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marker-assisted selection in wheat - ictsd

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270Marker-<strong>assisted</strong> <strong>selection</strong> – Current status and future perspectives <strong>in</strong> crops, livestock, forestry and fishFigure 6Rapid decay of <strong>in</strong>tragenic l<strong>in</strong>kage disequilibrium estimated as r 2 <strong>in</strong> the c<strong>in</strong>namyl-alcoholdehydrogenase (cad) gene <strong>in</strong> two different Eucalyptus species10.90.80.70.60.50.40.30.20.1E. urophyllar 2 000 500 1000 1500 2000 2500 3000Distance <strong>in</strong> basepairs10.9E. grandis0.80.7r 20.60.50.40.30.20.10 500 1000 1500 2000 2500Distance <strong>in</strong> basepairsSource: Faria et al., 2006.LD <strong>marker</strong>s, i.e. polymorphisms that aresufficiently close to the functional mutation(Neale and Savola<strong>in</strong>en, 2004). The challenge,however, is considerable, as LD <strong>in</strong>outcross<strong>in</strong>g forest trees such as p<strong>in</strong>es decaysvery rapidly, <strong>in</strong> general with<strong>in</strong> 1 500 to 2 000bp (Neale and Savola<strong>in</strong>en, 2004), and similarbehaviour has been seen <strong>in</strong> the few Eucalyptusgenes analysed to date with significant LDextend<strong>in</strong>g for only a few hundred base pairs(Thumma et al., 2005; Kirst, Marques andSederoff, 2005; Faria et al., 2006) (Figure 6).Genome-wide association studies for LD<strong>marker</strong>-trait discovery <strong>in</strong> tress will requirevery high SNP <strong>marker</strong> densities that are currentlystill impracticable (but see below), sothat the only alternative left is a candidategene approach. F<strong>in</strong>ally, direct <strong>marker</strong>s (i.e.polymorphisms that code for the functionalmutations) would be the most valuable and

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