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marker-assisted selection in wheat

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Chapter 12 – Marker-<strong>assisted</strong> <strong>selection</strong> <strong>in</strong> dairy cattle 205scored from the centromere. Chromosomesare denoted with the prefix “BTA” (B.taurus). Similar to other mammals, thebov<strong>in</strong>e DNA <strong>in</strong>cludes 3×10 9 base pairs(bp), and the map length is approximately3 000 cM. The human genome is estimatedto encode 20 000–25 000 prote<strong>in</strong>-cod<strong>in</strong>ggenes (International Human GenomeSequenc<strong>in</strong>g Consortium, 2004), and it canbe assumed that the number of genes <strong>in</strong>other mammals, <strong>in</strong>clud<strong>in</strong>g cattle, should bequite similar. Thus, a s<strong>in</strong>gle map unit, onaverage, <strong>in</strong>cludes approximately eight genesand one million bp.As <strong>in</strong> other animal species, microsatellitesare still the <strong>marker</strong> of choice for mapconstruction due to their prevalence andhigh polymorphism. Although s<strong>in</strong>glenucleotide polymorphisms (SNPs) aremuch more prevalent, genetic maps basedon SNPs are still <strong>in</strong> the future. Morethan 50 000 SNPs have been identified <strong>in</strong>humans, but only several thousand havebeen validated <strong>in</strong> cattle (www.afns.ualberta.ca/Hosted/Bov<strong>in</strong>e%20Genomics/), andrates of polymorphism are generallyunknown. With the completion of the sixfoldcoverage of the bov<strong>in</strong>e genome bythe Bov<strong>in</strong>e Genome Sequenc<strong>in</strong>g Project atBaylor College of Medic<strong>in</strong>e (www.hgsc.bcm.tmc.edu/projects/bov<strong>in</strong>e/) many moreSNPs will be identified.Several genetic maps are available on the<strong>in</strong>ternet. The United States Meat AnimalResearch Center (MARC) (www.marc.usda.gov/) <strong>in</strong>cludes thousands of <strong>marker</strong>s,chiefly microsatellites. The ArkDB databasesystem, hosted at Rosl<strong>in</strong> Institute,<strong>in</strong>cludes data from several published maps(www.thearkdb.org/). The CommonwealthScientific and Industrial Research Organization(CSIRO) livestock <strong>in</strong>dustries cattlegenome <strong>marker</strong> map is built upon dataprovided by the University of Sydney’scomparative location database (www.livestockgenomics.csiro.au/perl/gbrowse.cgi/cattlemap/). This map comb<strong>in</strong>ed all publicly-availablemaps <strong>in</strong>to a s<strong>in</strong>gle <strong>in</strong>tegratedmap that currently <strong>in</strong>cludes 9 400 <strong>marker</strong>s.Methods of QTL detectionsuitable for commercial dairycattle populationsDetection of QTL requires generation ofl<strong>in</strong>kage disequilibrium (LD) between thegenetic <strong>marker</strong>s and QTL. In plants, this isgenerally accomplished by crosses between<strong>in</strong>bred l<strong>in</strong>es but, for the reasons noted <strong>in</strong>the <strong>in</strong>troduction, this is not a viable optionfor dairy cattle <strong>in</strong> developed countries, <strong>in</strong>which all analyses must be based on analysisof the exist<strong>in</strong>g population. Detection ofQTL <strong>in</strong> develop<strong>in</strong>g countries is consideredbelow. For advanced commercial populations,the “daughter” and “granddaughter”designs, which make use of the existenceof large half-sib families, are most appropriatefor QTL analysis (Weller, Kashi andSoller, 1990). These designs are presented<strong>in</strong> Figures 2 and 3.Both designs are similar to the backcrossdesign for crosses between <strong>in</strong>bred l<strong>in</strong>es <strong>in</strong>that only the alleles of one parent are followed<strong>in</strong> the progeny. Thus, similar to thebackcross design, dom<strong>in</strong>ance cannot beestimated. These designs differ from crossesbetween <strong>in</strong>bred l<strong>in</strong>es <strong>in</strong> that several familiesare analysed <strong>in</strong> which the l<strong>in</strong>kage phasebetween QTL and genetic <strong>marker</strong>s maydiffer. In addition, any specific QTL will beheterozygous <strong>in</strong> only a fraction of the families<strong>in</strong>cluded <strong>in</strong> the analysis. Thus, QTLeffects must be estimated with<strong>in</strong> families,and these designs are therefore less powerfulper <strong>in</strong>dividual genotyped than designsbased on crosses between <strong>in</strong>bred l<strong>in</strong>es.The granddaughter design has theadvantage of greater statistical power per

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