SlYABBY2b基因对番茄果实心室数和内源赤霉素含量的影响（英文）
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摘要
目的:探讨赤霉素对SlYABBY2b基因调控番茄心室形成过程中的作用,为进一步研究番茄畸形果发生机理提供了理论基础。创新点:首次明确了SlYABBY2b基因与赤霉素的关系,且筛选出SlYABBY2b调控赤霉素合成的关键基因方法:利用Gateway技术法构建SlYABBY2b基因超表达和沉默载体,并通过农杆菌介导转化法获得转基因植株。用酶联免疫吸附测定(ELISA)试剂盒检测转基因植株中赤霉素的含量,用实时荧光定量分析(qRT-PCR)赤霉素突变体中SlYABBY2b基因表达水平和SlYABBY2b转基因植株中赤霉素相关基因的转录水平。结论:本实验中赤霉素突变体中SlYABBY2b基因表达量显示,赤霉素短缺导致番茄植株体内SlYABBY2b基因的升高。ELISA实验结果显示,SlYABBY2b基因也能够反馈调控赤霉素的合成。qRT-PCR结果显示,SlYABBY2b基因抑制GA20ox1和GA3ox2基因的表达,促进GA2ox1和GA2ox5基因的表达。综上所述,研究结果明确了SlYABBY2b基因位于赤霉素信号转导的途径上,反馈调节赤霉素的合成,感受外源赤霉素的信号,从而进一步调控番茄心室的形成。
Tomato is an ideal model species for fleshy fruit development research. Sl YABBY2 b regulates the ovary locule number, which is increased by gibberellins, in tomato. However, the relationship between Sl YABBY2 b and endogenous gibberellin is poorly understood. In this study, Sl YABBY2 b-overexpressing and RNA interference(RNAi) transgenic tomato plants were used to elucidate the mechanism by which Sl YABBY2 b regulates the ovary locule number and endogenous gibberellin content in tomato. Sl YABBY2 b-overexpressing plants showed fewer locules and lower gibberellin content than the control plants. Contrasting results were found in the RNAi lines. Therefore, the Sl YABBY2 b gene negatively regulates tomato ovary locule number and endogenous gibberellin content. Furthermore, the expression of Sl YABBY2 b gene was remarkably higher than that of the wild type in the apical shoots of gibberellindeficient mutants. This showed that the gibberellins can inhibit the expression of Sl YABBY2 b gene negative regulation. Further study revealed that Sl YABBY2 b suppressed the expression of Sl GA20 ox1 and Sl GA3 ox2, but increased that of Sl GA2 ox1 and Sl GA2 ox5 in the apical shoots of Sl YABBY2 b-overexpressing plants, thereby reducing gibberellin content. Contrasting results were found in the RNAi lines. Our results showed that the Sl YABBY2 b gene was located on gibberellin signal transduction pathways, fed back regulation of the synthesis of gibberellin, and felt exogenous gibberellin signal to further regulate the formation of tomato locule.
Tomato is an ideal model species for fleshy fruit development research. Sl YABBY2 b regulates the ovary locule number, which is increased by gibberellins, in tomato. However, the relationship between Sl YABBY2 b and endogenous gibberellin is poorly understood. In this study, Sl YABBY2 b-overexpressing and RNA interference(RNAi) transgenic tomato plants were used to elucidate the mechanism by which Sl YABBY2 b regulates the ovary locule number and endogenous gibberellin content in tomato. Sl YABBY2 b-overexpressing plants showed fewer locules and lower gibberellin content than the control plants. Contrasting results were found in the RNAi lines. Therefore, the Sl YABBY2 b gene negatively regulates tomato ovary locule number and endogenous gibberellin content. Furthermore, the expression of Sl YABBY2 b gene was remarkably higher than that of the wild type in the apical shoots of gibberellindeficient mutants. This showed that the gibberellins can inhibit the expression of Sl YABBY2 b gene negative regulation. Further study revealed that Sl YABBY2 b suppressed the expression of Sl GA20 ox1 and Sl GA3 ox2, but increased that of Sl GA2 ox1 and Sl GA2 ox5 in the apical shoots of Sl YABBY2 b-overexpressing plants, thereby reducing gibberellin content. Contrasting results were found in the RNAi lines. Our results showed that the Sl YABBY2 b gene was located on gibberellin signal transduction pathways, fed back regulation of the synthesis of gibberellin, and felt exogenous gibberellin signal to further regulate the formation of tomato locule.
引文
Asahira T,Hosoki T,Shinya K,1982.Regulation of low temperature-induced malformation of tomato fruit by plant growth regulators.J Jpn Soc Hort Sci,50(4):468-474.https://doi.org/10.2503/jjshs.50.468
Barrero LS,Tanksley SD,2004.Evaluating the genetic basis of multiple-locule fruit in a broad cross section of tomato cultivars.Theor Appl Genet,109(3):669-679.https://doi.org/10.1007/s00122-004-1676-y
Barrero LS,Cong B,Wu F,et al.,2006.Developmental characterization of the fasciated locus and mapping of Arabidopsis candidate genes involved in the control of floral meristem size and carpel number in tomato.Genome,49(8):991-1006.https://doi.org/10.1139/g06-059
Chakrabarti M,Zhang N,Sauvage C,et al.,2013.A cytochrome P450 regulates a domestication trait in cultivated tomato.Proc Natl Acad Sci USA,110(42):17125-17130.https://doi.org/10.1073/pnas.1307313110
Chen JG,Zhou X,Zhang YZ,1998.Gibberellin-responding and non-responding dwarf mutants in foxtail millet.Plant Growth Reg,26(1):19-24.https://doi.org/10.1023/A:100609160
Cong B,Barrero LS,Tanksley SD,2008.Regulatory change in YABBY-like transcription factor led to evolution of extreme fruit size during tomato domestication.Nat Genet,40(6):800-804.https://doi.org/10.1038/ng.144
Dai M,Zhao Y,Ma Q,et al.,2007.The rice YABBY1 gene is involved in the feedback regulation of gibberellin metabolism.Plant Physiol,144(1):121-133.https://doi.org/10.1104/pp.107.096586
Fernández-Lozano A,Yuste-Lisbona FJ,Pérez-Martín F,et al.,2015.Mutation at the tomato EXCESSIVE NUMBER OFFLORAL ORGANS(ENO)locus impairs floral meristem development,thus promoting an increased number of floral organs and fruit size.Plant Sci,232:41-48.https://doi.org/10.1016/j.plantsci.2014.12.007
Foolad MR,2007.Genome mapping and molecular breeding of tomato.Int J Plant Genomics,2007:64358.https://doi.org/10.1155/2007/64358
Fukazawa J,Sakai T,Ishida S,et al.,2000.REPRESSION OFSHOOT GROWTH,a b ZIP transcriptional activator,regulates cell elongation by controlling the level of gibberellins.Plant Cell,12(6):901-915.https://doi.org/10.1105/tpc.12.6.901
Gazzarrini S,Tsuchiya Y,Lumba S,et al.,2004.The transcription factor FUSCA3 controls developmental timing in Arabidopsis through the hormones gibberellin and abscisic acid.Dev Cell,7(3):373-385.https://doi.org/10.1016/j.devcel.2004.06.017
Hay A,Kaur H,Phillips A,et al.,2002.The gibberellin pathway mediates KNOTTED1-type homeobox function in plants with different body plans.Curr Biol,12(18):1557-1565.https://doi.org/10.1016/S0960-9822(02)01125-9
Hedden P,Phillips AL,2000.Gibberellin metabolism:new insights revealed by the genes.Trends Plant Sci,5(12):523-530.https://doi.org/10.1016/S1360-1385(00)01790-8
Hernández-Bautista A,Lobato-Ortiz R,Cruz-Izquierdo S,et al.,2015.Fruit size QTLs affect in a major proportion the yield in tomato.Chilean J Agric Res,75(4):402-409.https://doi.org/10.4067/S0718-58392015000500004
Huang Z,van der Knaap E,2011.Tomato fruit weight 11.3maps close to fasciated on the bottom of chromosome 11.Theor Appl Genet,123(3):465-474.https://doi.org/10.1007/s00122-011-1599-3
Huang Z,Houten JV,Gonzalez G,et al.,2013.Genome-wide identification,phylogeny and expression analysis of SUN,OFP and YABBY gene family in tomato.Mol Genet Genomics,288(3-4):111-129.https://doi.org/10.1007/s00438-013-0733-0
Illa-Berenguer E,van Houten J,Huang Z,et al.,2015.Rapid and reliable identification of tomato fruit weight and locule number loci by QTL-seq.Theor Appl Genet,128(7):1329-1342.https://doi.org/10.1007/s00122-015-2509-x
Imai R,Yang YY,Tahar AA,et al.,1996.Cloning and light-regulated expression of two c DNAs for ent-kaurene synthase A from tomato.Pant Cell Physiol,37:143.
Ishida S,Fukazawa J,Yuasa T,et al.,2004.Involvement of14-3-3 signaling protein binding in the functional regulation of the transcriptional activator REPRESSION OFSHOOT GROWTH by gibberellins.Plant Cell,16(10):2641-2651.https://doi.org/10.1105/tpc.104.024604
Jain M,Kaur N,Tyagi AK,et al.,2006.The auxin-responsive GH3 gene family in rice(Oryza sativa).Funct Integr Genomics,6(1):36-46.https://doi.org/10.1007/s10142-005-0142-5
Kumar R,Khurana A,Sharma AK,2014.Role of plant hormones and their interplay in development and ripening of fleshy fruits.J Exp Bot,65(16):4561-4575.https://doi.org/10.1093/jxb/eru277
Kumaran MK,Bowman JL,Sundaresan V,2002.YABBY polarity genes mediate the repression of KNOX homeobox genes in Arabidopsis.Plant Cell,14(11):2761-2770.https://doi.org/10.1105/tpc.004911
Li H,Qi MF,Sun MH,et al.,2017.Tomato transcription factor Sl WUS plays an important role in tomato flower and locule development.Front Plant Sci,8:457.https://doi.org/10.3389/fpls.2017.00457
Li Y,Li TL,Wang D,2008.Correlation between endogenous hormones of stem apices and fruit locule numbers in tomatoes during floral bud differentiation stages.Agric Sci China,7(4):447-454.https://doi.org/10.1016/S1671-2927(08)60088-7
Lippman Z,Tanksley SD,2001.Dissecting the genetic pathway to extreme fruit size in tomato using a cross between the small-fruited wild species Lycopersicon pimpinellifolium and L.esculentum var.Giant Heirloom.Genetics,158(1):413-422.
Liu S,Li TL,2012.Regulation effects of exogenous gibberellin acid(GA3)on the formation of tomato(Solanum lycoperscium)ovary locule and fasciated transcription.Arf J Biotechnol,11(72):13732-13738.https://doi.org/10.5897/AJB12.2244
Magome H,Yamaguchi S,Hanada A,et al.,2004.dwarf and delayed-flowering 1,a novel Arabidopsis mutant deficient in gibberellin biosynthesis because of overexpression of a putative AP2 transcription factor.Plant J,37(5):720-729.https://doi.org/10.1111/j.1365-313X.2003.01998.x
Mu?os S,Ranc N,Botton E,et al.,2011.Increase in tomato locule number is controlled by two single-nucleotide polymorphisms located near WUSCHEL.Plant Physiol,156(4):2244-2254.https://doi.org/10.1104/pp.111.173997
Olszewski N,Sun TP,Gubler F,2002.Gibberellin signaling biosynthesis,catabolism,and response pathways.Plant Cell,14(Suppl 1):S61-S80.https://doi.org/10.1105/tpc.010476
Pesaresi P,Mizzotti C,Colombo M,et al.,2014.Genetic regulation and structural changes during tomato fruit development and ripening.Front Plant Sci,5:124.https://doi.org/10.3389/fpls.2014.00124
Rebers M,Kaneta T,Kawaide H,et al.,1999.Regulation of gibberellin biosynthesis genes during flower and early fruit development of tomato.Pant J,17(3):241-250.
Rodríguez GR,Mu?os S,Anderson C,et al.,2011.Distribution of SUN,OVATE,LC,and FAS in the tomato germplasm and the relationship to fruit shape diversity.Plant Physiol,156(1):275-285.https://doi.org/10.1104/pp.110.167577
Rosin FM,Hart JK,Horner HT,et al.,2003.Overexpression of a knotted-like homeobox gene of potato alters vegetative development by decreasing gibberellin accumulation.Plant Physiol,132(1):106-117.
Sakamoto T,Miura K,Itoh H,et al.,2004.An overview of gibberellin metabolism enzyme genes and their related mutants in rice.Plant Physiol,134(4):1642-1653.https://doi.org/10.1104/pp.103.033696
Sawhney V,Greyson R,1971.Induction of multilocular ovary in tomato by gibberellic acid.J Am Soc Hort Sci,96(2):196-198.
Sawhney VK,Dabbs DH,1978.Gibberellic acid induced multilocular fruits in tomato and the role of locule number and seed number in fruit size.Can J Bot,56(22):2831-2835.https://doi.org/10.1139/b78-339
Schoof H,Lenhard M,Haecker A,et al.,2000.The stem cell population of Arabidopsis shoot meristems is maintained by a regulatory loop between the CLAVATA and WUSCHEL genes.Cell,100(6):635-644.https://doi.org/10.1016/S0092-8674(00)80700-X
Serrani JC,Sanjuán R,Ruiz-Rivero O,et al.,2007.Gibberellin regulation of fruit set and growth in tomato.Plant Physiol,145(1):246-257.https://doi.org/10.1104/pp.107.098335
Serrani JC,Ruiz-Rivero O,Fos M,et al.,2008.Auxin-induced fruit-set in tomato is mediated in part by gibberellins.Plant J,56(6):922-934.https://doi.org/10.1111/j.1365-313X.2008.03654.x
Seymour GB,?stergaard L,Chapman NH,et al.,2013.Fruit development and ripening.Annu Rev Plant Biol,64(1):219-241.https://doi.org/10.1146/annurev-arplant-050312-120057
Sponsel VM,Hedden P,2010.Gibberellin biosynthesis and inactivation.In:Davies PJ(Ed.),Plant Hormones.Springer,Dordrecht,p.63-94.https://doi.org/10.1007/978-1-4020-2686-7_4
Tanksley SD,2004.The genetic,developmental,and molecular bases of fruit size and shape variation in tomato.Plant Cell,16(Suppl_1):S181-S189.https://doi.org/10.1105/tpc.018119
Tomer E,Moshkovits H,Rosenfeld K,et al.,1998.Varietal differences in the susceptibility to pointed fruit malformation in tomatoes:histological studies of the ovaries.Sci Hortic,77(3-4):145-154.https://doi.org/10.1016/S0304-4238(98)00170-8
van der Knaap E,Chakrabarti M,Chu YH,et al.,2014.What lies beyond the eye:the molecular mechanisms regulating tomato fruit weight and shape.Front Plant Sci,5:227.https://doi.org/10.3389/fpls.2014.00227
Wang H,Caruso LV,Downie AB,et al.,2004.The embryo MADS domain protein AGAMOUS-Like 15 directly regulates expression of a gene encoding an enzyme involved in gibberellin metabolism.Plant Cell,16(5):1206-1219.https://doi.org/10.1105/tpc.021261
Xu C,Liberatore KL,Macalister CA,et al.,2015.A cascade of arabinosyltransferases controls shoot meristem size in tomato.Nat Genet,47(7):784-792.https://doi.org/10.1038/ng.3309
Yamaguchi S,2008.Gibberellin metabolism and its regulation.Annu Rev Plant Biol,59(1):225-251.https://doi.org/10.1146/annurev.arplant.59.032607.092804
Zheng Z,Zhou X,1995.A monoclonal antibody recognizing nonderivative 13-hydroxy gibberellins and their glucosides.Acta Bot Sin,37(10):761-769(in Chinese).
Barrero LS,Tanksley SD,2004.Evaluating the genetic basis of multiple-locule fruit in a broad cross section of tomato cultivars.Theor Appl Genet,109(3):669-679.https://doi.org/10.1007/s00122-004-1676-y
Barrero LS,Cong B,Wu F,et al.,2006.Developmental characterization of the fasciated locus and mapping of Arabidopsis candidate genes involved in the control of floral meristem size and carpel number in tomato.Genome,49(8):991-1006.https://doi.org/10.1139/g06-059
Chakrabarti M,Zhang N,Sauvage C,et al.,2013.A cytochrome P450 regulates a domestication trait in cultivated tomato.Proc Natl Acad Sci USA,110(42):17125-17130.https://doi.org/10.1073/pnas.1307313110
Chen JG,Zhou X,Zhang YZ,1998.Gibberellin-responding and non-responding dwarf mutants in foxtail millet.Plant Growth Reg,26(1):19-24.https://doi.org/10.1023/A:100609160
Cong B,Barrero LS,Tanksley SD,2008.Regulatory change in YABBY-like transcription factor led to evolution of extreme fruit size during tomato domestication.Nat Genet,40(6):800-804.https://doi.org/10.1038/ng.144
Dai M,Zhao Y,Ma Q,et al.,2007.The rice YABBY1 gene is involved in the feedback regulation of gibberellin metabolism.Plant Physiol,144(1):121-133.https://doi.org/10.1104/pp.107.096586
Fernández-Lozano A,Yuste-Lisbona FJ,Pérez-Martín F,et al.,2015.Mutation at the tomato EXCESSIVE NUMBER OFFLORAL ORGANS(ENO)locus impairs floral meristem development,thus promoting an increased number of floral organs and fruit size.Plant Sci,232:41-48.https://doi.org/10.1016/j.plantsci.2014.12.007
Foolad MR,2007.Genome mapping and molecular breeding of tomato.Int J Plant Genomics,2007:64358.https://doi.org/10.1155/2007/64358
Fukazawa J,Sakai T,Ishida S,et al.,2000.REPRESSION OFSHOOT GROWTH,a b ZIP transcriptional activator,regulates cell elongation by controlling the level of gibberellins.Plant Cell,12(6):901-915.https://doi.org/10.1105/tpc.12.6.901
Gazzarrini S,Tsuchiya Y,Lumba S,et al.,2004.The transcription factor FUSCA3 controls developmental timing in Arabidopsis through the hormones gibberellin and abscisic acid.Dev Cell,7(3):373-385.https://doi.org/10.1016/j.devcel.2004.06.017
Hay A,Kaur H,Phillips A,et al.,2002.The gibberellin pathway mediates KNOTTED1-type homeobox function in plants with different body plans.Curr Biol,12(18):1557-1565.https://doi.org/10.1016/S0960-9822(02)01125-9
Hedden P,Phillips AL,2000.Gibberellin metabolism:new insights revealed by the genes.Trends Plant Sci,5(12):523-530.https://doi.org/10.1016/S1360-1385(00)01790-8
Hernández-Bautista A,Lobato-Ortiz R,Cruz-Izquierdo S,et al.,2015.Fruit size QTLs affect in a major proportion the yield in tomato.Chilean J Agric Res,75(4):402-409.https://doi.org/10.4067/S0718-58392015000500004
Huang Z,van der Knaap E,2011.Tomato fruit weight 11.3maps close to fasciated on the bottom of chromosome 11.Theor Appl Genet,123(3):465-474.https://doi.org/10.1007/s00122-011-1599-3
Huang Z,Houten JV,Gonzalez G,et al.,2013.Genome-wide identification,phylogeny and expression analysis of SUN,OFP and YABBY gene family in tomato.Mol Genet Genomics,288(3-4):111-129.https://doi.org/10.1007/s00438-013-0733-0
Illa-Berenguer E,van Houten J,Huang Z,et al.,2015.Rapid and reliable identification of tomato fruit weight and locule number loci by QTL-seq.Theor Appl Genet,128(7):1329-1342.https://doi.org/10.1007/s00122-015-2509-x
Imai R,Yang YY,Tahar AA,et al.,1996.Cloning and light-regulated expression of two c DNAs for ent-kaurene synthase A from tomato.Pant Cell Physiol,37:143.
Ishida S,Fukazawa J,Yuasa T,et al.,2004.Involvement of14-3-3 signaling protein binding in the functional regulation of the transcriptional activator REPRESSION OFSHOOT GROWTH by gibberellins.Plant Cell,16(10):2641-2651.https://doi.org/10.1105/tpc.104.024604
Jain M,Kaur N,Tyagi AK,et al.,2006.The auxin-responsive GH3 gene family in rice(Oryza sativa).Funct Integr Genomics,6(1):36-46.https://doi.org/10.1007/s10142-005-0142-5
Kumar R,Khurana A,Sharma AK,2014.Role of plant hormones and their interplay in development and ripening of fleshy fruits.J Exp Bot,65(16):4561-4575.https://doi.org/10.1093/jxb/eru277
Kumaran MK,Bowman JL,Sundaresan V,2002.YABBY polarity genes mediate the repression of KNOX homeobox genes in Arabidopsis.Plant Cell,14(11):2761-2770.https://doi.org/10.1105/tpc.004911
Li H,Qi MF,Sun MH,et al.,2017.Tomato transcription factor Sl WUS plays an important role in tomato flower and locule development.Front Plant Sci,8:457.https://doi.org/10.3389/fpls.2017.00457
Li Y,Li TL,Wang D,2008.Correlation between endogenous hormones of stem apices and fruit locule numbers in tomatoes during floral bud differentiation stages.Agric Sci China,7(4):447-454.https://doi.org/10.1016/S1671-2927(08)60088-7
Lippman Z,Tanksley SD,2001.Dissecting the genetic pathway to extreme fruit size in tomato using a cross between the small-fruited wild species Lycopersicon pimpinellifolium and L.esculentum var.Giant Heirloom.Genetics,158(1):413-422.
Liu S,Li TL,2012.Regulation effects of exogenous gibberellin acid(GA3)on the formation of tomato(Solanum lycoperscium)ovary locule and fasciated transcription.Arf J Biotechnol,11(72):13732-13738.https://doi.org/10.5897/AJB12.2244
Magome H,Yamaguchi S,Hanada A,et al.,2004.dwarf and delayed-flowering 1,a novel Arabidopsis mutant deficient in gibberellin biosynthesis because of overexpression of a putative AP2 transcription factor.Plant J,37(5):720-729.https://doi.org/10.1111/j.1365-313X.2003.01998.x
Mu?os S,Ranc N,Botton E,et al.,2011.Increase in tomato locule number is controlled by two single-nucleotide polymorphisms located near WUSCHEL.Plant Physiol,156(4):2244-2254.https://doi.org/10.1104/pp.111.173997
Olszewski N,Sun TP,Gubler F,2002.Gibberellin signaling biosynthesis,catabolism,and response pathways.Plant Cell,14(Suppl 1):S61-S80.https://doi.org/10.1105/tpc.010476
Pesaresi P,Mizzotti C,Colombo M,et al.,2014.Genetic regulation and structural changes during tomato fruit development and ripening.Front Plant Sci,5:124.https://doi.org/10.3389/fpls.2014.00124
Rebers M,Kaneta T,Kawaide H,et al.,1999.Regulation of gibberellin biosynthesis genes during flower and early fruit development of tomato.Pant J,17(3):241-250.
Rodríguez GR,Mu?os S,Anderson C,et al.,2011.Distribution of SUN,OVATE,LC,and FAS in the tomato germplasm and the relationship to fruit shape diversity.Plant Physiol,156(1):275-285.https://doi.org/10.1104/pp.110.167577
Rosin FM,Hart JK,Horner HT,et al.,2003.Overexpression of a knotted-like homeobox gene of potato alters vegetative development by decreasing gibberellin accumulation.Plant Physiol,132(1):106-117.
Sakamoto T,Miura K,Itoh H,et al.,2004.An overview of gibberellin metabolism enzyme genes and their related mutants in rice.Plant Physiol,134(4):1642-1653.https://doi.org/10.1104/pp.103.033696
Sawhney V,Greyson R,1971.Induction of multilocular ovary in tomato by gibberellic acid.J Am Soc Hort Sci,96(2):196-198.
Sawhney VK,Dabbs DH,1978.Gibberellic acid induced multilocular fruits in tomato and the role of locule number and seed number in fruit size.Can J Bot,56(22):2831-2835.https://doi.org/10.1139/b78-339
Schoof H,Lenhard M,Haecker A,et al.,2000.The stem cell population of Arabidopsis shoot meristems is maintained by a regulatory loop between the CLAVATA and WUSCHEL genes.Cell,100(6):635-644.https://doi.org/10.1016/S0092-8674(00)80700-X
Serrani JC,Sanjuán R,Ruiz-Rivero O,et al.,2007.Gibberellin regulation of fruit set and growth in tomato.Plant Physiol,145(1):246-257.https://doi.org/10.1104/pp.107.098335
Serrani JC,Ruiz-Rivero O,Fos M,et al.,2008.Auxin-induced fruit-set in tomato is mediated in part by gibberellins.Plant J,56(6):922-934.https://doi.org/10.1111/j.1365-313X.2008.03654.x
Seymour GB,?stergaard L,Chapman NH,et al.,2013.Fruit development and ripening.Annu Rev Plant Biol,64(1):219-241.https://doi.org/10.1146/annurev-arplant-050312-120057
Sponsel VM,Hedden P,2010.Gibberellin biosynthesis and inactivation.In:Davies PJ(Ed.),Plant Hormones.Springer,Dordrecht,p.63-94.https://doi.org/10.1007/978-1-4020-2686-7_4
Tanksley SD,2004.The genetic,developmental,and molecular bases of fruit size and shape variation in tomato.Plant Cell,16(Suppl_1):S181-S189.https://doi.org/10.1105/tpc.018119
Tomer E,Moshkovits H,Rosenfeld K,et al.,1998.Varietal differences in the susceptibility to pointed fruit malformation in tomatoes:histological studies of the ovaries.Sci Hortic,77(3-4):145-154.https://doi.org/10.1016/S0304-4238(98)00170-8
van der Knaap E,Chakrabarti M,Chu YH,et al.,2014.What lies beyond the eye:the molecular mechanisms regulating tomato fruit weight and shape.Front Plant Sci,5:227.https://doi.org/10.3389/fpls.2014.00227
Wang H,Caruso LV,Downie AB,et al.,2004.The embryo MADS domain protein AGAMOUS-Like 15 directly regulates expression of a gene encoding an enzyme involved in gibberellin metabolism.Plant Cell,16(5):1206-1219.https://doi.org/10.1105/tpc.021261
Xu C,Liberatore KL,Macalister CA,et al.,2015.A cascade of arabinosyltransferases controls shoot meristem size in tomato.Nat Genet,47(7):784-792.https://doi.org/10.1038/ng.3309
Yamaguchi S,2008.Gibberellin metabolism and its regulation.Annu Rev Plant Biol,59(1):225-251.https://doi.org/10.1146/annurev.arplant.59.032607.092804
Zheng Z,Zhou X,1995.A monoclonal antibody recognizing nonderivative 13-hydroxy gibberellins and their glucosides.Acta Bot Sin,37(10):761-769(in Chinese).