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Results and discussion

Pollination of 143 emasculated Chinese Spring florets by
D. villosum yielded 1 seed, a ca. 0.7% seed set in 1993, whereas of 102 florets pollinated, 3 seeds were obtained, seed set was 2.9% in 1994. The t-test indicates there is no significant difference in two years, which inferred the crossing rates were identical in this study.

The results of cross were listed in
Table 1, including the pollinated florets and percentages of seed set. The percentage of seed set varies between 0.0% and 22.6%. With regard to the female parents of monosomics, two distinct classes were found when D. villosum was used as pollinator. In the combinations with homoeologous group 5 of monosomics (5A, 5B and 5D) 17.1%, 22.6% and 10.8% of seed set were obtained, which demonstrated significantly higher seed set than that in cross with normal Chinese Spring by t-test, whereas in those with other homoeologous groups of monosomics, the percentages of seed set were below 10%, which did not exhibit remarkable differences in comparison to that with normal Chinese Spring. These results suggested the import ant role of maternal genotypes in wheat-D. villosum hybridization. We have known that three kr genes, which located on chromosomes 5A, 5B and 5D, respectively, control crossability between Chinese Spring wheat and rye species. The crossabilities of Chinese Spring with other wild species controlled by the kr genes have been reported in Elymus giganteus (Mujeeb-Kazi et al. 1983), Hordeum vulgare (Fedak and Jui 1982), and Hordeum bulbosum (Falk and Kasha 1981). Dai et al. (1988) investigated the relationship between the crossability of common wheat cultivars with D. villosum and rye, Secale cereale, by using common wheat cultivars having high and low crossable rates with rye, and found that the crossabilities of common wheat cultivars with D. villosum and rye are controlled by the same genetic system. Koba and Shimada (1998) reported the similar result using Aegilops squarrosa and Secale cereale. These findings support the traditional concept that the crossability between Chinese Spring wheat with wild species was suppressed by Kr genes, while was promoted by the recessive loci. Halloran (1966) reported a 1.2% seed set in his work using the monosomic 5B of Chinese Spring as female parent crossed with D. villosum, while using Chinese Spring as female parent, no seed was obtained. He has suggested that there might be a gene(s) on chromosome 5B of Chinese Spring which prevented successful cross in disomic condition, but in monosomic state, was ineffective. In the present study, it was found a high percentage of seed set in the cross between monosomic lines of 5A, 5B and 5D and D. villosum. With the results in the present study, we suggested that the three pairs of kr genes for crossability would be the recessive mutants of Kr gene, which expressed as reducing the effect on preventing cross between wheat and its wild species. Different kr gene would exhibit different degree of effect an reducing to prevent wide cross. The kr1 gene is the strongest mutant to reduce the effect of preventing wide cross and kr3 is the weakest one. The results would have demonstrated the dosage effect of hr genes on preventing wide cross. The lack of kr genes in monosomics resulted in a higher seed set in their crosses with D. villosum.
Many obtained hybrid seeds failed to germinate on moist paper in petri dishes in the laboratory due mainly to their extremely shrivelling endosperm. Chen et al. (1982) and Dai et al. (1988) found that the percentages of wed set in crossing T. tauschii as female parent, with D. villosum were considerably high (up to 80%), but the hybrid seeds almost did not germinate due to the lack of endosperm.
T. tauschii is the donor species of D genome in hexaploid wheat. These facts may provide an evidence that a gene(s), carried on a certain D genome chromosome in T. tauschii and hexaploid wheat, refrains from development of endosperm in the process of seed formation after hybridization.

Table 1 shows the seedling frequencies per 100 florets pollinated is relatively low. For obtaining more hybrid plants, it is necessary to employ the cultural embryo rescue technique in the cross as von Bothmer and Claesson (1990) did in their work.

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