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marker-assisted selection in wheat - ictsd

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120Marker-<strong>assisted</strong> <strong>selection</strong> – Current status and future perspectives <strong>in</strong> crops, livestock, forestry and fish<strong>in</strong> the nuclear genome. As described below,the methods used to detect RFLPs were<strong>in</strong>compatible with the magnitude, speedand efficiency of all but a few aspects of<strong>selection</strong> <strong>in</strong> maize breed<strong>in</strong>g programmes.Gradually, however, the methods used todetect DNA polymorphisms and to createmean<strong>in</strong>gful <strong>in</strong>formation from DNA <strong>marker</strong>and phenotypic data sets have evolved to thepo<strong>in</strong>t where they are rout<strong>in</strong>e componentsof some maize breed<strong>in</strong>g programmes <strong>in</strong> theprivate sector.Selection occurs at various stages <strong>in</strong>maize breed<strong>in</strong>g programmes. The firstopportunity arises when choos<strong>in</strong>g <strong>in</strong>bredl<strong>in</strong>es to mate as parents of new populations.In some programmes, all such <strong>in</strong>breds aregenotyped systematically at DNA <strong>marker</strong>loci (Smith and Smith, 1992). If the <strong>marker</strong>loci are sufficiently close on genetic orphysical maps then reasonably good <strong>in</strong>ferencesmay be made about the <strong>in</strong>bred’shaplotype. Such <strong>in</strong>formation is used toestablish identity, resolve disagreementsrelated to germplasm ownership and acquisition,enforce laws <strong>in</strong>tended to encouragegenetic diversity of the hybrids and avoidus<strong>in</strong>g <strong>in</strong>breds that conta<strong>in</strong> transgenes whichmay violate regulatory considerations andrestrictions. These <strong>selection</strong> practices, whileadmittedly not conventional MAS, haveled to improvements <strong>in</strong> the maize crop byenabl<strong>in</strong>g more <strong>in</strong>formed stewardship anddeployment of genetic resources and byprovid<strong>in</strong>g a degree of protection of <strong>in</strong>tellectualproperty and related <strong>in</strong>vestments <strong>in</strong>maize breed<strong>in</strong>g.Unquestionably, the most pervasive anddirect use of MAS <strong>in</strong> maize by the privatesector has been with backcross<strong>in</strong>g oftransgenes <strong>in</strong>to elite <strong>in</strong>bred l<strong>in</strong>es, the directparents of the commercial hybrids (Ragotet al., 1995; Crosbie et al., 2006). Currently,the most widely deployed transgenes andcomb<strong>in</strong>ations thereof (i.e. gene stacks) arefor resistance to herbicides or <strong>in</strong>sects (e.g.Ostr<strong>in</strong>ia and Diabrotica). As the commercialmaize crop of any region, maturityzone, market or country is not yet uniformor homogeneous for any transgene,maize breeders have elected to developnear-isogenic versions (transgenic and nontransgenic)of elite <strong>in</strong>breds and commercialhybrids <strong>in</strong> order to satisfy comb<strong>in</strong>ations oflicens<strong>in</strong>g agreements, agronomic practices,regulatory requirements, market demandsand product development schemes. Thishas required companies to have twoparallel maize breed<strong>in</strong>g programmes, transgenicand non-transgenic. In this manner,<strong>marker</strong>-<strong>assisted</strong> backcross<strong>in</strong>g (MABC) oftransgenes, and to a lesser degree, of nativegenes and quantitative trait loci (QTL) forother traits, has expedited the developmentof commercial hybrids.More recently, <strong>marker</strong>-<strong>assisted</strong> recurrent<strong>selection</strong> (MARS) schemes and <strong>in</strong>frastructurehave been developed for “forwardbreed<strong>in</strong>g” of native genes and QTL forrelatively complex traits such as diseaseresistance, abiotic stress tolerance and gra<strong>in</strong>yield (Ribaut and Betrán, 1999; Ragot et al.,2000; Ribaut, Jiang and Hois<strong>in</strong>gton, 2000;Eath<strong>in</strong>gton, 2005; Crosbie et al., 2006).Simulation studies suggested that MAScould be effective for such traits under certa<strong>in</strong>conditions (Edwards and Page, 1994;Gimelfarb and Lande, 1994), but the <strong>in</strong>itialempirical attempts at such <strong>selection</strong> werenot successful (Stromberg, Dudley andRufener, 1994; Openshaw and Frascaroli,1997; Holland, 2004; Moreau, Charcossetand Gallais, 2004) except <strong>in</strong> the specialcase of sweetcorn (Edwards and Johnson,1994; Yousef and Juvik, 2001). The successreported for sweetcorn is due to the fact thatthe genetic base of sweetcorn is extremelynarrow relative to dent or fl<strong>in</strong>t maize; thus

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