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marker-assisted selection in wheat - ictsd

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Chapter 5 – Marker-<strong>assisted</strong> <strong>selection</strong> for improv<strong>in</strong>g quantitative traits of forage crops 63Figure 2Strategy applied for <strong>marker</strong> <strong>selection</strong>Co-dom<strong>in</strong>ant scor<strong>in</strong>g of<strong>marker</strong>s associated withQTLsGenotypeNumber of plus allelesQTL1QTL2QLT3QTL4QTL5SumSelectioncriterionF 21 2 1 0 0 2 52 1 1 0 0 2 43 0 2 2 1 1 64 2 2 2 1 1 85 2 2 1 1 1 7...195 1 1 2 2 2 8196 1 1 2 0 2 6197 0 2 0 1 2 5198 0 2 0 1 1 4199 2 2 1 1 1 7200 2 1 2 2 1 8fragments) turned out to be straightforward.Figure 3 shows the F 2 frequency distributionfor the number of “plus alleles”.The population mean is somewhat belowthe expected number of five ow<strong>in</strong>g to thefact that the AFLP <strong>marker</strong> on LG1 showeda skewed segregation. This is likely dueto gametophytic <strong>selection</strong> <strong>in</strong> favour of thenegative QTL allele, perhaps due to l<strong>in</strong>kagewith an <strong>in</strong>compatibility locus.The <strong>in</strong>tensities of <strong>selection</strong> were setat about 25 percent, represent<strong>in</strong>g about50 genotypes per <strong>selection</strong> (Figure 3). The<strong>selection</strong> pressure was kept fairly lowbecause of the need to have sufficient seedsfor measur<strong>in</strong>g <strong>selection</strong> responses. In thisway, the <strong>in</strong>fluence of genetic drift accompany<strong>in</strong>g<strong>marker</strong> <strong>selection</strong> was m<strong>in</strong>imized. Thecut-off po<strong>in</strong>t for the top <strong>selection</strong> was sixpositive alleles and three for the opposite<strong>selection</strong> (Figure 3). The frequency of theplus alleles was on average 0.66 and 0.27,respectively. Selection showed a positiveresponse for all NUE loci. However, thebetween-<strong>selection</strong> difference <strong>in</strong> allele frequencyof the loci ranged from 0.18 to 0.77,show<strong>in</strong>g that <strong>in</strong>dex <strong>selection</strong> did not affectall NUE loci to the same degree. The differenceswere probably ma<strong>in</strong>ly due to chance.Indirect response to <strong>marker</strong><strong>selection</strong>The <strong>selection</strong>s were then multiplied us<strong>in</strong>ga polycross scheme (after vegetative propagation)to obta<strong>in</strong> sufficient seeds forevaluation on hydroponics and under variousfield conditions. The <strong>marker</strong> <strong>selection</strong>swere evaluated for NUE <strong>in</strong> a replicated trialwith two cuts on hydroponics at two Nlevels, be<strong>in</strong>g 2.5 and 5 percent N <strong>in</strong> leaves(van Loo et al., 2003). The same set of plantcharacteristics as <strong>in</strong> the orig<strong>in</strong>al mapp<strong>in</strong>gstudies was monitored after each cut. Leafarea expansion rate, leaf length and width,as well as tiller number, were determ<strong>in</strong>edone week after cutt<strong>in</strong>g. The determ<strong>in</strong>ationof shoot and root dry weight followedthree weeks later. The <strong>in</strong>direct responsesto <strong>marker</strong> <strong>selection</strong> are summarized <strong>in</strong>Figure 4. At low N supply, the NUEplus

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