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The endosperm development in all hybrids, except hybrid kernels produced by 3D (3B) and 6D(6B), was very poor and all of the kernels were shrivelled. However, hybrid kernels produced by 3D(3B) germinated (5.9%), while no hybrid kernels produced by other substitution lines germinated. These data indicate that the 3D chromosome may contribute to the development of embryo. At the same time, Miller et al. (1983) reported that 3D chromosome effected embryo development in Chinese Spring x Hordeum bulbosum crosses.

From the results of this study, it was demonstrated that not only chromosome 5B, 5A and 5D, but also chromosome 3A and 3 B were involved in the crossability of wheat with Secale cereale . Also, 3D chromosome may be contributing to the development of embryo.

Aegilops squarrosa x D-genome substitution lines
Crossability of D genome substitution lines of Langdon with Aegilops squarrosa is presented in Table 2.

As can be seen in Table 2, a wide range in crossability percentage was obtained by crosses of Aegilops sguarrosa x LDN-D genome substitution lines. When the crossability percentages are compared, it was shown that in 3D(3A), 6D(6A), 3D(3B) and 6D(6B) disomic substitution lines it was higher than other disomic substitution lines. The crossability percentage of these lines ranged from 15.0 % to 46.2 %, while in six substitution lines viz. lD(lA), 2D(2A), lD(lB), 2D(2B), 4D(4B)and 7D(7B), it ranged from 3.6 % to 9.4 %. The crossability of the control and 5D(5A), 7D(7A) and 5D(5B) lines were zero. This result suggested that 3A, 6A, 3B and 6B chromosomes had a gene/ genes that inhibit crosses between Aegilops squarrosa and durum wheat. It has been clearly shown that chromosome 5D is also responsible for the crossability of wheat with Aegilops squarrosa. At the same time, Koba and Shimada (1993) reported that, like 5A and 5B, 5D chromosome carried genetic factor(s) controlling the crossability with Aegilops squarrosa.

The endosperm development in all hybrid kernels, except hybrid kernels produced by 6D(6A), 6D(6B) and 2D(2B), was very poor and all kernels were shriveled. The hybrid kernels produced by 6D(6B) germinated (6.7 %), while no hybrid kernels produced by other substitution lines germinated. This results suggested that 6D chromosome may be partly responsible for the embryo development of Aegilops squarrosa x durum wheat crosses.

However, we could have had a permissive accession of Aegilops squarrosa because most researchers have found it very difficult to cross Aegilops squarrosa with wheat when Aegilops squarrosa used as the female (Dr. Joppa, pers. com.). At the same time, if we can find a permissive accessions of Aegilops squarrosa, this accession(s) could be used to improve new durum or bread wheat in future.

Also, main reasons for low and high crossability of Langdon D-genome substitution lines with Aegilops squarrosa can be the technique used, and climatic conditions during crossing. Similar results were obtained in wheat x Aegilops crosses by Ozgen (1983).

Acknowledgements

We acknowledge Dr. L.R. Joppa, North Dakota State University, for supplying the seed of Langdon D genome chromosome substitution lines. Also, We thanks Dr. L.R. Joppa and Dr. S.M. Reader (John Innes Centre, Cereals Research Department, UK) for their valuable critical renew of the manuscript.

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