番茄果实形状的调控机制研究进展

doi:10.16420/j.issn.0513-353x.2022-0786

摘要/Abstract

摘要：

番茄是果实研究的模式植物，番茄果形研究有助于理解果实形态发生的机制。本文主要从QTL基因调控、转录调控、转录后调控和植物激素调控等方面综述了番茄果形建成的研究进展，旨在更好理解调控番茄果形的分子机制，为番茄优质育种提供见解。

关键词: 番茄, 果形, QTL基因, 转录调控, 转录后调控, 植物激素

Abstract:

Tomato is a model plant for fruit research. The study of fruit shape formation is helpful to understand the regulatory mechanisms of fruit morphogenesis. This review summarizes the progress of fruit shape formation in tomato from the aspects of QTL gene regulation，transcriptional and post-transcriptional regulation，and plant hormones. This work contributes to understanding the molecular mechanisms underlying tomato fruit shape formation，and provides new insights for tomato quality breeding.

Key words: tomato, fruit shape, QTL gene, transcriptional regulation, post-transcriptional regulation, plant hormone

姬雅静, 李金焱, 张沛宇, 马力群, 朱鸿亮. 番茄果实形状的调控机制研究进展[J]. 园艺学报, 2023, 50(9): 2015-2030.

JI Yajing, LI Jinyan, ZHANG Peiyu, MA Liqun, ZHU Hongliang. Research Progress on the Regulatory Mechanism of Shape Formation in Tomato Fruit[J]. Acta Horticulturae Sinica, 2023, 50(9): 2015-2030.

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类型 Type	基因号 Gene ID	对果形的影响 Effect on fruit shape	调控果形的分子机制 Molecular mechanisms that regulate fruit shape	参考文献 Reference
QTL基因 QTL gene	SUN Solyc10g079240	sun果实伸长 Tomato fruit elongates in sun	SUN通过招募CAM影响微管重排来调节细胞分裂 SUN regulates cell division by recruiting CAM to influence microtubule rearrangement	Bürstenbinder et al.,2017；Jiang et al.,2009；Xiao et al.,2008
	OVATE Solyc02g085500	ovate番茄为梨形 ovate mutant is a pear-shaped tomato	OVATE和SlOFP20与TRM相互作用改变调节细胞分裂和器官生长 Interaction between OVATE and SlOFP20 and TRM regulates cell division and organ growth	Liu et al.,2002； van de Knaap et al.， 2002；Wu et al.,2018
	WUSCHEL/LC Solyc02g083950	lc番茄心室数增多，果实更大更扁平 The number of ventricles in lc mutant tomato increases，and the fruit is larger and flattened	lc中两个SNP的出现抑制了转录抑制因子AGAMOUS与WUS的结合，影响了WUS-CLV途径 Two SNPs in lc are believed to inhibit the binding of transcription inhibitor AGAMOUS to WUS and affect the WUS-CLV pathway	Liu et al.,2011；Muños et al.,2011；Tilly et al.,1998
	GLOBE Solyc12g006860	扁平型番茄敲除GLOBE变为圆果形 Knock out GLOBE in flat tomato to make round fruit	GLOBE编码油菜素内酯羟化酶，调控的分子机制未知 GLOBE encodes a brassinoidin-lipid hydroxylase，but its molecular mechanism is unknown	Sierra-Orozco et al.,2021
	CLV3/FAS Solyc11g071380	fas番茄心室数增多，果实更大更扁平 The number of ventricles was increased and the fruit was larger and flattened in fas	影响了WUS-CLV途径 It affects the WUS-CLV pathway	Chu et al.,2019
	CLV1/FAB Solyc04g081590	fab番茄心室数增多，果实更大 The carpel number increased and the fruit was larger and flattened in fab	影响了WUS-CLV途径 It affects the WUS-CLV pathway	Xu et al.,2015
	HAPT3/FIN Solyc11g064850	fin番茄心室数增多，果实更大 fin had more carpels and larger fruits	HAPT3的突变导致CLV3不能完全阿拉伯糖基化，影响了WUS-CLV途径 Mutations in HAPT3 lead to incomplete arabinylation of CLV3，which affects the WUS-CLV pathway	Xu et al.,2015
转录因子 Transcription factors	ENO Solyc03g117230	eno形成更大的多心室番茄 eno forms larger and multi-ventricular tomatoes	ENO调节WUS的表达，影响了WUS-CLV途径 ENO regulates the expression of WUS and affects the WUS-CLV pathway	Fernández-Lozano et al.,2015；Yuste- Lisbona et al.,2020
	BZR1.7 Solyc10g076390	超表达BZR1.7形成长果表型 Overexpression of BZR1.7 forms a long-fruit phenotype	BZR1.7通过调控SUN的表达来改变果实形状 BZR1.7 regulates SUN expression to change fruit shape	Yu et al.,2022
	BES1.8 Solyc10g076390	过表达 BES1.8，番茄心室数增多 Overexpression of BES1.8 increases the number of tomato ventricles	SlBES1.8与WUS互作抑制SlWUS和SlCLV3启动子的结合，影响了WUS-CLV途径 The interaction between SlBES1.8 and WUS inhibits the binding of SlWUS and SlCLV3 promoters which affecting the WUS-CLV pathway	Su et al.,2022
	KNU Solyc02g094428	knu心室增多 The number of carpel increases in knu	影响了WUS-CLV途径 It affects the WUS-CLV pathway	Bollier et al.,2018
锌指蛋白 Zinc finger protein	IMA Solyc02g087970	ima心室增多 The number of carpel increases in ima	影响了WUS-CLV途径 It affects the WUS-CLV pathway	Bollier et al.,2018； Sicard et al.,2008
转录后调控 Post- transcriptional regulation	miR159	抑制Sly-miR159表达导致果实更扁更大 Inhibition of Sly-miR159 expression results in flattened and larger fruit	Sly-miR159通过其靶基因SlGAMYB2调控赤霉素生物合成影响果实形状 Sly-miR159 regulates GA biosynthesis and affects fruit shape through its target gene SlGAMYB2	Zhao et al.,2021
转录后调控 Post- transcriptional regulation	miR160	圆果敲除Sly-miR160形成梨形果 Knock out Sly-miR160 in round fruit to form pear-shaped tomato	miR160通过靶向3种生长素反应因子SlARF10、SlARF16和SlARF17调控生长素信号 miR160 regulates auxin signaling by targeting SlARF10、SlARF16 and SlARF17	Damodharan et al.,2016
	miR319	miR319基因敲除产生类似ovate的梨形果 Knockout of the miR319 gene produces a pear-shaped fruit similar to ovate	Sly-miR319调节OVATE的表达以及雌蕊发育期间的生长素水平 Sly-miR319 regulates OVATE expression and auxin homeostasis during pistil development	Carvalho et al.,2022
	miR396	miR396下调导致果实更大 Downregulation of miR396 results in larger tomato fruit	miR396的下调导致靶向的生长调节因子GRF表达上调，导致果实明显变大 The downregulation of miR396 leads to upregulation of the targeted growth regulator GRF，resulting in significantly larger fruits	Cao et al.,2016
生长素 Auxin	IAA17 Solyc06g008590	SlIAA17- RNAi株系产生更大果实 The SlIAA17- RNAi produces larger tomato fruit	抑制SlIAA17的表达后，通过倍性水平的增加影响果皮细胞的大小，最终影响水果的最终大小 Inhibition of SlIAA17 affects the size of epidermal cells by increasing ploidy levels and ultimately the final fruit size	Su et al.,2014
生长素 Auxin	ARF7 Solyc07g042260	SlARF7-RNAi为心形果实 SlARF7-RNAi line is heart-shaped fruit	SlARF7通过调节生长素和GA反应，调控细胞分裂和扩张 SlARF7 regulates cell division and expansion by regulating auxin and GA responses	de Jong et al.,2011
赤霉素 GA	PRE2 Solyc02g067380	SlPRE2过表达系的果实略增大，功能丧失系果实变小 The diameter of SlPRE2 overexpression fruit increases slightly，while the fruit diameter of functional loss line decreases	通过GA调节反应参与细胞伸长 It is involved in cell elongation through GA regulation	Zhu et al.,2017, 2019
	GRAS2 Solyc07g063940	SlGRAS2-RNAi株系果实变小 The fruit becomes smaller in the SlGRAS2-RNAi line	SlGRAS2参与GA的生物合成和信号转导，调节细胞扩张，影响果实的大小和质量 SlGRAS2 participates in regulating GA biosynthesis and signal transduction，cell expansion，and then influences fruit size and weight	Li et al.,2018b
	CDF4 Solyc02g067230	PPC2启动SlCDF4表达增加，通过细胞层数和细胞大小的增加，产生更大果实 PPC2 activated SlCDF4 expression is increased，which results in larger fruit with increased number of cell layers and cell size	SlCDF4可能影响生长素和GA合成，调节细胞分裂和扩张 SlCDF4 may affect synthesis of auxin and GA，and regulate cell division and expansion	Renau-Morata et al., 2020
脱落酸 ABA	FLACCA Solyc07g066480	ABA缺失双突变体not/flc 果皮细胞更小，果实更小 ABA-deficient double mutants not/flc shows smaller pericarp cells and fruit	ABA通过抑制乙烯合成从而促进细胞扩张和果实生长 ABA promotes cell expansion and fruit growth by inhibiting ethylene synthesis	Nitsch et al.,2012
脱落酸 ABA	NCED1 Solyc07g056570			Nitsch et al.,2012
乙烯 Ethylene	EBF2-like Solyc07g008250	过表达SlEBF2-like导致果实纵向伸长Overexpression of SLEBF2-like results in longitudinal elongation of fruit	SlEBF2-like通过调节乙烯反应参与果实发育 SlEBF2-like is involved in fruit development by regulating ethylene reaction	Guo et al.,2018
翻译后修饰 Post- translational modification	ORF9/MEL Solyc05g012790	圆形番茄敲除ORF9果实拉长 Knocking out of ORF9 elongates fruit shape in round tomato	ORF9与MAP65-2和CAM1互作，推测CAM1在MEL高度可逆的棕榈酰化修饰中与MAP65-1结合或解离，进而对生长旺盛部位动态调控细胞状态 ORF9 interacts with MAP65-2 and CAM1 that suggest binds to or dissociates between CAM1 and MAP65-1 during highly reversible palmitoylation modification of MEL，thereby dynamically regulates cell state of vigorous growth sites	杨其洪,2020

类型 Type	基因号 Gene ID	对果形的影响 Effect on fruit shape	调控果形的分子机制 Molecular mechanisms that regulate fruit shape	参考文献 Reference
QTL基因 QTL gene	SUN Solyc10g079240	sun果实伸长 Tomato fruit elongates in sun	SUN通过招募CAM影响微管重排来调节细胞分裂 SUN regulates cell division by recruiting CAM to influence microtubule rearrangement	Bürstenbinder et al.,2017；Jiang et al.,2009；Xiao et al.,2008
	OVATE Solyc02g085500	ovate番茄为梨形 ovate mutant is a pear-shaped tomato	OVATE和SlOFP20与TRM相互作用改变调节细胞分裂和器官生长 Interaction between OVATE and SlOFP20 and TRM regulates cell division and organ growth	Liu et al.,2002； van de Knaap et al.， 2002；Wu et al.,2018
	WUSCHEL/LC Solyc02g083950	lc番茄心室数增多，果实更大更扁平 The number of ventricles in lc mutant tomato increases，and the fruit is larger and flattened	lc中两个SNP的出现抑制了转录抑制因子AGAMOUS与WUS的结合，影响了WUS-CLV途径 Two SNPs in lc are believed to inhibit the binding of transcription inhibitor AGAMOUS to WUS and affect the WUS-CLV pathway	Liu et al.,2011；Muños et al.,2011；Tilly et al.,1998
	GLOBE Solyc12g006860	扁平型番茄敲除GLOBE变为圆果形 Knock out GLOBE in flat tomato to make round fruit	GLOBE编码油菜素内酯羟化酶，调控的分子机制未知 GLOBE encodes a brassinoidin-lipid hydroxylase，but its molecular mechanism is unknown	Sierra-Orozco et al.,2021
	CLV3/FAS Solyc11g071380	fas番茄心室数增多，果实更大更扁平 The number of ventricles was increased and the fruit was larger and flattened in fas	影响了WUS-CLV途径 It affects the WUS-CLV pathway	Chu et al.,2019
	CLV1/FAB Solyc04g081590	fab番茄心室数增多，果实更大 The carpel number increased and the fruit was larger and flattened in fab	影响了WUS-CLV途径 It affects the WUS-CLV pathway	Xu et al.,2015
	HAPT3/FIN Solyc11g064850	fin番茄心室数增多，果实更大 fin had more carpels and larger fruits	HAPT3的突变导致CLV3不能完全阿拉伯糖基化，影响了WUS-CLV途径 Mutations in HAPT3 lead to incomplete arabinylation of CLV3，which affects the WUS-CLV pathway	Xu et al.,2015
转录因子 Transcription factors	ENO Solyc03g117230	eno形成更大的多心室番茄 eno forms larger and multi-ventricular tomatoes	ENO调节WUS的表达，影响了WUS-CLV途径 ENO regulates the expression of WUS and affects the WUS-CLV pathway	Fernández-Lozano et al.,2015；Yuste- Lisbona et al.,2020
	BZR1.7 Solyc10g076390	超表达BZR1.7形成长果表型 Overexpression of BZR1.7 forms a long-fruit phenotype	BZR1.7通过调控SUN的表达来改变果实形状 BZR1.7 regulates SUN expression to change fruit shape	Yu et al.,2022
	BES1.8 Solyc10g076390	过表达 BES1.8，番茄心室数增多 Overexpression of BES1.8 increases the number of tomato ventricles	SlBES1.8与WUS互作抑制SlWUS和SlCLV3启动子的结合，影响了WUS-CLV途径 The interaction between SlBES1.8 and WUS inhibits the binding of SlWUS and SlCLV3 promoters which affecting the WUS-CLV pathway	Su et al.,2022
	KNU Solyc02g094428	knu心室增多 The number of carpel increases in knu	影响了WUS-CLV途径 It affects the WUS-CLV pathway	Bollier et al.,2018
锌指蛋白 Zinc finger protein	IMA Solyc02g087970	ima心室增多 The number of carpel increases in ima	影响了WUS-CLV途径 It affects the WUS-CLV pathway	Bollier et al.,2018； Sicard et al.,2008
转录后调控 Post- transcriptional regulation	miR159	抑制Sly-miR159表达导致果实更扁更大 Inhibition of Sly-miR159 expression results in flattened and larger fruit	Sly-miR159通过其靶基因SlGAMYB2调控赤霉素生物合成影响果实形状 Sly-miR159 regulates GA biosynthesis and affects fruit shape through its target gene SlGAMYB2	Zhao et al.,2021
转录后调控 Post- transcriptional regulation	miR160	圆果敲除Sly-miR160形成梨形果 Knock out Sly-miR160 in round fruit to form pear-shaped tomato	miR160通过靶向3种生长素反应因子SlARF10、SlARF16和SlARF17调控生长素信号 miR160 regulates auxin signaling by targeting SlARF10、SlARF16 and SlARF17	Damodharan et al.,2016
	miR319	miR319基因敲除产生类似ovate的梨形果 Knockout of the miR319 gene produces a pear-shaped fruit similar to ovate	Sly-miR319调节OVATE的表达以及雌蕊发育期间的生长素水平 Sly-miR319 regulates OVATE expression and auxin homeostasis during pistil development	Carvalho et al.,2022
	miR396	miR396下调导致果实更大 Downregulation of miR396 results in larger tomato fruit	miR396的下调导致靶向的生长调节因子GRF表达上调，导致果实明显变大 The downregulation of miR396 leads to upregulation of the targeted growth regulator GRF，resulting in significantly larger fruits	Cao et al.,2016
生长素 Auxin	IAA17 Solyc06g008590	SlIAA17- RNAi株系产生更大果实 The SlIAA17- RNAi produces larger tomato fruit	抑制SlIAA17的表达后，通过倍性水平的增加影响果皮细胞的大小，最终影响水果的最终大小 Inhibition of SlIAA17 affects the size of epidermal cells by increasing ploidy levels and ultimately the final fruit size	Su et al.,2014
生长素 Auxin	ARF7 Solyc07g042260	SlARF7-RNAi为心形果实 SlARF7-RNAi line is heart-shaped fruit	SlARF7通过调节生长素和GA反应，调控细胞分裂和扩张 SlARF7 regulates cell division and expansion by regulating auxin and GA responses	de Jong et al.,2011
赤霉素 GA	PRE2 Solyc02g067380	SlPRE2过表达系的果实略增大，功能丧失系果实变小 The diameter of SlPRE2 overexpression fruit increases slightly，while the fruit diameter of functional loss line decreases	通过GA调节反应参与细胞伸长 It is involved in cell elongation through GA regulation	Zhu et al.,2017, 2019
	GRAS2 Solyc07g063940	SlGRAS2-RNAi株系果实变小 The fruit becomes smaller in the SlGRAS2-RNAi line	SlGRAS2参与GA的生物合成和信号转导，调节细胞扩张，影响果实的大小和质量 SlGRAS2 participates in regulating GA biosynthesis and signal transduction，cell expansion，and then influences fruit size and weight	Li et al.,2018b
	CDF4 Solyc02g067230	PPC2启动SlCDF4表达增加，通过细胞层数和细胞大小的增加，产生更大果实 PPC2 activated SlCDF4 expression is increased，which results in larger fruit with increased number of cell layers and cell size	SlCDF4可能影响生长素和GA合成，调节细胞分裂和扩张 SlCDF4 may affect synthesis of auxin and GA，and regulate cell division and expansion	Renau-Morata et al., 2020
脱落酸 ABA	FLACCA Solyc07g066480	ABA缺失双突变体not/flc 果皮细胞更小，果实更小 ABA-deficient double mutants not/flc shows smaller pericarp cells and fruit	ABA通过抑制乙烯合成从而促进细胞扩张和果实生长 ABA promotes cell expansion and fruit growth by inhibiting ethylene synthesis	Nitsch et al.,2012
脱落酸 ABA	NCED1 Solyc07g056570			Nitsch et al.,2012
乙烯 Ethylene	EBF2-like Solyc07g008250	过表达SlEBF2-like导致果实纵向伸长Overexpression of SLEBF2-like results in longitudinal elongation of fruit	SlEBF2-like通过调节乙烯反应参与果实发育 SlEBF2-like is involved in fruit development by regulating ethylene reaction	Guo et al.,2018
翻译后修饰 Post- translational modification	ORF9/MEL Solyc05g012790	圆形番茄敲除ORF9果实拉长 Knocking out of ORF9 elongates fruit shape in round tomato	ORF9与MAP65-2和CAM1互作，推测CAM1在MEL高度可逆的棕榈酰化修饰中与MAP65-1结合或解离，进而对生长旺盛部位动态调控细胞状态 ORF9 interacts with MAP65-2 and CAM1 that suggest binds to or dissociates between CAM1 and MAP65-1 during highly reversible palmitoylation modification of MEL，thereby dynamically regulates cell state of vigorous growth sites	杨其洪,2020