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marker-assisted selection in wheat - ictsd

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Chapter 12 – Marker-<strong>assisted</strong> <strong>selection</strong> <strong>in</strong> dairy cattle 217ferentially expressed <strong>in</strong> the mammary glanddur<strong>in</strong>g lactation, as compared with the liver.Furthermore, anti-sense SPP1 transgenicmice displayed abnormal mammary glanddifferentiation and milk secretion (Nemiret al., 2000).Schnabel et al. (2005) found concordancebased on four heterozygous and fourhomozygous sires for the United StatesHolste<strong>in</strong> population, as determ<strong>in</strong>ed by agranddaughter design, while Cohen-Z<strong>in</strong>deret al. (2005) found concordance for threeheterozygous and 15 homozygous siresfrom both the United States and IsraeliHolste<strong>in</strong> populations. Cohen-Z<strong>in</strong>der et al.(2005) also analysed the site proposed bySchnabel et al. (2005), and found that thissite was hyper-variable <strong>in</strong> that at leastfour s<strong>in</strong>gle nucleotide changes were foundwith<strong>in</strong> the 20 bp region centred on thepoly-A sequence. Eight of n<strong>in</strong>e Israelisires analysed by the daughter design wereheterozygous for at least one of thesepolymorphisms.Many studies have found a QTLaffect<strong>in</strong>g all five milk production traits andSCS near the middle of BTA 20. Blott et al.(2003) claimed that a mis-sense mutation<strong>in</strong> the bov<strong>in</strong>e growth hormone receptorwas responsible for the QTL affect<strong>in</strong>gmilk yield and composition on BTA 20,but did not f<strong>in</strong>d concordance for the bullsheterozygous for the QTL. Thus, this polymorphismmay be responsible for onlypart of the observed effect on BTA 20, ormay be a physiologically neutral mutation<strong>in</strong> LD with the QTN.For both the QTL on BTA 6 and 14,the polymorphisms analysed apparently donot account for the entire effect observed <strong>in</strong>these chromosomal regions (Bennewitz etal., 2004a; Kuhn et al., 2004; Cohen-Z<strong>in</strong>deret al., 2005). The effect associated with themis-sense mutation <strong>in</strong> ABCG2 expla<strong>in</strong>s theentire effect observed on milk yield and fatand prote<strong>in</strong> concentration, but does notexpla<strong>in</strong> the effects associated with fat andprote<strong>in</strong> yield. It is likely that <strong>in</strong> the nearfuture additional QTN will be resolved.As noted, the meta-analysis (Khatkar et al.,2004) found significant effects on BTA 1, 3,9 and 10, <strong>in</strong> addition to the effects describedon BTA 6, 14 and 20.Methods and theory for MAS <strong>in</strong>dairy cattleConsider<strong>in</strong>g the long generation <strong>in</strong>terval,the high value of each <strong>in</strong>dividual, the verylimited female fertility and the fact thatnearly all economic traits are expressedonly <strong>in</strong> females, it would seem that dairycattle should be a nearly ideal species forapplication of MAS. However, most theoreticalstudies have been rather pessimisticwith respect to the expected ga<strong>in</strong>s that canbe obta<strong>in</strong>ed by MAS. As noted by Weller(2001), MAS can potentially <strong>in</strong>crease annualgenetic ga<strong>in</strong> by <strong>in</strong>creas<strong>in</strong>g the accuracy ofevaluation, <strong>in</strong>creas<strong>in</strong>g the <strong>selection</strong> <strong>in</strong>tensityand decreas<strong>in</strong>g the generation <strong>in</strong>terval.The follow<strong>in</strong>g dairy cattle breed<strong>in</strong>gschemes that <strong>in</strong>corporate MAS have beenproposed:• a standard PT system, with <strong>in</strong>formationfrom genetic <strong>marker</strong>s be<strong>in</strong>g used to<strong>in</strong>crease the accuracy of sire evaluations<strong>in</strong> addition to phenotypic <strong>in</strong>formationfrom daughter records (Meuwissen andvan Arendonk, 1992);• a MOET nucleus breed<strong>in</strong>g scheme <strong>in</strong>which <strong>marker</strong> <strong>in</strong>formation is used toselect sires for service <strong>in</strong> the MOETpopulation, <strong>in</strong> addition to phenotypic<strong>in</strong>formation on half-sisters (Meuwissenand van Arendonk, 1992);• PT schemes <strong>in</strong> which <strong>in</strong>formation ongenetic <strong>marker</strong>s is used to preselect youngsires for entrance <strong>in</strong>to the PT (Kashi,

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