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marker-assisted selection in wheat

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Chapter 3 – Molecular <strong>marker</strong>s for use <strong>in</strong> plant molecular breed<strong>in</strong>g and germplasm evaluation 41Yoshimura et al., 1998; Wang et al., 1999;Bryan et al., 2000; Sun et al., 2004) andmany other genes for disease resistancehave been tagged with l<strong>in</strong>ked <strong>marker</strong>s. Thisopens the door for targeted approaches toMAS (Valent et al., 2001). While the diseaseresistance literature is too vast to summarizehere, it is important to note that advances <strong>in</strong>this area are hav<strong>in</strong>g an impact on varietalimprovement programmes (www.syix.com/rrb/98rpt/MarkerAssist.htm). Pyramid<strong>in</strong>gof resistance genes <strong>in</strong>to a s<strong>in</strong>gle varietyand the construction of multil<strong>in</strong>e varieties,each with one or more R genes (resistancegenes) that can be used <strong>in</strong> various comb<strong>in</strong>ations,are under way to develop moredurable forms of disease and <strong>in</strong>sect resistance(Yoshimura et al., 1992; Yoshimura etal., 1995; Hittalmani et al., 1995; Blair andMcCouch, 1997; Ndjiondjop et al., 1999;Davierwala et al., 2001; Su et al., 2002;Conaway-Bormans et al., 2003; Lorieux etal., 2003; Hayashi et al., 2004).Marker-based <strong>selection</strong> is also helpful<strong>in</strong> attempts to transfer genes from exoticgermplasm <strong>in</strong>to cultivated l<strong>in</strong>es. In rice,several workers have used RFLP andSSR <strong>marker</strong>s to monitor <strong>in</strong>trogression ofbrown planthopper resistance from O.offic<strong>in</strong>alis (Kochert, Jena and Zhao, 1990),bacterial blight resistance from O. longistam<strong>in</strong>ata(Ronald et al., 1992), alum<strong>in</strong>umtolerance or yield and quality-related traitsfrom O. rufipogon (Nguyen et al., 2002;Thomson et al., 2003; Sept<strong>in</strong><strong>in</strong>gsih et al.,2003a, b) or from other wild species such asO. glumaepatula (Brondani et al., 2002) orO. glaberrima (Jones et al., 1997; Lorieuxet al., 2003) <strong>in</strong>to cultivated O. sativa backgrounds.Marker-<strong>assisted</strong> <strong>in</strong>trogressionstrategies have also been used <strong>in</strong> a numberof livestock breed<strong>in</strong>g programmes but,because of longer generation <strong>in</strong>tervals andlower reproductive rates, this is generallyfeasible for genes of large effect (Dekkers,2004; Chapter 10). Identify<strong>in</strong>g the recomb<strong>in</strong>antswith the least amount of donorDNA flank<strong>in</strong>g the genes of <strong>in</strong>terest isenhanced by the use of molecular <strong>marker</strong>s(Monna et al., 2002; Takeuchi et al., 2003;Blair, Panaud and McCouch, 2003). Inthese examples, MAS offers a powerfulstrategy for mak<strong>in</strong>g efficient use of thewealth of useful genetic variation that exists<strong>in</strong> the early landraces and wild speciesof cultivated food crops (Tanksley andMcCouch, 1997).As this k<strong>in</strong>d of <strong>in</strong>formation accumulates,MAS permits rapid identification of <strong>in</strong>dividualsthat may conta<strong>in</strong> only one geneticcomponent of a complex trait. Once identified,such an <strong>in</strong>dividual can be crossedwith another <strong>in</strong>dividual <strong>in</strong> a breed<strong>in</strong>g programmeso that multiple, complementarygenes are comb<strong>in</strong>ed to optimize a quantitatively<strong>in</strong>herited trait. Individuals conta<strong>in</strong><strong>in</strong>gonly one gene of <strong>in</strong>terest often defy accuratephenotypic identification wherepolygenic traits are concerned because varioustypes of epistasis, or gene <strong>in</strong>teraction,may be required to generate the phenotypeof <strong>in</strong>terest (Yamamoto et al., 2000; Zhenget al., 2000).L<strong>in</strong>kage disequilibrium (LD) mapp<strong>in</strong>gis another <strong>marker</strong>-<strong>assisted</strong> approachthat provides important <strong>in</strong>formationthat is immediately relevant to breed<strong>in</strong>gprogrammes (Rem<strong>in</strong>gton, Ungererand Purugganan, 2001; Fl<strong>in</strong>t-Garcia,Thornsberry and Buckler, 2003). Us<strong>in</strong>gcollections of distantly related germplasmaccessions rather than populations derivedfrom bi-parental crosses allows researchersto explore the relationship betweenphenotype and genotype <strong>in</strong> materials thathave been amply tested over years andenvironments, often as part of an appliedbreed<strong>in</strong>g programme. This provides

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