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marker-assisted selection in wheat - ictsd

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Chapter 7 – Marker-<strong>assisted</strong> <strong>selection</strong> <strong>in</strong> common beans and cassava 83Common beans: importance andgeneticsCommon beans (Phaseolus vulgaris L.)are the most important gra<strong>in</strong> legume fordirect human consumption, especially <strong>in</strong>Lat<strong>in</strong> America and eastern and southernAfrica. They are seed-propagated, true diploids(2n = 22) and have a relatively smallgenome (650 Mb) (Broughton et al., 2003).Orig<strong>in</strong>at<strong>in</strong>g <strong>in</strong> the Neotropics, commonbeans were domesticated <strong>in</strong> at least twomajor centres <strong>in</strong> Mesoamerica and theAndes (Gepts, 1988) and possibly <strong>in</strong> athird m<strong>in</strong>or centre <strong>in</strong> the northern Andes(Islam et al., 2002). Wide DNA polymorphismis expressed between the twomajor gene pools. Mesoamerican beanstypically have small to medium size seedsand can be classed <strong>in</strong>to four races that aredist<strong>in</strong>guished by randomly amplified polymorphicDNA (RAPD) polymorphisms(Beebe et al., 2000). Andean beans usuallyhave medium to large seeds, and landraceshave been classed <strong>in</strong>to three races basedon plant morphology and agro-ecologicaladaptation (S<strong>in</strong>gh, Gepts and Debouck,1991). These can be differentiated by microsatellites(M. Blair, unpublished data) butthe genetic distance among Andean races isnarrower than that among Mesoamericanraces (Beebe et al., 2001). A large numberof gene tagg<strong>in</strong>g studies have been conducted<strong>in</strong> common beans, predom<strong>in</strong>antlywith RAPD <strong>marker</strong>s, some of which havebeen converted subsequently to sequencecharacterized amplified regions (SCARs;reviewed most recently by Miklas et al.,2006).Beans display a wide range of growthhabits (Van Schoonhoven and Pastor-Corrales, 1987), from determ<strong>in</strong>ate bushtypes, to <strong>in</strong>determ<strong>in</strong>ate upright or v<strong>in</strong>ybush types, to vigorous climbers. Bushtypes are the most widely grown, and are arelatively short season crop, matur<strong>in</strong>g <strong>in</strong> aslittle as 60 days from seed<strong>in</strong>g <strong>in</strong> a tropicalclimate and yield<strong>in</strong>g from 700 to 2 000 kg/ha on average. On the other hand, <strong>in</strong> smallholderagriculture where land is scarce,labour-<strong>in</strong>tensive, high-yield<strong>in</strong>g climb<strong>in</strong>gbeans enjoy cont<strong>in</strong>u<strong>in</strong>g or even expand<strong>in</strong>gpopularity. Climb<strong>in</strong>g beans can mature <strong>in</strong>100 to 120 days at mid-elevations, but candelay as long as ten months at higher elevationsand can produce the highest yields forthe crop, up to 5 000 kg/ha. These featureshave significant implications for breed<strong>in</strong>gprogrammes. In bush types it is possibleto obta<strong>in</strong> up to three cycles per year <strong>in</strong>the field, or even four cycles <strong>in</strong> greenhouseconditions. Breed<strong>in</strong>g bush beans isthus quite agile with regard to advance ofgenerations, although seed harvest of <strong>in</strong>dividualplants is sometimes limited. Withclimb<strong>in</strong>g beans, on the other hand, at bestit is possible to obta<strong>in</strong> two cycles per yearwith field grown plants, while manag<strong>in</strong>gclimb<strong>in</strong>g beans <strong>in</strong> the greenhouse is logisticallydifficult. However, while bush beansproduce on average 20 to 50 seeds/plant,<strong>in</strong>dividual plants of climb<strong>in</strong>g beans oftenproduce enough seeds to plant several rows(100 to 150 seeds).Beans are self-poll<strong>in</strong>at<strong>in</strong>g and thusbreed<strong>in</strong>g methods for autogamous cropsare employed. Pedigree <strong>selection</strong> or someadaptation thereof is most common, andboth recurrent (Muñoz et al., 2004) andadvanced (or <strong>in</strong>bred) backcross<strong>in</strong>g (Sullivanand Bliss, 1983; Buendia et al., 2003; Blair,Iriarte and Beebe, 2003) have been used.Recurrent <strong>selection</strong> has also been employed(Kelly and Adams, 1987; Beaver et al.,2003) but seldom <strong>in</strong> a formal sense with adef<strong>in</strong>ed population structure. S<strong>in</strong>gh et al.(1998) suggested a system that they calledgamete <strong>selection</strong> <strong>in</strong> which <strong>in</strong>dividual F 1plants of multiple parent crosses give rise

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