番茄果实品质形成及其分子机理研究进展
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摘要
番茄营养丰富、风味独特,是世界上消耗量最大的蔬菜之一,近年来人们对番茄品质有着越来越高的要求。影响番茄品质的化学组分包括糖、有机酸、VC、类胡萝卜素等代谢物,它们决定番茄果实独特的风味、营养和外观品质,影响番茄产品的商品性。本文论述了这些物质的代谢途径及其内在调控机制,旨在为番茄品质改良提供新的思路和方法。
Tomato is rich in nutrition and unique in flavor.It is one of the largest consumption vegetables in the world.Recently,people have increasing requirements on tomato quality.Chemical components affecting tomato quality include sugar,organic acids,vitamin C and carotenoids,etc. metabolites,determining the unique flavor,nutrients and appearance quality of tomato fruits and affecting the commodity quality of tomato products.This paper expounded the metabolic pathway and internal regulatory mechanism of these metabolites,aiming at providing new ideas and methods for improving tomato quality.
Tomato is rich in nutrition and unique in flavor.It is one of the largest consumption vegetables in the world.Recently,people have increasing requirements on tomato quality.Chemical components affecting tomato quality include sugar,organic acids,vitamin C and carotenoids,etc. metabolites,determining the unique flavor,nutrients and appearance quality of tomato fruits and affecting the commodity quality of tomato products.This paper expounded the metabolic pathway and internal regulatory mechanism of these metabolites,aiming at providing new ideas and methods for improving tomato quality.
引文
程孙亮,周宝利,马迎杰.2007.红果番茄果实成熟过程中类胡萝卜素含量动态变化研究.安徽农业科学,(22):6728-6750.
Acree T E,Teranishi R.1993.Flavor science:sensible principles and techniques.Washington,DC:American Chemical Society.
Agarwal S,Rao A V.2000.Tomato lycopene and its role in human health and chronic diseases.Canadian Medical Association Journal,163:739-744.
Agius F,Gonzalez-Lamothe R,Caballero J L,Munoz-Blanco J,Botella M A,Valpuesta V.2003.Engineering increased vitamin C levels in plants by overexpression of a D-galacturonic acid reductase.Nat Biotechnol,21:177-181.
Alseekh S,Tohge T,Wendenberg R,Scossa F,Omranian N,Li J,Kleessen S,Giavalisco P,Pleban T,Mueller-Roeber B,Zamir D,Nikoloski Z,Fernie A R.2015.Identification and mode of inheritance of quantitative trait loci for secondary metabolite abundance in tomato.Plant Cell,27:485-512.
Badejo A A,Wada K,Gao Y,Maruta T,Sawa Y,Shigeoka S,Ishikawa T.2012.Translocation and the alternative D-galacturonate pathway contribute to increasing the ascorbate level in ripening tomato fruits together with the D-mannose/L-galactose pathway.J Exp Bot,63:229-239.
Bahrami A R,Chen Z H,Walker R P,Leegood R C,Gray J E.2001.Ripening-related occurrence of phosphoenolpyruvate carboxykinase in tomato fruit.Plant Molecular Biology,47:499-506.
Bianchetti R E,Cruz A B,Oliveira B S,Demarco D,Purgatto E,Peres L E P,Rossi M,Freschi L.2017.Phytochromobilin deficiency impairs sugar metabolism through the regulation of cytokinin and auxin signaling in tomato fruits.Sci Rep,7:7822.
Bramley P,Teulieres C,Blain I,Bird C,Schuch W.1992.Biochemical-characterization of transgenic tomato plants in which carotenoid synthesis has been inhibited through the expression of Antisense RNA to pTOM5.Plant Journal,2:343-349.
Bucheli P,Voirol E,de la Torre R,López J,Rytz A,Tanksley S D,Pétiard V.1999.Definition of nonvolatile markers for flavor of tomato(Lycopersicon esculentum Mill.)as tools in selection and breeding.Journal of Agricultural and Food Chemistry,47:659-664.
Cai X,Zhang C,Ye J,Hu T,Ye Z,Li H,Zhang Y.2015.Ectopic expression of FaGalUR leads to ascorbate accumulation with enhanced oxidative stress,cold,and salt tolerance in tomato.Plant Growth Regulation,76:187-197.
Carrari F,Baxter C,Usadel B,Urbanczyk-Wochniak E,Zanor MI,Nunes-Nesi A,Nikiforova V,Centero D,Ratzka A,Pauly M.2006.Integrated analysis of metabolite and transcript levels reveals the metabolic shifts that underlie tomato fruit development and highlight regulatory aspects of metabolic network behavior.Plant Physiology,142:1380-1396.
Centeno D C,Osorio S,Nunes-Nesi A,Bertolo A L,Carneiro RT,Araújo W L,Steinhauser M C,Michalska J,Rohrmann J,Geigenberger P.2011.Malate plays a crucial role in starch metabolism,ripening,and soluble solid content of tomato fruit and affects postharvest softening.The Plant Cell,23:162-184.
Chamovitz D,Pecker I,Sandmann G,B?ger P,Hirschberg J.1990.Cloning a gene coding for norflurazon resistance in cyanobacteria.Zeitschrift für Naturforschung,45:482-486.
Chen Z,Gallie D R.2004.The ascorbic acid redox state controls guard cell signaling and stomatal movement.The Plant Cell,16:1143-1162.
Conklin P L,Saracco S A,Norris S R,Last R L.2000.Identification of ascorbic acid-deficient Arabidopsis thaliana mutants.Genetics,154:847-856.
Davies J,Maw G.1972.Metabolism of citric and malic acids during ripening of tomato fruit.Journal of the Science of Food and Agriculture,23:969-976.
Dogbo O,Laferriere A,D’Harlingue A,Camara B.1988.Carotenoid biosynthesis:isolation and characterization of a bifunctional enzyme catalyzing the synthesis of phytoene.PNAS,85:7054-7058.
Fridman E,Carrari F,Liu Y S,Fernie A R,Zamir D.2004.Zooming in on a quantitative trait for tomato yield using interspecific introgressions.Science,305:1786-1789.
Fulton T M,Bucheli P,Voirol E,Lopez J,Petiard V,Tanksley S D.2002.Quantitative trait loci(QTL)affecting sugars,organic acids and other biochemical properties possibly contributing to flavor,identified in four advanced backcross populations of tomato.Euphytica,127:163-177.
Gallie D R.2013.L-ascorbic acid:a multifunctional molecule supporting plant growth and development.Scientifica,2013:795-964.
Giovannoni J J.2007.Fruit ripening mutants yield insights into ripening control.Current Opinion In Plant Biology,10:283-289.
Hackel A,Schauer N,Carrari F,Fernie A R,Grimm B,Kuhn C.2006.Sucrose transporter LeSUT1 and LeSUT2 inhibition affects tomato fruit development in different ways.Plant J,45:180-192.
Hemavathi,Upadhyaya C P,Akula N,Kim H S,Jeon J H,Ho O M,Chun S C,Kim D H,Park S W.2011.Biochemical analysis of enhanced tolerance in transgenic potato plants overexpressing D-galacturonic acid reductase gene in response to various abiotic stresses.Molecular Breeding,28:105-115.
Hu T,Ye J,Tao P,Li H,Zhang J,Zhang Y,Ye Z.2016.The tomato HD-Zip I transcription factor SlHZ24 modulates ascorbate accumulation through positive regulation of the D-mannose/L-galactose pathway.Plant J,85:16-29.
Isaacson T,Ronen G,Zamir D,Hirschberg J.2002.Cloning of tangerine from tomato reveals a carotenoid isomerase essential for the production ofβ-carotene and xanthophylls in plants.The Plant Cell,14:333-342.
Klann E M,Hall B,Bennett A B.1996.Antisense acid invertase(TIV1)gene alters soluble sugar composition and size in transgenic tomato fruit.Plant Physiol,112:1321-1330.
Lambeth V N,Fields M L,Huecker D E.1964.The sugar-acid ratio of selected tomato varieties.Columbia:University of Missouri.
Levin I,Gilboa N,Yeselson E,Shen S,Schaffer A.2000.Fgr,a major locus that modulates the fructose to glucose ratio in mature tomato fruits.Theoretical and Applied Genetics,100:256-262.
Locato V,Cimini S,Gara L D.2013.Strategies to increase vitamin C in plants:from plant defense perspective to food biofortification.Frontiers in Plant Science,4:152.
Lorence A,Chevone B I,Mendes P,Nessler C L.2004.Myoinositol oxygenase offers a possible entry point into plant ascorbate biosynthesis.Plant Physiol,134:1200-1205.
Luo Z,Zhang J,Li J,Yang C,Wang T,Ouyang B,Li H,Giovannoni J,Ye Z.2013.A STAY-GREEN protein SlSGR1regulates lycopene and beta-carotene accumulation by interacting directly with SlPSY1 during ripening processes in tomato.New Phytol,198:442-452.
Martel C,Vrebalov J,Tafelmeyer P,Giovannoni J J.2011.The tomato MADS-box transcription factor RIPENING INHIBITORinteracts with promoters involved in numerous ripening processes in a COLORLESS NONRIPENING-dependent manner.Plant Physiology,157:1568-1579.
Mayne S T.1996.Beta-carotene,carotenoids,and disease prevention in humans.The FASEB Journal,10:690-701.
N’Tchobo H,Dali N,Nguyen-Quoc B,Foyer C H,Yelle S.1999.Starch synthesis in tomato remains constant throughout fruit development and is dependent on sucrose supply and sucrose synthase activity.Journal of Experimental Botany,50:1457-1463.
Osorio S,Vallarino J G,Szecowka M,Ufaz S,Tzin V,Angelovici R,Galili G,Fernie A R.2013.Alteration of the interconversion of pyruvate and malate in the plastid or cytosol of ripening tomato fruit invokes diverse consequences on sugar but similar effects on cellular organic acid,metabolism,and transitory starch accumulation.Plant Physiology,161:628-643.
Pandey A,Misra P,Choudhary D,Yadav R,Goel R,Bhambhani S,Sanyal I,Trivedi R,Trivedi P K.2015.At MYB12 expression in tomato leads to large scale differential modulation in transcriptome and flavonoid content in leaf and fruit tissues.Sci Rep,5:12412.
Penfield S,Clements S,Bailey K J,Gilday A D,Leegood R C,Gray J E,Graham I A.2012.Expression and manipulation of phosphoenolpyruvate carboxykinase 1 identifies a role for malate metabolism in stomatal closure.The Plant Journal,69:679-688.
Petreikov M,Shen S,Yeselson Y,Levin I,Bar M,Schaffer A A.2006.Temporally extended gene expression of the ADP-Glc pyrophosphorylase large subunit(AgpL1)leads to increased enzyme activity in developing tomato fruit.Planta,224:1465-1479.
Ruggieri V,Bostan H,Barone A,Frusciante L,Chiusano M L.2016.Integrated bioinformatics to decipher the ascorbic acid metabolic network in tomato.Plant Mol Biol,91:397-412.
Sagar M,Chervin C,Mila I,Hao Y,Roustan J P,Benichou M,Gibon Y,Biais B,Maury P,Latche A,Pech J C,Bouzayen M,Zouine M.2013.Sl ARF4,an auxin response factor involved in the control of sugar metabolism during tomato fruit development.Plant Physiol,161:1362-1374.
Smirnoff N,Wheeler G L.2000.Ascorbic acid in plants:biosynthesis and function.Critical Reviews In Biochemistry and Molecular Biology,35:291-314.
Tanksley S D,Ganal M W,Prince J P,de Vicente M C,Bonierbale M W,Broun P,Fulton T M,Giovannoni J J,Grandillo S,Martin G B.1992.High density molecular linkage maps of the tomato and potato genomes.Genetics,132:1141-1160.
Watanabe K,Suzuki K,Kitamura S.2006.Characterization of a GDP-D-mannose 3’,5’-epimerase from rice.Phytochemistry,67:338-346.
Wolucka B A,Persiau G,van Doorsselaere J,Davey M W,Demol H,Vandekerckhove J,van Montagu M,Zabeau M,Boerjan W.2001.Partial purification and identification of GDP-mannose 3’,5’-epimerase of Arabidopsis thaliana,a key enzyme of the plant vitamin C pathway.PNAS,98:14843-14848.
Xiong C,Luo D,Lin A,Zhang C,Shan L,He P,Li B,Zhang Q,Hua B,Yuan Z.2018.A tomato B-box protein Sl BBX20 modulates carotenoid biosynthesis by directly activating PHYTOENE SYNTHASE 1,and is targeted for 26S proteasomemediated degradation.New Phytologist,221:279-294.
Ye J,Wang X,Hu T,Zhang F,Wang B,Li C,Yang T,Li H,Lu Y,Giovannoni J J,Zhang Y,Ye Z.2017.An InDel in the promoter of Al-ACTIVATED MALATE TRANSPORTER9selected during tomato domestication determines fruit malate contents and aluminum tolerance.Plant Cell,29:2249-2268.
Yelle S,Chetelat R T,Dorais M,Deverna J W,Bennett A B.1991.Sink metabolism in tomato fruit:Ⅳ.Genetic and biochemical analysis of sucrose accumulation.Plant Physiol,95:1026-1035.
Zhang C,Liu J,Zhang Y,Cai X,Gong P,Zhang J,Wang T,Li H,Ye Z.2011.Overexpression of SlGMEs leads to ascorbate accumulation with enhanced oxidative stress,cold,and salt tolerance in tomato.Plant Cell Reports,30:389-398.
Zhu G,Wang S,Huang Z,Zhang S,Liao Q,Zhang C,Lin T,Qin M,Peng M,Yang C,Cao X,Han X,Wang X,van der Knaap E,Zhang Z,Cui X,Klee H,Fernie A R,Luo J,Huang S.2018.Rewiring of the fruit metabolome in tomato breeding.Cell,172:249-261.
Zou L,Li H,Ouyang B,Zhang J,Ye Z.2006.Cloning and mapping of genes involved in tomato ascorbic acid biosynthesis and metabolism.Plant Science,170:120-127.
Acree T E,Teranishi R.1993.Flavor science:sensible principles and techniques.Washington,DC:American Chemical Society.
Agarwal S,Rao A V.2000.Tomato lycopene and its role in human health and chronic diseases.Canadian Medical Association Journal,163:739-744.
Agius F,Gonzalez-Lamothe R,Caballero J L,Munoz-Blanco J,Botella M A,Valpuesta V.2003.Engineering increased vitamin C levels in plants by overexpression of a D-galacturonic acid reductase.Nat Biotechnol,21:177-181.
Alseekh S,Tohge T,Wendenberg R,Scossa F,Omranian N,Li J,Kleessen S,Giavalisco P,Pleban T,Mueller-Roeber B,Zamir D,Nikoloski Z,Fernie A R.2015.Identification and mode of inheritance of quantitative trait loci for secondary metabolite abundance in tomato.Plant Cell,27:485-512.
Badejo A A,Wada K,Gao Y,Maruta T,Sawa Y,Shigeoka S,Ishikawa T.2012.Translocation and the alternative D-galacturonate pathway contribute to increasing the ascorbate level in ripening tomato fruits together with the D-mannose/L-galactose pathway.J Exp Bot,63:229-239.
Bahrami A R,Chen Z H,Walker R P,Leegood R C,Gray J E.2001.Ripening-related occurrence of phosphoenolpyruvate carboxykinase in tomato fruit.Plant Molecular Biology,47:499-506.
Bianchetti R E,Cruz A B,Oliveira B S,Demarco D,Purgatto E,Peres L E P,Rossi M,Freschi L.2017.Phytochromobilin deficiency impairs sugar metabolism through the regulation of cytokinin and auxin signaling in tomato fruits.Sci Rep,7:7822.
Bramley P,Teulieres C,Blain I,Bird C,Schuch W.1992.Biochemical-characterization of transgenic tomato plants in which carotenoid synthesis has been inhibited through the expression of Antisense RNA to pTOM5.Plant Journal,2:343-349.
Bucheli P,Voirol E,de la Torre R,López J,Rytz A,Tanksley S D,Pétiard V.1999.Definition of nonvolatile markers for flavor of tomato(Lycopersicon esculentum Mill.)as tools in selection and breeding.Journal of Agricultural and Food Chemistry,47:659-664.
Cai X,Zhang C,Ye J,Hu T,Ye Z,Li H,Zhang Y.2015.Ectopic expression of FaGalUR leads to ascorbate accumulation with enhanced oxidative stress,cold,and salt tolerance in tomato.Plant Growth Regulation,76:187-197.
Carrari F,Baxter C,Usadel B,Urbanczyk-Wochniak E,Zanor MI,Nunes-Nesi A,Nikiforova V,Centero D,Ratzka A,Pauly M.2006.Integrated analysis of metabolite and transcript levels reveals the metabolic shifts that underlie tomato fruit development and highlight regulatory aspects of metabolic network behavior.Plant Physiology,142:1380-1396.
Centeno D C,Osorio S,Nunes-Nesi A,Bertolo A L,Carneiro RT,Araújo W L,Steinhauser M C,Michalska J,Rohrmann J,Geigenberger P.2011.Malate plays a crucial role in starch metabolism,ripening,and soluble solid content of tomato fruit and affects postharvest softening.The Plant Cell,23:162-184.
Chamovitz D,Pecker I,Sandmann G,B?ger P,Hirschberg J.1990.Cloning a gene coding for norflurazon resistance in cyanobacteria.Zeitschrift für Naturforschung,45:482-486.
Chen Z,Gallie D R.2004.The ascorbic acid redox state controls guard cell signaling and stomatal movement.The Plant Cell,16:1143-1162.
Conklin P L,Saracco S A,Norris S R,Last R L.2000.Identification of ascorbic acid-deficient Arabidopsis thaliana mutants.Genetics,154:847-856.
Davies J,Maw G.1972.Metabolism of citric and malic acids during ripening of tomato fruit.Journal of the Science of Food and Agriculture,23:969-976.
Dogbo O,Laferriere A,D’Harlingue A,Camara B.1988.Carotenoid biosynthesis:isolation and characterization of a bifunctional enzyme catalyzing the synthesis of phytoene.PNAS,85:7054-7058.
Fridman E,Carrari F,Liu Y S,Fernie A R,Zamir D.2004.Zooming in on a quantitative trait for tomato yield using interspecific introgressions.Science,305:1786-1789.
Fulton T M,Bucheli P,Voirol E,Lopez J,Petiard V,Tanksley S D.2002.Quantitative trait loci(QTL)affecting sugars,organic acids and other biochemical properties possibly contributing to flavor,identified in four advanced backcross populations of tomato.Euphytica,127:163-177.
Gallie D R.2013.L-ascorbic acid:a multifunctional molecule supporting plant growth and development.Scientifica,2013:795-964.
Giovannoni J J.2007.Fruit ripening mutants yield insights into ripening control.Current Opinion In Plant Biology,10:283-289.
Hackel A,Schauer N,Carrari F,Fernie A R,Grimm B,Kuhn C.2006.Sucrose transporter LeSUT1 and LeSUT2 inhibition affects tomato fruit development in different ways.Plant J,45:180-192.
Hemavathi,Upadhyaya C P,Akula N,Kim H S,Jeon J H,Ho O M,Chun S C,Kim D H,Park S W.2011.Biochemical analysis of enhanced tolerance in transgenic potato plants overexpressing D-galacturonic acid reductase gene in response to various abiotic stresses.Molecular Breeding,28:105-115.
Hu T,Ye J,Tao P,Li H,Zhang J,Zhang Y,Ye Z.2016.The tomato HD-Zip I transcription factor SlHZ24 modulates ascorbate accumulation through positive regulation of the D-mannose/L-galactose pathway.Plant J,85:16-29.
Isaacson T,Ronen G,Zamir D,Hirschberg J.2002.Cloning of tangerine from tomato reveals a carotenoid isomerase essential for the production ofβ-carotene and xanthophylls in plants.The Plant Cell,14:333-342.
Klann E M,Hall B,Bennett A B.1996.Antisense acid invertase(TIV1)gene alters soluble sugar composition and size in transgenic tomato fruit.Plant Physiol,112:1321-1330.
Lambeth V N,Fields M L,Huecker D E.1964.The sugar-acid ratio of selected tomato varieties.Columbia:University of Missouri.
Levin I,Gilboa N,Yeselson E,Shen S,Schaffer A.2000.Fgr,a major locus that modulates the fructose to glucose ratio in mature tomato fruits.Theoretical and Applied Genetics,100:256-262.
Locato V,Cimini S,Gara L D.2013.Strategies to increase vitamin C in plants:from plant defense perspective to food biofortification.Frontiers in Plant Science,4:152.
Lorence A,Chevone B I,Mendes P,Nessler C L.2004.Myoinositol oxygenase offers a possible entry point into plant ascorbate biosynthesis.Plant Physiol,134:1200-1205.
Luo Z,Zhang J,Li J,Yang C,Wang T,Ouyang B,Li H,Giovannoni J,Ye Z.2013.A STAY-GREEN protein SlSGR1regulates lycopene and beta-carotene accumulation by interacting directly with SlPSY1 during ripening processes in tomato.New Phytol,198:442-452.
Martel C,Vrebalov J,Tafelmeyer P,Giovannoni J J.2011.The tomato MADS-box transcription factor RIPENING INHIBITORinteracts with promoters involved in numerous ripening processes in a COLORLESS NONRIPENING-dependent manner.Plant Physiology,157:1568-1579.
Mayne S T.1996.Beta-carotene,carotenoids,and disease prevention in humans.The FASEB Journal,10:690-701.
N’Tchobo H,Dali N,Nguyen-Quoc B,Foyer C H,Yelle S.1999.Starch synthesis in tomato remains constant throughout fruit development and is dependent on sucrose supply and sucrose synthase activity.Journal of Experimental Botany,50:1457-1463.
Osorio S,Vallarino J G,Szecowka M,Ufaz S,Tzin V,Angelovici R,Galili G,Fernie A R.2013.Alteration of the interconversion of pyruvate and malate in the plastid or cytosol of ripening tomato fruit invokes diverse consequences on sugar but similar effects on cellular organic acid,metabolism,and transitory starch accumulation.Plant Physiology,161:628-643.
Pandey A,Misra P,Choudhary D,Yadav R,Goel R,Bhambhani S,Sanyal I,Trivedi R,Trivedi P K.2015.At MYB12 expression in tomato leads to large scale differential modulation in transcriptome and flavonoid content in leaf and fruit tissues.Sci Rep,5:12412.
Penfield S,Clements S,Bailey K J,Gilday A D,Leegood R C,Gray J E,Graham I A.2012.Expression and manipulation of phosphoenolpyruvate carboxykinase 1 identifies a role for malate metabolism in stomatal closure.The Plant Journal,69:679-688.
Petreikov M,Shen S,Yeselson Y,Levin I,Bar M,Schaffer A A.2006.Temporally extended gene expression of the ADP-Glc pyrophosphorylase large subunit(AgpL1)leads to increased enzyme activity in developing tomato fruit.Planta,224:1465-1479.
Ruggieri V,Bostan H,Barone A,Frusciante L,Chiusano M L.2016.Integrated bioinformatics to decipher the ascorbic acid metabolic network in tomato.Plant Mol Biol,91:397-412.
Sagar M,Chervin C,Mila I,Hao Y,Roustan J P,Benichou M,Gibon Y,Biais B,Maury P,Latche A,Pech J C,Bouzayen M,Zouine M.2013.Sl ARF4,an auxin response factor involved in the control of sugar metabolism during tomato fruit development.Plant Physiol,161:1362-1374.
Smirnoff N,Wheeler G L.2000.Ascorbic acid in plants:biosynthesis and function.Critical Reviews In Biochemistry and Molecular Biology,35:291-314.
Tanksley S D,Ganal M W,Prince J P,de Vicente M C,Bonierbale M W,Broun P,Fulton T M,Giovannoni J J,Grandillo S,Martin G B.1992.High density molecular linkage maps of the tomato and potato genomes.Genetics,132:1141-1160.
Watanabe K,Suzuki K,Kitamura S.2006.Characterization of a GDP-D-mannose 3’,5’-epimerase from rice.Phytochemistry,67:338-346.
Wolucka B A,Persiau G,van Doorsselaere J,Davey M W,Demol H,Vandekerckhove J,van Montagu M,Zabeau M,Boerjan W.2001.Partial purification and identification of GDP-mannose 3’,5’-epimerase of Arabidopsis thaliana,a key enzyme of the plant vitamin C pathway.PNAS,98:14843-14848.
Xiong C,Luo D,Lin A,Zhang C,Shan L,He P,Li B,Zhang Q,Hua B,Yuan Z.2018.A tomato B-box protein Sl BBX20 modulates carotenoid biosynthesis by directly activating PHYTOENE SYNTHASE 1,and is targeted for 26S proteasomemediated degradation.New Phytologist,221:279-294.
Ye J,Wang X,Hu T,Zhang F,Wang B,Li C,Yang T,Li H,Lu Y,Giovannoni J J,Zhang Y,Ye Z.2017.An InDel in the promoter of Al-ACTIVATED MALATE TRANSPORTER9selected during tomato domestication determines fruit malate contents and aluminum tolerance.Plant Cell,29:2249-2268.
Yelle S,Chetelat R T,Dorais M,Deverna J W,Bennett A B.1991.Sink metabolism in tomato fruit:Ⅳ.Genetic and biochemical analysis of sucrose accumulation.Plant Physiol,95:1026-1035.
Zhang C,Liu J,Zhang Y,Cai X,Gong P,Zhang J,Wang T,Li H,Ye Z.2011.Overexpression of SlGMEs leads to ascorbate accumulation with enhanced oxidative stress,cold,and salt tolerance in tomato.Plant Cell Reports,30:389-398.
Zhu G,Wang S,Huang Z,Zhang S,Liao Q,Zhang C,Lin T,Qin M,Peng M,Yang C,Cao X,Han X,Wang X,van der Knaap E,Zhang Z,Cui X,Klee H,Fernie A R,Luo J,Huang S.2018.Rewiring of the fruit metabolome in tomato breeding.Cell,172:249-261.
Zou L,Li H,Ouyang B,Zhang J,Ye Z.2006.Cloning and mapping of genes involved in tomato ascorbic acid biosynthesis and metabolism.Plant Science,170:120-127.