2009 Major and minor QTL and epistasis contribute to fatty acid
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第四届“QTL作图和育种模拟研讨会”,2009年8月18-20日,山东泰安育种模拟的原理和遗传模拟工具QuLineCIMMYT’s headquarter in MexicoDr. Borlaug and green revolution ¾¾¾¾¾¾¾Mexico CityEl BatanToluca, 19º N, 2640 masl.High rainfall (800-900 mm)Cd. Obregon, 27º N, 39 masl. 8-11 t/ha under irrigation; 1-2t/ha under reduced irrigationCIMMYT’s Shuttle breedingMay to NovemberBreeding methods with self-pollinated cropsBreeding methods in CIMMYT’s wheat breeding program¾¾¾About 40% ofafter F7About 30% of the crosses are discarded in F1About 20% ofof yield trialsWhy do we need tools in breeding?¾¾¾••••Why do we need tools in breeding?¾¾Questions that can be studied by QuLine: A genetic and breeding simulation tool 1.2.3.4.Questions that can be studied by QuLine (mainly for inbred line development) 5.6.QuLine: A simulation tool for genetics and breeding¾QU-GENE (QUantitative GENEtics)A simulation platform for quantitative analysis of genetic models, developed byThe University of Queensland, Australia¾QuCim (funded by GRDC 2000-2004)A QU-GENE application breeding simulation module, specifically designed forCIMMYT’s wheat breeding programsSimulate most breeding programs for developing inbred linesVersion 1.1 released on July, 2003 (Workshop in Brisbane, Australia)More than 100 global requests for QuCim 1.1¾Renamed as QuLine (currently funded by GCP, H+, and Research Programs of China, i.e. 973, 863, and NSF)Landscape representation of a complex GE system (the real GE system is multi-dimensional)What can QuLine do?¾¾¾¾In genetics(implemented by the QU-GENE engine)¾¾¾¾In breeding (implemented by the QuLine module)How does QuLine work?¾•(= input for QuLine•(= input for QuLine(= input for QuLineDefine the QMP file for the selected bulk selection method: an exampleGeneral simulation parameters¾¾¾¾General simulation parameters ¾¾¾¾¾The number of models in the GE system and the number of runs for breeding strategy¾•9»¾Parameters to describe aset of breeding strategies¾¾Definition of a generation ¾¾¾Practical breeding small plot evaluation atVirtual breedingF6F7F8 (T), F8 (B)F8 (YT)F8 small plotFamilies selected14,760 3,8683,868 2,163 1,9741,974 779779An example for seed source indicator 0Bulk in F3 PYEI, F4PYEII, F5Pedigree in F4Families selected 4000 12001200 600600 100100 1000An example (LRC, Toowoomba, Australia) for seed source indicator 1Definition of each selection round¾¾Definition of each selection round ¾pedigree:bulk:Definition of each selection round¾••Definition of each selection round ¾•••9T for top, e.g. yield, tillering, grains per spike and 1000-kernel weight9B for bottom, e.g. lodging and rusts9M for middle, e.g. height and heading9R for random, for some special studies9Roundsof selectionSeedsourceindicatorGenerationtitleSeedpropagationtypeGenerationadvancemethodReplicationsPlotsizeTestlocationsEnvironmenttype10F6self pedigree175012, Toluca 40F7self bulk17011, Obregon F8(T)self bulk17012, TolucaF8(B)self bulk17013, El BatanF8(YT)self bulk110011, Obregon 10F8(SP)self bulk13011, Obregon An example of generation definitionTraitYieldLodg-ingStem rustLeaf rustStripe rustHeightTilleringHeadingGrains per spike1000 kernel weightTotalSelection mode T B B B B M T M T TF6, among 0.990.960.950.90F6, within 0.900.700.900.950.980.100.05F7, among 0.850.700.980.850.960.700.750.25F8(T), among 0.550.700.990.980.990.900.55F8(B), among 0.900.90F8(YT), among 0.400.40Traits, their selection modes andselected proportionsSteps to run QuLineBreeding strategiesGE systemPopulationInput information about the GxE system and populationsQUGENEQuLine*.fit*.var*.fre*.ham*.cro*.his*.rog*.pou *.fixMajor outputs from QuLineWhat has been done using QuLine?¾Crop Science(2003)¾Crop Science(2004)¾Aust. J. Agri. Sci.(2005)¾Crop Science(2007)Comparison of two breeding strategies: modified pedigree (MODPED) andselected bulk (SELBLK)Breeding methods with self-pollinated cropsBreeding methods in CIMMYT’s wheat breeding program¾¾¾methodsTrait, segregating gene number, gene effects and trait heritabilityTrait Genes Gene effecttypeAA Aa aa Trait range h b2(Indiv. plant)Yield20, 40E0, E1, E2Random value from UD (0, 1)0.05 Lodging3additive05100-300.10 Stem rust5additive00.510-50.30 Leaf rust5additive05100-500.30 Yellow rust5additive05100-500.30 Height3additive403020120-600.45 Tillers/plant 3additive53115-30.35 Heading5additive201612100-600.30 Grains/spike5additive1410670-300.35Trait correlation and pleiotropyTraitYieldLodgingStem rustLeaf rust YellowrustHeightTillers/plantHeadingGrains/spikeSeed weightYield-0.50-0.20-0.10-0.10-0.500.400.300.500.40Lodging-0.56Stem rust-0.25Leaf rust-0.05Yellow rust-0.09Height-0.62Tillers/plant-0.08-0.20-0.40Heading0.60Grains/spike 0.09-0.17-0.30Seed weight-0.07-0.30-0.07Estimated by CIMMYT breedersEstimated from the defined genetic modelExperiment design¾12 Genotype and environment (GE) systems¾Initial population¾¾Result 1: Genetic gain in yield from SELBLK is 3.3% higher than MODPED. SELBLK is slightly more efficient.For gains per spike and 1000-kernel weight, SELBLK has a faster genetic gain. For tillers/plant, MODPED has a faster genetic gain.Result 2: SELBLK retained 25% more crosses in the final selected population (more genetic diversity retained)Result 3: SELBLK required 1/3 less land from F1 to F8 than MODPED. SELBLK is more cost-effective.Result 4: SELBLK produced 40% less families (plots) to be planted from F1 to F8 (less labor required)Modeling of the Single Backcrossing Breeding Strategy(SBBS)(Theor. Appl. Genet., 2009, 118: 683-694)Category% favorablegenes Example% totalparentallinesElite adapted lines (EAL)80-85Major released cultivars in targeted mega-environments (MEs) either developed byCIMMYT or by partners10Adapted lines (AL)75-80Elite advanced lines from CIMMYT’sInternational Nursery and Yield Trials60Intermediate adapted lines (IAL)65-75Advanced lines from CIMMYT’s Yield Trialsin Ciudad Obregón and Toluca, Mexico10Un-adapted (or non-adapted) lines (UAL)20-40Land races 2Second generation of re-synthesized wheat (SYNII)40-60Derived lines between the first generation ofre-synthesized wheat derivatives andadapted lines10First generation of re-synthesized wheat (SYNI)20-40Derived lines between primary re-synthesized wheat and adapted lines5Estimated percentages of favourable alleles or gene combinations in different parental lines in wheat breeding at CIMMYTTwo traits defined in QU-GENE •––––•–––。
ORIGINAL PAPERQTL analysis of soybean seed weight across multi-genetic backgrounds and environmentsYingpeng Han•Dongmei Li•Dan Zhu•Haiyan Li•Xiuping Li•Weili Teng•Wenbin LiReceived:27December2011/Accepted:21March2012ÓSpringer-Verlag2012Abstract Seed weight,measured as mass per seed,is an important yield component of soybean and is generally positively correlated with seed yield(Burton et al,Crop Sci 27:1093,1987).In previous reports,quantitative trait loci (QTL)associated with seed weight,were identified in single genetic background.The objective of the present study was to identify QTL and epistatic QTL underlying soybean seed weight in three RIL populations(with one common male parent‘Hefeng25’)and across three differ-ent environments.Overall,18,11,and17seed weight QTL were identified in HC(‘Hefeng25’9‘Conrad’),HM (‘Hefeng25’9‘Maple Arrow’),and HB(‘Hefeng25’9‘Bayfield’)populations,respectively.The amount of phe-notypic variation explained by a single QTL underlying seed weight was usually less than10%.The environment and background-independent QTL often had higher addi-tive(a)effects.In contrast,the environment or back-ground-dependent QTL were probably due to weak expression of QTL.QTL by environment interaction effects were in the opposite direction of a effects and/or epistasis effects.Four QTL and one QTL could be identi-fied(2.0\LOD\9.06)in the HC and HB populations,respectively,across three environments(swHCA2-1, swHCC2-1,swHCD1b-1,swHCA2-2(linked to Satt233, Satt424,Satt460,Satt428,respectively)and swHBA1-1(Satt449).Seven QTL could be identified in all three RIL populations in at least one location.Two QTL could be identified in the three RIL populations across three envi-ronments.These two QTL may have greater potential for use in marker-assisted selection of seed weight in soybean. IntroductionSeed weight,measured as mass per seed,is an important yield component of soybean and was generally positively correlated with seed yield(Burton et al.1987).Seed weight is a quantitative trait,which is mainly inherited with the additive effect(Brim and Cockerham1961).Most of the breeding strategies for developing soybean cultivars take advantage of additive gene actions(Hartwig1973;Fehr 1987;Cooper1990).However,traditional plant improve-ment has relied on phenotypic selection of populations from crosses between commercial cultivars and/or experimental lines(Stuber et al.1992;Specht et al.1999;Stefaniak et al. 2005).The development of consensus molecular maps in soy-bean(Song et al.2004;Choi et al.2007;Hyten et al.2010; Song et al.2010)has facilitated the identification of common quantitative trait loci(QTL)controlling important quantitative traits in many soybean populations(Hyten et al.2004).Through molecular genetic linkage maps and QTL analysis,it is possible to estimate the number of QTL controlling genetic variation.Statistical analyses,such as ANOVA and interval mapping,can estimate the gene actions and phenotypic effects(Lander et al.1987;Basten et al.1996).Combined analyses allowed pleiotropicand/or Communicated by H.T.Nguyen.Y.HanÁD.LiÁD.ZhuÁH.LiÁX.LiÁW.TengÁW.LiKey Laboratory of Soybean Biology in Chinese Ministryof Education,(Key Laboratory of Biology and Genetics&Breeding for Soybean in Northeast China,Ministry ofAgriculture),Northeast Agricultural University,Harbin150030,ChinaW.Li(&)Soybean Research Institute(Key Laboratory of Soybean Biologyin Chinese Ministry of Education),Northeast AgriculturalUniversity,Harbin150030,Chinae-mail:wenbinli@123 Theor Appl GenetDOI10.1007/s00122-012-1859-xepistatic interactions to be estimated(Lark et al.1995;Yan 2001).Through selection with molecular markers,the results of QTL analyses can be used in soybean breeding, referred as marker-assisted selection(MAS).A common approach in MAS is to map QTL in a small sample of progeny from a cross,choose a marker linked to the tar-geted QTL,and apply MAS in a larger set of progeny from the same cross(Prabhu et al.1999;Ken et al.2005).This has been done for many traits,including agronomic traits in barley(Zhu et al.1999),disease resistance in soybean (Prabhu et al.1999),and for blight resistance in chickpea (Millan et al.2003)and rice(Davierwala et al.2001). However,many studies have shown that the effects of QTL underlying agronomic traits are not always detected in other genetic backgrounds for soybean(Specht et al.1999; Hyten et al.2004)and many other crops(McKendry et al. 1996;Toojinda et al.1998;Ken et al.2005).This phe-nomenon is a major obstacle to the efficient use of MAS in plant breeding.In the past two decades,many studies have focused on mapping of QTL influencing soybean seed weight [reviewed by Hyten et al.(2004)].In most of these reports, the phenotypic values of traits in single genetic background were used for QTL analysis.Few of these studies have used common parents among their multiple genetic backgrounds (Orf et al.1999;Hyten et al.2004),which is a very important factor influencing quantitative traits like yield components(Specht et al.1999;Liao et al.2001;Hyten et al.2004;Zhang et al.2004).Many results indicated that yield of soybean is controlled by a series of QTL with different expression in different genetic populations or environments. Though genetic background effects on quantitative traits have been well documented in many crops including tomato (Tanksley and Hewitt1988),rice(Li et al.1997,1998),soy-bean(Lark et al.1995;Orf et al.1999),and maize(Doebley et al.1995),the QTL associated with seed weight had been less frequently reported in multi-genetic background by2011 (Orf et al.1999;Hyten et al.2004).The objective of this study was to identify stable QTL and epistatic QTL effects for seed weight of soybean in different genetic backgrounds and environments,using three popula-tions with one common male parent,‘Hefeng25’,to compare the number and location of QTL and epistatic QTL mapped across different populations and environments,and to deter-mine QTL9environment interactions.Materials and methodsPlant materials‘Hefeng25’9‘Conrad’(HC,140RILs of F4:5),‘Hefeng 25’9‘Maple Arrow’(HM,149RILs of F5:6),and‘Hefeng 25’9‘Bayfield’(HB,144RILs of F5:6)were used to evaluate seed weight across multiple environments.‘Hef-eng25’,a local variety in Northeastern China,was selected as the common parent of the three soybean populations due to higher seed weight and other good agronomic traits.The RILs and their parents were grown in a randomized com-plete block design at Harbin in three locations during2007, 2008,and2009in four row plots.The rows were3m long, 90cm apart,and there was a space of about6cm between two plants in a row.In each plot,20plants from one genotype were grown as seed donors that were later used to analyze seed weight after maturity.Seeds were dried for 30min in oven at105°C and then continuously dried at 50–70°C until the seed weight was stable.All dried samples were weighed.Genetic linkage mapThe genetic map of the HC RIL population constructed with164SSR markers encompassed12linkage groups (LGs)(Li et al.2010a).The HC map covered about 3160.28cM of the soybean genome with a mean distance of19.27cM between markers.The HM RIL population map is composed of19LGs constructed with109SSRs with a mean distance of16.96cM between markers(Li et al.2009).The HB RIL population map included107 SSRs in20LGs with a mean distance of14.21cM between markers(Li et al.2010b).Data analysisQTL were identified by single-factor analysis of variance (PROC.GLM.SAS),interval mapping(Lander et al.1987) and composite interval mapping(Basten et al.1996)as described by Primomo et al.(2005).Significant QTL (LOD[2.0)were recorded based on the mean value of individual environment.For interactions locus main effects were considered for linear models if they were significant at P B0.01.Signif-icant loci on the same LG were tested by two-factor analysis of variance without interactions.If both loci were significant at P B0.05in the two-factor model,they would both be considered for linear models.Otherwise,the locus with the larger individual R2value was chosen to represent the effect of the putative QTL on the LG.Two-way analysis of variance was also used to detect significant(P\0.01)epistatic interactions between markers.The nomenclature of QTL included four parts. For example,QTL swHCA1-1is composed of sw(seed weight),HC(the RIL population),A1(the linkage group), and one(thefirst QTL in the LG).GT(Genotype by Trait)biplot methodology(Yan2001) was employed to analyze the interactions between QTLTheor Appl Genet123and different environments in the HC,HM,and HB popula-tions,based on the formula:T ij-T j/S j=k1f i1s j1?k2f i2s-j2?e ij,where T ij is the average value of QTL i for environment j;T j is the average value of environment j over all QTL of single population,S j is the standard deviation of environment j among the QTL average;f i1and f i2are the PC1(first principle component)and PC2(second principle component)scores,respectively.For QTL i;s j1and s j2are the PC1and PC2scores,respectively,for environment j;and e ij is the residual of the model associated with the QTL i in environment j.ResultsPhenotypic variationPhenotypic values of100-seed weight among different populations across three environments are shown in Table1.The differences between the two parents were significant in the three different populations across three environments.The seed weight for‘Hefeng25’was2–5g higher than those of‘Conrad’,‘Maple Arrow’and‘Bay-field’.In contrast,100-seed weight variations among the RIL populations were not significantly different.Both skewness and kurtosis values of100-seed weight distribu-tions were less than1.0for the three RIL populations in most environments,suggesting that the segregations of this traitfit a normal distribution model.Analysis of QTL in different environments and genetic backgroundsEighteen,11,and17QTL were found to be associated with seed weight in HC,HM,and HB populations,respectively (Table2).The QTL in HC,HM,and HB populations could explain3.86–31.09,5.75–23.89,and3.23–32.76%of the phenotypic variations at Harbin in3years,respectively.QTL swHCB1-1(Sat_123),swHMK-1(Satt555),and swHBD1a-1 (Satt168)explained the largest amount of phenotypic varia-tion(31.09,23.89,and32.76%).Four QTL swHCA2-1; swHCC2-1;swHCD1b-1;swHCA2-2(linked to Satt233, Satt424,Satt460,Satt428,respectively)in the HC population, and one QTL swHBA1-1(Satt449)in the HB population could be identified in all three environments.In the present work,eight QTL that derived beneficial alleles from‘Hefeng25’,could be detected in three different populations[Satt233located in LG A2(Chromosome(Gm) 8),Satt460in LG C2(Gm6),Satt428in LG D1b(Gm2), Satt302in LG H(Gm12),Satt354in LG I(Gm20),Satt555in LG K(Gm9),Satt527in LG L(Gm19),and Satt153in LG O (Gm10)].Six QTL of Conrad origin associated with Sat_181in LG A2(Gm8),Sat_123in LG B1(Gm11),Satt489in LG C2(Gm6),Satt277in LG C2(Gm6),Satt301in LG D2 (Gm17),and Satt380in LG J(Gm16).One QTL(Satt150in LG M,Gm7)from‘Maple Arrow’was detected,andfive QTL from‘Bayfield’[Satt449in LG A1(Gm 5),Satt596in LG B2(Gm14),Satt190in LG C1(Gm4),Satt277 in LG C2(Gm6),and Satt168in LG D1a(Gm1)]were found. QTL9environment interaction acrossmulti-environments and genetic backgrounds Eighteen,11,and17QTL in HC,HM,and HB population had additive main effect(a)and/or additive9environ-ment interaction effect(ae)at some specific environments in14,11,and17LGs(Table3).Eight QTL(swHCA2-1,swHCA2-2,swHCC2-1, swHCD1b-1,swHCD2-1,swHCH-1,swHCK-1,swHCO-Table1Statistical analysis of seed weight(100-seed weight)for parents and the three RIL populations at different environments Populations Environments Parents Recombinant inbred linesRange Mean CV Skewness KurtosisHefeng259Conrad Hefeng25Conrad2007Harbin20.5717.0411.24–23.9218.7454.39-1.290.782008Harbin21.0118.1313.76–24.3219.1237.890.860.592009Harbin20.3417.5611.29–22.3918.8447.890.300.77 Hefeng259Maple Arrow Hefeng25Maple Arrow2007Harbin20.5718.4914.67–22.5817.8938.97 1.030.942008Harbin21.0116.0210.87–22.3918.8757.390.970.672009Harbin20.3416.9812.39–21.2817.7644.760.82-0.74 Hefeng259Bayfield Hefeng25Bayfield2007Harbin20.5715.4711.78–22.2818.2057.300.760.952008Harbin21.0118.5312.73–22.7617.3262.29-0.840.962009Harbin20.3416.9814.39–23.3718.3258.390.890.79 Theor Appl Genet123T a b l e 2Q T L a s s o c i a t e d w i t h s e e d w e i g h t (100-s e e d w e i g h t )i n d i f f e r e n t p o p u l a t i o n s a c r o s s d i f f e r e n t e n v i r o n m e n t sQ T L s o u r c e Q T LL G (C h r o m o s o m e )M a r k e r i n t e r v a l P o p u l a t i o n E 1a E 2b E 3cL O DD R 2(%)L O D D R 2(%)L O D D R 2(%)H e f e n g 25s w H C A 2-1A 2(C h d 8)S a t t 233–S a t t 538H C 6.6712.4528.764.368.698.796.4014.5912.29H e f e n g 25s w H M A 2-1S a t t 233–S a t t 177H M 6.4323.7621.63H e f e n g 25s w H B A 2-1S a t t 538–S a t t 233H B 3.4516.488.605.4920.3010.29H e f e n g 25s w H C C 2-1C 2(C h 6)S a t t 100–S a t t 460H C 3.402.4018.894.498.8922.306.875.8413.59H e f e n g 25s w H M C 2-1S a t t 100–S a t t 460H M 5.5415.546.679.7627.985.76H e f e n g 25s w H B C 2-1S a t t 460–S a t t 202H B 3.872.1710.875.766.788.86H e f e n g 25s w H C D 1b -1D 1b (C h 2)S a t t 579–S a t t 428H C 2.6515.6527.863.9713.8310.657.6412.745.76H e f e n g 25s w H M D 1b -1S a t t 282–S a t t 428H M 5.456.6712.64H e f e n g 25s w H B D 1b -1S a t t 282–S a t t 428H B 3.6514.657.898.7613.6418.86H e f e n g 25s w H C H -1H (C h 12)S a t t 181–S a t t 302H C 5.9721.0815.86H e f e n g 25s w H M H -1S a t t 253–S a t t 302H M 3.9719.8921.066.8612.6414.45H e f e n g 25s w H B H -1S a t t 293–S a t t 302H B 3.760.8912.76H e f e n g 25s w H C I -1I (C h 20)S a t t 354–S a t t 571H C 6.7510.315.792.0511.634.43H e f e n g 25s w H M I -1S a t t 354–S a t t 419H M 5.7615.7615.65H e f e n g 25s w H B I -1S a t t 354–S a t t 440H B 3.7637.6515.97H e f e n g 25s w H C K -1K (C h 9)S a t t 349–S a t t 555H C 4.780.547.746.650.6415.76H e f e n g 25s w H M K -1S a t t 555–S a t t 046H M 3.065.8623.89H e f e n g 25s w H B K -1S a t t 555–S a t t 046H B 7.971.874.654.651.655.53H e f e n g 25s w H C L -1L (C h 19)S a t t 527–S a t t 561H C 5.984.7621.75H e f e n g 25s w H M L -1S a t t 527–S c t _010H M 3.760.546.683.890.436.42H e f e n g 25s w H B L -1S a t t 527–S a t t 561H B 9.750.9810.76H e f e n g 25s w H C O -1O (C h 10)S a t t 479–S a t t 153H C 3.7517.546.904.8612.674.54H e f e n g 25s w H M O -1S a t t 153–S a t t S a t _106H M 6.5921.865.757.6526.9714.76H e f e n g 25s w H B O -1S a t t 153–S a t t 109H B 8.755.764.74H e f e n g 25s w H C A 1-1A 1(C h 5)S a t t 382–S a t t 211H C 5.6410.875.384.0817.647.77H e f e n g 25s w H B A 1-2S a t t 454–S a t t 382H B 3.604.6114.563.454.7513.29H e f e n g 25s w H C A 2-2A 2(C h 8)S a t t 424–S a t t 390H C 4.218.766.655.548.7610.657.654.868.70H e f e n g 25s w H B A 2-2S a t t 177–S a t t 424H B 4.6714.4510.765.568.897.87H e f e n g 25s w H M B 1-1B 1(C h 11)S a t _123–S a t t 583H M 2.8638.9712.53H e f e n g 25s w H B B 1-1S a t t 583–S a t _096H B 5.322.106.23H e f e n g 25s w H C D 1a -1D 1a (C h 1)S a t t 383–S a t t 468H C 3.976.8721.05H e f e n g 25s w H M D 1a -1S a t t 502–S a t t 383H M 4.6517.7612.86H e f e n g 25s w H C E -1E (C h 15)S a t t 263–S a t t 491H C 6.8710.4028.76H e f e n g 25s w H B E -1S a t t 263–S a t t 268H B 9.0632.985.98C o n r a d s w H C A 2-3A 2(C h 8)S a t _181–S a t t 409H C 2.373.4510.32C o n r a d s w H C B 1-1B 1(C h 11)S a t t 453–S a t _123H C 4.4514.455.877.658.7631.09C o n r a ds w H C C 2-2C 2(C h 6)S a t t 489–S a t t 557H C6.5413.653.863.097.878.54Theor Appl Genet1231)from HC population,four QTL (swHMC2-1,swHMI-1,swHML-1and swHMM-1)from HM population,and nine QTL (swHBA1-2,swHBA2-1,swHBC1-1,swHBC2-1,swHBC2-2,swHBD1b-1,swHBH-1,swHBK-1,swHBL-1)from HB population,contributed the allele that increased seed weight through significant a effects.Three QTL (swHCB1-1,swHCC2-2,swHCI-1)originating from HC population,four QTL (swHMB1-1,swHMD1a-19,swHMH-1and swHMO-1)origininating from HM population,and two QTL (swHBA2-2and swHBB2-1)originating from HB pop-ulation,contributed the allele that decreased seed weight through significant a effects,respectively.The impact of ae effects on QTL was different at dif-ferent years for these three populations.For example,QTL swHCA2-1of HC population,increased seed weight at Harbin in 2007and 2009,but decreased seed weight at Harbin in 2008.Six QTL (swHCA2-3,swHCC2-3,swHCD1a-1,swHCE-1,swHCJ-1,swHCL-1)of HC pop-ulation,three QTL (swHMA2-1,swHMD1b-1and swHMK-1)of HM population,and five QTL (swHBB1-1,swHBD1a-1,swHBE-1,swHBI-1,swHBO-1)of HB pop-ulation,had only significant ae effects,but no significant a effects.Other QTL in these three populations had both significant a effects and significant ae effects.Epistatic analysis of QTL across multi-environments Forty,16.and 20epistatic pairwise QTL were detected in HC,HM,and HB population in different environments,respectively (Table 4).Of them,18,5and 5epistatic pairs of QTL,derived from HC,HM and HB populations,were beneficial for the increase of seed weight through signifi-cant aa effects.Seven,five and four epistatic pairs of QTL trended to decrease seed weight through significant aa effects in HC,HM,and HB population,respectively (Table 4).The epistasis 9environment interaction effect (aae )was an important component of the total QTL 9environ-ment (QE)interaction effects.Six,four,and five pairs of QTL were found with only epistatic effects (aa )in HC,HM,and HB populations,respectively.Fifteen,five,and eleven pairs had only aae effects in HC,HM,and HB populations,and other pairs of epistatic QTL had both aa and aae effects (Table 4).Stability evaluation of QTL derived from ‘Hefeng 25’In this study,additive effects of eight QTL,linked to Satt233,Satt460,Satt428,Satt302,Satt354,Satt555,Satt527,and Satt153)from ‘Hefeng25’,were detected in three different populations.(Table 3).The scale of a effect of QTL,derived from ‘Hefeng 25’,was diverse in different populations.For example,a effect of QTL,associated withT a b l e 2c o n t i n u e dQ T L s o u r c eQ T L L G (C h r o m o s o m e )M a r k e r i n t e r v a l P o p u l a t i o nE 1aE 2bE 3cL O DDR 2(%)L O D DR 2(%)L O D DR 2(%)C o n r a d s w H C C 2-3C 2(C h 6)S a t t 307–S a t t 277H C 3.395.787.52C o n r a d s w H CD 2-1D 2(C h 17)S a t t 615–S a t t 301H C 5.860.8610.86C o n r a d s w H C J -1J (C h 16)S a t t 380–S a t t 183H C 2.764.6717.72M a p l e A r r o w s w H M M -1M (C h 7)S a t t 150–S a t t 201H M 2.062.788.762.984.5615.83B a y fie l d s w H B A 1-1A 1(C h 5)S a t t 449–S a t t 454H B 2.702.4520.703.471.8916.675.793.2920.28B a y fie l d s w H B B 2-1B 2(C h 14)S a t t 070–S a t t 596H B 5.5412.837.537.867.649.65B a y fie l d s w H B C 1-1C 1(C h 4)S a t t 190–S a t t 195H B 4.478.8721.676.6512.433.23B a y fie l d s w H B C 2-2C 2(C h 6)S a t t 277–S a t t 376H B 5.874.7615.983.872.068.76B a y fie l ds w H B D 1a -1D 1a (C h 1)S a t t 198–S a t t 168H B5.748.8632.76aa t H a rb i n i n 2007ba t H a rb i n i n 2008ca t H a rb i n i n 2009dC h r o m o s o m e n u m b e rTheor Appl Genet123Theor Appl GenetTable3Additive and additive9environment interaction effect of QTL associated with seed weight at different populationsQTL source QTL LG(Chromosome)Marker interval Population a d a9E1a a9E2b a9E3cHefeng25swHCA2-1A2(Ch8)Satt233–Satt538HC0.27*0.11*-0.17*0.32* Hefeng25swHMA2-1Satt233–Satt177HM0.39* Hefeng25swHBA2-1Satt538–Satt233HB0.62**0.76**0.17* Hefeng25swHCC2-1C2(Ch6)Satt100–Satt460HC 2.01**0.40*-0.89*0.87* Hefeng25swHMC2-1Satt100–Satt460HM0.69**0.39*Hefeng25swHBC2-1Satt460–Satt202HB0.79**0.26*-0.27*Hefeng25swHCD1b-1D1b(Ch2)Satt579–Satt428HC 1.64**-1.02**0.19*0.76** Hefeng25swHMD1b-1Satt282–Satt428HM0.30* Hefeng25swHBD1b-1Satt282–Satt428HB 1.34**0.65**0.76**Hefeng25swHCH-1H(Ch12)Satt181–Satt302HC0.30*-0.37*Hefeng25swHMH-1Satt253–Satt302HM-0.19*0.38*-0.69** Hefeng25swHBH-1Satt293–Satt302HB0.49*Hefeng25swHCI-1I(Ch20)Satt354–Satt571HC-0.87**0.88**-0.32*Hefeng25swHMI-1Satt354–Satt419HM0.22*0.16*Hefeng25swHBI-1Satt354–Satt440HB0.58**Hefeng25swHCK-1K(Ch9)Satt349–Satt555HC 1.34**0.67** 1.20** Hefeng25swHMK-1Satt555–Satt046HM-0.28*Hefeng25swHBK-1Satt555–Satt046HB0.78**-0.89**-1.25** Hefeng25swHCL-1L(Ch19)Satt527–Satt561HC0.59**Hefeng25swHML-1Satt527–Sct_010HM 1.04**0.45**-1.02** Hefeng25swHBL-1Satt527–Satt561HB0.55**Hefeng25swHCO-1O(Ch10)Satt479–Satt153HC0.79**-0.87**-0.95**Hefeng25swHMO-1Satt153–SattSat_106HM-0.67**0.20*Hefeng25swHBO-1Satt153–Satt109HB-0.17*Hefeng25swHCA1-1A1(Ch5)Satt382–Satt211HC-0.76**-0.48** Hefeng25swHBA1-2Satt454–Satt382HB0.97**0.28*-0.59**Hefeng25swHCA2-2A2(Ch8)Satt424–Satt390HC 1.34**0.21*0.56**-0.67** Hefeng25swHBA2-2Satt177-Satt424HB-0.30*-0.49**0.89**Hefeng25swHMB1-1B1(Ch11)Sat_123–Satt583HM-0.20*Hefeng25swHBB1-1Satt583–Sat_096HB-0.87**Hefeng25swHCD1a-1D1a(Ch1)Satt383–Satt468HC0.89** Hefeng25swHMD1a-1Satt502–Satt383HM-0.31*Hefeng25swHCE-1E(Ch15)Satt263–Satt491HC0.30*Hefeng25swHBE-1Satt263–Satt268HB-0.98**Conrad swHCA2-3A2(Ch8)Sat_181–Satt409HC-0.15*Conrad swHCB1-1B1(Ch11)Satt453–Sat_123HC-0.79**-0.88** 1.09**Conrad swHCC2-2C2(Ch6)Satt489–Satt557HC-0.97*-0.66**-0.59** Conrad swHCC2-3C2(Ch6)Satt307–Satt277HC0.33*Conrad swHCD2-1D2(Ch17)Satt615–Satt301HC0.32*-0.49*Conrad swHCJ-1J(Ch16)Satt380–Satt183HC-0.30*Maple Arrow swHMM-1M(Ch7)Satt150–Satt201HM0.98**-1.17*Bayfield swHBA1-1A1(Ch5)Satt449–Satt454HB 1.29** 1.21**-0.98**0.63** Bayfield swHBB2-1B2(Ch14)Satt070–Satt596HB-0.27*-0.83**-1.07* Bayfield swHBC1-1C1(Ch4)Satt190–Satt195HB0.20*0.67**0.80** Bayfield swHBC2-2C2(Ch6)Satt277–Satt376HB0.95**0.94**-0.91** Bayfield swHBD1a-1D1a(Ch1)Satt198–Satt168HB0.25*a at Harbin in2007b at Harbin in2008c at Harbin in2009d additive effect123Satt460,was2.01in the HC population,0.69in the HM population,and0.79in the HB population,respectively (Table3).The impact of different environments on ae effect of QTL was higher than that of different populations.In the present study,both the aa and aae effects of the QTL derived from‘Hefeng25’were identified in the3or2 of the three populations,and the number of QTL epistatic with other QTL was different across different populations (Table4).For example,six,three,and three epistatic pairwise QTL interacting with the QTL linked to Satt233 was found in HC,HM,and HB populations,respectively. The aa and aae effect of the epistatis QTL were0.23–0.45, 0.16–0.87,0.25–1.07,and0.17–0.45,0.17–0.37,0.19–0.89, respectively.GT biplot analysis(Yan2001)showed the13QTL with beneficial alleles from‘Hefeng25’detected in three or two populations explained73%of the total variation.Figure1 indicated that two QTL(Satt428and Satt460)had highest performance and stability across multiple environments and populations.The QTL linked to Satt555,Satt424, Satt382,Satt354,and Satt183also had above-average performance and stability across multiple environments and populations.The QTL linked to Satt383,Satt583, Satt233,Satt263,Satt302,and Satt527had the lowest-average performance and stability across multi-environments and multi-populations.Furthermore,the performance of different QTL on each environment or population was evaluated.With the QTL linked to Satt460,Satt555,Satt383, Satt263,Satt527,and Satt153as the corner QTL,three environments and two populations(HC and HM)fell in the sector in which two QTL(linked to Satt460and Satt428) were the strongest for these three environments and two populations,respectively(Fig.2).Two QTL(linked to Satt383and Satt555)were the strongest for the HB population.DiscussionRapid advances in genomics have identified and analyzed about36,000genes in soybean(Schmutz et al.2010; Severin et al.2010).However,the usefulness of genomics data for crop breeding and improvement was dependent upon the identification of candidate gene markers for MAS via map-based cloning or association analyses(Varshney et al.2005;Lightfoot2008;Guttierez-Gonzalez et al. 2010).Seed weight is an important yield component of soybean(Burton et al.1987).Hence,there was interest in understanding the genetic control of seed weight in soy-bean,especially the beneficial alleles from‘Hefeng25’,a superior cultivar in Heilongjiang Province of China. Because seed weight of soybean was affected by environ-ments and genetic backgrounds,increasing the genotype selection intensity by MAS could improve the selection efficiency.To be useful to a breeding program,QTL need to be stable across environments and genetic backgrounds (Brummer et al.1997).Stability can be assessed by eval-uating many populations across several environments to determine if a particular QTL is detected in each popula-tion or environment.In this study,three sets of RIL pop-ulations with one common male parent(‘Hefeng25’)were analyzed for QTL that associated with seed weight across multi-environments.A total of18,11,and17QTL were identified in HC,HM,and HB populations,respectively. An individual QTL explained phenotypic variations ranged from3to28%for seed weight in different environments and populations.However,most loci explained less than 10%of the variation(Table2).The low level of pheno-typic variation explained by these QTL indicated the quantitative nature of seed weight,which was similar to the findings of other studies(Mansur et al.1996;Mian et al. 1996;Hyten et al.2004;Zhang et al.2004).QTL specific to one environment were also reported by other studies for different traits(Price et al.2002;Li et al.2003).Many instable QTL detected in different environments or genetic backgrounds were detected(Table2)in this study,which was due to weak expression of the QTL,QTL by envi-ronment interaction in the opposite direction to a effects and/or epistasis(Table3,4).For example,swHCE-1and swHCA2-2were subject to these effects.Therefore,the information of QTL by environment interaction should be considered if MAS was to be applied to the manipulation of quantitative traits.Four QTL swHCA2-1,swHCA2-2,swHCC2-1,and swHCD1b-1linked to Satt233,Satt424,Satt460,and Satt428,respectively,in the HC population across all three environments.The QTL swHCA2-1was reported previ-ously in V71-3709PI407162,Noir19Archer(Hyten et al.2004).The QTL swHCA2-1was reported previously in Minsoy9Noir1(Hyten et al.2004).The QTL swHCC2-1was also reported previously in Young9 PI416937;Noir19Archer;and Essex9Williams (Hyten et al.2004).The QTL swHCD1b-1was also reported previously in Essex9Williams(Hyten et al. 2004).The sole QTL swHBA1-1(linked to Satt449)in the HB population that could be found in all three environments was reported previously in Noir19Archer(Hyten et al. 2004).Therefore,all the QTL stable across environments have been confirmed,which adds to their usefulness in MAS even across multi-environments.Seven QTL(Satt460,Satt428,Satt302,Satt354,Satt555, Satt527,Satt153)could be detected in all three populations in some environments.These QTL might be useful for MAS in a particular location.Theor Appl Genet123T a b l e 4Q T L e p i s t a t i c e f f e c t s a s s o c i a t e d w i t h s e e d w e i g h t (100-s e e d w e i g h t )a t d i f f e r e n t p o p u l a t i o n sQ T L i S o u r c eQ T L iL G (C h r o m o s o m e )M a r k e r i n t e r v a lM a r k e r L G (C h r o m o s o m e )Q T L j S o u r c e Q T L j M a r k e r i n t e r v a l P o p u l a t i o n a a d a a 9E 1a a a 9E 2b a a 9E 3cH e f e n g 25s w H C A 2-1A 2(C h 8)s a t t 233–s a t t 538s a t t 233A 2(C h 8)‘H e f e n g 25’s w H C A 2-2s a t t 424–s a t t 390H C 0.45*0.87**-0.36*0.17*C 2(C h 6)‘H e f e n g 25’s w H C C 2-1s a t t 100–s a t t 460H C -0.89**C 2(C h 6)‘C o n r a d ’s w H C C 2-2s a t t 489–s a t t 557H C 0.23*D 2(C h 17)‘C o n r a d ’s w H C D 2-1s a t t 615–s a t t 301H C 0.48*0.59**I (C h 20)‘H e f e n g 25’s w H C I -1s a t t 354–s a t t 571H C-0.49*O (C h 10)‘H e f e n g 25’s w H C O -1s a t t 479–s a t t 153H C 0.23*-0.16*H e f e n g 25s w H M A 2-1A 2(C h 8)s a t t 233–s a t t 177s a t t 233C 2(C h 6)‘H e f e n g 25’s w H M C 2-1s a t t 100–s a t t 460H M 1.07**0.43*K (C h 9)‘H e f e n g 25’s w H M K -1s a t t 555–s a t t 046H M 0.17*O (C h 10)‘H e f e n g 25’s w H M O -1s a t t 153–s a t t s a t _106H M 0.25*H e f e n g 25s w H B A 2-1A 2(C h 8)s a t t 538–s a t t 233s a t t 233A 2(C h 8)‘H e f e n g 25’s w H B A 2-2s a t t 177–s a t t 424H B0.45*-0.29*C 2(C h 6)‘H e f e n g 25’s w H B C 2-1s a t t 460–s a t t 202H B 0.17*0.89**0.19*0.21*O (C h 10)‘H e f e n g 25’s w H B O -1s a t t 153–s a t t 109H B0.37*H e f e n g 25s w H C C 2-1C 2(C h 6)s a t t 100–s a t t 460s a t t 460C 2(C h 6)‘C o n r a d ’s w H C C 2-3s a t t 307–s a t t 277H C0.56**0.40*0.49*-0.59**J (C h 16)‘C o n r a d ’s w H C J -1s a t t 380–s a t t 183H C-0.67**H e f e n g 25s w H M C 2-1C 2(C h 6)s a t t 100–s a t t 460s a t t 460D 1b (C h 2)‘H e f e n g 25’s w H M D 1b -1s a t t 282–s a t t 428H M0.54**-0.76**M (C h 7)‘M a p l e A r r o w ’s w H M M -1s a t t 150–s a t t 201H M1.23**-0.59**H e f e n g 25s w H B C 2-1C 2(C h 6)s a t t 460–s a t t 202s a t t 460C 2(C h 6)‘B a y fie l d ’s w H B C 2-2s a t t 277–s a t t 376H B0.17*-0.90**H e f e n g 25s w H C D 1b -1D 1b (C h 2)s a t t 579–s a t t 428s a t t 428E (C h 15)‘H e f e n g 25’s w H C E -1s a t t 263–s a t t 491H C0.47*0.65**0.97**0.76**H e f e n g 25s w H M D 1b -1D 1b (C h 2)s a t t 282–s a t t 428s a t t 428L (C h 19)‘H e f e n g 25’s w H M L -1s a t t 527–s c t _010H M0.67**H e f e n g 25s w H B D 1b -1D 1b (C h 2)s a t t 282–s a t t 428s a t t 428E (C h 15)‘H e f e n g 25’s w H B E -1s a t t 263–s a t t 268H B0.82**H e f e n g 25s w H C H -1H (C h 12)s a t t 181–s a t t 302s a t t 302J (C h 16)‘C o n r a d ’s w H C J -1s a t t 380–s a t t 183H C0.86**0.67**O (C h 10)‘H e f e n g 25’s w H C O -1s a t t 479–s a t t 153H C0.47*H e f e n g 25s w H M H -1H (C h 12)s a t t 253–s a t t 302s a t t 302I (C h 20)‘H e f e n g 25’s w H M I -1s a t t 354–s a t t 419H M0.78**0.89**0.45*M (C h 7)‘M a p l e A r r o w ’s w H M M -1s a t t 150–s a t t 201H M-0.56**H e f e n g 25s w H B H -1H (C h 12)s a t t 293–s a t t 302s a t t 302I (C h 20)‘H e f e n g 25’s w H B I -1s a t t 354–s a t t 440H B0.22*H e f e n g 25s w H C I -1I (C h 20)s a t t 354–s a t t 571s a t t 354K (C h 9)‘H e f e n g 25’s w H C K -1s a t t 349–s a t t 555H C0.56**0.75**0.43*H e f e n g 25s w H M I -1I (C h 20)s a t t 354–s a t t 419s a t t 354K (C h 9)‘H e f e n g 25’s w H M K -1s a t t 555–s a t t 046H M0.65**H e f e n g 25s w H B I -1I (C h 20)s a t t 354–s a t t 440s a t t 354K (C h 9)‘H e f e n g 25’s w H B K -1s a t t 555–s a t t 046H B0.97**Theor Appl Genet123。
PART I DICTATIONNew Year’s EveFor many people in the west, New Year’s Eve is the biggest party of the year./ It’s time to get together with friends or family/ and welcome in the coming year./ New Year’s parties can take place in different places./ Some people hold a house party;/ others attend street parties,/ while some just go for a few drinks with their friends./ Big cities have large and spectacular fireworks displays./ There is one thing that all New Year’s Eve parties have in common--- the countdown to midnight./ When the clock strikes 12, people give a loud cheer and sing songs./ It’s also popular to make a promise in the New Year./ This is called a New Year’s resolution./ Typical resolutions include giving up smoking and keeping fit./ However, the promise is often broken quite quickly/ and people are back into their bad habits within weeks or days.PART II LISTENING COMPREHENSIONSETION A CONVERSATIONSConversation OneW: Hi, Mark. How are you?M: Actually, I am really fed up, Linda. It’s Jane (Q1).W: Jane? Who’s Jane?M: No, nobody really. Just a most stunningly attractive girl in my year school (Q1).W: Oh, is that all? So, what’s the problem?M: Well, the thing is I just don’t know how to make her notice me or….W: Wait a minute. I’ve got a brilliant idea. Why don’t you try talking to her?M: But I wouldn’t know what to say. I …W: Look, she is in your chemistry class, isn’t she? You are good at chemistry. You could offer to help her with her chemistry homework. How about that?M: Not that. Just one problem (Q3).W: What?M: She is better than me at chemistry.W: OK, then. W ell, there’s that party at John’s on Friday night. You could invite her.M: Just another small problem (Q3). John’s her boyfriend (Q2).Conversation TwoW: Yes. Can I help you?M: Yes. You see, I’ve bought this personal stereo at your shop three days ago. And I’m afraid it hasn’t really matche d up to what I was told about.W: I see. What exactly is the matter?M: Well, first of all, there’s this large scratch across the fron t of it (Q6).W: But you should have noticed that when you bought it (Q4).M: But it was in the box and all sealed up.W: Well, I’m sorry. But it really is your responsibility to check the goods when you buy them (Q4). How are we to know that it wasn’t y ou who made the scratch?M: But t hat’s ridiculous. But anyway, it’s not the most important thing. I really am not happy about this other thing (Q6).W: And what is that?M: Look, it says here that the noise from it should undetectable by other people.W: Y es, that’s right.M: But people can hear it, a nd it’s really embarrassing on the bus and the underground.W: Well, I’m sorry, but it must be the way you are wearing the headphones.M: Look, I know how to put earphones in my ears. Thank you very much (Q5). But what I want to know is what you are going to do about it at all.W: Well, I suppose we could exchange it for another model, if you really aren’t happy with it.M: No, I certainly am not.W: Well, if I could just have the receipt (Q7).M: Oh yes. Well, there is a slight problem about the receipt.Conversation ThreeW: Hello. Happy Time Catering Services. Victoria’s speaking. How can I help you?M: Hello, Victoria. This is Joe Smith from Country Holidays. I wonder if you could do some catering for us next week (Q9). We’re having a small reception (Q8). It’s to launch our summer holiday advertising campaign. Will you be free?W: When exactly is it, Mr. Smith?M: Apr.21st, t hat’s Thursday. Oh, sorry, no. It should be Friday (Q9).W: Oh, yes, I can do that. Where will you be holding it?M: We thought we’d have that at head office and use the conference room, because there is enough room for every one there.W: Ok. What sort of things would you like?M: Just a light lunch, I think, so that people can eat while they move around and talk to each other. You did something similar for us last year. We’d be happy to have the same menu again (Q10). W: Right. I’ll look at my diary and see what you had last time. Oh, I nearly forgot to ask you. How many should I cater for?M: Well, I think most people will be able to come, perhaps around 30. No, let’s say 35 to be sure. W: Right. Thank you for getting in touch, Mr. Smith. I’ll send you confirmation of the arrangements by the end of this week.M: OK.Section B P ASSAGESPassage AFor shopaholics, the post-Christmas period means only one thing —sales! Across the country, prices are slashed on clothing, electronics, home furnishings and more. But London is the place for serious shopping, and you can certainly pick up some amazing bargains (Q11). The sales start on Boxing Day — 26th December, and continue for the month of January, but the keenest bargain hunters get there early to be first through the doors. In Oxford Street, queues formed outside shops ahead of predawn openings for the start of the sales. At Brent Cross in north London, more than 1,000 people were queuing at 3:30 a.m. for the “n ext” clothing store’s sale which began at 4 a.m. Some people even camped outside the shops to be the first in line (Q12). Some people are taking their friends shopping with them and buying their Christmas presents in the sales (Q13A)— a practical but an unromantic way of making sure you get the gift you really want. For a less exciting but less stressful shopping experience, online retailers are also getting in on the actwith January sales of their own (Q13B). The most organized of all are those who are already doing their present shopping for next Christmas (Q13C), in the January sales!Passage BBallroom dancing used to be seen as something rather unfashionable that old people might do (Q14). For the past five years though, the popularity of ballroom dancing has soared thanks to a reality TV show (Q15). “Strictly Come Dancing” is one of the big TV shows. Millions of people tune in every Saturday night to watch the show, which airs from September right up to Christmas (Q16D). In the show, a number of professional ballroom dancers each dance with a celebrity (Q16B). Every week they have to learn a different ballroom dance and perform it live on TV on Saturday night (Q16C). The show demonstrates how glamorous ballroom dancing is. The celebrities get to wear colourful dresses and suits to dance in (Q16A), and it looks like a lot of fun. The TV programme also shows what good exercise it can be to ballroom dance and what hard work is involved in learning the dances and performing them properly.Dance schools around the country have seen a boost in the number of people wanting to learn how to da nce. And it’s not only old people who’re intereste d. Lots of children and young people in their twenties are keen to learn (Q17).Passage CRecently a couple in New Zealand were forbidden from naming their baby son 4Real. Even though New Zealand has quite liberal rules about naming children, names beginning with a number are not allowed (Q18). They decided to call him Superman instead.In many countries around the world, unusual names for children are becoming more popular, especially since the increasing trend for celebrities to give their children unusual names. Some parents choose names which come from popular culture (Q19). For example, there have been six boys named Gandalf after the character in the Lord of the Rings novels and films. Equally, names related to sports are fairly common (Q19). Since 1984, 36 children have been called Arsenal after the football team. Other parents like to make up names or combine names to make their own unique version (Q19), a method demonstrated by Jordan, the British model, who recently invented the name Theaamy for her daughter by combining the names Thea and Amy (the two grandmothers). She was quoted as saying that the accent and double letters were added to make the name “more exotic”.Other countries have much stricter rules when it comes to naming children. Countries including Japan, Denmark, Spain, Germany and Argentina have an approved list of names from which parents must choose (Q20).Section C NEWS BROADCASTNews Item 1 (For Questions 21 to 23)11 fishing boat crew who had been stranded since October in a r emote part of Russia’s Far East have been rescued after sheltering nearly three months at an abandoned military base. The eight men and three women took refuge at the base after their small boats collided on Oct. 10 (Q21). Their attempts to fix one of the boats did not succeed and they had to remain at the abandoned base where there were only flour and cooking fat (Q22). Other supplies at the base, which was abandoned in 2003, included Christmas ornaments, and the crew members put them upon a small tree inside their quarters. But supplies began running low and early this week, five set off on foot across snow fields. On Friday, after four days of trudging, they reached a working military radio station. The center called rescuers, and helicopters were sent to take the 11 to the regional capital (Q23).News Item 2 (For Questions 24 and 25)Spain’s King Juan Carlos, with 32 years on the throne (Q24), turns 70 Saturday. But after years of undeniable adulation among Spaniards for putting down an attempted coup i n 1981, he’s recently faced more difficult times. Small groups of leftists have burned his photo, and fiery criticism has also come from the right with one leading conservative radio host calling for him to step down. Juan Carlos fired back with a rare pub lic defense of his reign in a recent speech. “It’s been the longest period of stability and prosperity in Spain ever in a parliamentary monarchy,” the king said (Q25).News Item 3 (For Questions 26 and 27)It goes against religious taboos in Iraq to involve women in fighting, but three recent suicide bombings carried out by women could indicate insurgents are growing increasingly desperate. On Nov. 4 (Q26), a woman detonated an explosives vest next to a US patrol in Diyala’s regional capital, Baqouba, 35 miles northeast of Baghdad, wounding seven US troops and five Iraqis (Q27). On Dec. 7 (Q26), a woman attacked the offices of a Diyala-based Sunni group fighting al-Qaida in Iraq, killing 15 people and wounding 35 (Q27). Then, on Dec. 31 (Q26), a bomber in Baqouba detonated her suicide vest close to a police patrol, wounding five policemen and four civilians (Q27).News Item 4 (For Questions 28 to 30)Chinese government is poised to conduct its first national survey of pollution sources in February to help control environmental deterioration in the country (Q28). The study will identify and collect data on sources of industrial, agricultural and residential pollution for two months. Last year, China’s environment was facing a grave situation, with several maj or rivers and lakes clogged by industrial waste. China’s environmental cleanup is compromised by more than two decades of rapid economic growth, and a lack of technology especially (Q29). Every province, autonomous region and municipality has set up a census office and will report to a main center staffed by officials from government departments (Q30). Data will be reviewed multiple times before being put into a database and will be analyzed in the second half of 2008. Findings will be examined and approved by mid-2009.。
Prüfung für das Germanistik-Grundstudium (PGG-Prüfung) 2009I.Diktat (10P) (15 Minuten)Sie hören den Text insgesamt viermal. Zuerst hören Sie sich den ganzen Text einm al an. Dann hören Sie sich den Text noch zweimal mit Satzzeichen an. Die längeren Sätze werden in Teilen vorgelesen. Zuletzt hören Sie den ganzen Text noch einmal. Nun fangen wir an!II. Hörverstehen (20P) (20 Minuten)Teil 1: AlltagssituationenSie hören im Folgenden 6 Minidialoge nur einmal. Markieren Sie die Lösungen auf d em Antwortbogen1.a.Im krankenhause b.Auf der Straße c.Auf dem Gericht2.a.Bei Dr.müller b.In der Mensa c.Im Proseminar3.a.Über zwei Kilo b.Zwei Kilo c.Unter zwei Kilo4.a.Nach München b.Nach Mailand c.Nach Trier5.a.Von 5 Uhr bis 17 Uhr b. .Von 7 Uhr bis 15 Uhr c.Von 5 Uhr nachmittags bis 5 Uhrmorgens6.a.Er hat gestern in seinem restaurant Bier getrunken.b.Er war gestern wieder als wilder Mann in einem Restaurant.c.Er hat einen lustigen,wilden Abend lange in seinem Restaurant verbracht.Teil 2Nun hören Sie vier Texte. Jeden Text hören Sie zuerst einmal, dann lesen Sie die A ufgaben und danach hören Sie ihn noch einmal. Markieren Sie die richtigen Lösunge n auf dem Antwortbogen(Für Text A und B:a=richtig b=falsch)Text A7.Christopher Küch hat sich selber ein Gymnasium gewält,als er inder vierten Klasse war.8. Christopher Küch geht jetzt in die achte Klasse und hat gute Leistungen auf dem Gymnasium.Text B9.Es fällt nicht nur in Sudbayern Schnee.10.Nachts ist die tiefste Temperatur unter null Grad.11.Um vierzehn Uhr wurde der Wetterbericht vom Wetteramt München heraus gege ben.Text C12.Was für Mitarbeiter brauchen die meisten Firmen?a.Solche,die Fremdsprachen können.b.Solche,die Englisch und Französisch sprechen.c.Solche,die gute Chancen in der wirtschaft haben.13.Wie viele Gymnasium lernen welche Sprache?a.35 Prozent der Gymnasium lernen Französisch.b.5 Prozent der Gymnasium lernen Russisch.c.2 Prozent der Gymnasium lernen Italienisch.14.Was meinen die Firmen?a.In den Schulen alle Schüler neben Englisch auch Französisch lernen.b.Den Schülern sollte die Möglichkeit gegeben weden,mehrere Fremdsprache lernen zu können.c.Ein Schüler sollte möglichst viele Fremdsprache beherrschen.Text D15.Welche Aussage ist richtig?a.Der Mann redet über seinen dreiseitigen Lebenslauf.b.Der Mann redet darüber ,warum er oft die Stelle gewachselt hat.c.Der Mann blickt auf sein berufsleben zuzück.16.Welche Aussage ist richtig?a.Er war einmal Arbeitslos.b.Er will seinen Arbeitsplatz wechselnc.Er arbeitet als pfleger in einem Altenheim.17.Wie finet der Mann seinen jetztigen Beruf ? Wrum?a.Schrecklich.Weil er dafür zu alt ist.b.Gut.Weil er in Rente weiter als Hausmeister arbeiten möchte.c.Gut.Weil er den Arbeitsplatz mag und bald in Rente geht.Text E18.kauft die Frau jetzt anders ein als früher?a.Ja,sie kauft jetzt keinen Joghurt mehr.b.Nein,sie kauft genau so siel Milch in Packungen wie Früher.c.Na ja,da hat sich nicht wiel geändert.19.Was macht die Frau bei der Mülltrennung?a.Glas und Papier sammelt sie getrennt.b.Die Abfälle bringt sie in die Küche.c.Dosen und Plastik bringt sie in der Garten.20.Was meint die Frau zum Thema Müll unt Mülltrennung?a.Den Müll und die Mülltrennung finder sie gut.b.der Müll ist ein Problem,deshalb muss man etwas tun.c.Die Mülltrennung ist ein Problem für sie.III. Leseverstehen (20P) (40 MINUTEN) Text A Deutsch lernen im Restaurant120 Menschen aus 33 Läandern arbeiten jeden tag in den Restaurant der HamburgerOktoberkette.Sie bedienen die Gäste, kochen oder schenken aus. Verstäandigungsprobleme gibt’s auch beim größten Andrang nicht: Geschäftssprache ist Deutsch. Darauf legt Inhaber Ömer Merdin den größten Wert.Mit 19 Jahren ist er selbst aus der Türkei nach Deutschland gekommen und weiß, wie wichtig ist es ,Deutsch zu sprechen. Darum finanziert er seinen Angestellten regelmaßig Sprachkurse, wenn ihre Deutschkenntnisse nicht ausreichend sind.” Ich will Chancengleichheit für alle schaffen”,sagt Merdin. So viel Engagement wur de 2008 mitden ersten Preis beim bundesweiten Wettbewerb”Kulterelle Vielfalt als Chance”belohnt. Soga rein eigenes Qualifiyierungsangebot für seine Beschäftigten hat sich der Gastronom ausgedacht.Mitarbeiter ohne Berufsabschluss können sich in senen Restaurant zur Fachkraft im Gastgewerbe im Bereich Service und Küche qualifizieren.Markieren Sie die richtigen Lösungen auf dem Antwortbogen(a=richtig b=falsch)21.Ömer Merdin ist erst 19 Jahre alt.22.In den Restaurant von Ömer Merdin warden Deutsch und Türkisch gesprochen.23. Merdin hat den ersten Preis beim bundesweiten Wettwerb …Kulturelle Vielfalt als Chance“ gewonnen.24. Die Angestellten von Merdin haben die Möglichkeit, sich in seinen Restaurants ausbilden zu lassen.Text B Unterschiedliche AnschauungenWer ein Fernsehgerät kaufen will, hat meist konkrete Vorstellungen, was der Apparat leisten soll. Doch sind die Ansprüche von Land zu Land durchaus unterschiedlich. Wie die Gesellschaft für Unterhaltungs- und Kommunikationselektronik meldet, sind sich die Europäer lediglich darin einig, dass der Preis für die gebotene Leistung des Apparat stimmen muss. Europäer legen mit 50% den größten Wert auf den Preis eines fernsehgeräts. Beim Design sieht es anders aus. Für 44% der Einfache Bedienung des Geräts: Während Österreicher (34%) und Deutsche (32%) gerade hierauf achten, interessieren das Engländer (13%) und Italiener (11%) recht wenig. Dagegen wünschen sich die Engländer hohe Zuverlässigkeit (46%). Auch der gute Name eines Fernsehapparats ist den Engländern mit 25% wichtig, wogegen die Deutschen nur mäßig (12%), Holländer und Schweden mit 3 bzw. 2 Prozent fast gar nicht interessiert. Markieren Sie bitte die richtigen Lösungen auf dem Antwortbogen!25. Beim Kauf eines Fernsehappratsa. achten die Europäer nur auf den Preis.b. legen due Schweden weniger Wert auf das Design als die Holländer.c. ist den Europäern nur das gute Design wichtig.26. Die Bedienung eines Fernsehgeräts sollte nach Meinunga. von Engländern sehr einfach seinb. von Österreichern und Deutschen leicht sein.c. von Italienern nicht interessant sein.27. Wer legt worauf Wert_a. Den guten Namen eines Geräts halten ein Viertel der Engländer für wichtig.b. Holländer und Schweden legen großen Wert auf den guten Namen eines Geräts.c. Die Deutschen sind überhaupt nicht am Namen eines Fernsehgeräts interessiert.Text C RingparabelVor vielen Jahren lebte ein König im Osten, die einen wertvollen Ring besaß. Dieser Ring hatte eine Eingenschaft: Wer ihn trug, wurde von Gott und allen Menschen ohne Ausnahme geliebt. Deshalb nahm der König ihn nie von seiner Hand.Eingentlich wurde der Ring immer vom Vatter an den Lieblingssohn weitgegeben. Der König jedoch hatte drei Söhne, die er alle gleich liebte. Er konnte sich nicht entscheiden, welchem Sohn er den Ring geben sollte. Also ließ er heimlich noch zwei gleiche Ringe machen und schenkte jedem Sohn einen. Als der König stab, kam es zum Streit zwischen den Söhnen. Jeder galubte, den echten Ring zu Richter, einem weisen Mann, und erzählten ihm die Geschichte.Der Richter konnte das Rätsel auch nicht lösen und wollte sie schon wegschicken. Plötzlich erinnerte er sich daran, dass der richtige Ring seinen Besitzer bei allen beliebt machen sollte. Er fragte die Söhne: …Welchen Bruder haben zwei von euch am liebsten?“ Als sie nicht gleich antworteten, sprach er: …Was, ihr sagt nichts? Die Ringe wirken nicht? Jeder von euch liebt sich selbst also an meisten. Dann sind viellecht alle drei Ringe falsch. Keiner von ihnen besitzt die Macht,sich bei allen Menschen beliebt zu machen. Vermutlich hat euer Vater den Ring verloren und drei neue machen lassen.“Der weise Richter gab ihnen den Rat, so zu leben, als hätte jeder von ihnen den echten Ring. Nur auf diese Weise würde es ihnen gelingen, die Macht des Ringes zu wecken. Markieren Sie bitte die richtigen Lösungen auf dem Antwortbogen!( a=richtig, b=falsch )28. Der König hat den Ring an den Lieblingssohn weitergegeben.29. Die Ringe, die die drei Söhne bekommen, sehen gleich aus.30. Die Söhne haben den Richter am liebsten.31. Die Söhne können nicht mehr die Macht des Ringes wecken.Text D Fachkräftemangel in DeutschlandViele Unternehmen suchen dringend nach Informatikern. An deutschen Haupt- und Berufsschulen fehlen rund 50 000 Lehrer. Eine Studie zu diesem Thema kommt nun dem Ergebnis, dass diese schlimme Lage teilweise von Hochschulen und Wirtschaft selbt gemacht ist.Schuld an der Lage ist der sogenannte …Schweinezyklus“ – ein Wirtschaftstrend, nach dem auf ein Überangebot an Studenten in einem Studienfach ein Rückgang der Studienfänger in dem überlaufenen Fach folgt. Einige Jahre später fehlt in den Unternehmen dann der Nachwuchs.So schränkten viele Betribe Anfang der 90er Jahre ihre Ausbildung ein. Ingenieure und Informatiker wurden bur noch selten eingestellt. Durch die abnehmende Personalnachfrage durch die Industrie ging die Zahl der Studienanfänger in den Bereichen Informatik, Elektrotechnik und Maschinenbau um mehr als die Hälfte zurück. Als die Wirtschaft wieder besser wurde, fehlten die Fachkräfte.Der Arbeitsmarktforscher Gerhard Bosch fordert, dass man unabhängig vonSchwankungen in der Bevölkerungsentwirklung und wirtschaftlichen Krisen nach einem Weg suchen müsse, um …Schweinezyklen“ künftig vorbeugen zu können.Wenn in einem Unternehmen qualifizierte Kräft fehlten, so mussten bisher meist die Mitarbeiter helfen. Doch längere Arbeitszeiten und Überstunden lassen sich nicht beliebig ausdehnen.Generhell wird der Bedarf an qualifizierten Fachkräften wach sen: …Ob es zu einem Fachkräftemangel kommt, hängt sehr stark von den Ausbildungszahlen und der Modernität des Ausbildungssystems ab“, so Bosch. Die Chancen für Arbeitskräfte mit geringer Qualifikation seien schlecht, auch wenn einfache Tätigkeiten weiterhin gefragt blieben. Markieren Sie bitte die richtigen Lösungen auf dem Antwortbogen!32. Wer ist am Mangel an Fachkräften schuld?a. Hochschulen und Wirtschaftb. Unternehmenc. Hochschulen33.Was versteht man unter dem Begriff …Schweinezyklus“?a. Auf ein Überangebot an Studienanfängern in einem Studienfach folgt normalerweise ein Rückgang der Absolventen in dem Fach.b. Auf ein Überangebot an Studien in einem Studienfach folgt ein Rückgang der Studienanfängern in dem Fach.c. Auf ein Überangebot an Studien in einem Studienfach folgt ein Rückgang der Studienfächer.34. Welche Aussage ist richtig?a. Anfang der 90er Jahre förderten viele Beriebe ihre Ausbildung.b. Anfang der 90er Jahre ging die Zahl der Studienanfänger zurück.c. Anfang der 90er Jahre schränkte die Industrie die Zahl der Fachkräfte ein.35. Was ist die Meinung von Gerhard Bosch?a. Der Bedarf an qualifizierten Fachkräften wird weiter sinken.b. Die Wirtschaftskrisen haben den Fachkräftemangel verursacht.c.Arbeitskräfte mit geringer Qualifikation hanben weniger Chancen.Text E Konsum mit guten GewissenDen Trend zum nachhaltigen Konsum beobachtet man fast überall in der Welt.Immer mehr leute wollen möglichste schadstoffreie Lebesmittel haben, also Bio.Und mit der Veröffentlichung des Weltklimaberichts im Frühjahr 2007 gewann die Bewegung noch einmal an Fahrt.Als Vorriter haben Soyiologen und Marktforscher die …Lohas“(…Lifestyle of heslth and Sustainability“)ausgemacht.Sie diskutieren über Solaryellen auf dem Dach,Naturko smetik auf der Haut,und wie man im Skiurlaub der Umwelt möglichst wenig schadet.Aber mit der Ökobewegung der 80er-Jahre haben sie wenig gemein.Sie wollen Öko mit Spaß verbinden und nicht mit Veryicht.Auch Lohas spielt Gesumdheit eine große Rolle.Was sie kaufen,muss frisch sein,einen hohen Nahrwert haben.Wer zu dieser Gruppe gehört,konsumer mit Lust.aber immer Ökologisch korrekt.Sie achten beim Kauf auf ökologische Aspeckte und wollen mit gutem Gewissen Konsumieren.Der Einfluss der Lohas gründet nicht auf politischer Macht,sondern auf ihrer kollektiven Kaufkraft.Man schätzt die Zahl der neuen Ökokonsumenten in Deutschland auf etwa achtmillionen und ihre Kaufkraft auf rund 200 milliarden Euro.Im Vergleich zu früheren Ökopropagandisten sind sie pragmatischer und kund sehnen sich einfach nach mehr Lebensqualität.Aber unpolitisch seien die Lohas keineswegs.“ Sie wollen ja gerade Konsumetscheidungen als Hebel nutzen,um eine andere politik der Unternehmen zu bewirken. Das ist politisch ,nur eben nicht im Rahmen de s politischen institutionen System“Markieren Sie bitte die richtigen Lösungen auf dem Antwortbogen!36.Was heißt …gewann die Bewegung noch einmal an Fahrt.“?a.Bei der Bewegung hat man wieder gewonnen.b.Die Bewegung entwickelt sich mit steigender Geschwindigkeit.c.Die Bewegung fährt immer noch fort.37.Die Lohasa.diskutieren nur über Solarzahl.b.sind gegen die Ökobewegung der 80-er Jahrec.versuchen Öko mit Spaß zu verknüpfen.38.Wie sieht Konsumverhalten der Lohas aus?a.Sie konsumieren immer ohne Lustb.Gesundheit hat einen großen Einfluss auf das Konsuverhalten.c.Der Preis ist das eizige Ziel der Lohas.39.Warum kann man den Einfluss der Lohas erkennen?a.Die Lohas haben einen großen Kaufkraft.b.Die Lohas demonstriren für die ökobewegung.c.Der politike legen großen Wert auf die Interessen der Lohas.40.Worin liegt der Unterschied zwischen Lohas und früheren Ökopropagandisten?a.Die früheren Propagandisten sind praktischer.b.Die Lohas wollen mit guten Gewissen konsumieren.c.Die Lohas verzichten oft auf Konsum,um die Umwelt zu schützen.IV.WORT,SATZ,UND TEXT(35P) (40 MINUTEN)A.Bitte markieren Sie die grammtisch richtigen Lösungen auf dem Antwortbogen.(7P)41.Die kinder werden _____ Toleranz erzogen.a.zub.aufc.mit42.Eure Eltern feierten________ Hochyeistag mit einem Fest.a.Ihrenb. eurenc.ihren43.Peter verabredte sich mit seiner neuen Freundin__________den nächsten Tag.a.fürb.zuc.an44.Ich bin ganz ihrer Meinung und möchte dazu noch einen Vorschlag_________a.gebenb.machenc.stellen45.Die kollegen hielen ihren_________für netten Menschen.a.Vorgesetzterb.Vorgesetetenc.Vorgesetzte46.Er______ das schon vorher gewusst haben,sonst würde er anders reagieren.a.kannb.mussc.will47.Sein rundes Gesicht wirkte plötzlich etwas blasser als_____a.vorherb.bevorc.früher48.Die Neueverheirateten konnten die lange Trennung nicht ________a.verbringenb.haltenc.ertragen49._________die moisten Studenten gegen die Ausschaltung des Stromes um 12 Uhr in der Nacht sind,wird es doch getan.a.Trotzdemb.Zwarc.Obwohl50.Können Sie feststellen,welche Änderung sich inden vergangenen Jahren ___________?a.ergenben hatb.ergibtc.ergeben wird51.Angebot und_______bestimmen die wirtschaft,heiß es oft.a.Bedarfb.Nachfragec.Bedürfnis52.Die Rolle einer Mutter ist __________.Sie ist auch Lehrerin.Von ihr lernen Kinder viel.a.universalb.insgesamtc.gemeinsam53.Die Deutschen Lieben frische Luft,und______sie endlich Ferien haben,fahren sie in die Natur.a.nachdemb.wennc.indem54.Wann fliegt den eure Maschine?---Unser Flug geht um 14 Uhr.Ganz__________.a.ordentlichb.pünktlichc.plannmässig B. Bitte ergänzen Sie die passenden Wörter! Schreiben Sie sie dann auf den Antwor tbogen. (10P)55.Sabine stammt _____einer kinderreichen Famlie.56.In dieser Dunkelheit schien die Zeit stil zu______________.57.Wir machen sine Pauschal-Reise.Das bedeutet ,ein _________für alles.58.Können Sie bitte_________zehn Minuten noch einmal anrufen?59.Auf diese _______ wird uns gelingen,das Problem zu lösen.60.Falls Sie weitere__________haben,wenden Sie sich bitter an meine Sekretärin.61.Nur weinige Autos können________ eine so lange Erfolgsgeschichte zurückblicken, wie der Käfer.62.Kinder,die von Lehrer als besonders inteligent einschätzt werden,weisen am Ende des Semesters einen bedeutend höheren Intelligenzquotienten___________.63.Der junge Mann versucht ein leben zu schaffen,die seinen Wünschen_________.64.Wenn ich nicht so viele Prüfungen__________,könnten wie selbstverständlich öfte r ausgehen.65.Schau mal.Da ist schon wieder der teure Wagen,_______hier jeden tag parkt.66.Wir wollen den Kopf auf keinen Fall________lassen, sondern die Prüfung nächste s Jahr nocheinmal machen.67.__________ins Ausland geht, muss die Welt und die Menschen dort mit anderen Augen sehen.68.Draußen war es ein grauer kalter Novembermorgen, und es regnete________strömen.69. Kommen Sie endlich herein und machen Sie die Tür zu! Es___________.70. Der Junge war noch zu klein, _______ in die Schule zu gehen.71. Das Kind wußte nicht, ________ es tun sollte, deshalb blieb es einfach stehen und wortlos zu.72. Die Diebe arbeiteten vermutlich zu weit: Einer begann ein Gespräch mit einem Fahrgast, der ________ nahm ihm heimlich die Geldbörse aus der Tasche.73. Was muss man eigentlich tun, um so _______ wie dich loszwerden, kannst du mir einmal verraten?74. Heraus mit der _________ ! Mit wem warst du gestern Abend in Disko?C. Bitte ergänzen Sie die passenden Wörter auf dem Antwortbogen. (10 P)Text A (Folgende Wörter stehen Ihnen zur Wahl,aber nicht alle Wörter werden hier passen: an, der, finanziert, für, jährlichen, jährigen, steht ,stattfindet,und, unterrichtet, war, veranstaltet,zur )…Jugend musiyiert“ ist ein Wettbewerb(75)________ Jugendliche, der 2009 yum 46. Mal(76)_________. Viele tausend junge Leute melden sich in jedem Jahr(77)______. Mitmachen kann jeder, (78)________ privaten Musikunterrcht erhält, an einer Musikschule oder an der allgemein bildenden Schule (79)_________ wird. Bei …Jugend musiyziert“(80)_____ der Team-Gedanke im Vordergrund: miteinander zu musizieren, gemeinsam ein Werk der Musikliteratur zu erarbeiten und es vor Publikum(81)________- Aufführung zu bringen. …Jugend misiziert“ wird (82)_________ vom Bundesministerium für Familie, Senioron, Frauen und jugend (83)_______ von der Sparkassen-Finanygruppe. Hinzu kommen im (84)__________ Wechsel der Austragungsorte weitere Förderer.Text B(Füllen Sie bitte die folgenden Lücken nach dem Kontext.)50 Kilometer lang war die Lichterkette, die am Donnerstag von einigen lokalen Initia tiven (85)________ wurde. 15 000 Menschen mit Fackeln und Lichtern(86)_______ der Hand standen auf der Straße. So drückten Sie ihren(87)________ gegen Atome nergie aus. (88)________ nur Anwohner waren dabei, auch aus Berlin und Humburgwaren Menschen angereist. Die Organisatoren waren zufrieden, weil sie vorher (89) _________ nur etwa 10 000 Menschen gerechnet hatten. Dieses Ereignis zeigt, (90) _________ die Anti-Atom-Bewegung wieder da ist. Dieses Jahr ist ein wichtiges Jahr im Kampf(91)_________ die Atomkraft, denn (92)__________ jeden Preis muss ver hindert werden, dass der Atomausstieg nach der Bundestagswahl wieder rückgängig (93)_________ wird. Atomkraft ist die Energieform (94)_________ vorgestern. Die Z ukunft gehört den erneuerbaren Energien!D. Formen Sie die unterstrichenen Satzteile und Partizipialkonstruktionen in Nebensätze um bzw. die unterstrichenen Nebensätze in Satzteile oder Partizipialkonstruktione n.Schreiben Sie die Lösung auf den Antwortbogen. (4 P)95. Mit der gestohlenen Waffe erschoss der junge Mann viele unschuldige Mensche n.96. Der Regisseur, der während des Kriegs in Frankreich im Exil lebte, wurde yum Filmfest eingeladen.97. Vor der Installation eines Antivirusprogramms stürzte mein Computer oft ab.98. Sich gany auf die Arbeit konzentrierend, schaffte er am Wochenende seinen Pr ojektbericht.E. Was sagt man da?(Wie reagieren Sie darauf?)Bitte vervollständigen Sie die Sätz e situationsangemessen und schreiben Sie die Lösungen auf den Antwortbogen. (4 P)99. A: Ist das Restaurant in der Nähe teuer?B: Ja,es ist viel teurer geworden,seitdem_______________________.100. A: Dein neues Handy ist toll!B: Ja. Es sieht nicht nur super aus, sondern________________________. 101. A: Fährt dein Nachbar täglich mit dem Fahrrad zur Arbeit?B: Natürlich. Er fährt bei jeden Wetter mit dem Fahrrad ins Büro,um ________ __________________.102. A: Was hast du morgen vor?B: Da habe ich noch keinen konkreten Plan. Bei schönem Wetter____________ _____________.V. SCHRIFTLICHER AUSDRUCK (15 P) (40 MINUTEN) Schreiben Sie einen Text zu einem der folgenden Aufgaben. Ihr Text sollte nicht we niger als 120 Wörter haben.A.Im Folgenden ist der Anfang einer Geschichte. Erzählen Sie bitte die Geschichtezu Ende.Ein merkwürdiges HausNicht weit von unserem Dorf war ein Wald, in dem ein altes verfallenes Haus stand. Ich spielte oft min anderen Kindern in diesem Wald, aber in das Haus gingen wir nie, denn es sah nicht gemütlich aus. Wir waren schon mit dem Wald zufrieden, in dem wir schaukelten, wir liefen und Blindekuh spielen.An einen späten Nachmittag im Sommer spielten wir wieder Blidekuh...........B.Beschreiben Sie bitte das folgende Schaubild.Immer mehr Leute nehmen einen Konsumentenkredit auf,um sich einen Luxusartikel wie z.B. Autos, Möbel, Musikanlagan usw. Anyuschaffen. Das folgende Schbild zeigt die steigende Tendeny der Konsumentenkredite im zeitraum von 1970 bis 1988 in Deutschland.。
2009年北京航空航天大学英语专业(英美文学)真题试卷(总分80,考试时间90分钟)3. 名词解释1. Robinson Crusoe2. Henry Fleming3. The Bible4. Samuel Taylor Coleridge5. Death of a Salesman6. The Gothic novel7. Santiago8. Samuel Beckett9. Uncle Tom10. Ideology8. 分析题1. Why is the Knight first in the General Prologue to tell a tale inCanterbury Tales?2. Moby-Dickfeatures several seemingly insane characters. How does insanity relate to this story? How do these characters contrast with one another?3. Analyze the theme of Social Class in Dickens"Great Expectations.To be, or not to be—that is the question:Whether this nobler in the mind to sufferThe slings and arrows of outrageous fortuneor to take arms against a sea of troublesAnd by opposing end them. To die, to sleep—No more...Questions:4. A. From which play are these lines taken from?5. B. Who is the playwright?6. C. Who is the speaker?7. D. What does this speech show?The apparition of these faces in the crowd; Petals on a wet, black bough. Questions:8. A. What is the title of this short poem?9. B. Who is the author?10. C. What two images are juxtaposed or placed next to each other in this poem?11. D. How do you appreciate this poem?It is a truth universally acknowledged that a single man in possession of a good fortune must be in want of a wife. However little known the feelings or views of such a man may be on his first entering a neighborhood , this truth is so well fixed in the minds of the surrounding families, that he is considered as the rightful property of someone or other of their daughters. Questions:12. A. From which novel is this passage taken from?13. B. Who is the author of this novel?14. C. What is the literary style of this novel?15. D. What is this story about?Young Goodman Brown came forth at sunset into the street at Salem village; but put his head back after crossing threshold, to exchange a parting kiss with his wife. And Faith, as the wife was aptly named, thrust her own pretty head into the street, letting the wind play with the pink ribbons on her cap while she called to Goodman Brown. Questions:16. A. This passage is taken fromYoung Goodman Brown, who is the author?17. B. What is the symbolic meaning of "pink ribbons"?18. C. What is a symbol in literature?O, my love" s like a red, red rose. That" s newly sprung in June; O, my love" s like the melodie That" s sweetly played in tune. As fair art thou, my bonnie lass. So deep in love am I; And I will love thee still, my dear. Till a" the seas gone dry.…Questions:19. A. Who wrote this poem?20. B. What is the title of the poem?21. C. The odd-numbered lines are iambic tetrameters, what about the even-numbered lines?22. D. What is the rhyme scheme?23. E. What do you know about the poem?EUNICE; What"s the matter, honey? Are you lost?BLANCHE: They told me to take a street-car named Desire. And then transfer to one called Cemeteries and ride six blocks and get off at—Elysian Fields!EUNICE: That" s where you are now. Questions:24. A. From which play are the conversations taken?25. B. Who is the playwright?26. C. How to define "Desire" in the play?We are what we pretend to be, so we must be careful what we pretend to be.Mother Night, 1961 Anyone who cannot understand how useful a religion based on lies can be will not understand this book either.Cat"s Cradle, 1963 This is a novel somewhat in the telegraphic schizophrenic manner of tales of the planet Tralfa-madore, where the flying **e from.Slaughterhouse-Five, 1969 Questions:27. A. The author of these novels is Kurt V onnegut, name some other novelists who employ black humor.28. B. Define the literary term Black Humor with reference to the above quotations.Tragedy is, then, an imitation of a noble **plete action, having the proper magnitude; it employs language that has been artistically enhanced by each of the kinds of linguistic adornment, applied separately in the various parts of the play; it is presented in dramatic, not narrative form, and achieves, through the representation of pitiable and fearful incidents, the catharsis of such pitiable and fearful incidents. Questions:29. A. This definition of Tragedy is quoted fromPoetics, who is the author ofPoetics?30. B. Highlight the chief contributions ofPoeticsto the theory of tragedy.。
作物学报ACTA AGRONOMICA SINICA 2013, 39(2): 198−206 /zwxb/ ISSN 0496-3490; CODEN TSHPA9E-mail: xbzw@DOI: 10.3724/SP.J.1006.2013.00198回交自交系(BIL)群体4对主基因加多基因混合遗传模型分离分析方法的建立王金社赵团结盖钧镒*南京农业大学大豆研究所 / 国家大豆改良中心 / 农业部大豆生物学与遗传育种重点实验室(综合) / 作物遗传与种质创新国家重点实验室, 江苏南京 210095摘要: 主基因加多基因混合遗传模型是用于分析数量性状表型数据的统计分析方法, 该方法便于育种工作者利用杂种分离世代的数据对育种性状的遗传组成初步判断,制定相应的育种策略,也可用于校验QTL定位所揭示的数量性状的性状遗传组成。
回交自交系(BIL)群体是永久性群体, 可以进行有重复的比较试验, 适用于受环境影响较大的复杂性状的遗传研究。
本研究以BIL群体为对象构建了4对主基因、主基因加多基因分离分析方法的遗传模型, 包括2类11个遗传模型。
利用基于IECM (iterative expectation conditional maximization)算法的极大似然分析方法估算各个混合遗传模型中的分布参数, 用AIC值和一组适合性测验结果选取最优模型, 并从入选模型的分布参数通过最小二乘法估计遗传参数。
由1个模拟的随机区组试验对模型进行验证, 模拟群体中遗传参数的估计值与设定值之间具有很好的一致性。
利用本文建立的模型重新分析大豆回交自交系群体(Essex×ZDD2315)及其亲本对胞囊线虫(Hetero-dera glycines Ichinohe) 1号生理小种的抗性数据后发现4对主基因模型优于原报道的3对主基因模型, 说明本方法的有效性和正确性。
关键词:回交自交家系群体(BIL); 主基因加多基因混合遗传; 分离分析Establishment of Segregation Analysis of Mixed Inheritance Model with Four Major Genes Plus Polygenes in Backcross Inbred Lines (BIL) Populations WANG Jin-She, ZHAO Tuan-Jie, and GAI Jun-Yi*Soybean Research Institute of Nanjing Agricultural University / National Center for Soybean Improvement / Key Laboratory for Biology and Genetic Improvement of Soybean (General), Minister of Agriculture / National Key Laboratory for Crop Genetics and Germplasm Enhancement, Nanjing 210095, ChinaAbstract: The segregation analysis of major genes plus polygenes is a statistical method for genetic analysis of quantitative traits. The method is particularly valuable for plant breeders to use their data accumulated from segregation populations to estimate the genetic system of target traits, which is necessary for designing breeding strategies and also useful for validating the results of QTL mapping. The backcross inbred line (BIL) population is one of the permanent populations, which is suitable for genetic analysis of complex traits and can be used in replicated experiments. For BIL population, the analytical procedures of three and less major genes plus polygenes mixed inheritance models have been established. The objective of the present study was to estab-lish the analytical procedures of segregation analysis for four major genes plus polygenes mixed inheritance models in BIL popu-lation. Eleven genetic models with four additive and (or) epistatic major genes including those without and with polygenes were established. The component distribution parameters were solved and estimated by using maximum likelihood method based on IECM (Iterative Expectation Conditional Maximization) algorithm. Among the possible models, the best one was chosen accord-ing to Akaike’s Information Criterion (AIC) and a set of tests for goodness of fit. Then the genetic parameters of the optimal model were estimated through the least square method. For demonstration of the established procedures, a simulated data set of a本研究由国家重点基础研究发展计划(973计划)项目(2009CB1184, 2010CB1259, 2011CB1093), 国家高新技术研究发展计划(863计划)项目(2011AA10A105, 2012AA101106), 国家自然科学基金资助项目(31071442, 32671266), 农业部公益性行业专项(200803060), 江苏省优势学科建设工程专项和国家重点实验室自主课题项目资助。
DOUGLAS A. KIBBEEDepartment of French Telephone: (217) 333-2020University of Illinois FAX: (217) 244-22232090 Foreign Languages Building E-MAIL: dkibbee@ 707 South Mathews AvenueUrbana, IL 61801USAEDUCATIONPhD 1979 Indiana University, Bloomington, French LinguisticsMinors: French Literature, Medieval StudiesThesis: The Establishment of the French Grammatical Tradition, 1530-1580.Certificate in Medieval Studies 1978 Indiana UniversityMA 1975 Indiana UniversityFrench LiteratureBA 1971 Colgate University, Hamilton NYMajor: French LiteratureMinor: AnthropologyWORK EXPERIENCE2005- Executive Coordinator, Foreign LanguagesUniversity of Illinois1999- ProfessorDepartment of FrenchUniversity of Illinois1994-2000 HeadDepartment of FrenchUniversity of Illinois1993-94 Visiting ProfessorDépartement de Lettres ModernesUniversité de NantesFrance1991-1999 Associate ProfessorDepartment of FrenchUniversity of Illinois1990-1991 Resident DirectorSummer Study Program at the Université Laval, QuébecCommittee for Institutional Cooperation1985-1991 Assistant ProfessorDepartment of FrenchUniversity of Illinois1979-1985 Assistant Professor/Associate ProfessorDepartment of Modern LanguagesWestern Kentucky UniversityBowling Green, KYCourses TaughtAt the University of IllinoisFrench 134 Enriched Intermediate French, IIFrench 207 Advanced Grammar and CompositionFrench 414 Advanced Composition and StylisticsFrench 417 Histoire de la langue françaiseFrench 418 Linguistic Minorities in EuropeFrench 419 Techniques in Translation IFrench 421 Techniques in Translation IIFrench 429 Langue, Langage et Linguistique au XVIe siècleFrench 429 History of French Language PolicyFrench 429 Language and Public PolicyFrench 529 Linguistic Minorities in FranceFrench 529 French LexicologyLinguistics 260 Language and the LawLinguistics 340 History of LinguisticsCampus Honors Program Senior Seminar: “Language and the Law”Ford Foundation Seminar: “Transnational Culture Industries and Local Sites of Artistic Production”: led section on “Globalization and Cultural Protection in France” (guest lecture)European Union Seminar, “The European Charter for Regional or Minority Languages” (guest lecture)At the Université de Nantes, 1994Histoire de la langue française (Licence)La langue de Pascal (Agrégation)Grammaire du français (DEUG II)Middle English (Licence)Courses taught on-line to students around the world:Techniques in Translation, I & IIFrench LexicologyHistory of the French LanguageFrequently included on the List of Teachers Rated Excellent by their StudentsPhD direction:PhD Thesis Director: Hugh Ortman (French Linguistics), Michel Kouassi Nguessan (French Linguistics) Patrick Reidenbaugh (French Linguistics); Samira El-Atia (Second Language Acquisition and Teacher Education = SLATE), Fallou Ngom (French Linguistics – co-director); Auguste Zoure (FrenchLinguistics); E-Jung Choi (French Linguistics), Geneviève Maheux-Pelletier (SLATE, co-director) PhD Committees, French: Ngame Matuku (SLATE), Kirk Hagen (SLATE), Marc Deneire (SLATE), Steven Daniell (French Literature), Sharon Michalove (Educational Policy Studies), Denise Koglin (SLATE);Moses Kambou (Educational Psychology), Elizabeth Martin (SLATE), Patricia Phillips-Batoma (French Literature); Wynne Wong (SLATE), Mylène Priam (French Literature)in progress: Samira Hassa (French Linguistics), Aimée Johansen Alnet (Linguistics), Hyunju Park(Linguistics), Antonio Reyes-Rodriguez (Spanish, Italian and Portuguese); Errol O’Neill (SLATE)RESEARCH AND PUBLICATIONSPUBLICATIONSBooks1991 For to Speke French Trewely The French Language in England, 1000-1600: Its Status, Description and Instruction. Amsterdam: John Benjamin.1991 New Analysis of Romance Languages: Proceedings of the Eighteenth Linguistic Symposium on Romance Languages. Amsterdam: John Benjamins. (Editor, with Dieter Wanner)1998 Language Legislation and Linguistic Rights. Amsterdam/Philadelphia: John Benjamins. (Editor)2005 Voices of the Elders: An Evening of Stories with the Baoulé (= Wortkunst und Dokumentartexte in Afrikanischen Sprache, 22). Hamburg: Rüdiger Köppe Verlag. (Translation of Baoulé folktales (Côte d’Ivoire))2006 French: A Linguistic Introduction. Cambridge: Cambridge University Press. (with Zsuzsanna Fagyal and Frederick Jenkins)Under contract with Librairie Champion (Paris), a critical edition of Liberté de la langue françoise dans sa pureté (1651) by Scipion Dupleix. (with Marcus Keller)Under contract with John Benjamins (Amsterdam/Philadelphia), Editor, Selected Papers from the Tenth International Conference on the History of the Language Sciences.Journal: Editor of special issue2003: Editor of special issue of Histoire Epistémologie Langage devoted to Language Policy and Linguistic TheoryArticles and Book Chapters1982 "Iço/ço: The i - Forms of the Demonstratives in the Chanson de Roland." SECOL Review 6: 122-30. 1985 "Progress in Renaissance Bilingual Lexicography." Dictionaries 7:21-31.1986 "John Palsgrave's Lesclarcissement de la langue francoyse (1530)." Historiographia Linguistica 12:27-62.1986 "The Humanist Period in Renaissance Bilingual Lexicography." The History of Lexicography. R. R. K.Hartmann, ed., 137-146. Amsterdam: John Benjamins.1987 "Bilingual Lexicography in the Renaissance: Palsgrave's English-French Lexicon." Papers in the History of Linguistics. Hans Aarsleff, L. G. Kelly & H.-J. Niederehe, eds., 179-188. Amsterdam: JohnBenjamins.1988 "L'enseignement du français en Angleterre au seizième siècle." La langue française au XVIe siècle: Usage, enseignement et approches descriptives (=La pensée linguistique, 2), Pierre Swiggers & Willy Van Hoecke, eds., 54-77. Louvain: Leuven University Press.1989 "Enseigner la prononciation du français au seizième siècle". Documents de la Société Internationale pour l'Histoire du Français Langue Étrangère ou Seconde 3:15-20.1989 "Les manuels anglais du XVIe siècle et l'imprimerie". Documents de la Société Internationale pour l'Histoire du Français Langue Étrangère ou Seconde 4:18-21.1990 "French Grammarians and Grammars of French in the Sixteenth Century". History and Historiography of Linguistics, Hans-Josef Niederehe & Konrad Koerner, eds. I, 301-314. Amsterdam: John Benjamins. 1990 "Sources négligées dans l'histoire du vocabulaire." Historical Linguistics,1987. Henning Andersen & Konrad Koerner, eds., 265-273. Amsterdam: John Benjamins.1990 "Language Variation and Linguistic Description in 16th-Century France". Historiographia Linguistics 18: 49-65.1992 "Renaissance notions of medieval language and the development of historical linguistics" Journal of Medieval and Renaissance Studies 22:41-54.1992 "16th-Century Bilingual Dictionaries (French-English): Organization and Access, Then and Now".Historical Dictionary Databases, T. Russon Woolridge, ed., 33-68. Toronto: Centre for Computing in the Humanities.1993 "Durkheim, Language and History: A Pragmatist Perspective". Sociological Theory 11:152-172. (Co-authored with Robert A. Jones)1993 "The First French Grammar in English? Ms Walter Rye 38". Historiographia Linguistica 19:415-418.1993 "World French Takes on World English: Competing Visions of National and International Languages".World Englishes 12:209-221.1994 "Renaissance Linguistics: The French Tradition " Encyclopedia of Language and Linguistics, R. E.Asher, ed., Volume 7, pp. 3536-3540. London/Cambridge: Pergamon Press. [Revised version reprinted in A Concise History of the Language Sciences from the Sumerians to the Cognitivists, E. F. K. Koerner, ed., 161-166. Cambridge: Pergamon, 1996]1994 “Claude de Sainliens” and “John Palsgrave”. Encyclopedia of Language and Linguistics, R. E. Asher, ed., Volume 7, p. 3640 and Volume 6, pp. 2915-2916. London/Cambridge: Pergamon Press.1994 "Baret's Alvearies: Challenges for the Scholar". Early Dictionary Databases, Ian Lancashire & T.Russon Wooldridge, eds. 99-108. Toronto: Centre for Computing in the Humanities.1994 "John Palsgrave". Lexicon Grammaticorum. Harro Stammerjohann, ed., pp. 689-690. Tübingen: Niemeyer.1995 "Emigrant Languages and Acculturation: The Case of Anglo-French" Rasmus Rask Quarterly 2:1-20. 1995 "Assertion/atténuation, subjectivité/objectivité en anglais et en français: ‘seem‘ / ‘sembler’" Relations discursives et traduction, ed. by Michel Ballard, 73-87. Lille: Presses universitaires.1995 "The 'New Historiography', the History of French and 'Le Bon Usage' in Nicot's Dictionary (1606)"Cultures, Ideologies and the Dictionary. Studies in Honor of Ladislav Zgusta. Braj Kachru & Henry Kahane, ed., 103-118. Tübingen: Niemeyer.1995 "Dictionaries and Usage in 17th-Century France: Le 'Naïf François", The History of the Language Sciences. Kurt Jankowsky, ed., 167-177. Amsterdam: John Benjamins.1998 “La syntaxe entre démarche linguistique et didactique”. Histoire de la diffusion et de l’enseignement du français dans le monde, special issue of Le français dans le monde, Willem Frijhoff & André Reboullet (eds.), 133-141.1998 “Legal and Linguistic Perspectives on Language Legislation”. In Language Legislation and Linguistic Rights, Douglas A. Kibbee, ed., 1-23. Amsterdam/Philadelphia: John Benjamins.1998 “Théorie linguistique et droits humains linguistiques: perspectives tirées de l’histoire des communautés linguistiques en France”. DiversCité, electronic journal of the Université du Québec à Montréal(http://www.teluq.uquebec.ca/diverscite/)1999 “The Dictionary in the Service of the State: the Dictionnaire de l’Académie Française and the Dictionnaire des termes officiels”. Dictionaries 20.23-36.1999“Language for Special Purposes Goes On-Line” (co-authored with Elizabeth Martin) Global Business Languages 1999: 102-121.1999 “The ‘People’ and their Language in 19th-century French Linguistic Thought” The Emergence of the Modern Language Sciences. Studies on the transition from historical-comparative to structurallinguistics in honour of E. F. K. Koerner, Sheila Embleton, Hans-Josef Niederehe & John E. Joseph,eds., I, 111-127. Amsterdam/Philadelphia: John Benjamins.2000 “Historical Perspectives on the Place of Anglo-Norman in the History of the French Language” French Studies 54.137-153.2000“Le dictionnaire de Trévoux et la politique de la langue française”. Actes du colloque de Trévoux, International Society for the History of Dictionaries(http://www.univ-lyon3.fr/siehldaweb/trevoux/Kibbee-T.htm)2000 “Language Instruction for Western European Travelers” in Trade, Travel, and Exploration in the Middle Ages, John Block Friedman & Kristen Mossler Figg (eds.), New York: Garland, pp. 330-331.2001 “From Holyband to Mauger: Teaching French in 17th-century England” Grammaire et enseignement du français 1500-1700, Jan de Clercq & Nino Lioce, eds. Leuven: Peeters, pp. 179-195.2001“Les patois et l’histoire de la langue française selon le dictionnaire de Littré”, L’Information grammaticale 90: 68-72.2001 “Language protection and cultural policy in France”, Endangered Languages and the Media.Proceedings of the fifth conference of the Foundation for Endangered Languages, Christopher Moseley, Nicholas Ostler & Hassan Ouzzate, eds., pp. 29-34. Bath, England: The Foundation for Endangered Languages. (co-authored with Samira Elatia)2001 “Les géostratégies des langues et la théorie linguistique”. Terminogramme 99-100 (automne 2001): 69-80.2002 “Les fondamentaux, ou Revisiter les classiques: The French Language in England de Kathleen Lambley (1920)”. La lettre de la SIHFLES 47.3-5.2003 “Language policy and linguistic theory” in Languages in a Globalising World edited by Jacques Maurais, and Michael A. Morris. Cambridge: Cambridge University Press, 47-57.2003 “Growing pains: Territorial expansion and language policy in France from Villers-Cotterêts to the Revolution” Interpreting the history of French. A Festschrift for Peter Rickard on the occasion of his eightieth birthday, Wendy Ayres-Bennett & Rodney Sampson, eds., Amsterdam/ New York: Rodopi Press, pp. 313-328.2003 “Les textes autoritaires, sources des normes linguistiques” Histoire Épistémologie Langage 24.5-27. 2003 “When Children’s Literature Transcends its Genre: Translating Alice in Wonderland” Meta 48.307-321. 2003 “New, and fine, wine in an old bottle: The Dictionnaire Étymologique de l’Ancien Français.” (Review Article) Romance Philology 55.103-110.2003 ““Le développement d’une pédagogie du français langue étrangère: les pronoms relatifs en qu-” Langue française.139.59-72.2003 “La Romania submersa dans les îles britanniques: après 1066” (Romanische Sprachgeschichte. Histoire linguistique de la Romania, Gerhard Ernst, Martin-Dietrich Gleßgen, Christian Schmitt, WolfgangSchweickard, eds., Tubingen: Niemeyer, pp. 717-726)2003 “Louis Meigret, le parler lyonnais et les politiques de la langue française à la Renaissance” In Lyon et la défense et illustration de la langue française, Gérard Dufaux, ed., Lyon: ENS Éditions, pp. 63-75.2004 “The boundaries of nature in the discipline of linguistics” Language Sciences 26.301-311 Forthcoming:(a) “L'Histoire de la langue et de la littérature française de Petit de Julleville: L'histoire de quelle langue?”,Volume to celebrate the centenary of the publication of Petit de Julleville’s Histoire de la langue et de la littérature française, Yannick Portebois & Jacques-Philippe Saint-Gérand, eds. Louvain: Peeters. (b) “Les images de l'autorité en matière de langue en France (1453-1647)” in Images of Authority and theAuthority of Images, K. Fresco, ed. (Co-authored with Philippe Caron.)(c) “Walter de Bibbesworth”, In Lexicon grammaticorum, Harro Stammerjohann, ed.(d) “Jacques Dubois”, “Louis Meigret”, “John Palsgrave”, “Linguistic Traditions in the Renaissance: French” –articles to appear in the Elsevier Encyclopedia of Language and Linguistics.(e) “Endangered Languages/ Language Endangered” in Global Futures, Charles Stewart, ed. Urbana:University of Illinois Press.Internet CreationsCentre Culturel VirtuelDictionnaire de l’Académie Française 1798 (part of international consortium to post full-text of all editions of the Dictionnaire de l’Académie Française): /ARTFL/projects/academie/ Kolb-Proust Archive (leader of team to post hyperlinked guide to the correspondence of Marcel Proust and to his era): /kolbp/Exercices pour accompagner Internef (with Samira Elatia): /internef/index.html ReviewsPerspectives psychomécaniques sur la syntaxe by Roch Valin. (Modern Language Journal 67 (1983): 85) Syntaxe comparée du français et de l'anglais by Jacqueline Guillemin-Flescher. (French Review 56 (1983): 808-9)Geschichte des französischen Kernwortschatzes by Arnulf Stefenelli. (French Review 57 (1984):922)The Oxford-Duden French-English Pictorial Dictionary (French Review 58 (1985):588-9)Dictionnaire inverse de l'ancien français by Douglas Walker, and Dictionnaire inverse de l'ancien français by Ralph de Gorog. (French Review 58 (1985): 769-70)Dictionnaire explicatif et combinatoire du francais contemporain. Recherches lexico-sémantiques by Igor Mel'cuk et al. (Modern Language Journal 69 (1985): 175)An Introduction to Old French by William Kibler. (French Review 59 (1986): 1015)Maurice Grevisse und die französische Grammatik. Zur Geschichte eines Phänomens by Marie Lieber (French Review 61 (187) 338-339)César-Pierre Richelet (1626-1698): Biographie et oeuvre lexicographique by Laurent Bray. (Dictionairies 9 (1987): 220-222)La linguistique, la grammaire et l'école, 1750-1914 by Jean-Claude Chevalier and Simone Delesalle. (French Review 62 (1988): 381-382)Acquisition linguistique et interférences by Jean Petit. (Modern Language Journal 72 (1988): 335-6)Grammaire vivante du français by Monique Callamand and I mparfait et compagnie by Jacques Montredon.(Modern Language Journal 72 (1988): 471-2)La langue volée. Histoire de la langue française by Jean-Jacques Thomas (The French Review 65 (1991): 355-356)Studies in Romance Linguistics edited by Carl Kirschner and Janet DeCesaris (The Modern Language Journal75 (1991): 533-534)Les Inventeurs du bon usage (1529-1647) by Danielle Trudeau (Historiographia Linguistica 19 (1992): 409-414)Historie de la langue: Du latin à l'ancien français by Peter Machonis. (Modern Language Journal 76(1992):416)Langue et identité. Le français et les francophones d'Amérique du Nord edited by Noël Corbett (Québec Studies 14 (1992)137-138)Linguistic Imperialism by Robert Phillipson (World Englishes 12 (1993): 342-347)The French Language Today by Adrian Battye and Marie-Anne Hintze. (Modern Language Journal 77(1993):380-381)Vers la réconciliation? La question linguistique au Canada dans les années1990 ed. by Daniel Bonin.(Québec Studies 15(1993):160-161)Authority and the French Language ed. by Rodney Sampson (Historiographia Linguistica 21 (1994):221-226) Hermeneutics and Translation in the Middle Ages: Academic Traditions and Vernacular Texts by Rita Copeland (Target An International Journal of Translation Studies 6 (1994):109-112)Historical Linguistics by Winfred Lehmann. (Modern Languages Journal 78(1994): 134-135)The French Language in the Seventeenth Century. Contemporary Opinion in France by Peter Rickard (The French Review 67 (1993): 370)La construction linguistique de l'Europe by Normand Labrie and Le plurilinguisme européen ed. by Claude Truchot (The French Review 69 (1996): 510-511)History of the French Language through Texts by Wendy Ayres-Bennett (The Modern Language Journal 82 (1998): 131-132)Language is Power. The Story of Standard English and its Enemies by John Honey (Journal of Linguistics 34 (1998): 525-530)John Palsgrave as Renaissance Linguist by Gabriele Stein (Dictionaries 20 (1999): 169-172)Histoire de la didactique des langues by Jean Caravolas (Forthcoming in the Bulletin of the Henry Sweet Society)Introduction à l’histoire de la langue française by Michèle Perret (forthcoming in the Modern Language Journal)Linguistics and the Third Reich. Mother-tongue fascism, race and the science of language by Chrstopher Hutton (World Englishes19 (2000): 245-249)État de la Francophonie dans le monde. Données 1995-1996 et 5 études inédites.and État de la Francophonie dans le monde. Données 1997-1998 et 6 études inédites, both by the Haut Conseil de la Francophonie (The French Review74 (2001): 611-613)Le langage et ses disciplines, XIXe – XXe siècles by Jean-Louis Chiss and Christian Puech (Historiographia Linguistica 28:414-417)L’expansion du français dans les Suds (XVe-XXe siècles) by C. Dubois, J-M. Kasbarian & A. Queffélec (The French Review 75 (2001): 425-426)Changements politiques et statut des langues. Histoire et epistémologie 1780-1945 ed. by Marie- Christine Kok Escalle & Francine Melka (The French Review 76 (2002): 422-423)French in and out of France. Language policies, intercultural antagonisms and dialogue., Kamal Salhi, Ed.(The French Review 78 (2004): 416-417).Contacts de langue. Modèles, typologies, interventions, Jacqueline Billiez, Ed. (The French Review 78 (2005): 1256-1257).CONFERENCE PRESENTATIONS1980 "Renaissance Vernacular Grammar and the Development of the Port-Royal Model." SAMLA-SECOL, Atlanta, GA.1981 "Vernacular Language Instruction in the Middle Ages." Southeastern Medieval Association, Morgantown, WV.1981 "Icil/icist: The i - forms of the Old French Demonstratives." SAMLA-SECOL, Louisville, KY.1982 "Je me souviens: An Historical Perspective on French-Canadian Society." University of Louisville Seminar on French Civilization.1984 "From Glossaries to Dictionaries: The Development of Bilingual Lexicography in the Later Middle Ages." 20th Annual Conference of Medieval Studies, Kalamazoo, MI.1984 "Bilingual Lexicography in the Renaissance: John Palsgrave's English-French Lexicon." 3rd International Conference on the History of the Language Sciences, Princeton, NJ.1985 "Progress in Renaissance Bilingual Lexicography." Dictionary Society of North America, Ann Arbor, MI.1986 "The Humanist Period in Renaissance Bilingual Lexicography." LEXeter Conference on the History of Lexicography, Exeter, England.1986 "Bilingual Lexicography in the Middle Ages." Le pluralisme linguistique dans la société médiévale.Montréal, P.Q.1987 "Plus ça change... 600 Years of French Dialogues", American Association of Teachers of French, San Francisco.1987 "Phonetic Transcription and Phonological Description in the Sixteenth Century." Kentucky Foreign Language Conference, Lexington, KY.1987 "Cross-Channel Influences in Phonetic Description in the Sixteenth Century." Fourth International Conference on the History of the Language Sciences. Trier, Germany.1987 "Source négligées dans l'histoire du vocabulaire." Seventh International Conference on Historical Linguistics, Lille, France.1988 "Ancients, Moderns and Lexicographers: Literary Sources for Renaissance Dictionaries." Central Renaissance Conference, Terre Haute, IN.1988 "Alice's Adventures in French: What the Grammarian Found There." American Association of Teachers of French, Boston, MA.1989 "Language Variation and Linguistic Description in 16th-Century France". Georgetown University Roundtable on Language and Linguistics.1989 Organizer of a session on comparative syntax (French-English) at the meeting of the American Association of Teachers of French, (Paris, July 10-13) with Jacqueline Guillemin-Flescher (Université de Paris VII), Anne-Marie Léonard (Université Paul Valéry, Montpellier), and Hélène Chuquet and Michel Paillard (Université de Poitiers).1989 Organizer of two sessions on the History of Linguistics at the joint meeting of the Linguistic Society of America and the North American Association for the History of Language Sciences.1990 "Reason, Usage, and the Development of Historical Linguistics in the 16th Century" Fifth International Conference on the History of the Language Sciences (Galway, Ireland) September 1-6, 1990.1990 "Multiple Translations, Comparative Syntax, and the Teaching of French" Annual Meeting ot the British Association of the French Language Studies, Liverpool, England September 10-12, 1990.1990 "Anglo-Norman Literature and its Public(s)" Modern Language Association Chicago, IL December 27-30, 1990.1991 Organizer of three sessions on the History of Linguistics at the joint meeting of the Linguistic Society of America and the North American Association for the History of the Language Sciences Chicago ILJanuary 4-6, 1991.1991 "Renaissance Perceptions of Medieval Language and the Origins of Historical Linguistics". For presentation at the annual meeting of the Renaissance Society of America, Duke University, April 1991. 1991 "Durkheim in Translation/Durkheim and Translation" (with Robert Jones); invited for presentation at a conference entitled: "Humanistic Dilemma: Translation in the Humanities and Social Sciences", SUNY-Binghamton, Sept. 26-28, 1991.1992 "Durkheim, Language and Linguistics", Linguistic Society of America, Philadelphia.1992 "Intention, interpretation, intelligibility: computerized research in canonical texts". Association for Literary and Linguistic Computing/Association for Computing in the Humanities, Oxford, England,April 1992 (with Robert A. Jones).1993 "Dictionaries and Usage in 17th-Century France: Le Naïf François" International Conference on the History of the Language Sciences, Washington, D.C., August 1993.1993 "The History of Disciplinary Vocabulary: A Computer-Based Approach to Concepts of 'Usage' in 17th-Century Works on Language" Association for Literary and Linguistic Computing/Association forComputing in the Humanities, Washington D.C. June, 1993.1994 "Male Language and Female Language in Translations" Association for Literary and Linguistic Computing/Association for Computing in the Humanities, Paris, France, April 1994.1995 “French Culture on the Internet”, American Association of Teachers of French, San Antonio, July 1995.1996 “Legal and Linguistic Perspectives on Language Legislation”, Language Legislation and Linguistic Rights, Urbana IL, March 19961996 “Une comparaison des lois relatives à l’emploi de la langue en France et aux États-Unis”, American Association of Teachers of French, Lyon France, July 1996.1996 “Perspectives pour l’exploitation des nouvelles technologies de l’information dans l’enseignement des langues”, American Association of Teachers of French, Lyon France, July 1996.1997 “Linguistics and Official Language Declarations”, Law and Society Association, May 1997.1997 “The Dictionary in the Service of the State: The French Dictionnaire des Termes Officiels and its Virtual Successors”, Dictionary Society of North America, May 1997.1997 “Language and dialect in the history of linguistics: A case study of the politics of Anglo-Norman”, annual meeting of the Henry Sweet Society for the History of Linguistic Ideas, Luton, England,September 19971997 “Linguistic Human Rights and the Nature of Language”, Linguistic Human Rights Conference and Workshop for NGO-Representatives, Budapest, Hungary October 1997.1997 “Teaching Language Teachers to Use the Internet”, 8th annual conference on Educational Technology and Language Learning, Newcastle England, November 19971998 “Le Dictionnaire de l’Académie Française dans son contexte: la création d’une base de données axée sur la cinquième édition du DAF”. International conference on dictionary databases, Limoges, France,November 19-21 1998.1999 “’Le peuple’, ‘les peuples’, ‘un peuple’ dans la pensée linguistique en France . 1800-1850”. Eighth International Conference on the History of the Language Sciences, Paris, France.1999“Le Dictionnaire de Trévoux et la politique de la langue française”, Journées Internationales d'Études Lexicographiques sur le Dictionnaire de Trévoux 1704 – 1771, Trévoux October 15-16.2000 “The equivalence of language and species, from organic linguistics to the ecology of language”, LSA/NAAHoLS, Chicago, Jnnuary.2000 “’I know what ‘it’ means when I say it: Étude sur la traduction d’Alice au pays des merveilles”, American Association of Teachers of French, Paris, France, July.2000“Patois in the histories of the French language in the 19th century”, Nineteenth Century French Studies Conference, Urbana-Champaign, October.2001“Language Protection and Cultural Policy in France” meeting of the Foundation for Endangered Languages in Agadir, Morocco. (Joint presentation with Samira El Atia)2002 “Les avant-coureurs de la norme 1453-1647”, UIUC-CNRS Conference: “The Authority of Images, the Images of Authority”. Joint presentation with Philippe Caron. April 29 - May 1.2002 “Language and the Politics of Decentralization in France”, International Academy of Language Law, Iasi Romania May 23-25.2002 “Le langage vicieux, l’enseignement et la théorie grammaticale: Le Dictionnaire critique et raisonné du langage vicieux ou réputé vicieux (1835)”. 9th International Conference on the History of theLanguage Sciences, São Paolo, Brazil, August 26-29.2004 “Prescriptive traditions in linguistics”, Linguistic Society of America/ North American Association for the History of the Language Sciences, January 9.2004 “Free trade, fair trade and the linguistic marketplace: economic metaphors and language legislation”, Ninth International Conference on Language andLaw, Beijing, China, Sept. 15.2005 “Prescriptive Behavior in Language and in Linguistics”, Tenth International Conference on the History of the Language Sciences, Urbana IL, Sept 1.INVITED LECTURES AND CONFERENCE PLENARY PRESENTATIONS1991 "16th-Century Bilingual Dictionaries (French-English): Organization and Access, Then and Now".University of Toronto Conference on Early Dictionary Databases.1992 "Durkheim and the Language of Secular Ethics", Program for the Study of Cultural Values and Ethics (with Robert A. Jones).1992 "The Politics of Language in France" Center for Advanced Study, University of Illinois.1993 "John Palsgrave: External Perspectives on the French Language in England in the Renaissance", Indiana University, Bloomington.1993 "Baret's Alvearies: Challenges for the Scholar" University of Toronto Conference on Historical Dictionary Databases.。
ORIGINAL PAPERMajor and minor QTL and epistasis contribute to fatty acid compositions and oil concentration in high-oil maizeXiaohong Yang •Yuqiu Guo •Jianbing Yan •Jun Zhang •Tongming Song •Torbert Rocheford •Jian-Sheng LiReceived:31December 2008/Accepted:30September 2009/Published online:25October 2009ÓSpringer-Verlag 2009Abstract High-oil maize is a useful genetic resource for genomic investigation in plants.To determine the genetic basis of oil concentration and composition in maize grain,a recombinant inbred population derived from a cross between normal line B73and high-oil line By804was phenotyped using gas chromatography,and genotyped with 228molecular markers.A total of 42individual QTL,associated with fatty acid compositions and oil concentra-tion,were detected in 21genomic regions.Five major QTL were identified for measured traits,one each of which explained 42.0%of phenotypic variance for palmitic acid,15.0%for stearic acid,27.7%for oleic acid,48.3%for linoleic acid,and 15.7%for oil concentration in the RIL population.Thirty-six loci were involved in 24molecular marker pairs of epistatic interactions across all traits,which explained phenotypic variances ranging from 0.4to 6.1%.Seven of 18mapping candidate genes related to lipidmetabolism were localized within or were close to identi-fied individual QTL,explaining 0.7–13.2%of the popula-tion variance.These results demonstrated that a few major QTL with large additive effects could play an important role in attending fatty acid compositions and increasing oil concentration in used germplasm.A larger number of minor QTL and a certain number of epistatic QTL,both with additive effects,also contributed to fatty acid com-positions and oil concentration.IntroductionHigh-oil maize developed by artificial selection is a value-added crop since its kernels have advantages as sources of animal feed for higher caloric energy and better protein quality (Han et al.1987;Benitez et al.1999;Lambert et al.2004).Additionally,maize oil is considered as high quality oil for human health due to the high proportion of poly-unsaturated fatty acids (Lambert 2001).The long-term selection experiment for maize oil and protein was initiated with the open-pollinated variety Burr’s White in 1896(Dudley and Lambert 1992,2004).The oil concentration of Illinois High Oil (IHO)reached about 20%after 100generations of selection.In the early 1980s,Song et al.began to develop Chinese synthetic populations with increased oil concentration by artificial selection (Song et al.1999;Lambert et al.2004).Beijing High Oil (BHO)originated from a normal maize synthetic known as Zhongzong No.2,which was synthesized with 12inbred lines of Lancast heterotic group.Its oil concentration had increased from 4.71to 15.55%after 18selection cycles (Song and Chen 2004).These materials are unique resources providing an opportunity to examine the genetic basis of high oil concentration in maize.Communicated by C.Scho¨n.Electronic supplementary material The online version of thisarticle (doi:10.1007/s00122-009-1184-1)contains supplementary material,which is available to authorized users.X.Yang ÁY.Guo ÁJ.Yan ÁJ.Zhang ÁT.Song ÁJ.-S.Li (&)Beijing Key Laboratory of Crop Genetic Improvement,National Maize Improvement Center of China,China Agricultural University,Yuanmingyuan West Road,Haidian,100193Beijing,Chinae-mail:lijiansheng@;lijs@J.YanInternational Maize and Wheat Improvement Center(CIMMYT),Apdo.Postal 6-641,06600Mexico,D.F.,Mexico T.RochefordDepartment of Agronomy,Purdue University,West Lafayette,IN 47907,USATheor Appl Genet (2010)120:665–678DOI 10.1007/s00122-009-1184-1A number of chromosomal regions and QTL controlling oil concentration were identified using segregating mapping populations through molecular markers in maize(Goldman et al.1994;Berke and Rocheford1995;Mangolin et al.2004; Song et al.2004;Wassom et al.2008a;Zhang et al.2008). Recently,Laurie et al.(2004)estimated the number of QTL for oil concentration to be about50in a larger randomly mated population,IHO(70)9ILO(70),with SNP markers, which was consistent with the estimates obtained using traditional quantitative genetic methods(Dudley1977). Thus,a number of studies suggested that oil concentration was controlled by a large number of genes with individually small effects and mainly additive gene action.Chemically,maize oil is a mixture,mainly(99%) comprisingfive fatty acids,viz.,palmitic(16:0),stearic (18:0),oleic(18:1),linoleic(18:2)and linolenic(18:3) acids(Lambert2001).Sixty-eight loci associated with the five major fatty acids were detected by single-factor ANOVA using80RFLP markers in an IHO(90)9ILO (19)population of200F2S1lines(Alrefai et al.1995). Wassom et al.(2008b)identified15QTL controlling these fatty acids using the same markers in the same populations described in Wassom et al.(2008a).Epistasis,the interaction between alleles from two to more genetic loci(Fisher1918),may play an important role in evolutionary and quantitative variation in crops (Doebley et al.1995;Yu et al.1997;Lukens and Doebley 1999;Gadau et al.2002;Malmberg et al.2005;Mei et al. 2005;Xu and Jia2007).QTL mapping is one experimental approach to explore the role of epistasis in the genetic basis of complex traits(Carlborg and Haley2004).However, there are practical problems in efficiently measuring epi-static genetic variances.One common method is to use multiple interval mapping(MIM;Kao et al.1999)in the Windows QTL Cartographer software(Wang et al.2005), but this program only detects epistatic interactions among identified QTL with main effects.Recently,several com-puter programs were developed that enable genome-wide scans for epistatic effects(Chase et al.1997;Holland1998; Manly et al.2001;Sen and Churchill2001;Borevitz et al. 2002;Yang et al.2007).Among these,the mixed linear model presented in QTLNetwork Version2.0,can integrate multiple QTL,epistasis,QTL-by-environment interactions, and epistasis-by-environment interactions into one map-ping system(Yang et al.2007).Using this software,epis-tasis contributing to genetic variation in oil concentration was identified in oilseed crops(Zhao et al.2005,2006).In maize,a certain number of epistatic interactions affecting oil concentration and fatty acid compositions were identi-fied in segregating populations(Dudley2008;Wassom et al.2008a,2008b).However,similar studies have not been undertaken in recombinant inbred line(RIL) populations.Biosynthesis of storage oil in plant seeds is a complex metabolic process(Ohlrogge and Browse1995).Most of the biochemical steps are well known and many of the genes involved in these reactions have been characterized in Arabidopsis(Beisson et al.2003).However,much less is known in maize,and only a few genes related to lipid metabolism were cloned(Lee and Huang1994;Egli et al. 1995;Berberich et al.1998;Schreiber et al.2000).Some loci for the key enzymes in lipid metabolism associated with QTL for fatty acid compositions were identified in Arabidopsis,sunflower and mustard(Lionneton et al.2002; Pe´rez-Vich et al.2002;Hobbs et al.2004).In maize,the map positions of two fatty acid desaturases,fad2and fad6, were compared with the positions of QTL for the ratio of oleic:linoleic acids to determine if they map to the same genomic region(Mikkilineni and Rocheford2003).The results suggested that comparison of the positions of can-didate genes and QTL was a suitable strategy to investigate the molecular basis of quantitative traits.Additionally,the positioned candidate genes can be used to develop func-tional markers for increasing selection efficiency by mar-ker-assisted selection in plant breeding(Jeppe and Thomas 2003).In the present study,a RIL population derived from a single cross with a high-oil line was used to:(1)map QTL for fatty acid compositions and oil concentration;(2) estimate the number and magnitude effects of QTL and epistatic interactions underlying fatty acid compositions and oil concentration;and(3)analyze the relationships between candidate genes involved in lipid metabolism and mapped QTL for fatty acid compositions and oil concentration.Materials and methodsGenetic materialsA RIL population was developed from a cross between B73(normal inbred)and By804(high-oil inbred),which was selected from a high-oil population,BHO Cycle13. Two hundred and forty-five F7:8RILs along with both parents were planted in a randomized complete block design with three replications on the Agronomy Farm, China Agricultural University,Beijing,in2005and2006, respectively.Each family line was grown in a single5m row,rows were0.67m apart,and planting density was 45,000plants/ha.In order to avoid xenia effects,more than six plants in each row were pollinated by bulked pollen within the row and pollinated ears were harvested at maturity.Equal amounts of grain from each harvested ear in each row among RIL population were bulked for chemical analysis.Grain samples from twofieldreplications over2years were used for fatty acid compo-sitions and oil concentration analysis.Genetic linkage and candidate gene mappingA total of208markers,including202SSR markers and six IDP markers developed by our laboratory,were used to build a genetic framework map.SSR markers were selected from public maize database(). Genomic DNA was extracted from all245F7RILs and the parents using the modified procedure of Murry and Thompson(1980).A SSR procedure was followed:tem-plate DNA50ng,primers0.67l M,109reaction buffer 19,MgCl22.5mM,dNTPs0.2mM,Taq DNA polymerase 0.5U,made thefinal volume of15l L with deionized double distilled water.The PCR reaction conditions were set as:denaturation at94°C for5min,followed by30cycles of denaturation at94°C for1min,annealing at58°C for 1min,elongation at72°C for1.5min,withfinal extension at72°C for5min and then stored in refrigerator at4°C.In order to analyze the co-locations of QTL for fatty acid compositions and oil concentration with the genes of lipid metabolism,18candidate genes(supplementary Table1)were chosen to develop molecular markers for mapping according to the Arabidopsis lipid metabolism pathway(Beisson et al.2003;http://www.plantbiology. /lipids/genesurvey).Because the cDNA or DNA sequences in maize were unknown,known protein sequences of Arabidopsis were submitted to a tBLASTn (Altschul et al.1997)search for maize homologues of candidate genes in the NCBI database(http://www. /BLAST),and the highest matching hits were chosen for developing candidate gene markers.Gene-specific primers(supplementary Table1)for developing SNP and IDP markers were designed from gene or EST sequences,using the Primer5.0software(Clarke and Gorley 2001)to amplify and sequence parts of the genes.MAP-MAKER/EXP3.0(Lincoln et al.1993)was used to con-struct a genetic linkage map and to integrate the candidate genes into the map.The Kosambi mapping function was used for converting recombination values to map distances.To amplify the candidate gene fragments for sequenc-ing,DNA was amplified with a PTC-200Peltier Ther-malcycler(Bio-Rad,USA)in a50-l l reaction volume containing100ng genomic DNA,5l l of109PCR reac-tion buffer, 1.5mM MgCl2,0.2mM of each dNTP, 0.2l M of each gene-specific primer,and1.25U of Taq polymerase(Tiangen).Thermal cycling conditions were 95°C for5min;35cycles of95°C for45s,optimal annealing temperature for45s,72°C for1.5min;followed by afinal extension of10min at72°C.PCR products were analyzed on agarose gels,purified using a multifunctional DNA purification kit(Bioteke Corporation,China),and cloned into pGEM-T easy vector(Promega,USA).Three positive clones were picked and sequenced.The DNA-MAN software(Lynnon Biosoft,Canada)was used to search for nucleotide polymorphisms between sequences of the two parents.Subsequently,three types of genotyping, insertion–deletion polymorphisms(IDP,Bhattramakki et al.2002),cleaved amplified polymorphic sequences (CAPS;Konieczny and Ausubel1993)and allele-compet-itive PCR(AC-PCR)SNP(Falque et al.2005)were employed to develop markers for mapping candidate genes. Measurements of fatty acid compositionsand oil concentrationFatty acid compositions of223lines of the245RILs were measured because some RILs did not have enough grain or their kernels did not develop very well.Fifty kernels ran-domly chosen from the bulked grain of each plot were ground and a HP6890gas chromatogram(GC)(Agilent Technologies,USA)was employed for fatty acid analysis. All samples were measured with the two sub-samples at the same time.Lipids were extracted as described by Sukhija and Palmquist(1988)with some modifications:200–300mg of each sample was weighed and transferred to 15ml culture tubes.To each tube4ml methyl alco-hol:acetyl chloride(10:1)were added,followed by5ml of internal reference(methyl nonadecanoate,Sigma,USA), with1mg/ml in hexane,and after general vortexing, heated for2h in a water bath at80°C.After the contents were cooled to room temperature,5ml of7%K2CO3were added to neutralize extracted fatty acids,and the upper phase was transferred to sample bottles.Samples at250°C were automatically injected(1l l)and separated in the GC system equipped with a HP-INNOWAX polyethylene glycol capillary column(30m9320l m90.5l m, Agilent Technologies).The GC was operated at constant flow pressure of140.9kPa with an initial oven temperature of220°C,13min isothermal,then oven temperatures were increased by20°C/min to240°C,5min isothermal.The FID temperature was250°C and the split ratio of nitrogen was20:1.Fatty acids were identified by comparison of their retention times with that of the internal reference.All data were acquired on ChemStation software(Agilent Technologies)and normalized to sample weight and to the internal reference.The oil concentration was calculated as the sum of all identified fatty acid concentrations with percentage(%)of kernel weight.Individual fatty acids were expressed as percentage(%)of oil concentration. Phenotypic data analysisThe variance of fatty acid compositions and oil concen-tration were estimated using PROC GLM in SAS8.02software(SAS Institute1999).The model for variance analysis was Y=l?a g?b y?(ab)gy?c yr?e gyr, where a g was the effect of the g th line,b y was the effect of the y th year,(ab)gy was the line9year interaction,c yr was the year9replicate interaction,and e gyr was the residual. All effects were considered to be random.Broad-sense heritability(H2)was estimated as H2= r g2/(r g2?r gy2/r?r e2/yr)(Knapp et al.1985),where r g2is the genetic variance,r gy2is the interaction of genotype with year,r e2is the residual error,r is the number of replications, and y is the number of years.All the variances were acquired from variance analysis.To quantify the relation-ships among measured traits,correlation coefficients were calculated for each trait pair using PROC CORR in SAS 8.02software(SAS Institute1999).QTL and epistasis analysisAs described as Yang et al.2007,QTLNetwork version2.0 was employed for QTL and epistasis analysis.A mixed linear model was used to identify QTL at1-cM intervals with a window size of10cM.10-cM windows were defined to distinguish two adjacent test statistic peaks whether or not they represented two QTL.Two-dimen-sional(2D)genome scans were used to search for multiple interacting QTL.For each trait,a genome-wide threshold value of the F-statistic(a=0.01)for declaring the pres-ence of a QTL was estimated by10,000random permu-tations(Doerge and Churchill1996).A Bayesian method with Gibbs sampling was used to estimate QTL effects (Wang et al.1994).Some marker intervals coming directly from QTLNetwork version2.0were modified if the posi-tion of adjacent QTL for different traits was less than 10cM.The sum of individual phenotypic variance explained by each QTL was calculated as the total phe-notypic variance explained by all QTL for each trait. ResultsVariation in fatty acid compositionsand oil concentrationEleven fatty acid compositions were determined using GC in RIL,and their sum was tabulated as oil concentration. Data for four major fatty acids,viz.palmitic(16:0),stearic (18:0),oleic(18:1),and linoleic(18:2)acids,and total oil concentration were used for subsequent analysis because the other seven fatty acids were present in only trace analyzed amounts.The means of the RIL population combined over years was close to the mid-parent value for all measured traits(Table1).A normal distribution with a wide range of variation was observed for each trait(data not shown).Oil concentration was positively correlated with the level of18:1,but was negatively correlated with levels of16:0and18:2(Table2).Oleic acid showed a highly negative correlation with18:2(r=-0.96)and low or moderate correlations were detected for other fatty acid pairwise comparisons(-0.43to0.38).The GLM indicated highly significant effects due to genotype,year and geno-type9year interaction for each trait(Table1).For all traits,broad-sense heritability(H2)estimates were gener-ally high,ranging from86.8%for18:0–97.8%for18:2. The relatively high heritabilities indicated that much of the phenotypic variance in the RIL population was genetically controlled.Table1Means,range,variances and broad-sense heritability(H2)for fatty acid compositions and oil concentrationTraits a16:018:018:118:2OilB73Mean13.45±0.17 2.30±0.1125.11±0.8255.85±0.72 4.08±0.20 By804Mean13.42±0.20 2.38±0.2135.21±0.3646.82±0.5011.18±0.58 RIL Mean13.47±0.06 2.09±0.0130.56±0.2251.29±0.21 6.64±0.04 Range9.17–18.42 1.10–3.4917.45–46.4737.49–63.41 3.71–10.41F values year9replicate 4.39*20.64**0.89 4.190.09Year214.34**210.69**26.66** 1.39** 2.52**Genotype44.00**19.38**105.04**114.21**24.08**Genotype9year 2.00** 2.56** 2.44** 2.57** 1.80**Heritability(H2)b95.486.897.797.892.5Confidence interval c94.3–96.483.3–89.597.1–98.297.2–98.290.6–94.1*,**Significant at P\0.05and0.01,respectivelya16:0,Palmitic acid;18:0,stearic acid;18:1,oleic acid;18:2,linoleic acidb Broad-sense heritability(H2)of fatty acid compositions and oil concentrationc90%Confidence intervals of broad-sense heritabilitySingle-locus QTL for fatty acid compositions and oil concentrationBased on a linkage map of 1,675cM with an average interval of 7.3cM between adjacent markers,a total of 42QTL controlling fatty acid compositions and oil concen-tration were detected,corresponding to 21genomic regions because some QTL for different traits clustered in the same regions and shared the same molecular markers (Table 3,Supplementary Fig.1).These loci were distributed across all ten chromosomes.Individual loci for any given trait explained 0.7–48.3%of the phenotypic variance.Among the mapping population,all additive effects of mapped QTL for each trait accounted for a medium proportion of the total phenotypic variance,ranging from 53.4to 65.8%(Table 3).For oil concentration,nine QTL distributed across all chromosomes,except chromosome 3and chromosome 7,were detected.The QTL,oil1-1,with the largest effect (15.7%of the phenotypic variance)was located on chro-mosome 1,flanked by umc2217and bnlg2086.The By804allele at this locus had an additive effect of 0.48%for increased oil concentration.Another important QTL for oil concentration,oil6,was located in the same region as ste6-1,ole6-1and lin6-1,and explained 8.4%of phenotypic variance with an additive effect of 0.35%on theTable 2Pearson correlation coefficients for trait pairs affecting fatty acid compositions and oil concentration in the RIL population Traits 16:018:018:118:2Oil16:0118:0-0.33**118:1-0.43**0.38**118:20.16**-0.40**-0.96**1Oil-0.31**0.070.43**-0.33**1**Significant at P =0.001Table 3QTL for fatty acid compositions and oil concentration in RIL population Traits QTL a Chr Markerinterval P b A ch 2(a)d (%)16:0pal11umc2232-umc1988134.5-0.338 2.7pal22phi96100-kt214.0-0.1850.8pal44bnlg2162-umc2286110.1-0.354 3.0pal55umc2373-umc122184.3-0.282 1.9pal66nc010-umc110563.4-0.6199.1pal88umc1130-umc156265.9-0.398 3.8pal99bnlg1401-umc221326.1 1.33242.0pal1010umc1367-umc201640.2-0.288 2.0Subtotal e 65.318:0ste22phi083-u4113.60.051 1.3ste33phi053-sad7004109.1-0.16213.2ste55umc1491-umc12530.00.055 1.5ste6-16Q8-nc01054.30.17315.0ste6-26phi299852-umc2059152-0.1379.3ste88umc1075-umc130417.0-0.1369.2ste99bnlg1401-umc221327.7-0.088 3.9Subtotal e 53.418:1ole2-12phi96100-kt216.20.4590.7ole2-22bnlg108-u4114.80.633 1.4ole45m1-bnlg229181.4 1.6289.0ole6-16Q8-nc01041.5 2.85927.7ole6-26nc010-umc110567.4 1.394 6.6ole6-36umc1614-acda600189.8 1.4947.6ole77dupssr13-CA33103.4-0.763 2.0ole88umc1130-umc156265.90.904 2.8ole99umc1743-umc165449.5-0.873 2.6Subtotal e60.418:2lin1-11umc2217-bnlg208674.80.707 1.6lin1-21umc1725-phi064264.0-0.4610.7lin22bnlg108-u4114.8-0.564 1.0lin44m1-bnlg229181.4-1.115 4.0lin6-16Q8-nc01042.5-3.88148.3lin6-26umc1614-acda600189.8-1.400 6.3lin6-36phi299852-umc2059147.00.786 2.0lin77dupssr13-CA33104.40.592 1.1lin99bnlg1401-umc221326.7-0.5120.8Subtotale65.8Table 3continued TraitsQTL aChr MarkerintervalP b A ch 2(a)d (%)oiloil1-11umc2217-bnlg208685.00.48015.7oil1-21ols1-phi308707215.20.200 2.7oil22phi96100-kt29.00.309 6.5oil44dupssr28-bnlg2162104.80.146 1.5oil55umc2373-umc122190.30.275 5.3oil66Q8-nc01045.50.3518.4oil88umc1130-umc156265.90.279 5.3oil99bnlg1401-umc221322.10.233 3.7oil1010umc1367-umc201640.20.303 6.2Subtotal e55.3aNumber following the three letters represents the chromosome location of the QTL.Different lowercase numbers following the dash indicate putatively different QTL located on the same chromosome bThe QTL location on chromosome was estimated by QTLNetwork cAdditive effects estimated by QTLNetwork.Positive (?)indicates that the By804allele increases trait expression,and negative (-)indicates that the B73allele increases trait expression,respectively dPercentage of phenotypic variance explained by individual additive effects of the mapped QTL eTotal percentage of phenotypic variance explained by all additive effects of the mapped QTL for each traitchromosome6.The other minor QTL for oil concentration each explained 1.5–6.5%of the phenotypic variance. Alleles from By804,a high-oil parent,at all of mapped loci had increasing effects for oil concentration.Eight QTL were associated with16:0and explained 65.3%of the total phenotypic variance.One major QTL for 16:0,pal9,terminally located on chromosome9S,and bordered by markers bnlg1401and umc2213,contributed 42.0%of the explained phenotypic variance.The additive effect of the By804alleles increased16:0by 1.332% absolutely.The remaining seven QTL were located on chromosomes1,2,4–6,8and10with explained variances ranging from0.8to9.1%;B73alleles increased16:0at these loci.Seven QTL were involved in18:0and accounted for 53.4%of the total phenotypic variance.Two QTL with large effects,ste6-1,flanked by the markers Q8and nc010 on chromosome6,and ste3,flanked by the marker phi053 and sad7004on chromosome3,accounted for15.0and 13.2%,respectively,of the phenotypic variance.The other QTL explained only1.3–9.3%of the phenotypic variance. By804alleles contributed to increased18:0at three loci, ste2,ste5and ste6-1,and B73alleles contributed increases at the other loci.Nine QTL were significantly associated with18:1.The strongest QTL,ole6-1,located in the same position as st6-1,explained27.7%of the phenotypic variance,and By804alleles had an additive effect of 2.85%for increasing18:1.The phenotypic variance explained by the other eight QTL ranged from0.7to9.0%.Alleles of these eight QTL with increasing effects came from By804with the exception of ole7and ole9.Collectively,the nine QTL explained60.4%of the total phenotypic variance.A total of nine QTL controlling18:2explained65.8% of the total phenotypic variance.One major QTL,lin6-1, with a very large effect localized to the same region as ste6-1,accounted for48.3%of the phenotypic variance. B73alleles at this locus had an additive effect of3.88% for increased18:2.Another eight QTL with small effects explained0.7–6.3%of the phenotypic variance. Among these eight loci,alleles with increasing effects at three loci were contributed by By804andfive came from B73.Epistatic QTL for fatty acid compositionsand oil concentrationTwenty-four pairs of epistatic QTL involving36loci were identified for all measured traits(Table4).The number of epistatic QTL pairs for each trait ranged from two to seven. The proportion of total phenotypic variance explained by all epistatic QTL ranged from5.2to12.6%for each trait. Based on genetic effects,mapped epistatic QTL comprised three types:interactions between two QTL with additive effects(AA),interactions between a QTL with additive effect and a locus without significant additive effect(AN or NA),and interactions between two loci with only epistatic effects(NN).There were both positive and negative epi-static interactions,indicating the combination of alleles at interacting loci with increasing contributions from the same and different parents,respectively.For16:0,six pairs of epistatic QTL were detected, explaining0.4–6.1%of the phenotypic variance in the RIL population.Of these,increased16:0from three pairs of loci came from the parental digenic combination,and three came from combinations of alleles from different parents.These QTL were partitioned into four NA inter-actions and two NN interactions.Five pairs of epistatic QTL(two AN,and three NN interactions)for18:0were detected,accounting for 1.5–2.0%of the phenotypic variance.All those with increasing effects came from the parental digenic combination,except for an interaction between one locusflanked by umc2408-bnlg197on chromosome3and anotherflanked by umc1241-umc1695 on chromosome7.For18:1,four pairs of epistatic QTL with explained variance from0.5to2.5%were partitioned into one AA interaction,two NA interactions and one NN interaction.One of four pairs with increasing effect came from combinations of alleles from different parents, whereas the remaining ones were from parental digenic combinations.Seven pairs of epistatic QTL for18:2(two AN or NA)interactions andfive NN interactions) explained0.5to1.6%of the phenotypic variance.The epistatic effects of these interactions came from different parents except one interaction between a locusflanked by umc1725-phi064on chromosome1and anotherflanked by umc1367-bnlg1655on chromosome10.Two pairs of epistatic QTL were detected for oil concentration.One accounted for1.3%of the phenotypic variance with an increased epistatic effect from different parents,and another explained3.8%of phenotypic variance with an increased epistatic effect coming from the parental digenic combination.Co-location of mapped candidate geneswith single-loci QTLA total of18candidate genes related to lipid metabolism, having either InDel or SNP polymorphisms between the parents,were mapped using20PCR-based markers cor-responding to the candidate genes(Table5;Fig.1). Among these markers,11were IDP,seven were CAPS and two were AC-PCR SNP.Four markers corresponding to two putative functional genes were located at different loci due to multiple copies.They were ketoacyl-ACP synthase II (KAS II)on chromosome bins 2.07–2.09and7.04,andStearoyl-ACP desaturase(SAD)on chromosome bins 3.05–3.06and8.05–8.06.Comparing map positions of candidate genes and detected QTL for fatty acid compositions and oil concen-tration(Fig.1),seven candidate genes were located in regions containing18mapped QTL with explained phe-notypic variance ranging from0.7%(ole2-1)to13.2% (ste3)(Table6).They were Acetyl-CoA carboxylase(AC-Case),KAS II,Ketoacyl-ACP synthase III(KAS III)and SAD for fatty acid synthesis,and Monoacylglycerol acyltrans-ferase(MAGAT),Acyl-CoA:diacylglycerol acyltransferase (DGAT)and Oil-body Oleosin17KD(OLE17)for oil synthesis and storage.ACCase(u4)fell within QTL,ste2, ole2-2and l in2,explaining1.3%of the phenotypic vari-ance for18:0,1.4%for18:1and1.0%for18:2,respec-tively.KAS II(kass5001B),located close to QTL ole7and lin7,contributed2.0%of the explained genetic variance for 18:1and1.1%for18:2.KAS III(kt2)fell within a cluster of QTL for16:0,18:1and oil concentration with explained variances ranging from0.7to6.5%.SAD(sad7004)co-located with the strong QTL,st3,accounting for up to 13.2%of the phenotypic variance for18:0.MAGAT(m1)Table4Epistatic QTL for fatty acid compositions and oil concentration in RIL populationTrait Chr i Marker interval i P i a Chr j Marker interval j P j a Interaction b AA c h2(aa)d(%)16:02phi083-phi092121.18phi119-bnlg208640.2N*N0.1260.4 2C9_3-mmc027138.28umc1562-umc114180.7NA-0.1600.62mmc027-kass2172.38umc1130-umc156265.9NA0.292 2.02mmc027-kass2172.38phi121-bnlg204660.9NA-0.507 6.16umc1614-acda6001100.37phi034-umc140944.9N*A-0.303 2.26phi299852-umc2059146.07phi091-atf264.5N*N0.236 1.3 Subtotal e12.618:06Q8-nc01054.39bnlg1401-umc221327.7AA0.054 1.5 1bnlg400-umc1955169.19Q2-umc1170 4.6NN0.057 1.62phi083-u4113.63bnlg1144-umc101234.0AN0.057 1.63phi036-bnlg163861.98umc2082-bnlg186354.2NN0.056 1.63umc2408-bnlg197127.37umc1241-umc1695 1.0NN-0.064 2.0 Subtotal e8.318:11umc2217-bnlg208680.03bnlg1144-umc101245.0N*N0.750 1.9 1umc1988-umc1122152.46umc1614-acda600189.8N*A0.862 2.51bnlg400-umc1955162.26nc010-umc110567.4NA-0.722 1.82phi96100-kt216.29umc1743-umc165449.5AA0.3820.5 Subtotal e 6.718:21umc2232-umc1988138.55umc2143-aca184.0N*N-0.704 1.6 1bnlg400-umc1955164.39umc1794-bnlg152572.7NN-0.3980.51umc1725-phi064264.010umc1367-bnlg165540.9AN*0.638 1.32apat5-phi101049216.39bnlg1401-umc221326.7NA-0.5250.93phi053-sad7004103.15bnlg1237-f22a129.7N*N-0.543 1.06phi126-umc175310.910umc1337-phi05023.9NN*-0.3940.58bnlg2082-bnlg186353.89umc1078-umc147166.5NN-0.77 1.9 Subtotal e7.7oil1umc2217-bnlg208685.01ols1-ph308707215.2AA-0.14 1.3 6Q8-nc01045.59bnlg1525-bnlg1904118.2AN0.238 3.9 Subtotal e 5.2*Without significant additive QTL for this trait but with significant additive QTL for other trait in this studya QTL location on chromosome was estimated by QTLNetworkb Types of epistatic interaction.AA interactions between two QTL with additive effects,NA(AN)interactions between a QTL with additive effects and a locus without significant additive effects,NN interaction between two loci with epistatic effects onlyc AA means the epistatic effects estimated by QTLNetwork.Positive(?)means that parental digenic genotypes increase trait expression,and negative(-)means that recombinant alleles from two parents increase trait expression,respectivelyd Percentage of phenotypic variance explained by individual epistatic effects of the mapped QTLe Total percentage of phenotypic variance explained by all epistatic effects of the mapped QTL for each trait。