Chapter 13 – Marker-<strong>assisted</strong> <strong>selection</strong> <strong>in</strong> sheep and goats 243and genetic evaluation are provided byexternal service agents. This situation is differentfrom that <strong>in</strong> poultry and pigs and tosome extent dairy, and more similar to that<strong>in</strong> beef cattle, <strong>in</strong> the sense that the bus<strong>in</strong>essunits that <strong>in</strong>vest <strong>in</strong> genetic <strong>in</strong>formation arenot the same as those provid<strong>in</strong>g geneticevaluation, and EBVs are available <strong>in</strong> thepublic doma<strong>in</strong>. Also, genotypic <strong>in</strong>formationis an explicit part of the market<strong>in</strong>g ofgenetic material. The result is that genotypic<strong>in</strong>formation is more likely to be usedoutside the usual EBV system, with thechance of be<strong>in</strong>g overvalued once the <strong>in</strong>vestmentis made. There is a place for MASand MAI based on genetic tests for clearlydemonstrated phenotypic effects with economicbenefit, for example for disease,fecundity and meat quality.The number of detected and confirmedQTL is low for sheep and goats and genemapp<strong>in</strong>g is less advanced than <strong>in</strong> otherlivestock species. There is significant <strong>in</strong>vestmentand progress be<strong>in</strong>g made <strong>in</strong> <strong>marker</strong>development and gene discovery, but itwill take some years before large amountsof genetic <strong>in</strong>formation become available atlittle cost, e.g. <strong>in</strong> the form of SNP chips.Until then, genotypic <strong>in</strong>formation will provideadditional <strong>selection</strong> criteria, mak<strong>in</strong>goptimal <strong>selection</strong> a greater challenge.Ultimately, the additional value of gene<strong>marker</strong>s will be greatest <strong>in</strong> breed<strong>in</strong>g programmesthat already use <strong>in</strong>tensive pedigreeand performance record<strong>in</strong>g, and it will helpto shift <strong>selection</strong> pressure towards traitsthat are hard to improve based on phenotypic(BLUP) <strong>selection</strong> (i.e. traits suchas fertility, disease resistance and carcassquality). It is not essential that genetictests are based on functional mutations,as gene <strong>marker</strong>s can have predictive valuedue to be<strong>in</strong>g <strong>in</strong> l<strong>in</strong>kage disequilibrium withfunctional genes. In breed<strong>in</strong>g programmeswithout extensive record<strong>in</strong>g, it is moreimportant to rely on direct <strong>marker</strong>s, butthis will only be valuable <strong>in</strong> practice ifgenes have very large economic effects.The same holds for genetic tests for dist<strong>in</strong>ctMendelian traits, but the overall valueof these traits <strong>in</strong> breed<strong>in</strong>g programmes islimited. In less-developed breed<strong>in</strong>g programmes,<strong>in</strong>vestments <strong>in</strong> pedigree andperformance record<strong>in</strong>g will most likely bemore profitable than <strong>in</strong>vestments <strong>in</strong> genetechnology.Application of MAS or MAI <strong>in</strong> manysheep and goat breed<strong>in</strong>g programmes <strong>in</strong>develop<strong>in</strong>g countries is not a priority, butopportunities exist, conditional on hav<strong>in</strong>g aclearly visible phenotypic effect and a programmebased on well-def<strong>in</strong>ed objectivesand performance based <strong>selection</strong>.AcknowledgementsThe author would like to thank the follow<strong>in</strong>gcolleagues for provid<strong>in</strong>g <strong>in</strong>formationabout breed<strong>in</strong>g and MAS programmes:Richard Apps, Alex Ball, Stephen Bishop,Didier Boichard, Joanne Con<strong>in</strong>gton, SchalkCloete, Oswald Matika, Eduardo Manfrediand John McEwan. Information from JillMaddox, her Web site and her commentson the <strong>marker</strong> maps were very helpful, andRobert Banks made useful comments andimprovements to the manuscript.ReferencesAc<strong>in</strong>, C., Mart<strong>in</strong>-Burriel, I., Monleon, E., Rodellar, C., Badiola, J. & Zaragoza, P. 2003.Characterization of the capr<strong>in</strong>e PrP-gene study of new polymorphisms and relationship with theresistance/susceptibility to the scrapie disease. Proc. Internat. Workshop on Major Genes <strong>in</strong> sheepand Goats. 8–11 December 2003. CD-ROM Communication No. 2-31. Toulouse, France.
244Marker-<strong>assisted</strong> <strong>selection</strong> – Current status and future perspectives <strong>in</strong> crops, livestock, forestry and fishBarbieri, M.E., Manfredi, E., Elsen, J.M., Ricordeau, G., Bouillon, J., Grosclaude, F., Mahe, M.F. &Bibe, B. 1995. Effects of the alpha-s1-case<strong>in</strong> locus on dairy performances and genetic parameters ofAlp<strong>in</strong>e goats. Genet. Sel. Evol. 27: 437–450.Barendse, W., Bunch, R., Thomas, M., Armitage, S., Baud, S. & Donaldson, N. 2004. The TG5 thyroglobul<strong>in</strong>gene test for a marbl<strong>in</strong>g quantitative trait loci evaluated <strong>in</strong> feedlot cattle. Austr. J. Exper.Agric. 44: 669 – 674.Barillet, F. 1997. Genetics of milk production. In L. Piper & A. Ruv<strong>in</strong>sky, eds. The genetics of sheep,pp 539–564. Wall<strong>in</strong>gford, UK, CABI.Barillet, F., Arranz, J.J. & Carta, A. 2005. Mapp<strong>in</strong>g quantitative trait loci for milk production andgenetic polymorphisms of milk prote<strong>in</strong>s <strong>in</strong> dairy sheep. Genet. Sel. Evol. 37 (Suppl. 1): S109–S123.Broad, T.E., Glass, B.C., Greer, G.J., Robertson, T.M., Ba<strong>in</strong>, W.E., Lord, E.A. & McEwan, J.C.2000. Search for a locus near to myostat<strong>in</strong> that <strong>in</strong>creases muscl<strong>in</strong>g <strong>in</strong> Texel sheep <strong>in</strong> New Zealand.Proc. New Zealand Soc. Anim. Prod. 60: 110–112.Carrick, M. 2005. Genotype by environment <strong>in</strong>teraction <strong>in</strong> wool sheep. Report SHGEN.036. NorthSydney, NSW, Australia, Meat & Livestock Australia.Cockett, N.E., Jackson, S.P., Shay, T.L., Farnir, F., Berghmans, S., Snowder, G.D., Nielsen, D.M. &Georges, M. 1996. Polar overdom<strong>in</strong>ance at the ov<strong>in</strong>e callipyge locus. Science 273: 236–238.Cockett, N.E, Shay, T.L., Beever, J.E., Nielsen, D., Albretsen, J., Georges, M., Peterson, K.,Stephens, A., Vernon, W., Timofeevskaia, O., South, S., Mork, J., Maciulis, A. & Bunch, T.D.1999. Localization of the locus caus<strong>in</strong>g Spider Lamb Syndrome to the distal end of ov<strong>in</strong>e chromosome6. Mamm. Genome 10: 35–38.Cockett, N.E., Smith, M.A., Bidwell, C.A., Segers, K., Hadfield, T.L., Snowder, G., Georges, M. &Charlier, C. 2005. The callipyge mutation and other genes that affect muscle hypertrophy <strong>in</strong> sheep.Genet. Sel. Evol. 37 (Suppl. 1): S65–S81.Con<strong>in</strong>gton, J., Bishop, S.C., Waterhouse, A. & Simm, G. 2004. A bioeconomic approach to deriveeconomic values for pasture-based sheep genetic improvement programmes. J. Anim. Sci. 82:1290–1304.Crawford, A.M., Dodds, K.G., Pierson, C.A., Ede, A.J., Montgomery, G.W., Garmonsway, H.G.,Beattie, A.E., Davies, K., Maddox, J.F., Kappes, S.W., Stone, R.T., Nguyen, T.C., Penty, J.M.,Lord, E.A., Broom, J.E., Buitkamp, J., Schwaiger, W., Epplen, J.T. Matthew, P., Matthews, M.E.,Hulme, D.J., Beh, K.J., McGraw, R.A. & Beattie, C.W. 1995. An autosomal genetic l<strong>in</strong>kage mapof the sheep genome. Genetics 140: 703–724.Davis, G.H. 2005. Major genes affect<strong>in</strong>g ovulation rate <strong>in</strong> sheep. Genet. Sel. Evol. 37 (Suppl. 1):S11–S23.Davis, G.H., Montgomery, G.W., Allison, A.J., Kelly R.W. & Bray, A.R. 1982. Segregation of amajor gene <strong>in</strong>fluenc<strong>in</strong>g fecundity <strong>in</strong> progeny of Booroola sheep. New Zealand J. Agric. Res. 25:525–529.Davis, G.H., Galloway, S.M., Ross, I.K., Gregan, S.M., Ward, J., Nimbkar, B.V., Ghalsasi, P.M.,Nimbkar, C., Gray, G.D., Subandriyo, I.I., Tiesnamurti, B., Martyniuk, E., Eythorsdottir, E.,Mulsant, P., Lecerf, F., Hanrahan, J.P., Bradford, G.E. & Wilson, T. 2002. DNA tests <strong>in</strong> prolificsheep from eight countries provide new evidence on orig<strong>in</strong> of the Booroola (FecB) mutation. Biol.Reprod. 66: 1869–1874.Dodds, K.G., Tate, M.L. & Sise, J.A. 2005. Genetic evaluation us<strong>in</strong>g parentage <strong>in</strong>formation fromgenetic <strong>marker</strong>s. J. Anim Sci. 83: 2271–2279.
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iiiContentsAcknowledgementsForeword
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Section v - marker-assisted selecti
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viiForewordSince almost the beginni
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ixAbbreviations and acronymsAATFAB-
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xiFIVIMSFNPFSCFSILGABIGASGCAGCPGDPG
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xiiiOBMOECDOIEOPVPAGEPBRsPCRPGRFAPI
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xvContributorsAmalia BaroneProfesso
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xviiElcio Perpétuo GuimarãesSenio
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xixAndrea SonninoSenior Agricultura
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Chapter 1Marker-assisted selection
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This book provides a comprehensive