The Development and Meiotic Behavior of Asymmetrical Isochromosomes in Wheat

To determine which segments of a chromosome arm are responsible for the initiation of chiasmate pairing in meiosis, a series of novel isochromosomes was developed in hexaploid wheat (Triticum aestivum L.). These isochromosomes are deficient for different terminal segments in the two arms. It is proposed to call them ``asymmetrical.' Meiotic metaphase I pairing of these asymmetrical isochromosomes was observed in plants with various doses of normal and deficient arms. The two arms of an asymmetrical isochromosome were bound by a chiasma in only two of the 1134 pollen mother cells analyzed. Pairing was between arms of identical length whenever such were available; otherwise, there was no pairing. However, two arms deficient for the same segment paired with a frequency similar to that of normal arms, indicating that the deficient arms retained normal capacity for pairing. Pairing of arms of different length was prevented not by the deficiency itself, but rather, by the heterozygosity for the deficiency. Whether two arms were connected via a centromere in an isochromosome or were present in two different chromosomes had no effect on pairing. This demonstrates that in the absence of homology in the distal regions of chromosome arms, even if relatively short, very long homologous segments may remain unrecognized in meiosis and will not be involved in chiasmate pairing.