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The single tetraploid hybrid obtained was between Ae. triuncialis and Ae. cylindrica. Observed were an average of 7.29 bivalents, of which only 2.25 were rings, 1.16 trivalents, 0.39 quadrivalents, and 0.15 larger multivalents (Table 2). Higher bivalent frequencies were reported in earlier hybrids between these two species. PERCIVAL (1930) recorded a range from three to eleven bivalents of which five to six were rings. KIHARA & LILIENFELD (1932) reported zero to twelve bivalents with a mode of ten to eleven ; three to six were rings. KAGAWA (1931, cited in KIHARA 1937) in the reciprocal combination reported six to eleven bivalents and no or one trivalent. The possible modification in the C genome of this accession of Ae. triuncialis discussed above could account for the apparent discrepancy between the pairing reported here for this tetraploid hybrid and that from earlier reports.

In the three pentaploid hybrid combinations, Ae. recta (Zhuk.) Chenn. was a common parent. The second parent in each case possessed the U genome in common with Ae. recta. Observed in Ae. variabilis Eig x Ae. recta were means per PMC of 6.63 bivalents, 2.27 of which were rings, 1.19 trivalents, and 0.11 quadrivalents (Table 3). KIHARA (1945) reported seven to eleven bivalents, as many as four of which were rings, and zero to two multivalents. He concluded that the two species had only one genome (U) in common. The chromosomes of the B and U genomes were shown to be able to produce at least two bivalents in the diploid hybrid discussed above. This affinity may be reflected in this pentaploid by the frequencies of both rod bivalents and trivalents.

In addition to the U genome, the other pentaploid hybrid combinations supposedly have versions of the M genome in common. Evidence for this is indirect and in no case has an unmodified M genome from Ae. comosa been demonstrated in an Aegilops polyploid. In the case of Ae. geniculata Roth (=Ae. ovata auct.) evidence for a modified M genome is very indirect. In fact, the extent of pairing in hybrids of Ae. geniculata and Ae. ventricosa (e.g., KIHARA 1937) suggests that these two species have a genome in common, in which case the genome formula of Ae. geniculata should be LU. There is even less direct evidence for an M genome in Ae. lorentii Hoch. (=Ae. biuncialis Vis.). Based on karytype analysis CHENNAVEERAIAH (1960) concluded that the second genome of Ae. lorentii might well be something other than a modified M.

Aegilops recta x Ae. geniculata hybrids have not been reported before. The three plants reported here had means of 8.88 bivalents, 2.76 of which were rings, 1.10 trivalents, 0.21 open quadrivalents, and 0.02 quinquevalents (Table 3). In the Ae. recta x Ae. lorentii hybrids, means of 7.94 bivalents, 2.04 of which were rings, 1.79 trivalents, 0.24 open quadrivalents, 0.15 quinquevalents, and 0.01 sexivalents were observed (Table 3). KIHARA (1945) reported two hybrids from this parental combination. One had about nine bivalents, as many as five of which were rings, and some trivalents and quadrivalents. The other had a range of six to eleven bivalents, many of which were rings, and an occasional quinquevalent. These results for these two hybrid combinations provide evidence for no more than one genome in common in each case as was found for the Ae. variabilis x Ae. recta combination just discussed. In conjunction with the karyotypic evidence of CHENNAVEERAIAH (1960), this must be the U genome. Thus, the identity of the non-U genomes in these polyploids is questionable.

The hexaploid hybrid, Ae. juvenalis (Thell.) Eig x Ae. recta, is also a combination reported for the first time. In the two plants there were combined means per PMC of 7.25 bivalents, with 1.17 rings, 2.10 trivalents, and 0.29 quadrivalents. On the basis of these data, especially the low frequency of ring bivalents, one might question if one genome could be considered common to both species. However, other evidence has shown the presence of the U genome in both Ae. juvenalis (MCGINNIS 1956 ; KIHARA 1963 ; KIMBER & ABU 1981) and Ae. recta (KIHARA 1937, 1957).

None of the data suggested any definitive changes in the current understanding of the identity of the component genomes of the polyploid taxa considered here. However, there is enough uncertainty in the cases of the polyploid species that presumably have modified M genomes to warrant some direct genome analytical work particularly with the diploid species Ae. comosa and Ae. uniaristata as testers as was done recently with Ae. uniaristata and Ae. ventricosa (KIMBER et al. 1983). In the cases here where comparisons could be made with earlier hybrids, the discrepancies underscored the need for wider sampling within species to assess the extent of genome modification and of genetic variability for the requlation of heterogenetic pairing. In addition, the level of pairing in the Ae. ventricosa x Ae. umbellulata hybrid plant when compared with pairing reported in haploids of Ae. ventricosa (FEDAK 1983) suggested promotion of heterogenetic pairing by Ae. umbellulata.

Acknowledgements

The author wishes to acknowledge review and discussion of the data by JAN DVORAK and GORDON KIMBER.



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