into early vitellarial stages. Recombinational response to heat, evidenced by enhancement or induction of exchange, is limited to the S-phase with a peak at 144 hr coinciding with maximal extension of the SC. Coincidence of synapsis and recombination response with S at premeiotic interphase is contrary to their conventional localization at meiotic prophase. The interrelationship between exchange and nondisjunction has been clarified by the Distributive Pairing Model of meiosis. Originally revealed through high frequencies of nonrandom assortment of nonhomologous chromosomes, distributive pairing has been shown to follow and to be noncompetitive with exchange, to be based on size-recognition, not homology, and as a raison d'etre, to provide a segregational mechanism for noncrossover homologs. Rearrangements, recombination mutants, and aneuploids may contribute noncrossover chromosomes to the distributive pool and so promote the nonhomologous associations responsible for nondisjunction of homologs and regular segregation of nonhomologs. Further information concerning early meiotic events and their relation to segregation has been revealed by studies of the ts rec-1(26) mutant. Application of the restrictive temperature (31 degrees C) at sequential times during development shows wild-type activity to be drastically reduced beginning at 126/132 hr, terminating at 162 hr, and so coinciding with S and the heat-sensitive period of the normal genome. EMs of serially sectioned oocyte nuclei maintained at the restrictive or control temperature reveal SCs to be indistinguishable, and implicate recombination rather than synapsis as the target of the mutant. Activity of rec-1(26) in the range 17 degrees to 31 degrees C reveals a sharp decline between 28+ degrees and 31 degrees C, typical of a denaturation curve. If denaturation of the rec protein by the restrictive temperature marks its active phase, it follows that recombination terminates at the end of S when the restrictive temperature becomes ineffective. The notion that synapsis ensures regular segregation (35) is invalidated by rec-1(26) whose normal synapsis at the restrictive temperature is followed by a 200-fold increase in nondisjunction at segregation, as compared to the control.(ABSTRACT TRUNCATED AT 400 WORDS)

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