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marker-assisted selection in wheat - ictsd

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126Marker-<strong>assisted</strong> <strong>selection</strong> – Current status and future perspectives <strong>in</strong> crops, livestock, forestry and fishMABC is certa<strong>in</strong>ly the form of MASwith the most immediate and obvious benefitsfor maize breed<strong>in</strong>g. MABC is usedfor three ma<strong>in</strong> purposes: <strong>selection</strong> of transgenes(or of native DNA sequences of themaize genome, whether genes or QTL),elim<strong>in</strong>ation of unwanted regions of thedonor-parent genome l<strong>in</strong>ked to the transgeneand <strong>selection</strong> of unl<strong>in</strong>ked regions ofthe recurrent-parent genome. With theexception of DNA <strong>marker</strong>s and transgenes,these have been the same goals of backcrossbreed<strong>in</strong>g s<strong>in</strong>ce the <strong>in</strong>ception of that methoddecades ago (Fehr, 1987). Of course, DNA<strong>marker</strong>s enable breeders to identify progenythat conta<strong>in</strong> the desired recomb<strong>in</strong>antchromosomes and donor-parent genome <strong>in</strong>a more direct manner. Also, MABC facilitatesthe process of comb<strong>in</strong><strong>in</strong>g more thanone transgene <strong>in</strong> a given <strong>in</strong>bred l<strong>in</strong>e (e.g.“gene or trait stack<strong>in</strong>g or pyramid<strong>in</strong>g”).This reduces the number of generationsneeded to reach certa<strong>in</strong> stages of a breed<strong>in</strong>gprogramme and reduces the time needed toproduce commercial hybrids for the market.Generic MABC schemes suitable for maizebreed<strong>in</strong>g programmes have been described<strong>in</strong> detail for s<strong>in</strong>gle genes (Hospital, Chevaletand Mulsant, 1992; Ragot et al., 1995;Frisch, Bohn and Melch<strong>in</strong>ger, 1999a, 1999b;Frisch and Melch<strong>in</strong>ger, 2001a; Hospital,2001; Ribaut, Jiang and Hois<strong>in</strong>gton, 2002),for QTL (Hospital and Charcosset, 1997;Bouchez et al., 2002) and for gene stacks(Frisch and Melch<strong>in</strong>ger, 2001b). Versions ofsuch schemes have been used <strong>in</strong> maize breed<strong>in</strong>gprogrammes <strong>in</strong> the private sector, oftenat their cont<strong>in</strong>uous nurseries (Ragot et al.,1995). Most recently, MABC has also beenadopted as a tool to develop sets of nearisogenicl<strong>in</strong>es (NILs) for genomics research(Peleman and van der Voort, 2003).Theoretical and simulation studies havebeen conducted to identify the most efficientMABC protocols. Parameters mostcommonly studied <strong>in</strong>clude the number of<strong>in</strong>dividuals genotyped at each generation,the number of <strong>marker</strong>s used, relative <strong>selection</strong>pressure for recomb<strong>in</strong>ation around thetarget locus or global recovery of recurrentparent genome and the number of <strong>in</strong>dividualsselected at any generation. Optimalvalues for each of the above depend onthe objective of the MABC approach <strong>in</strong>terms of quality (required level of recurrentparent genome recovery), speed(fastest possible conversion or set numberof generations) and resources (unlimitedor limited). While the fastest and highestquality MABC approaches have themost expensive protocols, less <strong>in</strong>tensiveapproaches can result <strong>in</strong> significant timesav<strong>in</strong>gs and quality improvements whencompared with conventional backcross<strong>in</strong>gapproaches and at a fraction of the cost ofthe most expensive MABC protocols.Frisch, Bohn and Melch<strong>in</strong>ger (1999b)showed that to m<strong>in</strong>imize l<strong>in</strong>kage dragaround the target locus (loci), <strong>selection</strong> ofrecomb<strong>in</strong>ation events close to the targetlocus (loci) should be conducted <strong>in</strong> theearly backcross generations. Frisch andMelch<strong>in</strong>ger (2001a) and Ribaut, Jiang andHois<strong>in</strong>gton (2002) further demonstratedthat m<strong>in</strong>imiz<strong>in</strong>g l<strong>in</strong>kage drag around thetarget locus requires very large numbersof <strong>in</strong>dividuals (possibly hundreds) to begenotyped. Hospital and Charcosset (1997)proposed a <strong>selection</strong> scheme based onselect<strong>in</strong>g a s<strong>in</strong>gle <strong>in</strong>dividual to be backcrossed.By contrast, Frisch and Melch<strong>in</strong>ger(2001a) proposed select<strong>in</strong>g several <strong>in</strong>dividualsand determ<strong>in</strong><strong>in</strong>g the family sizeof their backcross progeny based on the<strong>in</strong>dividuals’ genotypes. By us<strong>in</strong>g vary<strong>in</strong>grather than constant numbers of <strong>in</strong>dividualsor <strong>marker</strong>s at the different backcrossgenerations, it was shown that the number

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