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marker-assisted selection in wheat

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254Marker-<strong>assisted</strong> <strong>selection</strong> – Current status and future perspectives <strong>in</strong> crops, livestock, forestry and fisheucalypt breed<strong>in</strong>g practice and the moleculartools available or <strong>in</strong> development.To set the stage for a realistic appraisal ofMAS for Eucalyptus, a brief <strong>in</strong>troductionis presented of the ma<strong>in</strong> features of moderneucalypt breed<strong>in</strong>g and clonal forestry <strong>in</strong>order to provide a better understand<strong>in</strong>gof the challenges and opportunities thatlie ahead for cost-efficient molecularbreed<strong>in</strong>g. Follow<strong>in</strong>g this section, some currentlow technological <strong>in</strong>put applicationsof molecular <strong>marker</strong>s <strong>in</strong> support of operationalbreed<strong>in</strong>g are presented, such as thequantification of genetic diversity and relationships,the analysis of mat<strong>in</strong>g patternsand paternity <strong>in</strong> seed orchards and f<strong>in</strong>gerpr<strong>in</strong>t<strong>in</strong>gfor quality assurance and qualitycontrol of clonal propagation. With<strong>in</strong> theframework of MAS for trait advancement,after review<strong>in</strong>g the status of QTL mapp<strong>in</strong>g<strong>in</strong> Eucalyptus, the challenges and somerealistic prospects for the application ofMAS to improve relevant traits are outl<strong>in</strong>ed.F<strong>in</strong>ally, with the expected availabilityof a draft of the whole Eucalyptus genomewith<strong>in</strong> the next years, a succ<strong>in</strong>ct summaryis presented on the prospects of advanc<strong>in</strong>ggenomic approaches for gene identificationand subsequent application of MAS.Eucalyptus breed<strong>in</strong>g and plantationforestryEucalyptus domesticationEucalypts spread rapidly around the worldfollow<strong>in</strong>g their discovery by Europeans<strong>in</strong> the late eighteenth century (Eldridgeet al., 1993). They were <strong>in</strong>troduced <strong>in</strong>tocountries such as Brazil, Chile, France,India, Portugal and South Africa <strong>in</strong> thefirst quarter of the 1800s (Doughty, 2000)and rapidly adopted <strong>in</strong> forest plantationsas their fast growth and good adaptabilitybecame known. Dur<strong>in</strong>g the n<strong>in</strong>eteenthand twentieth centuries, large quantitiesof seeds were collected and distributeddirectly from Australia through a numberof seed collection expeditions carried outboth by government organizations andprivate forestry companies throughout theworld.Eucalyptus species have a mixed mat<strong>in</strong>gsystem, but are predom<strong>in</strong>antly outcrossersand animal poll<strong>in</strong>ated. High levels of outcross<strong>in</strong>gare ma<strong>in</strong>ta<strong>in</strong>ed by protandry andvarious <strong>in</strong>complete pre- and post-zygoticbarriers to self-fertilization <strong>in</strong>clud<strong>in</strong>g strong<strong>selection</strong> aga<strong>in</strong>st the products of <strong>in</strong>breed<strong>in</strong>g(Pryor, 1976). Although the major eucalyptsubgenera do not hybridize <strong>in</strong> nature,hybridization among species with<strong>in</strong> thesame subgenus has been detected, oftenmak<strong>in</strong>g separation of species difficult (Pryorand Johnson, 1971). Hybridization becomesmore frequent <strong>in</strong> exotic conditions outsidethe natural species range. In fact, this propertyhas been widely exploited by eucalyptbreeders who take advantage of the naturallyoccurr<strong>in</strong>g genetic variation for growthand wood properties among species (deAssis, 2000). Several artificial hybrid comb<strong>in</strong>ationshave been produced, althoughhybrid <strong>in</strong>viability tends to <strong>in</strong>crease with<strong>in</strong>creas<strong>in</strong>g taxonomic distance between theparents (Griff<strong>in</strong>, Burgess and Wolf, 1988;Potts and Dungey, 2004).In several countries the cont<strong>in</strong>ued plantationfrom local seed sources gave riseto landraces adapted to the specific environmentof the country (Eldridge et al.,1993). Seed collections from such localexotic plant<strong>in</strong>gs of multiple species becamecommon and where plant<strong>in</strong>gs occurred, F 1hybrids were derived (Potts and Dungey,2004). While several of these F 1 hybrids performedwell, especially when deployed asclones, seed collection from hybrid standsoften resulted <strong>in</strong> plantations that performedpoorly <strong>in</strong> subsequent generations and were

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