Regulatory Genes for Tomato Flavor and Their Application in Quality Improvement

doi:10.16420/j.issn.0513-353x.2024-0614

Acta Horticulturae Sinica ›› 2025, Vol. 52 ›› Issue (8): 2099-2113.doi: 10.16420/j.issn.0513-353x.2024-0614

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Regulatory Genes for Tomato Flavor and Their Application in Quality Improvement

LI Dandan¹, GE Pingfei¹, LI Fangman¹, YANG Yang¹, XU Haobo¹, XIONG Chunhui³, and ZHANG Yuyang¹^,²^,⁴^,⁵^,^*()

¹ National Key Laboratory for Germplasm Innovation & Utilization of Horticultural Crops， Wuhan 430070， China
² Hubei Hongshan laboratory， Wuhan 430070， China
³ Ganzhou Vegetable and Flower Research Institute， Ganzhou， Jiangxi 341401， China
⁴ Shenzhen Institute of Nutrition and Health， Huazhong Agricultural University， Wuhan 430070， China
⁵ Agricultural Genomics Institute at Shenzhen， Chinese Academy of Agricultural Sciences（CAAS）and Lingnan Modern Agricultural Science and Technology Laboratory of Guangdong Province，Shenzhen Sub-center，Shenzhen， Guangdong 518800， China

Received:2024-10-25 Revised:2025-04-24 Online:2025-08-25 Published:2025-08-19
Contact: and ZHANG Yuyang

Abstract

Abstract:

Tomato is rich in nutrients and has a unique flavor，and in recent years，the improvement of tomato flavor quality has gradually attracted the attention of breeders. Modern cultivated tomatoes exhibit the advantages of high resistance and yield，but their taste and flavor fails to meet consumers’ demand. Soluble sugar，organic acid and volatile compounds in tomato are important components affecting tomato flavor. The type，content and proportion of sugars determine the sweetness of tomatoes，and the content of organic acids such as citric acid and malic acid affects the acidity of the fruit. The composition and concentration of volatile compounds in the fruit contributes to the aroma of tomatoes. This paper reviews the metabolic composition，genetic loci and regulatory genes of tomato flavor，summarizes the approaches of tomato fruit flavor improvement，to provide reference for breeding high-quality and tasty tomato cultivars.

Key words: tomato, flavor quality, soluble sugar, organic acid, volatile compound, genetic improvement

LI Dandan, GE Pingfei, LI Fangman, YANG Yang, XU Haobo, XIONG Chunhui, and ZHANG Yuyang. Regulatory Genes for Tomato Flavor and Their Application in Quality Improvement[J]. Acta Horticulturae Sinica, 2025, 52(8): 2099-2113.

Figures/Tables 3

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风味组分 Flavor component	基因 Gene	基因名称 Gene name	作用 Function	品质改良方法 Quality improvement method	背景材料 Background material	改良效果 Improving effect		参考文献 Reference
可溶性糖 Sugar	TIV1	反义蔗糖酶Antisense sucrase	调节果实中蔗糖积累 Regulates sucrose accumulation in fruits	反义RNA Antisense RNA	T5、H100	总糖含量提高约50% Total sugar increase by about 50%		Klann et al.,1996
	SlSWEET7a/14	糖转运蛋白 Sugar transport protein	参与果实中蔗糖、果糖、葡萄糖转运Involved in transport of sucrose，fructose and glucose in fruits	RNA干扰 RNAi	MicroTom	果糖、葡萄糖含量增加约30% ~ 60% Fructose and glucose content increased by about 30% to 60%		Zhang et al.,2021
	SlNAP2	NAC家族转录因子 NAC family transcription factors	参与调控植株叶片衰老 Involved in the regulation of plant leaf senescence	RNA干扰 RNAi	Moneymaker	糖含量增加约22.5% Sugar content increased by about 22.5%		Ma et al.,2018
	Lin5	细胞壁转化酶 Cell wall invertase	催化质外体中蔗糖裂解及其从番茄源到库供应 Catalyzing sucrose cleavage in apoplast and its supply from tomato sources to tank	超量表达 Overexpression	自交系FLA8059	葡萄糖、果糖含量提高约3.3% Glucose and fructose content increased by about 3.3%		Tieman et al.,2017
	SlVIF	液泡转化酶抑制剂 Vacuolar invertase inhibitor	抑制VIN活性，调控果实中蔗糖积累 Inhibition of VIN activity and regulation of sucrose accumulation in fruits	超量表达 Overexpression	Ailsa Craig	蔗糖含量提高10倍，己糖含量降低约40% Sucrose content increased by 10 times and hexose reduced by about 40%		Qin et al.,2016
	SlFgr	葡萄糖外排转移蛋白 Glucose exporter	影响果实中己糖的分配，改变果糖、葡萄糖比例 Influencing distribution of hexoses in the fruit，altering the ratio of fructose and glucose	超量表达 Overexpression	S. habrochaites	葡萄糖含量降低约41%，果糖葡萄糖比例提高约73% Glucose content reduced by about 41%，fructose-glucose ratio increased by about 73%		Shammai et al.,2018
	SlGLK2	MYB类转录因子 MYB-like transcription factors	调控果实中叶绿素含量 Regulation of chlorophyll content in fruits	超量表达 Overexpression	T63	葡萄糖、果糖含量提高约40% Glucose and fructose content increased by about 40%		Powell et al.,2012
	SlVPE5	番茄液泡加工酶 Tomato vacuolar processing enzyme	促进酸性转化酶合成，提高己糖水平 Promotes acid converting enzyme synthesis and increases hexose levels	CRISPR/Cas9	M82	葡萄糖、果糖含量增加约30% ~ 40% Glucose and fructose content increased by about 30% to 40%		Wang et al.,2021
	SlINVINH1	酸性转化酶抑制剂 Acid converting enzyme inhibitors	特异性抑制细胞壁转化酶活性 Specific inhibition of cell wall convertase activity	CRISPR/Cas9	M82	葡萄糖、果糖含量增加约35% ~ 45% Glucose and fructose content increased by about 35% to 45%		Wang et al.,2021
有机酸 Organic acids	SlAco	乌头酸酶 Aconitase	催化柠檬酸转化为异柠檬酸 Catalyzes the conversion of citric acid to isocitric acid	RNA干扰 RNAi	Moneymaker	柠檬酸含量提高40% 40% increase in citric acid content		Morgan et al.,2012
有机酸 Organic acids	SlPEPCK	番茄磷酸烯醇丙酮酸羧激酶 Tomato phosphoenolpyruvate carboxykinase	调节糖异生，提高糖酸比 Regulates gluconeogenesis and increases the sugar-acid ratio	RNA干扰 RNAi	Micro-Tom	柠檬酸含量提高约83% Citric acid content of fruits increases by about 83%		Osorio et al.,2013
有机酸 Organic acids	SlTDT	二羧酸转运蛋白 Dicarboxylic acid transporter protein	调控液泡中苹果酸和柠檬酸积累 Regulation of malic and citric acid accumulation in vacuole	超量表达 Overexpression	Ailsa Craig		苹果酸提高含量提高约58%，柠檬酸含量降低约37% Malic acid content increased by about 58%，citric acid content reduced by about 37%	Liu et al.,2017
	SlALMT9	铝激活苹果酸转运蛋白Aluminum- activated malate transporter	调控番茄果实苹果酸积累 Regulation of malic acid accumulation in tomato fruit	超量表达 Overexpression	Ailsa Craig		苹果酸含量提高了约50% Malic acid content of fruits increased by about 50%	Ye et al.,2017
	SlAREB1	ABA响应元件结合因子 ABA response element binding factor	调控果实柠檬酸和苹果酸积累 Regulation of fruit citric and malic acid accumulation	超量表达 Overexpression	Moneymaker		柠檬酸、苹果酸、氨基酸等代谢物含量显著增加 Fruit metabolites such as citric acid，malic acid and amino acids increased significantly in fruits	Bastias et al., 2014
	SlMIR164A	编码番茄miR164 Encodes tomato miR164	参与调控叶绿体发育和果实品质 Involved in the regulation of chloroplast development and fruit quality	CRISPR/Cas9	MicroTom		苹果酸含量提高约66% Fruit malic acid content increased by about 66%	Lin et al.,2022
	SlSAMT	番茄水杨酸甲基转移酶 Tomato salicylic acid methyltransferase	调控水杨酸甲酯积累 Regulation of methyl salicylate accumulation	超量表达 Overexpression	M82、 Pearson		水杨酸甲酯的释放量提高约33% The release of methyl salicylate increased by about 33%	Tieman et al.,2010
	LeAADC	芳香氨基酸脱羧酶 Aromatic amino acid decarboxylase	调控苯乙醛积累 Regulation of phenylacetaldehyde accumulation	超量表达 Overexpression	M82		2-苯乙醇、苯乙醛含量增加约10倍 About 10-fold increase in 2-phenylethanol and phenylacetaldehyde content	Tieman et al.,2006
挥发性化合物 Volatile Organic Compounds	SlMYB75	MYB转录因子 MYB transcription factor	转录调控多种代谢途径的下游基因，果实品质的关键调控因子 Regulating genes downstream multiple metabolic pathways，key regulators of fruit quality	超量表达 Overexpression	MicroTom		萜烯挥发物的含量提高约10倍 Terpene volatiles content increases by about 10 times	Jian et al.,2019
	FLORAL4	3-甲基-2-氧代丁酸脱氢酶 3-Methyl-2-oxobutyric acid dehydrogenase	改变番茄果实中酚类挥发物含量 Change the content of phenolic volatiles in tomato fruits	CRISPR/Cas9	RIL携带C085纯合等位基因 RIL carries a homozygous allele of C085		类黄酮和苯丙醇含量增加 Flavonoids and phenylpropanoid increased	Tikunov et al.,2020
	SlPSY1	植物烯合成酶 Phytoene synthase	调控类胡萝卜素衍生挥发物的合成 Regulation of the synthesis of carotenoid-derived volatiles	CRISPR/Cas9	Moneymaker		戊醛和2-异丁基噻唑挥发物含量增加 Glutaraldehyde and 2-isobutylthiazole volatiles increased	Cao et al.,2024

风味组分 Flavor component	基因 Gene	基因名称 Gene name	作用 Function	品质改良方法 Quality improvement method	背景材料 Background material	改良效果 Improving effect		参考文献 Reference
可溶性糖 Sugar	TIV1	反义蔗糖酶Antisense sucrase	调节果实中蔗糖积累 Regulates sucrose accumulation in fruits	反义RNA Antisense RNA	T5、H100	总糖含量提高约50% Total sugar increase by about 50%		Klann et al.,1996
	SlSWEET7a/14	糖转运蛋白 Sugar transport protein	参与果实中蔗糖、果糖、葡萄糖转运Involved in transport of sucrose，fructose and glucose in fruits	RNA干扰 RNAi	MicroTom	果糖、葡萄糖含量增加约30% ~ 60% Fructose and glucose content increased by about 30% to 60%		Zhang et al.,2021
	SlNAP2	NAC家族转录因子 NAC family transcription factors	参与调控植株叶片衰老 Involved in the regulation of plant leaf senescence	RNA干扰 RNAi	Moneymaker	糖含量增加约22.5% Sugar content increased by about 22.5%		Ma et al.,2018
	Lin5	细胞壁转化酶 Cell wall invertase	催化质外体中蔗糖裂解及其从番茄源到库供应 Catalyzing sucrose cleavage in apoplast and its supply from tomato sources to tank	超量表达 Overexpression	自交系FLA8059	葡萄糖、果糖含量提高约3.3% Glucose and fructose content increased by about 3.3%		Tieman et al.,2017
	SlVIF	液泡转化酶抑制剂 Vacuolar invertase inhibitor	抑制VIN活性，调控果实中蔗糖积累 Inhibition of VIN activity and regulation of sucrose accumulation in fruits	超量表达 Overexpression	Ailsa Craig	蔗糖含量提高10倍，己糖含量降低约40% Sucrose content increased by 10 times and hexose reduced by about 40%		Qin et al.,2016
	SlFgr	葡萄糖外排转移蛋白 Glucose exporter	影响果实中己糖的分配，改变果糖、葡萄糖比例 Influencing distribution of hexoses in the fruit，altering the ratio of fructose and glucose	超量表达 Overexpression	S. habrochaites	葡萄糖含量降低约41%，果糖葡萄糖比例提高约73% Glucose content reduced by about 41%，fructose-glucose ratio increased by about 73%		Shammai et al.,2018
	SlGLK2	MYB类转录因子 MYB-like transcription factors	调控果实中叶绿素含量 Regulation of chlorophyll content in fruits	超量表达 Overexpression	T63	葡萄糖、果糖含量提高约40% Glucose and fructose content increased by about 40%		Powell et al.,2012
	SlVPE5	番茄液泡加工酶 Tomato vacuolar processing enzyme	促进酸性转化酶合成，提高己糖水平 Promotes acid converting enzyme synthesis and increases hexose levels	CRISPR/Cas9	M82	葡萄糖、果糖含量增加约30% ~ 40% Glucose and fructose content increased by about 30% to 40%		Wang et al.,2021
	SlINVINH1	酸性转化酶抑制剂 Acid converting enzyme inhibitors	特异性抑制细胞壁转化酶活性 Specific inhibition of cell wall convertase activity	CRISPR/Cas9	M82	葡萄糖、果糖含量增加约35% ~ 45% Glucose and fructose content increased by about 35% to 45%		Wang et al.,2021
有机酸 Organic acids	SlAco	乌头酸酶 Aconitase	催化柠檬酸转化为异柠檬酸 Catalyzes the conversion of citric acid to isocitric acid	RNA干扰 RNAi	Moneymaker	柠檬酸含量提高40% 40% increase in citric acid content		Morgan et al.,2012
有机酸 Organic acids	SlPEPCK	番茄磷酸烯醇丙酮酸羧激酶 Tomato phosphoenolpyruvate carboxykinase	调节糖异生，提高糖酸比 Regulates gluconeogenesis and increases the sugar-acid ratio	RNA干扰 RNAi	Micro-Tom	柠檬酸含量提高约83% Citric acid content of fruits increases by about 83%		Osorio et al.,2013
有机酸 Organic acids	SlTDT	二羧酸转运蛋白 Dicarboxylic acid transporter protein	调控液泡中苹果酸和柠檬酸积累 Regulation of malic and citric acid accumulation in vacuole	超量表达 Overexpression	Ailsa Craig		苹果酸提高含量提高约58%，柠檬酸含量降低约37% Malic acid content increased by about 58%，citric acid content reduced by about 37%	Liu et al.,2017
	SlALMT9	铝激活苹果酸转运蛋白Aluminum- activated malate transporter	调控番茄果实苹果酸积累 Regulation of malic acid accumulation in tomato fruit	超量表达 Overexpression	Ailsa Craig		苹果酸含量提高了约50% Malic acid content of fruits increased by about 50%	Ye et al.,2017
	SlAREB1	ABA响应元件结合因子 ABA response element binding factor	调控果实柠檬酸和苹果酸积累 Regulation of fruit citric and malic acid accumulation	超量表达 Overexpression	Moneymaker		柠檬酸、苹果酸、氨基酸等代谢物含量显著增加 Fruit metabolites such as citric acid，malic acid and amino acids increased significantly in fruits	Bastias et al., 2014
	SlMIR164A	编码番茄miR164 Encodes tomato miR164	参与调控叶绿体发育和果实品质 Involved in the regulation of chloroplast development and fruit quality	CRISPR/Cas9	MicroTom		苹果酸含量提高约66% Fruit malic acid content increased by about 66%	Lin et al.,2022
	SlSAMT	番茄水杨酸甲基转移酶 Tomato salicylic acid methyltransferase	调控水杨酸甲酯积累 Regulation of methyl salicylate accumulation	超量表达 Overexpression	M82、 Pearson		水杨酸甲酯的释放量提高约33% The release of methyl salicylate increased by about 33%	Tieman et al.,2010
	LeAADC	芳香氨基酸脱羧酶 Aromatic amino acid decarboxylase	调控苯乙醛积累 Regulation of phenylacetaldehyde accumulation	超量表达 Overexpression	M82		2-苯乙醇、苯乙醛含量增加约10倍 About 10-fold increase in 2-phenylethanol and phenylacetaldehyde content	Tieman et al.,2006
挥发性化合物 Volatile Organic Compounds	SlMYB75	MYB转录因子 MYB transcription factor	转录调控多种代谢途径的下游基因，果实品质的关键调控因子 Regulating genes downstream multiple metabolic pathways，key regulators of fruit quality	超量表达 Overexpression	MicroTom		萜烯挥发物的含量提高约10倍 Terpene volatiles content increases by about 10 times	Jian et al.,2019
	FLORAL4	3-甲基-2-氧代丁酸脱氢酶 3-Methyl-2-oxobutyric acid dehydrogenase	改变番茄果实中酚类挥发物含量 Change the content of phenolic volatiles in tomato fruits	CRISPR/Cas9	RIL携带C085纯合等位基因 RIL carries a homozygous allele of C085		类黄酮和苯丙醇含量增加 Flavonoids and phenylpropanoid increased	Tikunov et al.,2020
	SlPSY1	植物烯合成酶 Phytoene synthase	调控类胡萝卜素衍生挥发物的合成 Regulation of the synthesis of carotenoid-derived volatiles	CRISPR/Cas9	Moneymaker		戊醛和2-异丁基噻唑挥发物含量增加 Glutaraldehyde and 2-isobutylthiazole volatiles increased	Cao et al.,2024

类别 Type	风味组分 Flavor component	候选基因名称 Candidate gene	基因号 Locus name	染色体 Chr	关联SNP标记 SNP	关联阈值 Meta P-value
可溶性糖 Sugar	果糖Fructose	糖基水解酶Glycosylhydrolase	Solyc01g009150	1	rs01_3327330	6.37 × 10⁻¹¹
		糖基转移酶蛋白Glycosyltransferase-likeprotein	Solyc08g081420	8	rs08_64470216	2.33 × 10⁻¹⁰
		葡萄糖基转移酶Glucosyltransferase	Solyc05g053400	5	rs05_63485334	4.68 × 10⁻¹⁰
		3-磷酸甘油醛脱氢酶Glyceraldehyde-3-phosphate dehydrogenase	Solyc10g005510	10	rs10_422707	6.27 × 10⁻¹⁰
		柠檬酸合成酶Citratesynthase	Solyc07g055840	7	rs07_63757414	4.28 × 10⁻⁰⁹
		果糖-1,6-二磷酸酶Fructose-16-bisphosphataseclass1	Solyc10g086720	10	rs10_65465775	6.84 × 10⁻⁰⁹
	葡萄糖Glucose	琥珀酰辅酶连接酶 Succinyl-CoAligase	Solyc01g007910	1	rs01_1998383	2.36 × 10⁻¹⁰
		α-葡萄糖苷酶α-glucosidase	Solyc04g007160	4	rs04_911809	6.62 × 10⁻⁰⁹
		β-1,3-半乳糖基转移酶6 β-1,3-galactosyltransferase 6	Solyc08g069060	8	rs08_58158082	4.99 × 10⁻⁰⁸
有机酸 Organic acids	柠檬酸Citrate	铝激活苹果酸转运蛋白 Aluminum-activated malate transporter	Solyc06g072920	6	rs06_44955568	7.46 × 10⁻²⁷
		糖原合酶 Glycogensynthase	Solyc03g083090	3	rs03_52998165	1.84 × 10⁻¹⁵
		糖基转移酶Glycosyl transferasegroup1	Solyc02g084820	2	rs02_47904426	4.30 × 10⁻¹³
		柠檬酸合酶 Citratesynthase	Solyc07g055840	7	rs07_63601724	4.70 × 10⁻¹²
	苹果酸Malate	肉桂酰辅酶A还原酶蛋白 Cinnamoyl CoAreductase-likeprotein	Solyc01g008550	1	rs01_2650772	2.08 × 10⁻¹⁵
		苹果酸酶 Malicenzyme	Solyc12g008430	12	rs12_1824226	1.75 × 10⁻¹⁹
		糖基转移酶类蛋白 Glycosyl transferase-likeprotein	Solyc11g072700	11	rs11_55879120	7.14 × 10⁻¹⁶
		蔗糖合成酶Sucrose synthase	Solyc09g098590	9	rs09_72364359	1.34 × 10⁻¹⁵
挥发性化合物 Volatile organic compounds	乙烯醛Hexenal	脂氧合酶Lipoxygenase	Solyc01g006540	1	rs01_1083181	1.45 × 10⁻¹⁰
	苯丙氨酸Phenylalanine	乙醇脱氢酶 Alcohol dehydrogenase	Solyc11g010960	11	rs11_4002767	9.57 × 10⁻⁰⁹
	脯氨酸Proline	丝氨酸整合因子Serine incorporator 1	Solyc03g117770	3	rs03_66798980	2.39 × 10⁻⁰⁹
	丝氨酸Serine	苏氨酸合成酶 Threonine synthase	Solyc03g121910	3	rs03_69913055	3.06 × 10⁻¹⁴
	水杨酸甲酯Methyl salicylate	1-氨基环丙烷-1-羧酸氧化酶1-Aminocyclopropane-1-carboxylic acid oxidaseoxidase-like protein	Solyc09g089580	9	rs09_69293875	2.34 × 10⁻¹⁹
	天门冬氨酸Asparagine	甲基转移酶11型Methyltransferase type 11	Solyc02g093550	2	rs02_54365596	3.72 × 10⁻¹⁰
	天门冬氨酸Asparagine	GDLS酯酶/脂肪酶GDSL esterase/lipase	Solyc12g089350	12	rs06_3502385	1.13 × 10⁻⁰⁹
	香叶基丙酮Geranylacetone	八氢番茄红素合酶 Phytoene synthase2	Solyc02g081330	2	rs03_4328514	6.00 × 10⁻¹⁵
	甲基庚烯酮6-Methyl-5-hepten-2-one	长链脂肪酸-辅酶A连接酶Long-chain-fatty-acid--CoA ligase	Solyc03g025720	3	rs03_3212583	6.76 × 10⁻²⁶
	乙基乙烯基酮1-Penten-3-one	脂磷酸磷酸酶 Lipid phosphate phosphatase 3	Solyc05g008800	5	rs05_3036212	7.07 × 10⁻⁰⁹

类别 Type	风味组分 Flavor component	候选基因名称 Candidate gene	基因号 Locus name	染色体 Chr	关联SNP标记 SNP	关联阈值 Meta P-value
可溶性糖 Sugar	果糖Fructose	糖基水解酶Glycosylhydrolase	Solyc01g009150	1	rs01_3327330	6.37 × 10⁻¹¹
		糖基转移酶蛋白Glycosyltransferase-likeprotein	Solyc08g081420	8	rs08_64470216	2.33 × 10⁻¹⁰
		葡萄糖基转移酶Glucosyltransferase	Solyc05g053400	5	rs05_63485334	4.68 × 10⁻¹⁰
		3-磷酸甘油醛脱氢酶Glyceraldehyde-3-phosphate dehydrogenase	Solyc10g005510	10	rs10_422707	6.27 × 10⁻¹⁰
		柠檬酸合成酶Citratesynthase	Solyc07g055840	7	rs07_63757414	4.28 × 10⁻⁰⁹
		果糖-1,6-二磷酸酶Fructose-16-bisphosphataseclass1	Solyc10g086720	10	rs10_65465775	6.84 × 10⁻⁰⁹
	葡萄糖Glucose	琥珀酰辅酶连接酶 Succinyl-CoAligase	Solyc01g007910	1	rs01_1998383	2.36 × 10⁻¹⁰
		α-葡萄糖苷酶α-glucosidase	Solyc04g007160	4	rs04_911809	6.62 × 10⁻⁰⁹
		β-1,3-半乳糖基转移酶6 β-1,3-galactosyltransferase 6	Solyc08g069060	8	rs08_58158082	4.99 × 10⁻⁰⁸
有机酸 Organic acids	柠檬酸Citrate	铝激活苹果酸转运蛋白 Aluminum-activated malate transporter	Solyc06g072920	6	rs06_44955568	7.46 × 10⁻²⁷
		糖原合酶 Glycogensynthase	Solyc03g083090	3	rs03_52998165	1.84 × 10⁻¹⁵
		糖基转移酶Glycosyl transferasegroup1	Solyc02g084820	2	rs02_47904426	4.30 × 10⁻¹³
		柠檬酸合酶 Citratesynthase	Solyc07g055840	7	rs07_63601724	4.70 × 10⁻¹²
	苹果酸Malate	肉桂酰辅酶A还原酶蛋白 Cinnamoyl CoAreductase-likeprotein	Solyc01g008550	1	rs01_2650772	2.08 × 10⁻¹⁵
		苹果酸酶 Malicenzyme	Solyc12g008430	12	rs12_1824226	1.75 × 10⁻¹⁹
		糖基转移酶类蛋白 Glycosyl transferase-likeprotein	Solyc11g072700	11	rs11_55879120	7.14 × 10⁻¹⁶
		蔗糖合成酶Sucrose synthase	Solyc09g098590	9	rs09_72364359	1.34 × 10⁻¹⁵
挥发性化合物 Volatile organic compounds	乙烯醛Hexenal	脂氧合酶Lipoxygenase	Solyc01g006540	1	rs01_1083181	1.45 × 10⁻¹⁰
	苯丙氨酸Phenylalanine	乙醇脱氢酶 Alcohol dehydrogenase	Solyc11g010960	11	rs11_4002767	9.57 × 10⁻⁰⁹
	脯氨酸Proline	丝氨酸整合因子Serine incorporator 1	Solyc03g117770	3	rs03_66798980	2.39 × 10⁻⁰⁹
	丝氨酸Serine	苏氨酸合成酶 Threonine synthase	Solyc03g121910	3	rs03_69913055	3.06 × 10⁻¹⁴
	水杨酸甲酯Methyl salicylate	1-氨基环丙烷-1-羧酸氧化酶1-Aminocyclopropane-1-carboxylic acid oxidaseoxidase-like protein	Solyc09g089580	9	rs09_69293875	2.34 × 10⁻¹⁹
	天门冬氨酸Asparagine	甲基转移酶11型Methyltransferase type 11	Solyc02g093550	2	rs02_54365596	3.72 × 10⁻¹⁰
	天门冬氨酸Asparagine	GDLS酯酶/脂肪酶GDSL esterase/lipase	Solyc12g089350	12	rs06_3502385	1.13 × 10⁻⁰⁹
	香叶基丙酮Geranylacetone	八氢番茄红素合酶 Phytoene synthase2	Solyc02g081330	2	rs03_4328514	6.00 × 10⁻¹⁵
	甲基庚烯酮6-Methyl-5-hepten-2-one	长链脂肪酸-辅酶A连接酶Long-chain-fatty-acid--CoA ligase	Solyc03g025720	3	rs03_3212583	6.76 × 10⁻²⁶
	乙基乙烯基酮1-Penten-3-one	脂磷酸磷酸酶 Lipid phosphate phosphatase 3	Solyc05g008800	5	rs05_3036212	7.07 × 10⁻⁰⁹

Regulatory Genes for Tomato Flavor and Their Application in Quality Improvement

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