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4,ü,,,,,ü0ëU+&ÿU‹ÿÀÀÀÝ ƒ;¤T!ÝÓ  ÓÝ  ÝÔ_ÔÓ@ÓSouthern€Illinois€University,ÌDepartment€of€Plant€and€Soil€Science,ÌMolecular€Science€Program,ÌCarbondale,€IL€62901.ÌÔ€6X"FXX!XÔÓ  ÓÌòòGenetic€analysis€of€resistance€to€manganese€toxicity€in€soybean€óóÐ ’ â Ѐà@LL'ìàòòusing€molecular€markersóó.ˆÐ Œ Ü ÐÌòòóóà  àManganese€toxicity€is€a€problem€associated€with€soybeans€grownÐ € Ð Ðon€acidic€soils€that€may€limit€soybean€yields€in€the€Southern€US.€ÏEvery€year€farmers€spend€thousands€applying€lime€to€reduce€the€pHÏlevel€in€acid€soils€and€thereby€reduce€the€effects€of€Mn€toxicity.ÏHowever,€the€application€of€lime€occurs€weather€permitting.€SoybeanÏcultivars€with€improved€manganese€toxicity€resistance€might€protectÏyields€in€acid€soils,€particularly€in€years€when€Spring€rainfallÏprecludes€the€application€of€lime.€€Many€studies€have€shown€thatÏsoybean€cultivars€vary€in€plant€growth€response€to€toxic€levels€ofÏMn€in€solution€(Carter€et€al.,€1975;€Ô_ÔHenanÔ_Ô€and€Campbell,€1980,€Ô_ÔOhkiÔ_ÔÏet€al.,€1987).€€The€molecular€map€of€Essex€and€Forrest€soybeanÏmakes€it€possible€to€map€manganese€tolerance€Quantitative€TraitÏLoci€(Ô_ÔQTLÔ_Ô)€in€adapted€Ô_ÔgermplasmÔ_Ô.€€Since€Essex€and€Forrest€contrastÏfor€their€resistance€to€Mn€toxicity€(Carter€et€al.,€1975)€and€SDSÏresistance€we€can€use€the€Ô_ÔExFÔ_Ô€Ô_ÔRILÔ_Ô€population€and€molecular€map€forÏcomparisons€with€Sudden€Death€Syndrome€resistance€Ô_ÔQTLÔ_Ô€(Ô_ÔHnetkovskyÔ_ÔÏet€al.,€1996;€Chang€et€al.,€1996).€€Forrest€is€resistant€to€SDS€butÏsensitive€to€Mn€toxicity€while€Essex€is€resistant€to€Mn€toxicityÏbut€sensitive€to€SDS.€ÌÌòòMaterials€and€MethodsóóÐ  X Ðà  àMn€resistance€was€analyzed€in€Essex,€Forrest€and€their€Fòò2óó€Ð !R Ðderived€Fòò5:9óó€progenies.€€Traits€were€determined€in€two€experimentsÐ ü!L Ð(four€replicates€per€experiment€with€plants€grown€€in€hydroponicsÏfor€28€days€containing€125€Ô_ÔððMÔ_Ô€€Mn,€the€toxic€concentration.€TraitsÏmeasured€included€Ô_ÔchlorosisÔ_Ô€(Mg1€at€14€days,€Mg6€at€28€days),€wiltÏ(Mg5),€height€(Mg2),€shoot€weight€(Mg3=fresh),€(Mg7=dry)€and€rootÏweight€(Mg4=fresh),€(Mg8=dry).€€The€relative€contributions€ofÏindividual€genes€to€total€manganese€tolerance€was€estimated€byÏtesting€for€associations€with€particular€chromosomal€segments€byÏÔ_ÔMapmakerÔ_Ô/€Ô_ÔQTLÔ_Ô€1.1€and€with€particular€markers€by€one€way€Ô_ÔANOVAÔ_Ô.ÌÌòòResultsóóÐ À+'( Ðà  àOne€hundred€and€thirty„five€markers€covering€more€than€2000ÏÔ_ÔcMÔ_Ô,€including€ten€linkage€groups,€were€tested€for€association.€Ð ´-)* ÐÔ_ÔTwenty€markers€were€associated€(P€<€0.05)€with€at€least€one€MnÏtrait.€The€twenty€markers€identified€at€least€€at€least€sixÏindependent€QTL€conditioning€Mn€resistance€as€judged€by€mapÏposition€(€at€P€<€0.005,€Table€1).€€Three€of€these€QTL€defined€by;ÏOC01,€(QTL€N),€€OEO4€and€OE02€(QTL€2G)€and€OG13€(QTL€1G)€were€lessÏthan€15€cM€from€the€map€positions€of€SDS€resistance€QTL€(HnetkovskyÏet€al.,€1996;€Chang€et€al.,€1996).€€Ì€€€€€€Markers€OCO1,€€OF04€and€OO04,€that€map€to€a€20€cM€interval€onÏlinkage€group€N€(Hnetkovsky€et€al.,€1996),€were€associatedÏsignificantly€with€most€Mn€toxicity€traits€including;€chlorosis,Ïwilt,€height,€root€and€shoot€€weight.€The€QTL€explained€up€to€36%Ïof€the€trait€variability€(Table€1).€€In€contrast€the€SDS€QTL€onÏlinkage€group€G€(Chang€et€al.,€1996)€was€only€associated€withÏchlorosis€(OE02)€or€chlorosis,€wilt€and€shoot€weight€(OG13).€ForÏeach€of€these€QTL€the€beneficial€allele€for€Mn€resistance€wasÏderived€from€Essex€whereas€the€SDS€resistance€allele€was€derivedÏfrom€Forrest.€Therefore,€QTL€for€Mn€toxicity€and€SDS€resistancesÏare€in€repulsion.Ì€€€€€€Three€unlinked€markers€found€associated€with€Mn€toxicityÏresistance€are€not€associated€with€SDS€resistance;€OU15€€wasÏassociated€with€the€chlorosis€and€height€Mn€toxicity€traits.€OC10€Ï€was€associated€with€the€wilt€Mn€toxicity€trait.€€OC11€wasÏassociated€with€the€wilt,€chlorosis,€root€and€shoot€dry€weight€MnÏtoxicity€traits.€The€QTL€explained€up€to€36%€of€the€traitÏvariability€(Table€1).€The€beneficial€alleles€for€these€QTL€wereÏderived€from€Essex.Ì€€€€€€€Jointly€the€QTL€can€explain€about€75%€of€each€trait.ÏTherefore,€resistance€to€Mn€toxicity€in€the€Essex€soybean€isÏcontrolled€by€3„4€major€QTL.€€Since€SDS€resistance€and€Mn€toxicityÏresistance€QTL€were€€in€repulsion,€either€linkage€exists€or€theÏphysiological€bases€of€the€Mn€and€SDS€responses€share€some€commonÏelements.€Leaf€Mn€deficiency€has€been€reported€to€be€associatedÏwith€increased€SDS€severity€in€the€field€(Rupe€et€al.,€1994).€ÏMarker€saturation€and€subline€population€analyses€may€allow€the€QTLÏto€be€separated.€Markers€capable€of€distinguishing€the€QTL€could€beÏused€to€break€linkage€between€SDS€resistance€and€Mn€toxicityÏsusceptibility.€The€greenhouse€Mn€resistance€reaction€may€predictÏpartial€SDS€resistance€better€than€assays€including€òòFusariumÐ Ò("$% Ðsolanióó,€the€incitant€of€SDS€(see€Torto€et€al.,€1996).Ð þ)N%& ÐÌÐ  $,t'( ÐTable€1:€Molecular€markers€associated€with€Mn€toxicity.Ìòò€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€óóÐ ªú ÐMarker€€€LGà ¸ ànà  àà h àTraità  àà p àòòRóóòò2óóà È àà  àòòPóó€€€€€€Ð ¤ô Ðòò€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€óóÐ Ð  ÐOF04òò1000óóà ` àNà ¸ à31à  àà h àMg1à À àà  àà p à0.28à È àà  à0.002Ð Ê Ðà  àà ` à€€€€€à  à€€€€€Mg2à À àà  àà p à0.14à È à€€€€€0.033Ð Ä  Ѐ€€€€€€€€€€€€€€€€€€€€€€€€Mg3à À àà  àà p à0.20à È à€€€€€0.011€€€€€€€€€€€Ð ¾  Ѐ€€€€€€€€€€€€€€€€€€€€€€€€Mg5à À àà  àà p à0.23à È àà  à0.006Ð ¸  Ѐ€€€€€€€€€€€€€€€€€€€€€€€€Mg7à À àà  àà p à0.25à È àà  à0.004Ð ²  Ѐ€€€€€€€€€€€€€€€€€€€€€€€€Mg8€€€€€€€€€€€€0.21€€€€€€0.009Ì€€€€€€€€€€€€€€€€€€€€€€€€€Mg4€€€€€€€€€€€€0.23€€€€€€0.004ÌOC01òò650óóà ` àNà ¸ à29à  àà h àMg1à À àà  àà p à0.20à È àà  à0.014Ð  ð  Ѐ€€€€€€€€€€€€€€€€€€€€€€€€Mg2à À àà  àà p à0.21à È àà  à0.013Ð šê  Ѐ€€€€€€€€€€€€€€€€€€€€€€€€Mg3à À àà  àà p à0.18à È àà  à0.021Ð ”ä  Ѐ€€€€€€€€€€€€€€€€€€€€€€€€Mg5à À àà  àà p à0.36à È àà  à0.006Ð ŽÞ  Ѐ€€€€€€€€€€€€€€€€€€€€€€€€Mg7à À àà  àà p à0.11à È à€€€€€0.076Ð ˆØ ÐOO04òò1075óóà ` àNà ¸ à32à  àà h àMg3à À àà  àà p à0.14à È àà  à0.032Ð ‚Ò Ѐ€€€€€€€€€€€€€€€€€€€€€€€€Mg5à À àà  àà p à0.21€€€€€€0.008Ð |Ì Ѐ€€€€€€€€€€€€€€€€€€€€€€€€Mg7à À àà  àà p à0.15à È à€€€€€0.027Ð vÆ Ѐ€€€€€€€€€€€€€€€€€€€€€€€€Mg8à À àà  àà p à0.10à È àà  à0.076Ð pÀ ÐOE04òò450óóà ` àG€€€€28€€€€€€€€Mg1€€€€€€€€€€€€0.26€€€€€€0.006Ð jº ÐOE02òò1000óóà ` àG€€€€16€€€€€€€€Mg1€€€€€€€€€€€€0.35€€€€€€0.016Ð d´ Ѐ€€€€€€€€€€€€€€€€€€€€€€€€Mg2€€€€€€€€€€€€0.34€€€€€€0.02Ì€€€€€€€€€€€€€€€€€€€€€€€€€Mg7€€€€€€€€€€€€0.30€€€€€€0.03Ì€€€€€€€€€€€€€€€€€€€€€€€€€Mg8€€€€€€€€€€€€0.30€€€€€€0.03ÌOG13òò490óóà ` àG€€€€32€€€€€€€€Mg3€€€€€€€€€€€€0.15€€€€€€0.003Ð Lœ Ѐ€€€€€€€€€€€€€€€€€€€€€€€€Mg6€€€€€€€€€€€€0.18€€€€€€0.008Ì€€€€€€€€€€€€€€€€€€€€€€€€€Mg5€€€€€€€€€€€€0.24€€€€€€0.003Ì€€€€€€€€€€€€€€€€€€€€€€€€€Mg7€€€€€€€€€€€€0.12€€€€€€0.04€€€Ô€!X"FXX6X"FÔÐ : Š ÐOU15òò900óóà ` à?€€€€30€€€€€€€€Mg6€€€€€€€€€€€€0.36€€€€€€0.005Ð 4!„ Ѐ€€€€€€€€€€€€€€€€€€€€€€€€Mg2€€€€€€€€€€€€0.36€€€€€€0.02Ì0C10òò600óóà ` à?€€€€32€€€€€€€€Mg5€€€€€€€€€€€€0.23€€€€€€0.005Ð (#x Ð0C11òò400óóà ` à?€€€€32€€€€€€€€Mg6€€€€€€€€€€€€0.26€€€€€€0.002Ð "$r  Ѐ€€€€€€€€€€€€€€€€€€€€€€€€Mg5€€€€€€€€€€€€0.25€€€€€€0.004Ì€€€€€€€€€€€€€€€€€€€€€€€€€Mg7€€€€€€€€€€€€0.13€€€€€€0.04Ì€€€€€€€€€€€€€€€€€€€€€€€€€Mg8€€€€€€€€€€€€0.16€€€€€€0.023Ìòò€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€€óóÐ  (Z#$ ÐÌòòReferencesóóÐ þ)N%& Ðà X àBrown€J.C€and€W.E€Jones€:€1974€Differential€response€of€oats€toÏmanganese€stress.€Agro.€J.€66:€624„626.Ìà X àCarter,€O.G.,€I.A.€Rose,€and€P.F.€Reading€1975.€€Variation€inÏsusceptibility€to€manganese€toxicity€in€30€soybean€genotypes.€€Ð æ-6)* ÐCrop€ÌSci.€15:€730„732.Ìà X àChang€S.J.C.,€T.W.€Doubler,€V.€Kilo,€R.€Suttner,€J.€Klein,€M.E.€Schmidt,ÏP.T.€Gibson€and€D.A.€Lightfootòòóó.€1996.€Two€additional€LociÐ žî 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àTortoòòóó,€G.A.,€V.€Njiti€and€D.A.€Lightfoot.€1996.€Loci€underlyingÐ 2‚ Ðresistance€to€sudden€death€syndrome€and€òòFusarium€solanióó€in€fieldÐ ,| Ðand€greenhouse€assays€do€not€correspond.€Soybean€GeneticsÏNewsletter.€(This€issue)ÌÒÀ°Òà  àà p àà È àV.€KiloÌà  àà p àà È àD.A.€LightfootÐ  F– Ð