吴爽;刘莉;王守海;王德正;孙传清;

• 1:安徽农业科学院水稻研究所
• 2:中国农业大学作物遗传育种系
• 3:中国种子集团公司

摘要(Abstract)：

文中主要从性状定位和基因克隆方面对产量构成三因子,即分蘖、穗粒数和粒重的研究成果进行了综述,并同时详细阐述了影响产量的其他因素,包括株高和杂种优势等的研究进展,并在最后就影响单株产量的分子基础、激素与产量的关系、水稻产量性状的定位策略3个方面对这些研究成果进行了总结和讨论。

关键词(KeyWords)： 水稻;产量性状;定位;克隆

Abstract:

Keywords:

基金项目(Foundation): 国家“863”计划(2010AA101304);; 农业部转基因专项(2011ZX08001-001);; 安徽省农业科学院院长青年创新基金(11B10105)

作者(Author): 吴爽;刘莉;王守海;王德正;孙传清;

Email:

DOI: 10.13989/j.cnki.0517-6611.2012.14.115

参考文献(References)：

• [1]KHUSH G.Productivity improvements in rice[J].Nutr Rev,2003,61(6):114-116.
• [2]程式华,胡培松.中国水稻科技发展战略[J].中国水稻科学,2008,22(3):223-226.
• [3]XIAO J H,GRANDILLO S,AHN S N,et al.Genes from wild rice improveyield[J].Nature,1996,384:223-224.
• [4]李德军,孙传清,付永彩,等.利用AB-QTL法定位江西东乡野生稻中的高产基因[J].科学通报,2002,47(11):854-858.
• [5]HE G M,LUO X J,TIAN F,et al.Haplotype variation in structure and ex-pression of a gene cluster associated with a quantitative trait locus for im-proved yield in rice[J].Genome Res,2006,16:618-626.
• [6]XING Y,ZHANG Q.Genetic and molecular bases of rice yield[J].AnnuRev Plant Biol,2010,61:421-442.
• [7]巩鹏涛,李迪.植物分枝发育的遗传控制[J].分子植物育种,2005,3(2):151-162.
• [8]LI X,QIAN Q,FU Z,et al.Control of tillering in rice[J].Nature,2003,422:618-621.
• [9]TAKEDA T,SUWA Y,SUZUKI M,et al.The OsTB1 gene negatively regu-lates lateral branching in rice[J].Plant J,2003,33:513-520.
• [10]ZOU J,CHEN Z,ZHANG S,et al.Characterizations and fine mapping of amutant gene for high tillering and dwarf in rice(Oryza sativa L.)[J].Planta,2005,222:604-612.
• [11]ZOU J,ZHANG S,ZHANG W,et al.The rice HIGH-TILLERING DWARF1encoding an ortholog of Arabidopsis MAX3 is required for negative regu-lation of the outgrowth of axillary buds[J].Plant J,2006,48:687-696.
• [12]江海湃,张淑英,包劲松,等.水稻多分蘖矮秆突变体htd1-2的遗传分析与基因定位[J].遗传,2009,31(5):531-539.
• [13]ISHIKAWA S,MAEKAWA M,ARITE T,et al.Suppression of tiller budactivity in tillering dwarf mutants of rice[J].Plant Cell Physiol,2005,46(1):79-86.
• [14]ZHOU Y,ZHU J,LI Z,et al.Fine mapping and cloning of MT1,a novelallele of D10[J].Progress Nat Sci,2009,19:1683-1689.
• [15]TONG H,JIN Y,LIU W,et al.DWARF AND LOW-TILLERING,a newmember of GRAS family,plays positive roles in brassinosteroid signalingin rice[J].Plant J,2009,58:803-816.
• [16]MEI H W,XU J L,LI Z K,et al.QTLs influencing panicle size detectedin two reciprocal introgressive line(IL)populations in rice(Oryza sativaL.)[J].Theor Appl Genet,2006,112:648-656.
• [17]ASHIKARI M,SAKAKIBARA H,LIN S,et al.Cytokinin oxidase regulatesrice grain production[J].Science,2005,309:741-745.
• [18]YU S B,LI J X,XU C G,et al.Importance of epistasis as the genetic basisof heterosis in an elite rice hybrid[J].Proc Natl Acad Sci USA,1997,94:9226-9231.
• [19]YU S B,LI J X,XU C G,et al.Identification of quantitative trait loci andepistatic interactions for plant height and heading date in rice[J].TheorAppl Genet,2002,104:619-625.
• [20]邢永忠,徐才国,华金平,等.水稻株高和抽穗期基因的定位和分离[J].植物学报,2001,43(7):721-726.
• [21]XING Y Z,TAN Y F,HUA J P,et al.Characterization of the maineffects,epistatic and their environmental interactions of QTLs on the ge-netic basis of yield traits in rice[J].Theor Appl Genet,2002,105:248-257.
• [22]XING Y Z,TANG W J,XUE W Y,et al.Fine mapping of a major quanti-tative trait loci,qSSP7,controlling the number of spikelets per panicles asa single Mendelian factor in rice[J].Theor Appl Genet,2008,116:789-796.
• [23]XUE W,XING Y,WENG X,et al.Natural variation in Ghd7 is an impor-tant regulator of heading date and yield potential in rice[J].Nat Genet,2008,40:761-767.
• [24]HUANG X,QIAN Q,LIU Z,et al.Natural variation at the DEP1 locus en-hances grain yield in rice[J].Nat Genet,2009,41:494-497.
• [25]LI S,QIAN Q,FU Z,et al.Short panicle1 encodes a putative PTR familytransporter and determines rice panicle size[J].Plant J,2009,58(4):592-605.
• [26]KURAKAWA T,UEDA N,MAEKAWA M,et al.Direct control of shootmeristem activity by a cytokinin-activating enzyme[J].Nature,2007,445:652-655.
• [27]KOMATSU M,MAEKAWA M,SHIMANOTO K,et al.The LAX1 andFRIZZY PANICLE 2 genes determine the inflorescence architecture ofrice by controlling rachis-branch and spikelet development[J].Dev Bi-ol,2001,231:364-373.
• [28]KOMATSU K,MAEKAWA M,UJIIE S,et al.LAX and SPA:Major regula-tors of shoot branching in rice[J].Proc Natl Acad Sci USA,2003,100:11765-11770.
• [29]TABUCHI H,ZHANG Y,HATTORI S,et al.LAX PANICLE2 of rice en-codes a novel nuclear protein and regulates the formation of axillary meri-stems[J].Plant Cell,2011,23:3276-3287.
• [30]KOMATSU M,CHUJO A,NAGATO Y,et al.FRIZZY PANICLE is re-quired to prevent the formation of axillary meristems and to establish flo-ral meristem identity in rice spikelets[J].Development,2003,130:3841-3850.
• [31]IKEDA K,NAGASAWA N,NAGATO Y.ABERRANT PANICLE ORGANI-ZATION 1 temporally regulates meristem identity in rice[J].Dev Biol,2005,282:349-360.
• [32]IKEDA K,ITO M,NAGASAWA N,et al.Rice ABERRANT PANICLE OR-GANIZATION 1,encoding an F-box protein,regulates meristem fate[J].Plant J,2007,51:1030-1040.
• [33]IKEDA-KAWAKATSU K,YASUNO N,OIKAWA T,et al.Expression lev-el of ABERRANT PANICLE ORGANIZATION1 determines rice inflores-cence form through control of cell proliferation in the meristem[J].PlantPhysiol,2009,150:736-747.
• [34]ANDO T,YAMANOTO T,SHIMIZU T,et al.Genetic dissection and pyra-miding of quantitative traits for panicle architecture by using chromosomalsegment substitution lines in rice[J].Theor Appl Genet,2008,116:881-890.
• [35]TERAO T,NAGATA K,MORINO K,et al.A gene controlling the numberof primary rachis branckes alse controls the vascular bundle formation andhence is responsible to increase the harvest index and grain yield in rice[J].Theor Appl Genet,2010,120:875-893.
• [36]NAKAGAWA M,SHIMAMOTO K,KYOZUKA J.Overexpression of RCN1and CN2,rice TERMINAL FLOWER1/CENTRORADIALIS homologs,con-fers delay of phase transition and altered panicle morphology in rice[J].Plant J,2002,29(6):743-750.
• [37]ZHA X,LUO X,QIAN X,et al.Over-expression of the rice LRK1 gene im-proves quantitative yield components[J].Plant Biotech J,2009,7:611-620.
• [38]TIAN F,ZHU Z,ZHANG B,et al.Fine mapping of a quantitative trait lo-cus for grain number per panicle from wild rice(Oryza rufipogon Griff.)[J].Theor Appl Genet,2006,113:619-629.
• [39]LI X,TIAN F,HUANG H,et al.Construction of the physical map of thegpa7 locus reveals that a large segment was deleted during rice domestica-tion[J].Plant Cell Reports,2008,27:1087-1092.
• [40]LIU T,MAO D,ZHANG S,et al.Fine mapping SPP1,a QTL controllingthe number of spikelets per panicle,to a BAC clone in rice(Oryza sativa)[J].Theor Appl Genet,2009,118:1509-1517.
• [41]SHAN J X,ZHU M Z,SHI M,et al.Fine mapping and candidate gene a-nalysis of spd6,responsible for small panicle and dwarfness in wild rice(Oryza rufipogon Griff.)[J].Theor Appl Genet,2009,119:827-836.
• [42]SONG X J,HUANG W,SHI M,et al.A QTL for rice grain width andweight encodes a previously unknown RING-type E3 ubiquitin ligase[J].Nat Genet,2007,39:623-630.
• [43]SHOMURA A,IZAWA T,EBANA K,et al.Deletion in a gene associatedwith grain size increased yields during rice domestication[J].Nat Genet,2008,40:1023-1028.
• [44]WENG J,GU S,WAN X,et al.Isolation and initial characterization ofGW5,a major QTL associated with rice grain width and weight[J].CellRes,2008,18:1199-1209.
• [45]TAN Y F,XING Y Z,LI J X,et al.Genetic bases of appearance quality ofrice grains in Shanyou 63,an elite rice hybrid[J].Theor Appl Genet,2000,101:823-829.
• [46]THOMSON M J,TAI T H,MCCLUNG A M,et al.Mapping quantitativetrait loci for yield,yield components and morphological trains in a ad-vanced backcross population between Oryza rufipogon and the Oryza sati-va cultivar Jefferson[J].Theor Appl Genet,2003,107:479-493.
• [47]LI J,THOMSON M,MCCOUCH S R.Fine mapping of a grain-weightquantitative trait locus in the pericentromeric region of rice chromosome 3[J].Genetics,2004,168:2187-2195.
• [48]FAN C,XING Y,MAO H,et al.GS3,a major QTL for grain length andweight and minor QTL for grain width and thickness in rice,encodes aputative transmembrane protein[J].Theor Appl Genet,2006,112:1164-1171.
• [49]FAN C,YU S,WANG C,et al.A causal C-A mutation in the second ex-on of GS3 highly associated with rice grain length and validated as a func-tional marker[J].Theor Appl Genet,2009,118:465-472.
• [50]MAO H,SUN S,YAO J,et al.Linking differential domain functions of theGS3 protein to natural variation of grain size in rice[J].Proc Natl AcadSci USA,2010,107:19679-19584.
• [51]LI Y,FAN C,XING Y,et al.Natural variation in GS5 plays an importantrole in regulating grain size and yield in rice[J].Nat Genet,2011,43:1266-1269.
• [52]WANG E,WANG J,ZHU X,et al.Control of rice grain-filling and yieldby a gene with a potential signature of domestication[J].Nat Genet,2008,40(11):1370-1374.
• [53]XIE X,SONG M H,JIN F,et al.Fine mapping of a grain weight quantita-tive trait locus on rice chromosome 8 using near-isogenic lines derivedfrom a cross between Oryza sativa and Oryza rufipogon[J].Theor ApplGenet,2006,113:885-894.
• [54]XIE X,JIN F,SONG M H,et al.Fine mapping of a yield-enhancing QTLcluster associated with transgressive variation in an Oryza sativa×O.rufipogon cross[J].Theor Appl Genet,2008,116:613-622.
• [55]MONNA L,KITAZAWA N,YOSHINO R,et al.Positional cloning of ricesemidwarfing gene,sd-1:rice"green revolution gene"encodes a mutantenzyme involved in gibberellin synthesis[J].DNA Res,2002,9(1):11-17.
• [56]SASAKI A,ASHIKARI M,UEGUCHI-TANAKA M,et al.A mutant gib-berellin-synthesis gene in rice[J].Nature,2002,416:701-702.
• [57]SPIELMEYER W,ELLIS M H,CHANDLER P M.Semidwarf(sd-1),“green revolution”'rice,contains a defective gibberellin 20-oxidasegene.Proc Natl Acad Sci USA,2002,99:9043-9048.
• [58]ASHIKARI M,WU J,YANO M,et al.Rice gibberellin-insensitive dwarfmutant gene Dwarf 1 encodes theα-subunit of GTP-binding protein[J].Proc Natl Acad Sci USA,1999,96:10284-10289.
• [59]FUJISAWA Y,KATO T,OHKI S,et al.Suppression of the heterotrimericG protein causes abnormal morphology,including dwarfism,in rice[J].Proc Natl Acad Sci USA,1999,99:7575-7580.
• [60]IKEDA A,UEGUCHI-TANAKA M,SONODA Y,et al.Slender rice,a con-stitutive gibberellin response mutant,is caused by a null mutation of theSLR1 gene,an ortholog of the height-regulating gene GAI/RGA/RHT/D8[J].Plant Cell,2001,13:999-1010.
• [61]UEGUCHI-TANAKA M,ASHIKARI M,NAKAJIMA M,et al.GIBBEREL-LIN INSENSITIVE DWARF1 encodes a soluble receptor for gibberellin[J].Nature,2005,437:693-698.
• [62]HATTORI Y,NAGAI K,FURUKAWA S,et al.The ethylene response fac-tors SNORKEL1 and SNORKEL2 allow rice to adapt to deep water[J].Nature,2009,460:1026-1031.
• [63]RUTGER J N,CARNAHAN H L.A fourth genetic element to facilitatehybrid cereal production-a recessive tall in rice[J].Crop Sci,1981,21:373-376.
• [64]LUO A,QIAN Q,YIN H,et al.EUI1,encoding a putative cytochromeP450 monooxygenase,regulates internode elongation by modulating gibber-ellin responses in rice[J].Plant Cell Physiol,2006,47(2):181-191.
• [65]杨蜀岚,杨仁崔,曲雪萍,等.水稻长穗颈高秆隐性基因eui2的遗传及其微卫星分析[J].植物学报,2001,43(1):67-71.
• [66]杨仁崔,张书标,黄荣华,等.高秆隐性杂交水稻(e-杂交水稻)的育种技术[J].中国农业科学,2002,35(3):233-237.
• [67]YAMAMURO C,IHARA Y,WU X,et al.Loss of function of a rice brassi-nosteroid insensitive1 homolog prevents internode elongation and bendingof the lamina joint[J].Plant Cell,2000,12:1591-1605.
• [68]HONG Z,UEGUCHI-TANAKA M,UMEMURA K,et al.A rice brassinos-teroid-deficient mutant,ebisu dwarf(d2),is caused by a loss of functionof a new member of cytochrome P450[J].Plant Cell,2003,15:2900-2910.
• [69]TANABE S,ASHIKARI M,FUJIOKA S,et al.A novel cytochrome P450 isimplicated in brassinosteroid biosynthesis via the characterization of a ricedwarf mutant,dwarf11,with reduced seed length[J].Plant Cell,2005,17:776-790.
• [70]HONG Z,UEGUCHI-TANAKA M,SHIMIZU-SATO S,et al.Loss-of-function of a rice brassinosteroid biosynthetic enzyme,C-6 oxidase,pre-vents the organized arrangement and polar elongation of cells in the leavesand stem[J].Plant J,2002,32:495-508.
• [71]HONG Z,UEGUCHI-TANAKA M,FUJIOKA F,et al.The rice brassinos-teroid-deficient dwarf2 mutant,defective in the rice homolog of Arabi-dopsis DIMINUTO/DWARF1,is rescued by the endogenously accumula-ted alternative bioactive brassinosteroid,dolichosterone[J].Plant Cell,2005,17:2243-2254.
• [72]HUA J,XING Y,WU W,et al.Single-locus heterotic effects and domi-nance by dominance interaction can adequately explain the genetic basisof heterosis in an elite rice hybrid[J].Proc Natl Acad Sci USA,2003,100(5):2574-2579.
• [73]LI Z K,LUO L J,MEI H W,et al.Overdominant epistatic loci are the pri-mary genetic basis of inbreeding depression and heterosis in rice.I.Bio-mass and grain yield[J].Genetics,2001,158:1737-1753.
• [74]LUO L J,LI Z K,MEI H W,et al.Overdominant epistatic loci are the pri-mary genetic basis of inbreeding depression and heterosis in rice.II.Grainyield components[J].Genetics,2001,158:1755-1771.
• [75]MEI H W,LUO L J,YING C S,et al.Gene actions of QTLs affecting sev-eral agronomic traits resolved in a recombinant inbred rice population andtwo testcross populations[J].Theor Appl Genet,2003,107:89-101.
• [76]LUO X,FU Y,ZHANG P,et al.Additive and over-dominant effects re-sulting from epistatic loci are the primary genetic basis of heterosis in rice[J].J Integr Plant Biol,2009,51(4):393-408.
• [77]LI Z K,PINSON S R M,PATERSON A H,et al.Epistasis for three grainyield components in rice(Oryza sativa L.)[J].Genetics,1997,145:453-465.
• [78]LI Z K,PINSON S R M,STANSEL J M,et al.Genetic dissection of thesource-sink relationship affecting fecundity and yield in rice(Oryza sa-tiva L.)[J].Mol Breed,1998,4:419-426.
• [79]LI Z K,LUO L J,MEI H W,et al.Overdominant epistatic loci are the pri-mary genetic basis of inbreeding depression and heterosis in rice.I.Bio-mass and grain yield[J].Genetics,2001,158:1737-1753.
• [80]KRIEGER U,LIPPMAN Z B,ZAMIR D.The flowering gene SINGLEFLOWER TRUSS drives heterosis for yield in tomato[J].Nat Genet,2010,42:459-463.
• [81]LUO X J,WU S,TIAN F,et al.Identification heterotic loci associated withyield-related traits from Chinese common wild rice(Oryza rufipogonGriff.)[J].Plant Sci,2011,181:14-22.
• [82]CHEN J,DING J,OUYANG Y,et al.A triallelic system of S5 is a majorregulator of the reproductive barrier and compatibility of indica-japonicahybrids in rice[J].Proc Natl Acad Sci USA,2008,105:11436-11441.
• [83]JIAO Y,WANG Y,XUE D,et al.Regulation of OsSPL14 by OsmiR156defines ideal plant architecture in rice[J].Nat Genet,2010,42:541-544.
• [84]MIURA K,IKEDA M,MATSUBARA A,et al.OsSPL14 promotes paniclebranching and higher grain productivity in rice[J].Nat Genet,2010,42:545-549.
• [85]SAKAMOTO T,MORINAKA Y,OHNISHI T,et al.Erect leaves caused bybrassinosteroid deficiency increase biomass production and grain yield inrice[J].Nat Biotech,2006,24:105-109.
• [86]WU C,TRIEU A,RADHAKRISHNAN P,et al.Brassinosteriods regulategrain filling in rice[J].Plant Cell,2008,20:2130-2145.
• [87]OOKAWA T,HOBO T,YANO M,et al.New approach for rice improve-ment using a pleiotropic QTL gene for lodging resistance and yield[J].Nat Commun,2010,1:132.
• [88]SATO Y,HONG S K,TAGIRI A,et al.A rice homeobox gene,OSH1,isexpressed before organ differentiation in a specific region during early em-bryogenesis[J].Proc Natl Acad Sci USA,1996,93:8117-8122.
• [89]PENG J,RICHARDS D E,HARTLEY N M,et al.‘Green revolution’genes encode mutant gibberellin response modulators[J].Nature,1999,400:256-261.
• [90]PATERSON A H,LAND E S,HEWITT J D,et al.Resolution of quantita-tive traits into Mendelian factors by using a complete linkage map of re-striction fragment length polymorphisms[J].Nature,1988,335:721-726.
• [91]PATERSON A H,DEVERNA J W,LANINI B,et al.Fine mapping ofquantitative trait loci using selected overlapping recombinant chromo-somes in an interspecies cross of tomato[J].Genetics,1990,124:735-742.
• [92]ESHED Y,ABU-ABIED M,SARANGA Y,et al.Lycopersicon esculentumlines containing small overlapping introgressions from L.pennellii[J].Theor Appl Genet,1992,83:1027-1034.
• [93]TANKSLEY S D.Mapping polygenes[J].Annu Rev Genet,1993,27:205-233.
• [94]TANKSLEY S D,NELSON J C.Advanced backcross QTL analysis:amethod for the simultaneous discovery and transfer of valuable QTLs fromunadapted germplasm into elite breeding lines[J].Theor Appl Genet,1996,92:191-203.
• [95]SOBRIZAL K,SANCHEZ P L,DOI K,et al.Development of Oryza glu-maepatula introgression lines in rice,O.sativa L[J].Rice Genet Newsl,1996,16:107-108.
• [96]KURAKAZU T,SOBRIZAL,IKEDA K,et al.Oryza meridionalis chromo-somal segment introgression lines in cultivated rice,O.sative L[J].RiceGenet Newsl,2001,18:81-82.
• [97]JIN J,HUANG W,GAO J P,et al.Genetic control of rice plant architec-ture under domestication[J].Nat Genet,2008,40:1365-1369.
• [98]TIAN F,LI DJ,FU Q,et al.Construction of introgression lines carryingwild rice(Oryza rufipogon Griff.)segments in cultivated rice(O.sativaL.)background and characterization of introgressed segments associatedwith yield-related traits[J].Theor Appl Genet,2006,112:570-580.
• [99]TAN L B,LIU F X,XUE W,et al.Development of Oryza rufipogon andOryza sativa introgression lines and assessment for yield-related quanti-tative trait loci[J].J Integr Plant Biol,2007,49:871-884.
• [100]TAN L,LI X,LIU F,et al.Control of a key transition from prostrate to e-rect growth in rice domestication[J].Nat Genet,2008,40:1360-1364.
• [101]LIN Z,GRIFFITH M E,LI X,et al.Origin of seed shattering in rice(Oryza stativa L.)[J].Planta,2007,226:11-20.