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marker-assisted selection in wheat

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86Marker-<strong>assisted</strong> <strong>selection</strong> – Current status and future perspectives <strong>in</strong> crops, livestock, forestry and fishl<strong>in</strong>es. MAS for this trait has also been practisedat the University of Puerto Rico andat the Biotechnology Institute of Cuba.Bean common mosaic virus and beancommon mosaic necrotic virusBean common mosaic virus (BCMV) andthe related necrotic stra<strong>in</strong>s (bean commonmosaic necrotic virus [BCMNV]) areaphid-transmitted potyviruses that arefound worldwide and are seed-borne fromseason to season. BCMNV resistance isvery important <strong>in</strong> Africa where necroticstra<strong>in</strong>s are prevalent and has become arenewed priority for parts of the Caribbeanwhere necrotic stra<strong>in</strong>s have been discovered.BCMV is also endemic <strong>in</strong> the Andeanregion where it persists <strong>in</strong> farmer-saved seedand long-season climb<strong>in</strong>g beans. Climb<strong>in</strong>gbeans are grown <strong>in</strong> both <strong>in</strong>tensive (trellised/staked monoculture) and extensive (<strong>in</strong>tercropp<strong>in</strong>gwith maize) farm<strong>in</strong>g systems. Inboth systems the need to protect the cropfrom easily transmitted viral diseases suchas BCMV or BCMNV is great; however,very few climb<strong>in</strong>g beans have been bred forresistance to BCMV. A number of BCMV/BCMNV resistance genes have been tagged<strong>in</strong>clud<strong>in</strong>g the dom<strong>in</strong>ant I gene (with whichthe necrotic stra<strong>in</strong>s <strong>in</strong>teract to producenecrosis) and the recessive bc-3, bc-2 andbc-1 2 genes (Haley, Afanador and Kelly,1994; Melotto, Afanador and Kelly, 1996;Johnson et al., 1997; Miklas et al., 2000a).The genes can be dist<strong>in</strong>guished by <strong>in</strong>oculationwith different viral isolates, and a rangeof molecular <strong>marker</strong> tags are available foreach gene (reviewed <strong>in</strong> Kelly et al., 2003;Miklas et al., 2006). The dom<strong>in</strong>ant I genewas <strong>in</strong>corporated <strong>in</strong>to a wide range of smallseeded bush beans at CIAT, while resistantbush beans of the bush bean resistant toblack root (BRB) series carry<strong>in</strong>g recessivegenes were developed <strong>in</strong> the 1990s andhave been widely distributed as breed<strong>in</strong>gparents. The need to reselect the recessivegenes with confidence from segregat<strong>in</strong>gpopulations makes MAS a priority.CIAT started a collaborative project withthe Colombian national bean programmebased at the Colombian AgriculturalResearch Corporation (CORPOICA) <strong>in</strong>2002 to <strong>in</strong>trogress BCMV resistance genesfrom BRB l<strong>in</strong>es <strong>in</strong>to local landraces andimproved genotypes of Andean climb<strong>in</strong>gbeans (CIAT, 2002, 2003, 2004; Santana etal., 2004). Dur<strong>in</strong>g the breed<strong>in</strong>g programmefor BCMV and over the course of fouryears, MAS was used extensively basedprimarily on the SCAR <strong>marker</strong> ROC11developed for the bc-3 gene (Johnson etal., 1997) and the SCAR <strong>marker</strong> SW13 forthe I gene (Melotto, Afanador and Kelly,1996) along with virus screen<strong>in</strong>g to confirmthe <strong>selection</strong> of resistant progeny.The programme was successful <strong>in</strong> mov<strong>in</strong>gbc-3 resistance <strong>in</strong>to a background of creammottled and red mottled seed types forboth highland areas (known as Cargamantocommercial class) as well as mid-altitudeareas through triple-, double- and backcrosses.Although virus resistance was alsoscreened phenotypically, the frequency ofescape, the complex <strong>in</strong>teraction of multiplegenes and the recessive nature of mostof these made MAS the best option forbreed<strong>in</strong>g resistant varieties rapidly. In addition,as climb<strong>in</strong>g bean breed<strong>in</strong>g is a moretime-consum<strong>in</strong>g and expensive endeavourthan bush bean breed<strong>in</strong>g due to the longerseason, wider plant spac<strong>in</strong>g and need forstak<strong>in</strong>g material, MAS was also found to bea very effective measure to reduce breed<strong>in</strong>gcosts and save on breed<strong>in</strong>g nursery space.The implementation of MAS for BCMVwas based on a comb<strong>in</strong>ation of the previouslydeveloped SCAR <strong>marker</strong>s previouslymentioned and techniques developed at

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