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marker-assisted selection in wheat

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Chapter 4 – Marker-<strong>assisted</strong> <strong>selection</strong> <strong>in</strong> <strong>wheat</strong>: evolution, not revolution 55FHBThe importance of FHB is less <strong>in</strong> its effecton yield reduction, but rather on the potentiallydamag<strong>in</strong>g reduction <strong>in</strong> gra<strong>in</strong> qualityassociated with <strong>in</strong>fected gra<strong>in</strong>, which canbe heavily contam<strong>in</strong>ated by the fungal tricothec<strong>in</strong>tox<strong>in</strong>s. An important source ofFHB resistance orig<strong>in</strong>ates from the Ch<strong>in</strong>esevariety Sumai 3, and a major component ofthis resistance (up to 50 percent) has beenassociated with a s<strong>in</strong>gle QTL (Waldron etal., 1999; Anderson et al., 2001; Buerstmayret al., 2002). While this QTL is largelyeffective <strong>in</strong> prevent<strong>in</strong>g the spread of thepathogen follow<strong>in</strong>g <strong>in</strong>fection, a further QTLthat gives a significant degree of protectionaga<strong>in</strong>st <strong>in</strong>itial <strong>in</strong>fection has been mapped toa different chromosome (Buerstmayr et al.,2003). Selection for FHB resistance by conventionalmeans is complicated both by thequantitative nature of the Sumai 3 resistanceand by difficulties <strong>in</strong> ensur<strong>in</strong>g evenand reliable artificial <strong>in</strong>fections <strong>in</strong> breed<strong>in</strong>gnurseries. However, SSR-based MAS protocolshave been developed for both QTL(see http://mas<strong>wheat</strong>.ucdavis.edu/ andBuerstmayr et al., 2003), and the urgencyof breed<strong>in</strong>g for resistance has ensured that<strong>in</strong>creas<strong>in</strong>g use is be<strong>in</strong>g made of such assays.Both these QTL <strong>in</strong> concert do not expla<strong>in</strong>all the genetic resistance of Sumai 3 to FHB,but the rema<strong>in</strong>der appears to be determ<strong>in</strong>edby QTL of m<strong>in</strong>or effects and/or pleiotropiceffects associated with an ear morphology,which is <strong>in</strong>consistent with a northwestEuropean w<strong>in</strong>ter <strong>wheat</strong> ideotype.STBSTB of <strong>wheat</strong> is caused by the fungusMycosphaerella gram<strong>in</strong>icola (syn. Septoriatritici), and <strong>in</strong> recent years has become themajor leaf disease of <strong>wheat</strong> <strong>in</strong> many regionsof the world. In past years, good levels ofcontrol were achieved by the application ofstrobilur<strong>in</strong> fungicides, but their heavy usehas led to the emergence of pathogen stra<strong>in</strong>sthat cannot be so easily controlled by chemicalmeans. A number both of major genesgiv<strong>in</strong>g near-complete resistance to specificraces of the pathogen and of quantitativerace non-specific resistances with polygenic<strong>in</strong>heritance have been def<strong>in</strong>ed, andone of the former, Stb6, which maps closeto the SSR locus Xgwm369 on chromosome3A (Chartra<strong>in</strong>, Brad<strong>in</strong>g and Brown,2004), is common <strong>in</strong> many gene pools. Thisensures that the gene has been reta<strong>in</strong>ed <strong>in</strong>elite materials, and its known map positionhas made it relatively straightforward touse a <strong>marker</strong> assay to track its presence <strong>in</strong>breed<strong>in</strong>g populations.BYDVSignificant gra<strong>in</strong> yield losses are attributableto natural <strong>in</strong>fections of BYDV, and no majorsource of resistance has been identified todate <strong>in</strong> <strong>wheat</strong>. Control is achieved <strong>in</strong> theabsence of genetic resistance by <strong>in</strong>secticidalspray, which is associated with both aneconomic and an environmental cost.However, a potent resistance is present <strong>in</strong> therelated species Th<strong>in</strong>opyrum <strong>in</strong>termedium.It is possible to generate sexual hybridsbetween <strong>wheat</strong> and this grass, but the F 1plants are self-sterile and either have tobe rescued by chromosome doubl<strong>in</strong>g orback-crossed to <strong>wheat</strong>. By this route, adistal segment of the grass chromosomethat carries the BYDV resistance geneBdv2 has been <strong>in</strong>troduced <strong>in</strong>to <strong>wheat</strong>. Asthis <strong>in</strong>trogression comprises a significantlength of non-<strong>wheat</strong> chromosome, it hasbeen relatively straightforward to generate<strong>marker</strong>s suitable for MAS use (Ayala et al.,2001a; Zhang et al., 2004). A MAS approachfor screen<strong>in</strong>g is attractive because artificial<strong>in</strong>oculation <strong>in</strong>volves the propagation ofvirus-bear<strong>in</strong>g aphids, while natural <strong>in</strong>fections

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