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The respective contributions of chiasmatic and achiasmatic (distributive) modes of chromosome segregation can be studied in organisms with naturally-occurring achiasmatic chromosomes [ 8 ] , in cells harboring non-recombined artificial chromosomes [ 9 ] , and in achiasmatic mutants [ 10 ] .
Rec12 and its active site tyrosine are required for chiasmatic meioses Models for recombination posit that gene conversion and reciprocal exchange (crossover recombination) are alternative outcomes of a common intermediate [ 11 ] .
Rec12 is required for chiasmatic and achiasmatic chromosome segregation during MI, and its active site tyrosine is required for chromosome segregation during MII Because each of the rec12 mutants had aberrant meiotic chromosome segregation (Figure 3), produced a high frequency of viable meiotic products (Figure 4A), and produced a high frequency of meiotic diploids (Figure 4B), we were able to monitor directly the segregation patterns of all three chromosome pairs in each of the mutants.
Rec12 has dual roles in chiasmatic and achiasmatic (distributive) chromosome segregation during Meiosis I In the absence of crossover recombination (chiasmata) and a backup distributive (achiasmatic) system for chromosome segregation, MI nondisjunction should occur and 100% of any resulting diploid spore colonies would be heterozygous for centromere linked markers (Figure 5B).
Thus, Rec12 protein has one or more functions in MI that are distinct from its functions to catalyze recombination and mediate chiasmatic segregation.
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