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In addition to the ten linear DAF primers utilized in the present study as above, ten additional mini-hairpin primers (Hirao et al. 1992; Caetano-Anolles and Gresshoff 1994) were also evaluated for their utility in detecting polymorphism. Mini-hairpin primers form a loop of 3-4 nucleotides, a stem of only two nucleotides and a core arbitrary sequence of variable length (1-8 nucleotides) at the 3' terminal end. This loop-stem region closes during early ramping (Caetano-Anolles and Gresshoff 1994; Gresshoff et al. 1997). In the present study, mini-hairpin primers are each made up of a core arbitrary sequence of 3 bases (in earlier studies, a core sequence of 3 bases was found most appropriate) at the 3' terminus and a loop stem of 8 bases. Each of the ten mini-hairpin primers listed in Table 2 could be tried, each only on a single pair of genotypes of bread wheat differing for one quality trait. Four of these primers (HPC 23, HPC 26, HPC 29 and HPC 44) gave excellent DAF profiles with distinct scorable amplified products (Fig. 2; Lanes 1, 2, 5 and 6). The remaining six mini-hairpin primers, however, gave only a few faint bands superimposed on high background smear, suggesting their limited utility in detecting polymorphism in a crop like bread wheat (Fig. 2; Lanes 3 and 4).

The mini-hairpin primers were also examined for their utility in relation to their GC/AT ratio as done above for linear DAF primers. In this case also, higher GC content in the core sequence seems to give a smear rather than distinct scorable DAF profile (
Fig. 2; Lanes 3 and 4). However, the presence of A or T nucleotide at the 3' end. of the core sequence of the mini-hairpin primer, despite high GC content (GC/AT=2: 1) gave excellent scorable DAF profiles (Fig. 2; Lanes 5 and 6). The number of amplification products (total number of bands of <2kb) obtained with mini-hairpin primers, was almost double to that obtained with a linear primer even at an annealing temperature of 55C (Table 1 and Table 2). This may be due to a smaller core sequence (3-mer) of the primer involved in annealing, which is provided with greater thermostability by the hairpin loop during annealing (Caetano-Anolles and Gresshoff 1994; Gresshoff et al. 1997).

While making an overall assessment of the utility of DAF primers in bread wheat in terms of producing polymorphic bands, during the present study, we found that neither the average number, nor the proportion of polymorphic DAF products obtained with the mini-hairpin primers was greater than those obtained with linear primers in bread wheat (
Table 1 and Table 2). These results thus do not support the earlier contention, that for detecting polymorphism, the mini-hairpin primers are superior over the linear primers. Since these earlier results pertain to smaller genome of centipedegrass (Caetano-Anolles and Gresshoff 1994), bermudagrass (Caetano-Anolles et al. 1995) and soybean (Caetano-Anolles and Gresshoff 1996), it is possible that the relative utility of mini-hairpin primers differs in different plant systems.

The genotypes of bread wheat utilized in the present study were also earlier analyzed by us using RAPDs and microsatellite primed PCR (MP-PCR), both of which failed to detect reproducible polymorphism among the above genotypes. DAF, on the other hand, revealed polymorphism between pairs of genotypes differing for individual traits e.g. protein content, grain size, preharvest sprouting tolerance and, leaf rust resistance (
Table 1 and Table 2). Similar conclusions were earlier drawn by Prabhu and Gresshoff (1994), while examining two species of Glycine. Using DAF they obtained three fold increase in polymorphism per primer between G. max and G. soja over the polymorphism revealed using RAPDs on the same Glycine species (Williams et al. 1990). We therefore, conclude that DAF technology may become a useful tool for detecting polymorphism and for developing molecular markers for tagging genes for molecular marker aided selection in plant breeding even for a difficult crop like bread wheat.


Acknowledgments

We gratefully acknowledge the financial support from the Department of Biotechnology, Government of India, for carrying out this study. Thanks are also due to Drs. G. Caetano-Anolles and PM Gresshoff for supplying DAF primers, and Drs. Harjit Singh and H S Dhaliwal for seed materials.

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