Identification of miRNA in Tomato and Its Application in Genetic Improvement

doi:10.16420/j.issn.0513-353x.2024-0948

Acta Horticulturae Sinica ›› 2026, Vol. 53 ›› Issue (10): 2983-2999.doi: 10.16420/j.issn.0513-353x.2024-0948

Identification of miRNA in Tomato and Its Application in Genetic Improvement

XUAN Xiaoxiao¹, XIE Yuxin¹, LI Fangman¹, GE Pingfei¹, YANG Yang¹, XU Haobo¹, ZHANG Xingyu¹, TAO Jinbao¹, WANG Ziyuan¹, ZHANG Yuyang¹^,²^,^*()

¹ National Key Laboratory for Germplasm Innovation & Utilization of Horticultural Crops， Huazhong Agricultural University， Wuhan 430070， China
² Hubei Hongshan Laboratory， Wuhan 430070， China

Received:2026-03-05 Revised:2026-07-09 Online:2026-10-25 Published:2026-10-26
Contact: ZHANG Yuyang

Abstract

Abstract:

MicroRNA（miRNA）is a kind of non-coding small RNA that is ubiquitous in organisms. Over the years，researchers have found tens of thousands of miRNAs in animals and plants，and explored their functions. More and more evidence show that miRNAs play key roles in regulating traits in crops. As a staple vegetable as well as model plant，miRNAs in tomato have been extensively investigated in growth and development，stress response and fruit quality. This article mainly reviews the research progress in tomato miRNA identification，function characterization and their applications in genetic improvement，and provides theoretical reference for miRNA in tomato genetic improvement.

Key words: tomato, miRNA, target gene, regulation mechanism, genetic improvement

CLC Number:

S 641.2

XUAN Xiaoxiao, XIE Yuxin, LI Fangman, GE Pingfei, YANG Yang, XU Haobo, ZHANG Xingyu, TAO Jinbao, WANG Ziyuan, ZHANG Yuyang. Identification of miRNA in Tomato and Its Application in Genetic Improvement[J]. Acta Horticulturae Sinica, 2026, 53(10): 2983-2999.

Figures/Tables 3

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性状 Trait	miRNA	靶基因 Target gene	调控生物途径 Regulating biological pathway	受体材料 Material	改良途径 Improving method	改良效果 Improvement effect	参考文献 Reference
株型 Plant type	miR171d/e	SlGRAS24	叶绿素生物合成 Chlorophyll biosynthesis	MicroTom	STTM	STTM沉默miR171d/e植株矮化、叶片颜色加深、叶片中总叶绿素含量比对照高5.36、7.6 μg · g^-1、花粉粒较饱满 The plant of STTM171d/e was dwarfed，the leaf color was deepened，the total chlorophyll content in the leaves was 5.36 and 7.6 μg · g^-1 higher than that of the control，and the pollen grains were fuller	蒲慧丽,2021
	miR156a	SPL13	激素合成和传导 Hormone synthesis and transmission	Ailsa Craig	超量表达Overex- pression	侧枝数量大约是对照的2倍 The number of lateral branches is about 2 times that of the control	Cui et al.,2020
	miR168	SlAGO1A	细胞分裂素等 Hormones such as cytokinin	JZ18/JZ34	超量表达Overex- pression	低钾条件下，根毛数增加、叶片发育更好、植株叶绿素含量增加 Under low K⁺，the number of root hairs increased，the leaf development was better，and the chlorophyll of plants increased	Liu et al.,2020
花期调控Flowering regulation	miR171b	/	乙烯生物合成 Ethylene biosynthesis	Ailsa Craig	CRISPR/Cas9	开花时间提前3 ~ 5d，单株果实产量提高21.3% ~ 22.4% The flowering time was 3-5 days earlier，and the fruit yield per plant increased by 21.3%-22.4%	潘燕婷,2020
	miR171d/e	SlGRAS24	乙烯生物合成、类胡萝卜素合成Ethylene biosynthesis，carotenoid synthesis	MicroTom	STTM	开花提前，开花至破色时间缩短6 ~ 8 d The flowering time was advanced and the flowering- breaking time was shortened by 6-8 days	蒲慧丽,2021
	miR1917	CPR4	乙烯生物合成 Ethylene biosynthesis	中蔬6号 Zhongshu6	超量表达Overex- pression	加速花梗脱落、果实成熟提前约3~ 4 d Accelerated pedicel abscission，fruit ripening 3-4 d in advance	Wang et al.,2018
	miR156	SPL	赤霉素生物合成 Gibberellin biosynthesis pathway	MicroTom	超量表达Overex- pression	超量表达番茄植株在种子萌发后的第10天，超过80%的顶端仍处于营养分生组织期，而MT顶端超过95%处于花序和花分生组织 At 10DPG，more than 80% of the tops of overexpressing tomato plants were still in vegetative meristem stage，while more than 95% of the tops of MT were in inflorescence and floral meristem	Silva et al.,2019
	miR319	LA	赤霉素生物合成 Gibberellin biosynthesis	MicroTom	靶基因同义突变 Synony- mous mutation	La-1突变体在种子萌发后的第10天，没有处于花序和花分生组织，而MT顶端超过95%处于花序和花分生组织 The La-1 mutant was not in the inflorescence and flower meristem at 10 DPG，while more than 95 % of the MT tip was in the inflorescence and flower meristem	Silva et al.,2019
果实发育 Fruit develop- ment	miR164a	SlNAM2/ SlNAM3	乙烯生物合成 Ethylene biosynthesis	MicroTom	CRISPR/Cas9	开花到成熟开始缩短约5d The time from flowering to maturity was shortened by about 5 days	Lin et al.,2022
	miR396	GRFs	激素途径、细胞壁相关途径Hormone pathway，cell wall related pathway	MicroTom	STTM	果实变大，质量增加约55%，果皮增厚32% ~ 35%，萼片长和宽分别增加约65%和20% The fruit became larger，the fruit weight increased by about 55%，the peel thickness increased by 32%-35%，and the sepal length and width increased by about 65 % and 20 %.	曹东艳,2018
	miR172a	AP2a	乙烯生物合成、类胡萝卜素合成Ethylene biosynthesis，carotenoid synthesis	Ailsa Craig	超量表达Overex- pression	β-胡萝卜素升高约两倍，果实成熟时间比野生型早4 ~ 5 d，萼片向花瓣转变 The β-carotene content increased by about two times，the fruit ripening time was 4-5 days earlier than WT，and sepals were transformed into petals	Chung et al.,2020
	miR159	SlGAMYB1/2	GA和IAA合成 GA and IAA synthesis	MicroTom	超量表达Overex- pression	诱导单性结实、果实100%无种子Induced parthenocarpy，fruit 100 % seedless	Da et al.,2017
	miR160a	SlARF10A/10B/17	IAA合成 IAA synthesis	M82	沉默突变Silent mutation	果实由圆形变为圆锥形，几乎无籽 The fruit changes from round to conical，and the fruit is almost seedless	Mekontso et al.,2023
果实品质 Fruit quality	miR164a	SlNAM2/ SlNAM3	碳水化合物代谢Carbohydrate metabolism	MicroTom	CRISPR/Cas9	淀粉、有机酸含量升高、葡萄糖和果糖含量在开花后30 ~ 35 d增加，成熟后期下降 The content of starch，organic acid，glucose and fructose increased at 30-35 DPA，and decreased at the later stage of maturity	Lin et al.,2022
	miR171b	/	糖酸生物合成 Sugar acid biosynthesis	Ailsa Craig	CRISPR/Cas9	葡萄糖含量上升45%，果糖含量增加2倍，柠檬酸和苹果酸含量下降20%左右 Glucose content increased by 45%，fructose content increased by 2 times，citric acid and malic acid content decreased by about 20%	潘燕婷,2020
	miR394	ZDS	类胡萝卜素代谢通路 Carotenoid metabolism	Micro Tom	STTM	STTM394株系番茄红素和β-类胡萝卜素含量为26.64和17.51 mg · kg^-1，是野生型2.17和1.39倍 The content of lycopene and β-carotene in the STTM394 line was 26.64 mg · kg^-1 and 17.51 mg · kg^-1，which was 2.17 and 1.39 times that of wild type	赵丽,2023
	miR159	MYB65-DFR	花青素合成 Anthocyanin biosynthesis	LA1996	超量表达Overex- pression	果实的花青素浓度比野生型增加1.452 nmol · L^-1 The anthocyanin concentration of fruit was 1.452 nmol · L^-1higher than that of wild type	马旭,2023
	miR160a	SlARF10A/10B/17	类胡萝卜素和番茄红素生物合成 Carotenoid and lycopene biosynthesis	MicroTom	超量表达Overex- pression	在破色期，果实类胡萝卜素含量比野生型高5.19 g · kg^-1，番茄红素、类胡萝卜素含量在整个成熟过程中呈增加趋势 At the color breaking stage，the content of carotenoids in fruits was 5.19 g · kg^-1 higher than that of WT，and the contents of lycopene and carotenoids increased during the whole ripening process	Mekontso et al.,2023
	mi9474	SlGMP/ SlGalUR	抗坏血酸合成 AsA synthesis	Ailsa Craig	超量表达Overex- pression	还原型抗坏血酸含量下降7.61% ~ 14.73%，总抗坏血酸含量下降5.61% ~ 8.04% The content of reduced ascorbic acid decreased by 7.61%-14.73%，and the total ascorbic acid decreased by 5.61%-8.04%.	朱立辉,2023
	miR319b	SlGME1/SlGME2	抗坏血酸合成 AsA synthesis	Ailsa Craig	超量表达Overex- pression	还原型抗坏血酸显著下降12.02% ~ 12.30%，总抗坏血酸下降17.47% ~ 17.87% The content of reduced ascorbic acid decreased significantly by 12.02%-12.30%，and the total ascorbic acid decreased by 17.47%-17.87%	朱立辉,2023
非生物胁迫 Abiotic stress	miR167	NF-YA1/ NF-YA2	叶绿素合成途径、脯氨酸和丙二醛相关途径 Chlorophyll synthesis，proline and malondialdehyde related pathways	中蔬4号Zhongshu 4	超量表达Overex- pression	与对照相比，超量表达植株叶绿素含量和最大光化学效率降低较少、脯氨酸含量增加1.5倍、MDA含量减少1.4倍、番茄对冷胁迫的耐受性提高 Compared with the control，the chlorophyll content and maximum photochemical efficiency of overexpressing plants decreased less，the proline content increased by 1.5 times，the MDA content decreased by 1.4 times，and the tolerance of tomato to cold stress increased	Gou et al.,2014
	miR319d	GAMYB- like1	活性氧系统关键酶 Key enzyme of reactive oxygen system	MicroTom	超量表达Overex- pression	SOD、CAT活性升高，耐冷性增强 The activities of SOD and CAT increased，and the cold tolerance enhanced	Shi et al.,2019
	miR169c	SIMRP1	内质网相关蛋白质降解途径 Endoplasmic reticulum-related protein degradation pathway	Ailsa Craig	超量表达Overex- pression	与WT相比，气孔导度降低33% ~ 45%，蒸腾速率降低45% ~ 62%，提高抗旱性和耐盐性 Compared with WT，the stomatal conductance decreased by 33%-45%，the transpiration rate decreased by 45%-62%，and the drought resistance and salt tolerance increased	Zhang et al.,2011
	miR157	SPL-CNR	侧根和根毛发育相关基因Genes related to lateral root and root hair development	Ailsa Craig	超量表达Overex- pression	高铁螯合物还原酶（FCR）活性显著提高，诱导缺Fe反应，维持根系Fe稳态 The activity of ferric chelate reductase（FCR）was significantly increased，which induced Fe deficiency reaction and maintained Fe homeostasis in roots	Zhu et al.,2022
非生物胁迫 Abiotic stress	miR168a	AGO1	侧根和根毛发育相关基因表达Genes related to lateral root and root hair development	JZ18/JZ34	超量表达Overex- pression	通过增多侧根和根毛、增加根表面积，增强耐低钾能力 By increasing the number of lateral roots and root hairs，increasing the root surface area，the ability of low potassium tolerance was enhanced	刘杨,2018
生物胁迫 Biotic stress	miR482b	NBS-LRR	触发免疫系统（ETI） Trigger the immune system（ETI）	L3708	STTM	致病性、叶片病斑明显较小，相对病斑直径显著降低，MDA水平降低，对番茄晚疫病抗性提高 The pathogenicity and leaf lesions were significantly smaller，the relative lesion diameter was significantly reduced，the MDA level was reduced，and the resistance to tomato late blight was improved	Jiang et al.,2018
	miR397	Lac12	SA和JA信号通路 SA and JA signaling pathway	MicroTom	STTM	STTM397株系病斑面积与叶面积的比值降低近3倍，叶片病斑较浅，对晚疫病抗性提高 The ratio of lesion area to leaf area of STTM397 lines was reduced by nearly 3 times，and the lesion area of leaves was less，and the resistance to late blight was improved	Guan et al.,2023
	miR172a/b	AR2/ERF	SOD、POD酶活性，MDA含量，REL（相对电解质渗漏）值 SOD，POD enzyme activity，MDA content，REL （relative electrolyte leakage）value	Zaofen 2	超量表达 Overex- pression	抗番茄晚疫病，平均病情指数下降17% Resistant to tomato late blight，the average disease index decreased by 17%	Luan et al.,2018
	miR1918	sl-TG2	触发PTI、ETI 免疫系统 Trigger PTI，ETI immune system	Zaofen 2	超量表达 Overex- pression	接种致病疫霉后6和10 d，超量表达株系相对病灶平均面积减少3.9%和25%，平均病情指数减少17%（10 dpi），对番茄晚疫病抗性提高 After inoculation with P.infestans，the average relative lesion area of overexpression lines decreased by 3.9%（6 dpi）and 25%（10 dpi），the average disease index decreased by 17%（10 dpi），and the resistance to tomato late blight increased	Luan et al.,2016
	miR166	SlHB	调控免疫系统 Regulating the immune system	H-88-78-1/Punjab Chhuhara	VIGS靶基因 VIGS target genes	ToLCNDV 感染后，沉默株系疾病严重程度降低、病毒滴度显著降低（约55%）、番茄对ToLCNDV抗性增强 After ToLCNDV infection，the disease severity of the silenced lines was reduced，the virus titer was significantly reduced（about 55%），and the resistance of tomato to ToLCNDV was enhanced	Prasad et al.,2022
	miR159	SlMYB33	HR相关调控 HR-related regulation	H-88-78-1/Punjab Chhuhara	VIGS靶基因 VIGS target genes	抗性材料VIGS沉默株系TYLCV感染后提前出现卷叶和生长迟缓（8 dpi），80%植株表现出典型卷叶，抗性品种中病毒滴度增加 > 4倍 After TYLCV infection，the VIGS silencing of resistant lines showed early leaf rolling and growth retardation（8 dpi），80% of the plants showed typical leaf rolling，and the virus titer in resistant varieties increased by more than 4 times	Sharma et al.,2021
	miR482f/miR5300	Solyc08g075630/Solyc08g076000;Solyc05g008650/Solyc09g018220	PTI、ETI免疫系统 PTI，ETI innate immune system	Motelle/ Money- maker	VIGS靶基因 VIGS target genes	植株生长缓慢，顶部叶片表现出枯萎，导致抗性Motelle品种对尖孢镰刀菌敏感，对番茄枯萎病抗性降低 The plants grew slowly，and the top leaves showed wilting，which led to the sensitivity of the resistant Motelle variety to Fusarium oxysporum and the decrease of resistance to tomato Fusarium wilt	Ouyang et al.,2014

性状 Trait	miRNA	靶基因 Target gene	调控生物途径 Regulating biological pathway	受体材料 Material	改良途径 Improving method	改良效果 Improvement effect	参考文献 Reference
株型 Plant type	miR171d/e	SlGRAS24	叶绿素生物合成 Chlorophyll biosynthesis	MicroTom	STTM	STTM沉默miR171d/e植株矮化、叶片颜色加深、叶片中总叶绿素含量比对照高5.36、7.6 μg · g^-1、花粉粒较饱满 The plant of STTM171d/e was dwarfed，the leaf color was deepened，the total chlorophyll content in the leaves was 5.36 and 7.6 μg · g^-1 higher than that of the control，and the pollen grains were fuller	蒲慧丽,2021
	miR156a	SPL13	激素合成和传导 Hormone synthesis and transmission	Ailsa Craig	超量表达Overex- pression	侧枝数量大约是对照的2倍 The number of lateral branches is about 2 times that of the control	Cui et al.,2020
	miR168	SlAGO1A	细胞分裂素等 Hormones such as cytokinin	JZ18/JZ34	超量表达Overex- pression	低钾条件下，根毛数增加、叶片发育更好、植株叶绿素含量增加 Under low K⁺，the number of root hairs increased，the leaf development was better，and the chlorophyll of plants increased	Liu et al.,2020
花期调控Flowering regulation	miR171b	/	乙烯生物合成 Ethylene biosynthesis	Ailsa Craig	CRISPR/Cas9	开花时间提前3 ~ 5d，单株果实产量提高21.3% ~ 22.4% The flowering time was 3-5 days earlier，and the fruit yield per plant increased by 21.3%-22.4%	潘燕婷,2020
	miR171d/e	SlGRAS24	乙烯生物合成、类胡萝卜素合成Ethylene biosynthesis，carotenoid synthesis	MicroTom	STTM	开花提前，开花至破色时间缩短6 ~ 8 d The flowering time was advanced and the flowering- breaking time was shortened by 6-8 days	蒲慧丽,2021
	miR1917	CPR4	乙烯生物合成 Ethylene biosynthesis	中蔬6号 Zhongshu6	超量表达Overex- pression	加速花梗脱落、果实成熟提前约3~ 4 d Accelerated pedicel abscission，fruit ripening 3-4 d in advance	Wang et al.,2018
	miR156	SPL	赤霉素生物合成 Gibberellin biosynthesis pathway	MicroTom	超量表达Overex- pression	超量表达番茄植株在种子萌发后的第10天，超过80%的顶端仍处于营养分生组织期，而MT顶端超过95%处于花序和花分生组织 At 10DPG，more than 80% of the tops of overexpressing tomato plants were still in vegetative meristem stage，while more than 95% of the tops of MT were in inflorescence and floral meristem	Silva et al.,2019
	miR319	LA	赤霉素生物合成 Gibberellin biosynthesis	MicroTom	靶基因同义突变 Synony- mous mutation	La-1突变体在种子萌发后的第10天，没有处于花序和花分生组织，而MT顶端超过95%处于花序和花分生组织 The La-1 mutant was not in the inflorescence and flower meristem at 10 DPG，while more than 95 % of the MT tip was in the inflorescence and flower meristem	Silva et al.,2019
果实发育 Fruit develop- ment	miR164a	SlNAM2/ SlNAM3	乙烯生物合成 Ethylene biosynthesis	MicroTom	CRISPR/Cas9	开花到成熟开始缩短约5d The time from flowering to maturity was shortened by about 5 days	Lin et al.,2022
	miR396	GRFs	激素途径、细胞壁相关途径Hormone pathway，cell wall related pathway	MicroTom	STTM	果实变大，质量增加约55%，果皮增厚32% ~ 35%，萼片长和宽分别增加约65%和20% The fruit became larger，the fruit weight increased by about 55%，the peel thickness increased by 32%-35%，and the sepal length and width increased by about 65 % and 20 %.	曹东艳,2018
	miR172a	AP2a	乙烯生物合成、类胡萝卜素合成Ethylene biosynthesis，carotenoid synthesis	Ailsa Craig	超量表达Overex- pression	β-胡萝卜素升高约两倍，果实成熟时间比野生型早4 ~ 5 d，萼片向花瓣转变 The β-carotene content increased by about two times，the fruit ripening time was 4-5 days earlier than WT，and sepals were transformed into petals	Chung et al.,2020
	miR159	SlGAMYB1/2	GA和IAA合成 GA and IAA synthesis	MicroTom	超量表达Overex- pression	诱导单性结实、果实100%无种子Induced parthenocarpy，fruit 100 % seedless	Da et al.,2017
	miR160a	SlARF10A/10B/17	IAA合成 IAA synthesis	M82	沉默突变Silent mutation	果实由圆形变为圆锥形，几乎无籽 The fruit changes from round to conical，and the fruit is almost seedless	Mekontso et al.,2023
果实品质 Fruit quality	miR164a	SlNAM2/ SlNAM3	碳水化合物代谢Carbohydrate metabolism	MicroTom	CRISPR/Cas9	淀粉、有机酸含量升高、葡萄糖和果糖含量在开花后30 ~ 35 d增加，成熟后期下降 The content of starch，organic acid，glucose and fructose increased at 30-35 DPA，and decreased at the later stage of maturity	Lin et al.,2022
	miR171b	/	糖酸生物合成 Sugar acid biosynthesis	Ailsa Craig	CRISPR/Cas9	葡萄糖含量上升45%，果糖含量增加2倍，柠檬酸和苹果酸含量下降20%左右 Glucose content increased by 45%，fructose content increased by 2 times，citric acid and malic acid content decreased by about 20%	潘燕婷,2020
	miR394	ZDS	类胡萝卜素代谢通路 Carotenoid metabolism	Micro Tom	STTM	STTM394株系番茄红素和β-类胡萝卜素含量为26.64和17.51 mg · kg^-1，是野生型2.17和1.39倍 The content of lycopene and β-carotene in the STTM394 line was 26.64 mg · kg^-1 and 17.51 mg · kg^-1，which was 2.17 and 1.39 times that of wild type	赵丽,2023
	miR159	MYB65-DFR	花青素合成 Anthocyanin biosynthesis	LA1996	超量表达Overex- pression	果实的花青素浓度比野生型增加1.452 nmol · L^-1 The anthocyanin concentration of fruit was 1.452 nmol · L^-1higher than that of wild type	马旭,2023
	miR160a	SlARF10A/10B/17	类胡萝卜素和番茄红素生物合成 Carotenoid and lycopene biosynthesis	MicroTom	超量表达Overex- pression	在破色期，果实类胡萝卜素含量比野生型高5.19 g · kg^-1，番茄红素、类胡萝卜素含量在整个成熟过程中呈增加趋势 At the color breaking stage，the content of carotenoids in fruits was 5.19 g · kg^-1 higher than that of WT，and the contents of lycopene and carotenoids increased during the whole ripening process	Mekontso et al.,2023
	mi9474	SlGMP/ SlGalUR	抗坏血酸合成 AsA synthesis	Ailsa Craig	超量表达Overex- pression	还原型抗坏血酸含量下降7.61% ~ 14.73%，总抗坏血酸含量下降5.61% ~ 8.04% The content of reduced ascorbic acid decreased by 7.61%-14.73%，and the total ascorbic acid decreased by 5.61%-8.04%.	朱立辉,2023
	miR319b	SlGME1/SlGME2	抗坏血酸合成 AsA synthesis	Ailsa Craig	超量表达Overex- pression	还原型抗坏血酸显著下降12.02% ~ 12.30%，总抗坏血酸下降17.47% ~ 17.87% The content of reduced ascorbic acid decreased significantly by 12.02%-12.30%，and the total ascorbic acid decreased by 17.47%-17.87%	朱立辉,2023
非生物胁迫 Abiotic stress	miR167	NF-YA1/ NF-YA2	叶绿素合成途径、脯氨酸和丙二醛相关途径 Chlorophyll synthesis，proline and malondialdehyde related pathways	中蔬4号Zhongshu 4	超量表达Overex- pression	与对照相比，超量表达植株叶绿素含量和最大光化学效率降低较少、脯氨酸含量增加1.5倍、MDA含量减少1.4倍、番茄对冷胁迫的耐受性提高 Compared with the control，the chlorophyll content and maximum photochemical efficiency of overexpressing plants decreased less，the proline content increased by 1.5 times，the MDA content decreased by 1.4 times，and the tolerance of tomato to cold stress increased	Gou et al.,2014
	miR319d	GAMYB- like1	活性氧系统关键酶 Key enzyme of reactive oxygen system	MicroTom	超量表达Overex- pression	SOD、CAT活性升高，耐冷性增强 The activities of SOD and CAT increased，and the cold tolerance enhanced	Shi et al.,2019
	miR169c	SIMRP1	内质网相关蛋白质降解途径 Endoplasmic reticulum-related protein degradation pathway	Ailsa Craig	超量表达Overex- pression	与WT相比，气孔导度降低33% ~ 45%，蒸腾速率降低45% ~ 62%，提高抗旱性和耐盐性 Compared with WT，the stomatal conductance decreased by 33%-45%，the transpiration rate decreased by 45%-62%，and the drought resistance and salt tolerance increased	Zhang et al.,2011
	miR157	SPL-CNR	侧根和根毛发育相关基因Genes related to lateral root and root hair development	Ailsa Craig	超量表达Overex- pression	高铁螯合物还原酶（FCR）活性显著提高，诱导缺Fe反应，维持根系Fe稳态 The activity of ferric chelate reductase（FCR）was significantly increased，which induced Fe deficiency reaction and maintained Fe homeostasis in roots	Zhu et al.,2022
非生物胁迫 Abiotic stress	miR168a	AGO1	侧根和根毛发育相关基因表达Genes related to lateral root and root hair development	JZ18/JZ34	超量表达Overex- pression	通过增多侧根和根毛、增加根表面积，增强耐低钾能力 By increasing the number of lateral roots and root hairs，increasing the root surface area，the ability of low potassium tolerance was enhanced	刘杨,2018
生物胁迫 Biotic stress	miR482b	NBS-LRR	触发免疫系统（ETI） Trigger the immune system（ETI）	L3708	STTM	致病性、叶片病斑明显较小，相对病斑直径显著降低，MDA水平降低，对番茄晚疫病抗性提高 The pathogenicity and leaf lesions were significantly smaller，the relative lesion diameter was significantly reduced，the MDA level was reduced，and the resistance to tomato late blight was improved	Jiang et al.,2018
	miR397	Lac12	SA和JA信号通路 SA and JA signaling pathway	MicroTom	STTM	STTM397株系病斑面积与叶面积的比值降低近3倍，叶片病斑较浅，对晚疫病抗性提高 The ratio of lesion area to leaf area of STTM397 lines was reduced by nearly 3 times，and the lesion area of leaves was less，and the resistance to late blight was improved	Guan et al.,2023
	miR172a/b	AR2/ERF	SOD、POD酶活性，MDA含量，REL（相对电解质渗漏）值 SOD，POD enzyme activity，MDA content，REL （relative electrolyte leakage）value	Zaofen 2	超量表达 Overex- pression	抗番茄晚疫病，平均病情指数下降17% Resistant to tomato late blight，the average disease index decreased by 17%	Luan et al.,2018
	miR1918	sl-TG2	触发PTI、ETI 免疫系统 Trigger PTI，ETI immune system	Zaofen 2	超量表达 Overex- pression	接种致病疫霉后6和10 d，超量表达株系相对病灶平均面积减少3.9%和25%，平均病情指数减少17%（10 dpi），对番茄晚疫病抗性提高 After inoculation with P.infestans，the average relative lesion area of overexpression lines decreased by 3.9%（6 dpi）and 25%（10 dpi），the average disease index decreased by 17%（10 dpi），and the resistance to tomato late blight increased	Luan et al.,2016
	miR166	SlHB	调控免疫系统 Regulating the immune system	H-88-78-1/Punjab Chhuhara	VIGS靶基因 VIGS target genes	ToLCNDV 感染后，沉默株系疾病严重程度降低、病毒滴度显著降低（约55%）、番茄对ToLCNDV抗性增强 After ToLCNDV infection，the disease severity of the silenced lines was reduced，the virus titer was significantly reduced（about 55%），and the resistance of tomato to ToLCNDV was enhanced	Prasad et al.,2022
	miR159	SlMYB33	HR相关调控 HR-related regulation	H-88-78-1/Punjab Chhuhara	VIGS靶基因 VIGS target genes	抗性材料VIGS沉默株系TYLCV感染后提前出现卷叶和生长迟缓（8 dpi），80%植株表现出典型卷叶，抗性品种中病毒滴度增加 > 4倍 After TYLCV infection，the VIGS silencing of resistant lines showed early leaf rolling and growth retardation（8 dpi），80% of the plants showed typical leaf rolling，and the virus titer in resistant varieties increased by more than 4 times	Sharma et al.,2021
	miR482f/miR5300	Solyc08g075630/Solyc08g076000;Solyc05g008650/Solyc09g018220	PTI、ETI免疫系统 PTI，ETI innate immune system	Motelle/ Money- maker	VIGS靶基因 VIGS target genes	植株生长缓慢，顶部叶片表现出枯萎，导致抗性Motelle品种对尖孢镰刀菌敏感，对番茄枯萎病抗性降低 The plants grew slowly，and the top leaves showed wilting，which led to the sensitivity of the resistant Motelle variety to Fusarium oxysporum and the decrease of resistance to tomato Fusarium wilt	Ouyang et al.,2014

Identification of miRNA in Tomato and Its Application in Genetic Improvement

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