Chromosome pairing and synaptonemal complex formation have been analyzed at the ultra_structural level in spread microsporocyte nuclei of allohexaploid wheat, nullisomic for chromosome 5B. Two plants were investigated, one was selected among the progeny ofselfed monosomic 5B wheat and the other was obtained by anther culture of wheat monosomic for 5B. The absence of chromosome 5B in the two plants was confirmed by N-banding of root tip complements and at metaphase I several of the cells contained multivalents. The ultrastructural analysis of 18 spread nuclei ranging in stage from the beginning of zygotene to early diplotene revealed an arrest of chromosome pairing and synaptonemal complex formation in this genotype when on the average 35-40% of the complement had synapsed. At zygotene there were several exchanges of pairing partners, the number of lateral components in multiple associations exceeding that observed in euploid wheat by a factor of two to three. Many of the pairing partner exchanges were dose to the telomeres. As crossing over between homoeologues can occur in nullisomic 5B wheat it is apparent that nullisomy for Chromosome 5B has a dual effect on meiosis, namely to reduce the stringency ofsynapsis and to permit crossing over between chromosomes of partial homology.
I N T R O D U C T I O NIn the absence of chromosome 5B, crossing over and ehiasma formation can occur between the homoeologues ofallohexaploid wheat (5, 6, 7). RILEY (5) thus reported that in nullisomic 5B wheat there were at metaphase I a mean of 1.41 univalents, 17.22 bivalents, 0.47 trivalents, 0.59 quadrivalents, 0.02 pentavalents and 0.05 hexavalents. That these multivalents had arisen by crossing over between homoeologous chromosomes was subsequently shown by RILEY and KEMPANA (7), who, by crossing plants nullisomic for 5 B with euhexaploid wheat, obtained plants that were heterozygous for translocations. The chromosomes involved in the translocations were identified by test crosses to lines in which one specific chromosome was monosomic and it was shown that the translocations had formed between homoeologous chromosomes, indicating crossing over between homoeologues in the nullisomic 5B wheat.The present paper is a part of a series, attempting at the ultrastructural level to investigate the effect of different dosages of chromosome 5B on synapsis and chiasma formation (1,2,3, 9,10,11). In this study the results of the analysis of plants nullisomic for chromosome 5B are reported. The significance of the combined results of these ultrastructural investigations for understanding the regulation of the disomic inheriSpringer-Vedag 0105-1938/88/0053/0091/$04.00