芝麻发芽期耐盐性鉴定方法研究及耐盐候选基因的挖掘
|
摘要
【目的】确定芝麻发芽期耐盐性鉴定适宜的Na Cl胁迫浓度和评价指标,发掘耐盐种质和耐盐相关重要基因,为芝麻耐盐性大规模鉴定、遗传改良和耐盐机理研究提供方法借鉴和优异基因资源。【方法】以8份耐盐性差异较大的芝麻种质为材料,在不同浓度的Na Cl(0、50、100、150、200和250 mmol·L-1)胁迫下发芽,测定其发芽势、成苗率、根长、芽长和鲜重等指标,通过对指标值的方差分析、主成分分析、隶属函数和相关性分析等,筛选芝麻发芽期耐盐性鉴定适宜的Na Cl处理浓度和评价指标。以100 mmol·L-1 Na Cl溶液为处理浓度,相对成苗率为鉴定指标对71份芝麻核心种质进行发芽期耐盐性鉴定和全基因组关联分析,并对获得的候选基因进行功能注释;通过Na Cl胁迫下芝麻幼苗叶片转录组测序和荧光定量PCR分析候选基因的表达模式,筛选耐盐相关候选基因。【结果】对不同浓度Na Cl胁迫下8份芝麻材料发芽各指标进行统计和方差分析表明,100 mmol·L-1的Na Cl处理下,除发芽势外,发芽指数、活力指数、成苗率、根长、芽长和苗鲜重的标准偏差值均较大,所有指标在0.05水平上均存在显著差异,100 mmol·L-1的Na Cl溶液可以作为芝麻发芽期耐盐性鉴定的适宜胁迫浓度;相对成苗率、相对发芽势、相对发芽指数、相对活力指数、相对芽长、相对根长和相对鲜重7个指标与芝麻发芽期耐盐性关系密切,贡献率较高,可以作为芝麻发芽期耐盐性鉴定的评价指标。71份芝麻核心种质材料的相对成苗率变异比较丰富,符合正态分布;通过对71份芝麻核心种质相对成苗率与SNP标记进行全基因组关联分析,检测到7个与芝麻发芽期耐盐性显著关联的SNP标记(LG5:688003、LG7:9582027、LG10:5274091、LG10:10788493、LG11:11924186、LG14:2128695和LG16:3930301),SNP标记上、下游各100 kb区间内共有基因67个,其中有功能注释的基因34个;通过耐盐材料S04在盐胁迫下的转录组测序和荧光定量PCR对预测的候选基因进行表达模式分析,共有21个候选基因受到盐胁迫诱导显著差异表达。【结论】芝麻发芽期耐盐性鉴定的适宜Na Cl胁迫浓度为100 mmol·L-1,相对成苗率等7个指标可以作为适宜的评价指标;检测到与芝麻发芽期耐盐性显著关联的SNP标记7个,并鉴定出耐盐候选基因21个。
【Objective】In order to evaluate salt tolerance of sesame germplasms and reveal the genetic basis of salt tolerance in sesame at the germination stage, optimum Na Cl concentration and best salt tolerance indexes were determined, salt-tolerant sesame genotypes as well as candidate genes were identified in this study. 【Method】 Eight sesame genotypes with different salt tolerance were germinated under different Na Cl concentrations(0, 50, 100, 150, 200 and 250 mmol·L-1). Seven germination indexes including germination potential, germination index, vigor index, seedling rate, radicle length, embryo length and fresh weight of seedling were measured. The optimum Na Cl concentration and salt tolerance indexes of sesame were determined using variance analysis, principal component analysis, membership function and correlation analysis based on the relative values of multiple indexes. In addition, genome-wide association analysis(GWAS) of salt tolerance related trait in 71 sesame germplasms was performed to find the SNP loci and candidate genes related to salt tolerance in sesame. Gene function annotation, transcriptome analyses and real-time quantitative reverse transcriptase PCR(q RT-PCR) were used to identify important candidate genes involved in salt tolerance. 【Result】The larger standard value deviations of germination indexes were observed at 100 mmol·L-1 Na Cl concentration, implying that 100 mmol·L-1 is the optimum Na Cl concentration for salt-tolerance screening in this study. The seven indexes were highly correlated with salt tolerance at germination stage, suggesting that, these indexes could be targeted for effective screening of salt-tolerance in large sesame germplasm. The genome-wide association analysis identified 7 SNP loci peaks(LG5:688003, LG7:9582027, LG10:5274091, LG10:10788493, LG11:11924186, LG14:2128695 and LG16:3930301) significantly associated with salt tolerance and 34 functional candidate genes. The transcriptome analysis and q RT-PCR showed that 21 candidate genes respond strongly to salt stress.【Conclusion】In this study, 100 mmol·L-1 is the optimum Na Cl concentration for salt-tolerance screening and seven indexes including relative seedling rate could be targeted for effective screening of salt-tolerance in large sesame germplasm. In addition, 7 SNP loci peaks significantly associated with salt tolerance were identified and 21 candidate salt-tolerant genes were discovered.
【Objective】In order to evaluate salt tolerance of sesame germplasms and reveal the genetic basis of salt tolerance in sesame at the germination stage, optimum Na Cl concentration and best salt tolerance indexes were determined, salt-tolerant sesame genotypes as well as candidate genes were identified in this study. 【Method】 Eight sesame genotypes with different salt tolerance were germinated under different Na Cl concentrations(0, 50, 100, 150, 200 and 250 mmol·L-1). Seven germination indexes including germination potential, germination index, vigor index, seedling rate, radicle length, embryo length and fresh weight of seedling were measured. The optimum Na Cl concentration and salt tolerance indexes of sesame were determined using variance analysis, principal component analysis, membership function and correlation analysis based on the relative values of multiple indexes. In addition, genome-wide association analysis(GWAS) of salt tolerance related trait in 71 sesame germplasms was performed to find the SNP loci and candidate genes related to salt tolerance in sesame. Gene function annotation, transcriptome analyses and real-time quantitative reverse transcriptase PCR(q RT-PCR) were used to identify important candidate genes involved in salt tolerance. 【Result】The larger standard value deviations of germination indexes were observed at 100 mmol·L-1 Na Cl concentration, implying that 100 mmol·L-1 is the optimum Na Cl concentration for salt-tolerance screening in this study. The seven indexes were highly correlated with salt tolerance at germination stage, suggesting that, these indexes could be targeted for effective screening of salt-tolerance in large sesame germplasm. The genome-wide association analysis identified 7 SNP loci peaks(LG5:688003, LG7:9582027, LG10:5274091, LG10:10788493, LG11:11924186, LG14:2128695 and LG16:3930301) significantly associated with salt tolerance and 34 functional candidate genes. The transcriptome analysis and q RT-PCR showed that 21 candidate genes respond strongly to salt stress.【Conclusion】In this study, 100 mmol·L-1 is the optimum Na Cl concentration for salt-tolerance screening and seven indexes including relative seedling rate could be targeted for effective screening of salt-tolerance in large sesame germplasm. In addition, 7 SNP loci peaks significantly associated with salt tolerance were identified and 21 candidate salt-tolerant genes were discovered.
引文
[1]杨真,王宝山.中国盐渍土资源现状及改良利用对策.山东农业科学,2015,47(4):125-130.YANG Z,WANG B S.Present status of saline soil resources and countermeasures for improvement and utilization in China.Shandong Agricultural Sciences,2015,47(4):125-130.(in Chinese)
[2]冯琛,党高兵,解建仓,孙博,汪妮.大米草玉米的耐盐效果及对盐渍化土壤的改良试验.水土保持通报,2011,31(6):47-50.FENG C,DANG G B,XIE J C,SUN B,WANG,N.Salt-tolerant effect of rice grass-maize and improvement on saline soil.Bulletin of Soil&Water Conservation,2011,31(6):47-50.(in Chinese)
[3]张鸿,林静,王立成,王阳,何威,李卫华.春小麦种质资源耐盐性鉴定及筛选.种子,2014,33(6):53-57.ZHANG H,LIN J,WANG L C,WANG Y,HE W,LI W H.The identification and screening of salt tolerance of spring wheat germplasm resources.Seed,2014,33(6):53-57.(in Chinese)
[4]田蕾,陈亚萍,刘俊,马晓刚,王娜,杨兵,李莹,郭海东,李娟,胡慧,张银霞,李培富.粳稻种质资源芽期耐盐性综合评价与筛选.中国水稻科学,2017,31(6):631-642.TIAN L,CHEN Y P,LIU J,MA X G,WANG N,YANG B,LI Y,GUO H D,LI J,HU H,ZHANG Y X,LI P F.Comprehensive evaluation and selection of rice(Oryza sativa japonica)germplasm for saline tolerance at germination stage.Chinese Journal of Rice Science,2017,31(6):631-642.(in Chinese)
[5]张海艳,赵海军.不同品种玉米萌发期和苗期耐盐性综合评价.玉米科学,2016,24(5):61-67.ZHANG H Y,ZHAO H J.Comprehensive evaluation of salt tolerance of different corn varieties at the germination and seedling stages.Journal of Maize Sciences,2016,24(5):61-67.(in Chinese)
[6]王俊娟,樊伟莉,叶武威,王德龙.陆地棉萌发出苗期耐盐鉴定方法的研究及应用.江苏农业科学,2010(2):87-88.WANG J J,FAN W L,YE W W,WANG D L.The research and application of the method of salt tolerance on upland cotton at the germination and seedling stages.Jiangsu Agricultural Sciences,2010(2):87-88.(in Chinese)
[7]沈义国,陈受宜.植物盐胁迫应答的分子机制.遗传,2001,23(4):365-369.SHEN Y G,CHEN S Y.Molecular mechanism of plant responses to salt stress.Hereditas,2001,23(4):365-369.(in Chinese)
[8]ZHU J K.Cell signaling under salt,water and cold stresses.Current Opinion in Plant Biology,2001,4(5):401-406.
[9]XU Y,LI S,LI L,ZHANG X,XU H,AN D.Mapping QTLs for salt tolerance with additive,epistatic and QTL×treatment interaction effects at seedling stage in wheat.Plant Breeding,2013,132(3):276-283.
[10]YU L X,LIU X,BOGE W,LIU X P.Genome-wide association study identifies loci for salt tolerance during germination in Autotetraploid alfalfa(Medicago sativa L.)using genotyping-by-sequencing.Frontiers in Plant Science,2016,7:956.
[11]KOBAYASHI Y,SADUKHAN A,TAZIB T,NAKANOY,KUSUNOKI K,KAMARA M,CHAFFAI R,IUCHI S,SAHOO L,KOBAYASHI M,HOEKENGA OA,KOYAMA H.Joint genetic and network analyses identify loci associated with root growth under Na Cl stress in Arabidopsis thaliana.Plant Cell&Environment,2016,39(4):918-934.
[12]DHINGRA A,DANIELL H.Plastid-expressed betaine aldehyde dehydrogenase gene in carrot cultured cells,roots,and leaves confers enhanced salt tolerance.Plant Physiology,2004,136(1):2843-2854.
[13]HMIDASAYARI A,GARGOURIBOUZID R,BIDANI A,JAOUA L,SAVOURE A,JAOUA S.Overexpression of delta1-pyrroline-5-carboxylate synthetase increases proline production and confers salt tolerance in transgenic potato plants.Plant Science,2005,169(4):746-752.
[14]QIAO W H,ZHAO X Y,LI W,LUO Y,ZHANG X S.Overexpression of Ae NHX1,a root-specific vacuolar Na+/H+antiporter from Agropyron elongatum,confers salt tolerance to Arabidopsis and Festuca plants.Plant Cell Reports,2007,26(9):1663-1672.
[15]陈凤莲,刘志斌,王健美,李旭锋,杨毅.油菜Bn RCH基因提高转基因拟南芥的耐盐性研究.四川大学学报(自然科学版),2013,50(3):643-648.CHEN F L,LIU Z B,WANG J M,LI X F,YANG Y.The studies of gene Bn RCH from Brassica napus enhances the tolerance to salt stress in transgenic Arabidopsis.Journal of Sichuan University(Natural Science),2013,50(3):643-648.(in Chinese)
[16]JING P,ZOU J,KONG L,HU S,WANG B,YANG J,XIE G.Os CCD1,a novel small calcium-binding protein with one EF-hand motif,positively regulates osmotic and salt tolerance in rice.Plant Science,2016,247:104-14.
[17]孙建,乐美旺,饶月亮,颜廷献,颜小文,周红英.芽期Na Cl胁迫下不同基因型芝麻的反应差异研究.干旱地区农业研究,2014,32(6):75-81.SUN J,LE M W,RAO Y L,YAN T X,YAN X W,ZHOU H Y.Genotypic variation in sesame salt tolerance during germination under Na Cl stress.Agricultural Research in the Arid Areas,2014,32(6):75-81.(in Chinese)
[18]徐芬芬,杜佳朋.干旱胁迫和盐胁迫对芝麻种子萌发的影响.种子,2013,32(11):85-86.XU F F,DU J P.Effect of drought and salt stress on seed germination of sesame.Seed,2013,32(11):85-86.(in Chinese)
[19]DIAS A S,LIMA G S D,GHEYI H R,NOBRE R G,SANTOS J B D.Emergence,growth and production of sesame under salt stress and proportions of nitrate and ammonium.Revista Caatinga,2017,30(2):458-467.
[20]FARISSI M,BOUIZGAREN A,FAGHIRE M,BARGAZ A,GHOULAM C.Agro-physiological and biochemical properties associated with adaptation of Medicago sativa populations to water deficit.Doga Turkish Journal of Botany,2013,37(6):1166-1175.
[21]吴兰荣,陈静,许婷婷,苗华荣,胡文广,禹山林.花生全生育期耐盐鉴定研究.花生学报,2005(1):20-24.WU L R,CHEN J,XU T T,MIAO H R,HU W G,YU S L.Identification of salt tolerance in peanut growth duration.Journal of Peanut Science,2005(1):20-24.(in Chinese)
[22]支巨振,毕辛华,杜克敏,常秀兰,杨淑惠,任淑萍,吴志行,李仁凤,赵菊英.GB/T 3543.4-1995,农作物种子检验规程发芽试验.北京:中国标准出版社,1995.ZHI J Z,BI X H,DU K M,CHANG X L,YANG S H,REN S P,WU Z H,LI R F,ZHAO J Y.GB/T 3543.4-1995,Rules for agricultural seed testing-Germination test.Beijing:China Standard Press,1995.(in Chinese)
[23]刘文萍,吕伟,黎冬华,任果香,张艳欣,文飞,韩俊梅,张秀荣.芝麻种质资源成株期抗旱性关联分析.中国农业科学,2017,50(4):625-639.LIU W P,LüW,LI D H,REN G X,ZHANG Y X,WEN F,HAN J M,ZHANG X R.Drought resistance of sesame germplasm resources and association analysis at adult stage.Scientia Agricultura Sinica,2017,50(4):625-639.(in Chinese)
[24]WEI X,LIU K,ZHANG Y,FENG Q,WANG L,ZHAO Y,LI D,ZHAO Q,ZHU X,ZHU X,LI W,FAN D,GAO Y,LU Y,ZHANG X,TANG X,ZHOU C,ZHU C,LIU L,ZHONG R,TIAN Q,WEN Z,WENG Q,HAN B,HUANG X,ZHANG X.Genetic discovery for oil production and quality in sesame.Nature Communications,2015,6:8609.
[25]BRADBURY P J,ZHANG Z,KROON D E,CASSTEVENS T M,RAMDOSS Y,BUCKLER E S.TASSEL:Software for association mapping of complex traits in diverse samples.Bioinformatics,2007,23(19):2633-2635.
[26]WEI L,MIAO H,ZHAO R,HAN X,ZHANG T,ZHANG H.Identification and testing of reference genes for Sesame gene expression analysis by quantitative real-time PCR.Planta,2013,237(3):873-889.
[27]SéNéCHAL F,WATTIER C,RUSTéRUCCI C,PELLOUX J.Homogalacturonan-modifying enzymes:Structure,expression,and roles in plants.Journal of Experimental Botany,2014,65(18):5125.
[28]OROZCOCáRDENAS M,NARVáEZVáSQUEZ J,RYAN C.Hydrogen peroxide acts as a second messenger for the induction of defense genes in tomato plants in response to wounding,systemin,and methyl jasmonate.The Plant Cell,2001,13(1):179-191.
[29]寇晓虹.番茄多聚半乳糖醛酸酶(PG)反义基因表达及相关功能研究[D].北京:中国农业大学,2003.KOU X H.Antisense gene expression and gene function of polygalacturonase(PG)in tomato[D].Beijing:China Agricultural University,2003.(in Chinese)
[30]O’DONNELL P J,TRUESDALE M R,CALVERT C M,DORANS A,ROBERTS M R,BOWLES D J.A novel tomato gene that rapidly responds to wound-and pathogen-related signals.The Plant Journal,1998,14(1):137-142.
[31]ROBERTS M R,WARNER S A J,DARBY R,LIM E K,DRAPER J,BOWLES D J.Differential regulation of a glucosyl transferase gene homologue during defence responses in tobacco.Journal of Experimental Botany,1999,50(332):407-410.
[32]李莹,柳参奎.红素氧还蛋白研究进展.草业科学,2016,33(6):1118-1125.LI Y,LIU S K.Research progress on rubredoxin.Pratacultural science,2016,33(6):1118-1125.(in Chinese)
[33]林凡云,陆琼娴,徐剑宏,史建荣.两个与盐和赤霉病菌胁迫相关的小麦糖基转移酶基因的克隆与表达.遗传,2008,30(12):1608-1614.LIN F Y,LU Q X,XU J H,SHI J R.Cloning and expression analysis of two salt and Fusarium graminearum stress associated UDPglucosyltransferases genes in wheat.Hereditas,2008,30(12):1608-1614.(in Chinese)
[34]ZHU J,FU X,KOO Y D,ZHU J K,FRANCIS E J J,MICHAEL W W A,ZHU Y,SHI H,YUN D J,HASEGAWA P M,BRESSAN R A.An enhancer mutant of arabidopsis salt overly sensitive 3 mediates both ion homeostasis and the oxidative stress response.Molecular&Cellular Biology,2007,27(14):5214.
[35]SAMANANI N,LISCOMBE D K,FACCHINI P J.Molecular cloning and characterization of norcoclaurine synthase,an enzyme catalyzing the first committed step in benzylisoquinoline alkaloid biosynthesis.The Plant Journal,2004,40(2):302-313.
[2]冯琛,党高兵,解建仓,孙博,汪妮.大米草玉米的耐盐效果及对盐渍化土壤的改良试验.水土保持通报,2011,31(6):47-50.FENG C,DANG G B,XIE J C,SUN B,WANG,N.Salt-tolerant effect of rice grass-maize and improvement on saline soil.Bulletin of Soil&Water Conservation,2011,31(6):47-50.(in Chinese)
[3]张鸿,林静,王立成,王阳,何威,李卫华.春小麦种质资源耐盐性鉴定及筛选.种子,2014,33(6):53-57.ZHANG H,LIN J,WANG L C,WANG Y,HE W,LI W H.The identification and screening of salt tolerance of spring wheat germplasm resources.Seed,2014,33(6):53-57.(in Chinese)
[4]田蕾,陈亚萍,刘俊,马晓刚,王娜,杨兵,李莹,郭海东,李娟,胡慧,张银霞,李培富.粳稻种质资源芽期耐盐性综合评价与筛选.中国水稻科学,2017,31(6):631-642.TIAN L,CHEN Y P,LIU J,MA X G,WANG N,YANG B,LI Y,GUO H D,LI J,HU H,ZHANG Y X,LI P F.Comprehensive evaluation and selection of rice(Oryza sativa japonica)germplasm for saline tolerance at germination stage.Chinese Journal of Rice Science,2017,31(6):631-642.(in Chinese)
[5]张海艳,赵海军.不同品种玉米萌发期和苗期耐盐性综合评价.玉米科学,2016,24(5):61-67.ZHANG H Y,ZHAO H J.Comprehensive evaluation of salt tolerance of different corn varieties at the germination and seedling stages.Journal of Maize Sciences,2016,24(5):61-67.(in Chinese)
[6]王俊娟,樊伟莉,叶武威,王德龙.陆地棉萌发出苗期耐盐鉴定方法的研究及应用.江苏农业科学,2010(2):87-88.WANG J J,FAN W L,YE W W,WANG D L.The research and application of the method of salt tolerance on upland cotton at the germination and seedling stages.Jiangsu Agricultural Sciences,2010(2):87-88.(in Chinese)
[7]沈义国,陈受宜.植物盐胁迫应答的分子机制.遗传,2001,23(4):365-369.SHEN Y G,CHEN S Y.Molecular mechanism of plant responses to salt stress.Hereditas,2001,23(4):365-369.(in Chinese)
[8]ZHU J K.Cell signaling under salt,water and cold stresses.Current Opinion in Plant Biology,2001,4(5):401-406.
[9]XU Y,LI S,LI L,ZHANG X,XU H,AN D.Mapping QTLs for salt tolerance with additive,epistatic and QTL×treatment interaction effects at seedling stage in wheat.Plant Breeding,2013,132(3):276-283.
[10]YU L X,LIU X,BOGE W,LIU X P.Genome-wide association study identifies loci for salt tolerance during germination in Autotetraploid alfalfa(Medicago sativa L.)using genotyping-by-sequencing.Frontiers in Plant Science,2016,7:956.
[11]KOBAYASHI Y,SADUKHAN A,TAZIB T,NAKANOY,KUSUNOKI K,KAMARA M,CHAFFAI R,IUCHI S,SAHOO L,KOBAYASHI M,HOEKENGA OA,KOYAMA H.Joint genetic and network analyses identify loci associated with root growth under Na Cl stress in Arabidopsis thaliana.Plant Cell&Environment,2016,39(4):918-934.
[12]DHINGRA A,DANIELL H.Plastid-expressed betaine aldehyde dehydrogenase gene in carrot cultured cells,roots,and leaves confers enhanced salt tolerance.Plant Physiology,2004,136(1):2843-2854.
[13]HMIDASAYARI A,GARGOURIBOUZID R,BIDANI A,JAOUA L,SAVOURE A,JAOUA S.Overexpression of delta1-pyrroline-5-carboxylate synthetase increases proline production and confers salt tolerance in transgenic potato plants.Plant Science,2005,169(4):746-752.
[14]QIAO W H,ZHAO X Y,LI W,LUO Y,ZHANG X S.Overexpression of Ae NHX1,a root-specific vacuolar Na+/H+antiporter from Agropyron elongatum,confers salt tolerance to Arabidopsis and Festuca plants.Plant Cell Reports,2007,26(9):1663-1672.
[15]陈凤莲,刘志斌,王健美,李旭锋,杨毅.油菜Bn RCH基因提高转基因拟南芥的耐盐性研究.四川大学学报(自然科学版),2013,50(3):643-648.CHEN F L,LIU Z B,WANG J M,LI X F,YANG Y.The studies of gene Bn RCH from Brassica napus enhances the tolerance to salt stress in transgenic Arabidopsis.Journal of Sichuan University(Natural Science),2013,50(3):643-648.(in Chinese)
[16]JING P,ZOU J,KONG L,HU S,WANG B,YANG J,XIE G.Os CCD1,a novel small calcium-binding protein with one EF-hand motif,positively regulates osmotic and salt tolerance in rice.Plant Science,2016,247:104-14.
[17]孙建,乐美旺,饶月亮,颜廷献,颜小文,周红英.芽期Na Cl胁迫下不同基因型芝麻的反应差异研究.干旱地区农业研究,2014,32(6):75-81.SUN J,LE M W,RAO Y L,YAN T X,YAN X W,ZHOU H Y.Genotypic variation in sesame salt tolerance during germination under Na Cl stress.Agricultural Research in the Arid Areas,2014,32(6):75-81.(in Chinese)
[18]徐芬芬,杜佳朋.干旱胁迫和盐胁迫对芝麻种子萌发的影响.种子,2013,32(11):85-86.XU F F,DU J P.Effect of drought and salt stress on seed germination of sesame.Seed,2013,32(11):85-86.(in Chinese)
[19]DIAS A S,LIMA G S D,GHEYI H R,NOBRE R G,SANTOS J B D.Emergence,growth and production of sesame under salt stress and proportions of nitrate and ammonium.Revista Caatinga,2017,30(2):458-467.
[20]FARISSI M,BOUIZGAREN A,FAGHIRE M,BARGAZ A,GHOULAM C.Agro-physiological and biochemical properties associated with adaptation of Medicago sativa populations to water deficit.Doga Turkish Journal of Botany,2013,37(6):1166-1175.
[21]吴兰荣,陈静,许婷婷,苗华荣,胡文广,禹山林.花生全生育期耐盐鉴定研究.花生学报,2005(1):20-24.WU L R,CHEN J,XU T T,MIAO H R,HU W G,YU S L.Identification of salt tolerance in peanut growth duration.Journal of Peanut Science,2005(1):20-24.(in Chinese)
[22]支巨振,毕辛华,杜克敏,常秀兰,杨淑惠,任淑萍,吴志行,李仁凤,赵菊英.GB/T 3543.4-1995,农作物种子检验规程发芽试验.北京:中国标准出版社,1995.ZHI J Z,BI X H,DU K M,CHANG X L,YANG S H,REN S P,WU Z H,LI R F,ZHAO J Y.GB/T 3543.4-1995,Rules for agricultural seed testing-Germination test.Beijing:China Standard Press,1995.(in Chinese)
[23]刘文萍,吕伟,黎冬华,任果香,张艳欣,文飞,韩俊梅,张秀荣.芝麻种质资源成株期抗旱性关联分析.中国农业科学,2017,50(4):625-639.LIU W P,LüW,LI D H,REN G X,ZHANG Y X,WEN F,HAN J M,ZHANG X R.Drought resistance of sesame germplasm resources and association analysis at adult stage.Scientia Agricultura Sinica,2017,50(4):625-639.(in Chinese)
[24]WEI X,LIU K,ZHANG Y,FENG Q,WANG L,ZHAO Y,LI D,ZHAO Q,ZHU X,ZHU X,LI W,FAN D,GAO Y,LU Y,ZHANG X,TANG X,ZHOU C,ZHU C,LIU L,ZHONG R,TIAN Q,WEN Z,WENG Q,HAN B,HUANG X,ZHANG X.Genetic discovery for oil production and quality in sesame.Nature Communications,2015,6:8609.
[25]BRADBURY P J,ZHANG Z,KROON D E,CASSTEVENS T M,RAMDOSS Y,BUCKLER E S.TASSEL:Software for association mapping of complex traits in diverse samples.Bioinformatics,2007,23(19):2633-2635.
[26]WEI L,MIAO H,ZHAO R,HAN X,ZHANG T,ZHANG H.Identification and testing of reference genes for Sesame gene expression analysis by quantitative real-time PCR.Planta,2013,237(3):873-889.
[27]SéNéCHAL F,WATTIER C,RUSTéRUCCI C,PELLOUX J.Homogalacturonan-modifying enzymes:Structure,expression,and roles in plants.Journal of Experimental Botany,2014,65(18):5125.
[28]OROZCOCáRDENAS M,NARVáEZVáSQUEZ J,RYAN C.Hydrogen peroxide acts as a second messenger for the induction of defense genes in tomato plants in response to wounding,systemin,and methyl jasmonate.The Plant Cell,2001,13(1):179-191.
[29]寇晓虹.番茄多聚半乳糖醛酸酶(PG)反义基因表达及相关功能研究[D].北京:中国农业大学,2003.KOU X H.Antisense gene expression and gene function of polygalacturonase(PG)in tomato[D].Beijing:China Agricultural University,2003.(in Chinese)
[30]O’DONNELL P J,TRUESDALE M R,CALVERT C M,DORANS A,ROBERTS M R,BOWLES D J.A novel tomato gene that rapidly responds to wound-and pathogen-related signals.The Plant Journal,1998,14(1):137-142.
[31]ROBERTS M R,WARNER S A J,DARBY R,LIM E K,DRAPER J,BOWLES D J.Differential regulation of a glucosyl transferase gene homologue during defence responses in tobacco.Journal of Experimental Botany,1999,50(332):407-410.
[32]李莹,柳参奎.红素氧还蛋白研究进展.草业科学,2016,33(6):1118-1125.LI Y,LIU S K.Research progress on rubredoxin.Pratacultural science,2016,33(6):1118-1125.(in Chinese)
[33]林凡云,陆琼娴,徐剑宏,史建荣.两个与盐和赤霉病菌胁迫相关的小麦糖基转移酶基因的克隆与表达.遗传,2008,30(12):1608-1614.LIN F Y,LU Q X,XU J H,SHI J R.Cloning and expression analysis of two salt and Fusarium graminearum stress associated UDPglucosyltransferases genes in wheat.Hereditas,2008,30(12):1608-1614.(in Chinese)
[34]ZHU J,FU X,KOO Y D,ZHU J K,FRANCIS E J J,MICHAEL W W A,ZHU Y,SHI H,YUN D J,HASEGAWA P M,BRESSAN R A.An enhancer mutant of arabidopsis salt overly sensitive 3 mediates both ion homeostasis and the oxidative stress response.Molecular&Cellular Biology,2007,27(14):5214.
[35]SAMANANI N,LISCOMBE D K,FACCHINI P J.Molecular cloning and characterization of norcoclaurine synthase,an enzyme catalyzing the first committed step in benzylisoquinoline alkaloid biosynthesis.The Plant Journal,2004,40(2):302-313.