拟南芥生态型Zu-0减少主茎分枝的遗传机理探究
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摘要
植物分枝发育很大程度上决定了植物地上部的形态结构,在植物的形态建成中占有非常重要的地位.它又与作物的生物量和结实量有着密切的关系,是作物的重要农艺性状之一.以实验室常用拟南芥生态型Col-0和减少主茎分枝(RSB)的生态型Zu-0为亲本,构建了包含192个株系的重组自交系群体.利用分离群体分组分析法(BSA)检测出与RSB性状连锁的13个分子标记.最终初定位到4个减少主茎分枝的数量性状位点(QTL).qRSBZu-2和qRSBZu-3区域分别含有已知的同时控制开花时间和减少主茎分枝表型的FRI和FLC基因.qRSBZu-1和qRSBZu-4区域尽管含有已知分枝调控基因AGL6和CUC2,但AGL6和CUC2测序结果表明这2个基因在Col-0和Zu-0中没有编码氨基酸的改变,而且AGL6的表达量也没有改变,表明这2个位点还存在着其他基因参与了RSB性状的调控.构建杂合自交家系(HIF)群体,对qRSBZu-1和qRSBZu-4的真实性进行了验证.
The development of shoot branching is a major determinant of plant architecture and plays a pivotal role in the morphogenesis of plant.Shoot branching is closely related to the accumulation of crop biomass and fruit yields,and is also considered as an important agronomic trait for crop.In this study,a recombinant inbred lines(RILs)population was generated from a cross between Col-0(the commonly used Arabidopsis accessions)and Zu-0 accessions which lacks most side shoots in the axils of cauline leaves.13 polymorphic molecular markers tightly linked to reduced stem branching phenotype(RSB)were screened out by bulked-segregant analysis(BSA).A total of 4 QTL for RSB were identified.The qRSBZu-2 and qRSBZu-3 regions respectively contain FRI and FLC genes that control flowering time and reduce the stem branch phenotype at the same time.Although qRSBZu-1 and qRSBZu-4 regions contain AGL6 and CUC2 genes that have been reported to involve the regulation of shoot branching,the sequencing results show there is no any amino acid change between Col-0 and Zu-0.In addition,there is no significant difference in the expression level of AGL6 between Col-0 and Zu-0.These results suggested there are other genes regulating the development of shoot branching in these two QTL regions.Using Heterogeneous inbred family(HIF)analysis,the qRSBZu-1 and qRSBZu-4 were validated.The results laid the foundation for further fine-mapping and map-based cloning of these two QTL.
The development of shoot branching is a major determinant of plant architecture and plays a pivotal role in the morphogenesis of plant.Shoot branching is closely related to the accumulation of crop biomass and fruit yields,and is also considered as an important agronomic trait for crop.In this study,a recombinant inbred lines(RILs)population was generated from a cross between Col-0(the commonly used Arabidopsis accessions)and Zu-0 accessions which lacks most side shoots in the axils of cauline leaves.13 polymorphic molecular markers tightly linked to reduced stem branching phenotype(RSB)were screened out by bulked-segregant analysis(BSA).A total of 4 QTL for RSB were identified.The qRSBZu-2 and qRSBZu-3 regions respectively contain FRI and FLC genes that control flowering time and reduce the stem branch phenotype at the same time.Although qRSBZu-1 and qRSBZu-4 regions contain AGL6 and CUC2 genes that have been reported to involve the regulation of shoot branching,the sequencing results show there is no any amino acid change between Col-0 and Zu-0.In addition,there is no significant difference in the expression level of AGL6 between Col-0 and Zu-0.These results suggested there are other genes regulating the development of shoot branching in these two QTL regions.Using Heterogeneous inbred family(HIF)analysis,the qRSBZu-1 and qRSBZu-4 were validated.The results laid the foundation for further fine-mapping and map-based cloning of these two QTL.
引文
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[14]LI X,QIAN Q,FU Z,et al.Control of tillering in rice[J].Nature,2003,422(6932):618-621.
[15]KOMATSU K,MAEKAWA M,UJIIE S,et al.LAX and SPA:Major regulators of shoot branching in rice[J].Proc Natl Acad Sci USA,2003,100(20):11765-11770.
[16]GALLAVOTTI A,ZHAO Q,KYOZUKA J,et al.The role of barren stalk1in the architecture of maize[J].Nature,2004,432(7017):630-635.
[17]STIRNBERG P,CHATFIELD S P,LEYSER H M.AXR1acts after lateral bud formation to inhibit lateral bud growth in Arabidopsis[J].Plant Physiol,1999,121(3):839-847.
[18]AGUILAR-MARTINEZ J A,POZA-CARRION C,CUBAS P.Arabidopsis BRANCHED1acts as an integrator of branching signals within axillary buds[J].Plant Cell,2007,19(2):458-472.
[19]BEVERIDGE C A,SYMONS G M,TURNBULL C G.Auxin inhibition of decapitation-induced branching is dependent on graft-transmissible signals regulated by genes Rms1and Rms2[J].Plant Physiol,2000,123(2):689-698.
[20]RAMEAU C,MURFET I C,LAUCOU V,et al.Pea RMS6mutants exhibit increased basal branching[J].Physiol Plant,2002,115(3):458-467.
[21]NAPOLI C.Highly Branched phenotype of the petunia dad1-1mutant is reversed by grafting[J].Plant Physiol,1996,111(1):27-37.
[22]SIMONS J L,NAPOLI C A,JANSSEN B J,et al.Analysis of the decreased apical dominance genes of petunia in the control of axillary branching[J].Plant Physiol,2007,143(2):697-706.
[23]ISHIKAWA S,MAEKAWA M,ARITE T,et al.Suppression of tiller bud activity in tillering dwarf mutants of rice[J].Plant Cell Physiol,2005,46(1):79-86.
[24]ZOU J,CHEN Z,ZHANG S,et al.Characterizations and fine mapping of a mutant gene for high tillering and dwarf in rice(Oryza sativa L.)[J].Planta,2005,222(4):604-612.
[25]ZOU J,ZHANG S,ZHANG W,et al.The rice HIGH-TILLERING DWARF1encoding an ortholog of Arabidopsis MAX3is required for negative regulation of the outgrowth of axillary buds[J].Plant J,2006,48(5):687-698.
[26]ARITE T,IWATA H,OHSHIMA K,et al.DWARF10,an RMS1/MAX4/DAD1ortholog,controls lateral bud outgrowth in rice[J].Plant J,2007,51(6):1019-1029.
[27]TAKEDA T,SUWA Y,SUZUKI M,et al.The OsTB1gene negatively regulates lateral branching in rice[J].Plant J,2003,33(3):513-520.
[28]TANTIKANJANA T,YONG J W,LETHAM D S,et al.Control of axillary bud initiation and shoot architecture in Arabidopsis through the SUPERSHOOT gene[J].Genes Dev,2001,15(12):1577-1588.
[29]WEIGEL D.Natural variation in Arabidopsis:From molecular genetics to ecological genomics[J].Plant Physiol,2012,158(1):2-22.
[30]UNGERER M C,HALLDORSDOTTIR S S,MODLISZEWSKI J L,et al.Quantitative trait loci for inflorescence development in Arabidopsis thaliana[J].Genetics,2002,160(3):1133-1151.
[31]EL-LITHY M E,REYMOND M,STICH B,et al.Relation among plant growth,carbohydrates and flowering time in the Arabidopsis Landsberg erecta×Kondara recombinant inbred line population[J].Plant Cell Environ,2010,33(8):1369-1382.
[32]HUANG X,DING J,EFFGEN S,et al.Multiple loci and genetic interactions involving flowering time genes regulate stem branching among natural variants of Arabidopsis[J].New Phytol,2013,199(3):843-857.
[33]KALININA A,MIHAJLOVIC N,GRBIC V.Axillary meristem development in the branchless Zu-0ecotype of Arabidopsis thaliana[J].Planta,2002,215(5):699-707.
[34]HUANG X,EFFGEN S,MEYER R C,et al.Epistatic natural allelic variation reveals a function of AGAMOUS-LIKE6in axillary bud formation in Arabidopsis[J].Plant Cell,2012,24(6):2364-2379.
[35]JOHANSON U,WEST J,LISTER C,et al.Molecular analysis of FRIGIDA,a major determinant of natural variation in Arabidopsis flowering time[J].Science,2000,290(5490):344-347.
[36]SHELDON C C,BURN J E,PEREZ P P,et al.The FLF MADS box gene:A repressor of flowering in Arabidopsis regulated by vernalization and methylation[J].Plant Cell,1999,11(3):445-458.
[37]MICHAELS S D,AMASINO R M.FLOWERING LOCUS C encodes a novel MADS domain protein that acts as a repressor of flowering[J].Plant Cell,1999,11(5):949-956.
[38]LONG J A,MOAN E I,MEDFORD J I,et al.A member of the KNOTTED class of homeodomain proteins encoded by the STM gene of Arabidopsis[J].Nature,1996,379(6560):66-69.
[2]TANG J,ZENG W,WANG W,et al.Genetic analysis and gene mapping of a rice few-tillering mutant in early backcross populations(Oryza sativa L.)[J].Sci China C:Life Sci,2001,44(6):570-575.
[3]DOEBLEY J,STEC A,HUBBARD L.The evolution of apical dominance in maize[J].Nature,1997,386(6624):485-488.
[4]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.
[5]BEVERIDGE C A.Axillary bud outgrowth:Sending a message[J].Curr Opin Plant Biol,2006,9(1):35-40.
[6]SCHUMACHER K,SCHMITT T,ROSSBERG M,et al.The Lateral suppressor(Ls)gene of tomato encodes a new member of the VHIID protein family[J].Proc Natl Acad Sci USA,1999,96(1):290-295.
[7]SCHMITZ G,TILLMANN E,CARRIERO F,et al.The tomato Blind gene encodes a MYB transcription factor that controls the formation of lateral meristems[J].Proc Natl Acad Sci USA,2002,99(2):1064-1069.
[8]BLEIN T,PULIDO A,VIALETTE-GUIRAUD A,et al.A conserved molecular framework for compound leaf development[J].Science,2008,322(5909):1835-1839.
[9]GREB T,CLARENZ O,SCHAFER E,et al.Molecular analysis of the LATERAL SUPPRESSOR gene in Arabidopsis reveals a conserved control mechanism for axillary meristem formation[J].Genes Dev,2003,17(9):1175-1187.
[10]MULLER D,SCHMITZ G,THERES K.Blind homologous R2R3 Myb genes control the pattern of lateral meristem initiation in Arabidopsis[J].Plant Cell,2006,18(3):586-597.
[11]TALBERT P B,ADLER H T,PARKS D W,et al.The REVOLUTA gene is necessary for apical meristem development and for limiting cell divisions in the leaves and stems of Arabidopsis thaliana[J].Development,1995,121(9):2723-2735.
[12]RAMAN S,GREB T,PEAUCELLE A,et al.Interplay of miR164,CUP-SHAPED COTYLEDONgenes and LATERAL SUPPRESSOR controls axillary meristem formation in Arabidopsis thaliana[J].Plant J,2008,55(1):65-76.
[13]YANG F,WANG Q,SCHMITZ G,et al.The bHLH protein ROX acts in concert with RAX1and LASto modulate axillary meristem formation in Arabidopsis[J].Plant J,2012,71(1):61-70.
[14]LI X,QIAN Q,FU Z,et al.Control of tillering in rice[J].Nature,2003,422(6932):618-621.
[15]KOMATSU K,MAEKAWA M,UJIIE S,et al.LAX and SPA:Major regulators of shoot branching in rice[J].Proc Natl Acad Sci USA,2003,100(20):11765-11770.
[16]GALLAVOTTI A,ZHAO Q,KYOZUKA J,et al.The role of barren stalk1in the architecture of maize[J].Nature,2004,432(7017):630-635.
[17]STIRNBERG P,CHATFIELD S P,LEYSER H M.AXR1acts after lateral bud formation to inhibit lateral bud growth in Arabidopsis[J].Plant Physiol,1999,121(3):839-847.
[18]AGUILAR-MARTINEZ J A,POZA-CARRION C,CUBAS P.Arabidopsis BRANCHED1acts as an integrator of branching signals within axillary buds[J].Plant Cell,2007,19(2):458-472.
[19]BEVERIDGE C A,SYMONS G M,TURNBULL C G.Auxin inhibition of decapitation-induced branching is dependent on graft-transmissible signals regulated by genes Rms1and Rms2[J].Plant Physiol,2000,123(2):689-698.
[20]RAMEAU C,MURFET I C,LAUCOU V,et al.Pea RMS6mutants exhibit increased basal branching[J].Physiol Plant,2002,115(3):458-467.
[21]NAPOLI C.Highly Branched phenotype of the petunia dad1-1mutant is reversed by grafting[J].Plant Physiol,1996,111(1):27-37.
[22]SIMONS J L,NAPOLI C A,JANSSEN B J,et al.Analysis of the decreased apical dominance genes of petunia in the control of axillary branching[J].Plant Physiol,2007,143(2):697-706.
[23]ISHIKAWA S,MAEKAWA M,ARITE T,et al.Suppression of tiller bud activity in tillering dwarf mutants of rice[J].Plant Cell Physiol,2005,46(1):79-86.
[24]ZOU J,CHEN Z,ZHANG S,et al.Characterizations and fine mapping of a mutant gene for high tillering and dwarf in rice(Oryza sativa L.)[J].Planta,2005,222(4):604-612.
[25]ZOU J,ZHANG S,ZHANG W,et al.The rice HIGH-TILLERING DWARF1encoding an ortholog of Arabidopsis MAX3is required for negative regulation of the outgrowth of axillary buds[J].Plant J,2006,48(5):687-698.
[26]ARITE T,IWATA H,OHSHIMA K,et al.DWARF10,an RMS1/MAX4/DAD1ortholog,controls lateral bud outgrowth in rice[J].Plant J,2007,51(6):1019-1029.
[27]TAKEDA T,SUWA Y,SUZUKI M,et al.The OsTB1gene negatively regulates lateral branching in rice[J].Plant J,2003,33(3):513-520.
[28]TANTIKANJANA T,YONG J W,LETHAM D S,et al.Control of axillary bud initiation and shoot architecture in Arabidopsis through the SUPERSHOOT gene[J].Genes Dev,2001,15(12):1577-1588.
[29]WEIGEL D.Natural variation in Arabidopsis:From molecular genetics to ecological genomics[J].Plant Physiol,2012,158(1):2-22.
[30]UNGERER M C,HALLDORSDOTTIR S S,MODLISZEWSKI J L,et al.Quantitative trait loci for inflorescence development in Arabidopsis thaliana[J].Genetics,2002,160(3):1133-1151.
[31]EL-LITHY M E,REYMOND M,STICH B,et al.Relation among plant growth,carbohydrates and flowering time in the Arabidopsis Landsberg erecta×Kondara recombinant inbred line population[J].Plant Cell Environ,2010,33(8):1369-1382.
[32]HUANG X,DING J,EFFGEN S,et al.Multiple loci and genetic interactions involving flowering time genes regulate stem branching among natural variants of Arabidopsis[J].New Phytol,2013,199(3):843-857.
[33]KALININA A,MIHAJLOVIC N,GRBIC V.Axillary meristem development in the branchless Zu-0ecotype of Arabidopsis thaliana[J].Planta,2002,215(5):699-707.
[34]HUANG X,EFFGEN S,MEYER R C,et al.Epistatic natural allelic variation reveals a function of AGAMOUS-LIKE6in axillary bud formation in Arabidopsis[J].Plant Cell,2012,24(6):2364-2379.
[35]JOHANSON U,WEST J,LISTER C,et al.Molecular analysis of FRIGIDA,a major determinant of natural variation in Arabidopsis flowering time[J].Science,2000,290(5490):344-347.
[36]SHELDON C C,BURN J E,PEREZ P P,et al.The FLF MADS box gene:A repressor of flowering in Arabidopsis regulated by vernalization and methylation[J].Plant Cell,1999,11(3):445-458.
[37]MICHAELS S D,AMASINO R M.FLOWERING LOCUS C encodes a novel MADS domain protein that acts as a repressor of flowering[J].Plant Cell,1999,11(5):949-956.
[38]LONG J A,MOAN E I,MEDFORD J I,et al.A member of the KNOTTED class of homeodomain proteins encoded by the STM gene of Arabidopsis[J].Nature,1996,379(6560):66-69.