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Specificity of these nucleo-cytoplasmic interactions was also indicated by the following observations : An addition of chromosome 1D to the T. durum genome was essential for viability of T. durum with the cytoplasm of Ae. squarrosa, Ae. cylindrica, or Ae. ventricosa. Therefore, chromosome 1D has critical nuclear genes controlling effects conditioned by these cytoplasms. These genes were not essential for viability of T. durum with the cytoplasms of Ae. uniaristata, Ae. crassa or Ae. juvenalis. Apparently, chromosome 1D or another D-genome chromosome has homoeoalleles which make T. aestivum fertile in the cytoplasm of Ae. uniaristata, Ae crassa, or Ae. juvenalis, even though alloplasmic plants of T. aestivum in these three cytoplasms are less productive than those with cytoplasms of Ae. squarrosa, Ae. cylindrica, or Ae. ventricosa. The critical gene(s) on an Ae. uniaristata chromosome arm which restored plant vigor to T. durum with Ae. uniaristata cytoplasm did not have male fertility restoring genes (MAAN, 1977a). These plants were male sterile, and the effect of the critical Ae. uniaristata chromosome remains to be examined in the cytoplasms of Ae. crassa or Ae. juvenalis. These results further indicate that polyploid nuclear genomes may be useful in the study of nucleo-cyioplasmic interactions involving specific nuclear genes or chromosome segrnents and alien cytoplasms.

In the studies of nucleo-cytoplasmic interactions between Triticum genomes and alien cytoplasms, the cytoplasm donor species are crossed as females, and Triticum species are used as the recurrent male parents. Two types of plants appear in the backcross progenies. In the first type of plant, all of the nuclear genes from the cytoplasm donor species are eliminated, and the nuclear genome of the recurrent male parent is substituted into the alien species cytoplasm. Plants so obtained may have cytoplasmically inherited male sterility or other cytoplasmic effects such as reduced plant vigor and enhanced sensitivity to environmental conditions. The male-sterile plants may be maintained by repeated backcrossing with the recurrent male parent. In the second type of plant, certain critical nuclear genes from the genome of the female cytoplasm donor are transferred to the genome of the recurrent male parent when parental chromosomes pair in the F1 or in the backcross plants. Alternatively, a chromosome or a chromosome arm with critical nuclear gene(s) is transferred to the genome of the recurrent male parent, if parental chromosomes do not pair in the F1. Plants of the second type have noticeably improved male fertility and/or plant vigor when compared to the first types and may be maintained by selfing, if fertile. Usually most of the fertile alloplasmic plants are less vigorous, and less productive than euplasmic controls used as the recurrent male parent ; either the critical nuclear gene(s) from the cytoplasm donor species are not fully expressed when transferred to the wheat genome, or not all of the genes necessary for complete control of cytoplasmic effects to produce a normal phenotype are transferred to the wheat genome by the backcrossing procedures. However, alloplasmic plants comparable to euplasmic controls in plant vigor or productivity can be produced by repeated selection and intercrossing of plants with maximum expression of the critical nuclear genes controlling cytoplasmic effects. In plants having normal vigor and fertility, certain genes with detrimental effects may be eliminated during backcrossing and selection procedures, or certain genes in T. aestivum may complement specific nuclear genes in improving nucleo-cytoplasmic interactions


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