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marker-assisted selection in wheat - ictsd

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274Marker-<strong>assisted</strong> <strong>selection</strong> – Current status and future perspectives <strong>in</strong> crops, livestock, forestry and fishMAS for that particular trait. Estimates ofheritability for a trait might be useful togive an <strong>in</strong>itial clue. Intuitively, the probabilityof major genes exist<strong>in</strong>g for traitsof low heritability is lower than for traitsof high heritability. However, this shouldnot be taken as a measure to discard possibleQTL mapp<strong>in</strong>g experiments as, evenwith low heritabilities, traits might still displaymajor QTL, and MAS would have thegreatest impact particularly <strong>in</strong> such cases.Conclusions and perspectivesThe successful application of molecularbreed<strong>in</strong>g <strong>in</strong> Eucalyptus will dependheavily on first demonstrat<strong>in</strong>g and validat<strong>in</strong>gthe clear-cut association between aDNA polymorphism and a quantitatively<strong>in</strong>herited phenotypic trait. In highly heterogeneouseucalypts, while conventionalQTL mapp<strong>in</strong>g can reveal useful <strong>marker</strong>sto be exploited <strong>in</strong> with<strong>in</strong>-family <strong>selection</strong>practices, only a more direct LD mapp<strong>in</strong>gapproach can uncover populationwide applicable <strong>marker</strong>-trait associations.Such studies based on candidate genes havebegun and the first candidate gene associationfor MFA was detected. However, thisassociation expla<strong>in</strong>s only a small proportion(3.4 percent) of the variation to bereally excit<strong>in</strong>g news to breeders (Thummaet al., 2005). One of the key issues whenembark<strong>in</strong>g on an association mapp<strong>in</strong>g experimentis the <strong>selection</strong> of candidate genes.Maximiz<strong>in</strong>g the probability of choos<strong>in</strong>g theproper genes requires levels of knowledgeof biochemistry, physiology and developmentthat are generally not yet availableeven for well def<strong>in</strong>ed phenotypes and/orknown metabolic pathways.Follow<strong>in</strong>g the path taken <strong>in</strong> humangenetics, co-localization of candidate genesand QTL for relevant traits on l<strong>in</strong>kagemaps together with <strong>in</strong>tegrative expression-QTL mapp<strong>in</strong>g (Kirst et al., 2004) could be apowerful way forward, although choos<strong>in</strong>gthe correct candidate depends heavily onthe precision of the QTL localization. Atthe moment, there are two possibilities forcircumvent<strong>in</strong>g the dilemma of choos<strong>in</strong>gcandidate genes correctly. The first is microarray-basedgenotyp<strong>in</strong>g with ultra-densearrays of short (25 nt) oligonucleotides(Borevitz et al., 2003; Hazen and Kay, 2003;West et al., 2006) that would allow sufficientthroughput for association geneticanalysis of thousands of genes at a time.Such an array format could later turn out tobe a useful <strong>in</strong>strument for MAS once validated<strong>marker</strong>-trait associations have beenestablished. The second would be to haveaccess to a whole genome sequence so thatcandidate genes <strong>in</strong> a f<strong>in</strong>e mapp<strong>in</strong>g <strong>in</strong>tervaldelimited by <strong>marker</strong>s flank<strong>in</strong>g a QTL withcentimorgan resolution could be m<strong>in</strong>ed,reannotated and then analysed <strong>in</strong> associationmapp<strong>in</strong>g experiments.A draft genome of E. camaldulensis is currentlybe<strong>in</strong>g sequenced at the Kazusa DNAResearch Institute <strong>in</strong> Japan (T. Hib<strong>in</strong>o, personalcommunication), and the possibilityexists that a fully public 4X draft of theE. grandis genome will be sequenced bythe Jo<strong>in</strong>t Genome Institute of the UnitedStates Department of Energy with<strong>in</strong> thenext years (J. Tuskan, personal communication)follow<strong>in</strong>g a proposal recentlysubmitted by an <strong>in</strong>ternational group ofEucalyptus geneticists (www.ieugc.up.ac.za/DOE%20proposal%20-%20f<strong>in</strong>al%20-%2026%20July%202006.pdf) who recentlyformed the International EucalyptusGenome Network (EUCAGEN) (www.ieugc.up.ac.za; Myburg, 2004). Such publiccollaborative efforts should contributegreatly to the advancement of Eucalyptusgenetics, genomics and molecular breed<strong>in</strong>gby br<strong>in</strong>g<strong>in</strong>g together exist<strong>in</strong>g private data-

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