木质素合成代谢调控及其积累对果蔬作物生长发育的影响

doi:10.16420/j.issn.0513-353x.2025-0008

摘要/Abstract

摘要：

木质素是植物细胞壁的关键组分，其合成与调控直接影响果蔬作物的抗逆性和品质。本文中系统综述了果蔬作物中木质素类型、合成代谢途径及调控机制。木质素积累受MYB/NAC转录因子、miRNA及植物激素的协同调控，并受温度、光照等环境因子显著影响。过量木质素会导致果蔬质地硬化，但适度积累可促进伤口愈合和成熟。未来研究应着重于解析木质素合成的生理生化及分子机制，扩大研究对象的多样性，通过对木质素合成与代谢过程的调控，提高其在果蔬品质提升及其他农业生产中的应用潜力。

关键词: 木质素, 果蔬作物, 调控机制, 品质, 抗逆性

Abstract:

Lignin，a pivotal constituent of plant cell walls，exerts a direct influence on the stress tolerance and quality attributes of fruit and vegetable crops through its synthesis and regulation. This paper presents a systematic review of lignin types，biosynthetic pathways，and regulatory mechanisms in fruit and vegetable crops. Studies have revealed that lignin accumulation is subject to the synergistic regulation of MYB/NAC transcription factors，miRNAs，and plant hormones，while also being significantly modulated by environmental cues such as temperature and light. Excessive lignin deposition leads to textural hardening in fruits and vegetables，whereas moderate accumulation facilitates wound healing and promotes ripening processes. Future research endeavors should prioritize the elucidation of the physiological，biochemical，and molecular mechanisms underlying lignin biosynthesis，expand the diversity of research subjects，and enhance the application potential of lignin in fruit and vegetable quality improvement and other agricultural contexts by manipulating lignin synthesis and metabolic pathways.

Key words: lignin, fruit and vegetable crops, regulatory mechanisms, quality, stress resistance

江慧萱, 徐燕萍, 赵彤, 余红瑞, 董正中, 任锡亮, 余小林. 木质素合成代谢调控及其积累对果蔬作物生长发育的影响[J]. 园艺学报, 2026, 53(6): 1803-1819.

JIANG Huixuan, XU Yanping, ZHAO Tong, YU Hongrui, DONG Zhengzhong, REN Xiliang, YU Xiaolin. Advances of Accumulation and Regulation of Lignin in Fruits and Vegetables[J]. Acta Horticulturae Sinica, 2026, 53(6): 1803-1819.

图/表 3

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种类 Type	名称 Name	物种 Species	对木质素含量影响 Influence on lignin content	参考文献 Reference
生长素 Auxins	NAA	梨 Pyrus L.	通过“PbrARF13-PbrNSC-PbrMYB132”调控途径，降低木质素含量 Reduce the lignin content via the“PbrARF13-PbrNSC- PbrMYB132”regulatory pathway	Xu et al.,2023
	IAA	甘薯 Ipomoea batatas （L.）Lam	上调IbSAUR36，抑制木质素相关基因，降低木质素含量 Up-regulate IbSAUR36，inhibit lignin-related genes，and reduce lignin content	Zhou et al.,2024
	IBA	胡萝卜Daucus carota L.	抑制木质素相关基因，降低木质素含量 Inhibit lignin-related genes and reduce lignin content	Khadr et al.,2020
	IAA	柚子 Citrus maxima （Burm.）Merr.	抑制CgMYB58及其靶基因，降低木质素含量 Inhibit CgMYB58 and its target genes，thereby reducing lignin content	Shi et al.,2020
赤霉素 Gibberellin		菜豆 Phaseolus vulgaris L.	上调木质素合成关键酶活性，增加木质素含量 Up-regulate the activity of key enzymes involved in lignin synthesis and increase lignin content	Cheng et al.,2024
		桃 Prunus persica（L.）Batsch	增加木质素含量Increase lignin content	Li et al.,2024b
		胡萝卜D. carota L.	增加木质素含量Increase lignin content	Jia et al.,2024
		樱桃 Cerasus pseudocerasus （Lindl.）G. Don	上调PaLectinL7，调节PaCAD1的酶活性促进木质素沉积 Up-regulate PaLectinL7，modulate the enzymatic activity of PaCAD1，and promote lignin deposition	Wu et al.,2023
细胞分裂素 Cytokinin		胡萝卜D. carota L.	上调木质素合成基因表达，增加木质素含量 Up-regulate the expression of lignin synthesis genes and increase lignin content	Khadr et al.,2020
脱落酸 Abscisic acid		苹果 Malus × domestica Mill.	上调MdSND1，激活MdMYB46和MdMYB83及其靶基因，增加木质素含量 Up-regulate MdSND1，activate MdMYB46 and MdMYB83 as well as their target genes，and increase lignin content	Chen et al.,2020
脱落酸 Abscisic acid		甜瓜 Cucumis melo L.	激活相关酶活性，增加木质素含量 Activate the activity of related enzymes and increase lignin content	Wang et al.,2024a
茉莉酸类 Jasmonates	MeJA	枇杷 Eriobotrya japonica （Thunb.）Lindl.	“EjbHLH14-EjHB1-EjPRX12”调控途径，降低木质素含量 Reduce lignin content via the “EjbHLH14-EjHB1-EjPRX12”regulatory pathway	Zhang et al.,2022b
	MeJA	猕猴桃 Actinidia chinensis Planch.	降低木质素含量 Reduce lignin content	Li et al.,2017
	JA	甜瓜 C. melo L.	上调相关基因，增加木质素含量 Up-regulate related genes and increase lignin content	Liu et al.,2021a
	MeJA	梨 Pyrus L.	促进类黄酮的积累，降低木质素含量 Promote flavonoid accumulation and reduce lignin content	Hong et al.,2024
油菜素内酯类 Brassinosteroids	BRs	葫芦 Lagenaria siceraria （Molina）Standl.	增加木质素含量 Increase lignin content	Liu et al.,2023a
	EBR	乌塌菜 Brassica rapa var. chinensis（L.）Kitam.	降低木质素含量Reduce lignin content	Yuan et al.,2021
	EBR	西瓜 Citrullus lanatus （Thunb.）Matsum. et Nakai	与施用浓度有关Be related to the application concentration	Liu et al.,2023b
乙烯 Ethylene		甘薯I. batatas（L.）Lam	增加木质素含量Increase lignin content	Liu et al.,2024a
乙烯 Ethylene		猕猴桃 A. chinensis Planch.	上调相关基因，增加木质素含量 Up-regulate related genes and increase lignin content	Li et al.,2017
褪黑素 Melatonin		杧果 Mangifera indica L.	激活相关酶活性，增加木质素含量 Activate the activity of related enzymes and increase lignin content	Lu et al.,2024
褪黑素 Melatonin		枇杷 E. japonica （Thunb.）Lindl.	抑制相关酶活性，降低木质素含量 Inhibit the activity of related enzymes and reduce lignin content	Wang et al.,2021b
其他 Others	NO	秋葵 Abelmoschus esculentus（L.）Moench	抑制ABA和乙烯合成，降低木质素含量 Inhibit ABA and ethylene synthesis and reduce lignin content	Sun et al.,2021
	硝普钠SNP	甜瓜 C. melo L.	激活相关酶活性，增加木质素含量 Activate the activity of related enzymes and increase lignin content	Wang et al.,2019a
	c-PTIO	甜瓜 C. melo L.	抑制相关酶活性，降低木质素含量 Inhibit the activity of related enzymes and reduce lignin content	Wang et al.,2019a
	H₂S	枇杷 E. japonica （Thunb.）Lindl.	抑制相关酶活性，降低木质素含量 Inhibit the activity of related enzymes and reduce lignin content	Shen et al.,2024

种类 Type	名称 Name	物种 Species	对木质素含量影响 Influence on lignin content	参考文献 Reference
生长素 Auxins	NAA	梨 Pyrus L.	通过“PbrARF13-PbrNSC-PbrMYB132”调控途径，降低木质素含量 Reduce the lignin content via the“PbrARF13-PbrNSC- PbrMYB132”regulatory pathway	Xu et al.,2023
	IAA	甘薯 Ipomoea batatas （L.）Lam	上调IbSAUR36，抑制木质素相关基因，降低木质素含量 Up-regulate IbSAUR36，inhibit lignin-related genes，and reduce lignin content	Zhou et al.,2024
	IBA	胡萝卜Daucus carota L.	抑制木质素相关基因，降低木质素含量 Inhibit lignin-related genes and reduce lignin content	Khadr et al.,2020
	IAA	柚子 Citrus maxima （Burm.）Merr.	抑制CgMYB58及其靶基因，降低木质素含量 Inhibit CgMYB58 and its target genes，thereby reducing lignin content	Shi et al.,2020
赤霉素 Gibberellin		菜豆 Phaseolus vulgaris L.	上调木质素合成关键酶活性，增加木质素含量 Up-regulate the activity of key enzymes involved in lignin synthesis and increase lignin content	Cheng et al.,2024
		桃 Prunus persica（L.）Batsch	增加木质素含量Increase lignin content	Li et al.,2024b
		胡萝卜D. carota L.	增加木质素含量Increase lignin content	Jia et al.,2024
		樱桃 Cerasus pseudocerasus （Lindl.）G. Don	上调PaLectinL7，调节PaCAD1的酶活性促进木质素沉积 Up-regulate PaLectinL7，modulate the enzymatic activity of PaCAD1，and promote lignin deposition	Wu et al.,2023
细胞分裂素 Cytokinin		胡萝卜D. carota L.	上调木质素合成基因表达，增加木质素含量 Up-regulate the expression of lignin synthesis genes and increase lignin content	Khadr et al.,2020
脱落酸 Abscisic acid		苹果 Malus × domestica Mill.	上调MdSND1，激活MdMYB46和MdMYB83及其靶基因，增加木质素含量 Up-regulate MdSND1，activate MdMYB46 and MdMYB83 as well as their target genes，and increase lignin content	Chen et al.,2020
脱落酸 Abscisic acid		甜瓜 Cucumis melo L.	激活相关酶活性，增加木质素含量 Activate the activity of related enzymes and increase lignin content	Wang et al.,2024a
茉莉酸类 Jasmonates	MeJA	枇杷 Eriobotrya japonica （Thunb.）Lindl.	“EjbHLH14-EjHB1-EjPRX12”调控途径，降低木质素含量 Reduce lignin content via the “EjbHLH14-EjHB1-EjPRX12”regulatory pathway	Zhang et al.,2022b
	MeJA	猕猴桃 Actinidia chinensis Planch.	降低木质素含量 Reduce lignin content	Li et al.,2017
	JA	甜瓜 C. melo L.	上调相关基因，增加木质素含量 Up-regulate related genes and increase lignin content	Liu et al.,2021a
	MeJA	梨 Pyrus L.	促进类黄酮的积累，降低木质素含量 Promote flavonoid accumulation and reduce lignin content	Hong et al.,2024
油菜素内酯类 Brassinosteroids	BRs	葫芦 Lagenaria siceraria （Molina）Standl.	增加木质素含量 Increase lignin content	Liu et al.,2023a
	EBR	乌塌菜 Brassica rapa var. chinensis（L.）Kitam.	降低木质素含量Reduce lignin content	Yuan et al.,2021
	EBR	西瓜 Citrullus lanatus （Thunb.）Matsum. et Nakai	与施用浓度有关Be related to the application concentration	Liu et al.,2023b
乙烯 Ethylene		甘薯I. batatas（L.）Lam	增加木质素含量Increase lignin content	Liu et al.,2024a
乙烯 Ethylene		猕猴桃 A. chinensis Planch.	上调相关基因，增加木质素含量 Up-regulate related genes and increase lignin content	Li et al.,2017
褪黑素 Melatonin		杧果 Mangifera indica L.	激活相关酶活性，增加木质素含量 Activate the activity of related enzymes and increase lignin content	Lu et al.,2024
褪黑素 Melatonin		枇杷 E. japonica （Thunb.）Lindl.	抑制相关酶活性，降低木质素含量 Inhibit the activity of related enzymes and reduce lignin content	Wang et al.,2021b
其他 Others	NO	秋葵 Abelmoschus esculentus（L.）Moench	抑制ABA和乙烯合成，降低木质素含量 Inhibit ABA and ethylene synthesis and reduce lignin content	Sun et al.,2021
	硝普钠SNP	甜瓜 C. melo L.	激活相关酶活性，增加木质素含量 Activate the activity of related enzymes and increase lignin content	Wang et al.,2019a
	c-PTIO	甜瓜 C. melo L.	抑制相关酶活性，降低木质素含量 Inhibit the activity of related enzymes and reduce lignin content	Wang et al.,2019a
	H₂S	枇杷 E. japonica （Thunb.）Lindl.	抑制相关酶活性，降低木质素含量 Inhibit the activity of related enzymes and reduce lignin content	Shen et al.,2024

环境因素 Environmental factor	处理 Treatment	物种 Species		对木质素含量的影响 Effect on lignin content	参考文献 Reference
温度 Temperature	低温 Low temperature	茭白Zizania latifoli（Griseb.）Turcz. ex Stapf		上调相关基因，增加木质素含量 Up-regulate related genes and increase lignin content	Qian et al.,2023
		血橙Citrus × sinensis‘Blood Orange’		上调相关基因，增加木质素含量 Up-regulate related genes and increase lignin content	Xie et al.,2022
		香蕉 Musa acuminate‘（AAA）’		上调相关基因，增加木质素含量 Up-regulate related genes and increase lignin content	Xiao et al.,2023
		大白菜 B. rapa subsp. pekinensis		上调相关基因，增加木质素含量 Up-regulate related genes and increase lignin content	Dai et al.,2022
		茭白Z. latifolia （Griseb.）Turcz. ex Stapf		抑制相关酶活性，降低木质素含量 Inhibit the activity of related enzymes and reduce lignin content	Yang et al.,2022a
		甘蔗Saccharum officinarum L.		下调相关基因，降低木质素含量 Down-regulate related genes and reduce lignin content	Santos et al.,2015
		甘蔗S. officinarum L.		上调相关基因，增加木质素含量 Up-regulate related genes and increase lignin content	Santos et al.,2015
	高温 Hight temperature	木瓜 Carica papaya L.		上调相关基因，增加木质素含量 Up-regulate related genes and increase lignin content	Rahman et al.,2024
	高温 Hight temperature	柑橘C. reticulata Blanco		增加木质素含量 Increase lignin content	Zandalinas et al.,2017
肥料 Fertilizer	Si	大豆 Glycine max（L.）Merr.		激活相关酶活性，增加木质素含量 Activate the activity of related enzymes and increase lignin content	Sajad et al.,2021
	Si	番茄 Solanum lycopersicum L.		上调相关基因，增加木质素含量 Up-regulate related genes and increase lignin content	Xie et al.,2024
	Si	甘蓝型油菜 B. napus L.		上调相关酶活性和基因表达，增加木质素含量 Up-regulate the activity of related enzymes and the expression of related genes, and increase lignin content	Hu et al.,2022
	Se	辣椒Capsicum annuum L.		上调相关基因，增加木质素含量 Up-regulate related genes and increase lignin content	Li et al.,2021
	Mn	黄瓜C. sativus L.		增加木质素含量Increase lignin content	Eskandari & Sharifnabi,2019
盐 Salt	不同浓度 Different concentrations		马齿苋 Portulaca oleracea L.	相关基因差异表达，调节木质素含量 Differential expression of related genes regulates lignin content	Li et al.,2023
	盐处理 Salt treatment		苹果 M. × domestica Borkh.	诱导MdSND1激活MdMYB46/83的转录，增加木质素含量 Induce MdSND1 to activate the transcription of MdMYB46/83, thereby increasing lignin content	Chen et al.,2020
	盐胁迫 Salt stress		胡萝卜 D. carota L.	上调相关基因，增加木质素含量 Up-regulate related genes and increase lignin content	陈皓炜等,2021
	盐胁迫 Salt stress		甜樱桃 P. avium L.	上调PaLectinL7表达，激活相关酶活性，增加木质素含量 Up-regulate the expression of PaLectinL7，activate the activity of related enzymes，and increase lignin content	Wu et al.,2023
	高盐环境 High-salt environment		葫芦 L. siceraria	激活相关酶活性，增加木质素含量 Activate the activity of related enzymes and increase lignin content	Liu et al.,2023a
	盐度增加Increase in salinity		叶用莴苣 Lactuca sativa	木质素含量随着盐度的增加而增加 Lignin content increased with the increase in salinity	Garrido et al.,2014
	盐胁迫 Salt stress		甜瓜 C. melo L.	激活相关酶活性，增加木质素含量 Activate the activity of related enzymes and increase lignin content	Liu et al.,2018
水 Water	干旱Drought		鹰嘴豆 Cicer arietinum L.	调控相关基因表达，最终导致增加木质素含量 Regulate the expression of related genes, ultimately leading to an increase in lignin content	Sharma et al.,2020
			甜瓜 C. melo L.	上调相关基因，增加木质素含量 Up-regulate related genes and increase lignin content	Liu et al.,2021b
			苹果 M. × domestica Borkh.	MdMRLK2受体样激酶激活相关酶活性，增加木质素含量 The receptor-like kinase MdMRLK2 activates the activity of related enzymes，thereby increasing lignin content	Jing et al.,2024
			甜瓜 C. melo L.	激活相关酶活性，增加木质素含量 Activate the activity of related enzymes and increase lignin content	Liu et al.,2021b
			猕猴桃 A. chinensis Planch.	激活相关酶活性，增加木质素含量 Activate the activity of related enzymes and increase lignin content	Wang et al.,2024c
	涝害Water logging injury		茄子 S. melongena L.	调控 miRNA，增加木质素含量 Regulate miRNAs and increase lignin content	Jiang et al.,2023
	涝害Water logging injury		玉米 Zea mays L.	降低木质素含量 Reduce lignin content	王倩等,2024
光 Light	遮荫 Shading		大豆 G. max（L.）Merr.	下调相关基因，降低木质素含量 Down-regulate related genes and reduce lignin content	Liu et al.,2018
	遮荫 Shading		芦笋 Asparagus officinalis L.	下调相关基因，降低木质素含量 Down-regulate related genes and reduce lignin content	Ma et al.,2023
	黑暗处理 Dark treatment		芹菜 Apium graveolens	下调相关基因，降低木质素含量 Down-regulate related genes and reduce lignin content	Zhu et al.,2024
	蓝光 Blue light		梨 Pyrus L.	上调相关基因，增加木质素含量 Up-regulate related genes and increase lignin content	Wang et al.,2022
	蓝光（低光强） Blue light (low light intensity)		大豆 G. max（L.）Merr.	上调相关基因，增加木质素含量 Up-regulate related genes and increase lignin content	He et al.,2024
	蓝光与红光（3:1） Blue light and red light (3:1)		甜瓜 C. melo L.	上调相关基因，增加木质素含量 Up-regulate related genes and increase lignin content	Jing et al.,2018
	红蓝光（3:1） Red and blue light (3:1)		叶用莴苣 L. sativa L.	下调相关基因，降低木质素含量 Down-regulate related genes and reduce lignin content	Sng et al.,2021
重金属 Heavy metals	Cu		芥菜 B. juncea（L.）Czern. et Coss.	增加木质素含量 Increase lignin content	Rather et al.,2022
	Cu		辣椒 C. annuum L.	抑制相关酶活性，降低木质素含量 Inhibit the activity of related enzymes and reduce lignin content	Dı́az et al.,2001
重金属 Heavy metal	Cd		叶用莴苣 L. sativa L.	增加木质素含量 Increase lignin content	Gao et al.,2022
	Cd		芥菜 B. juncea（L.） Czern. et Coss.	激活相关酶活性，增加木质素含量Activate the activity of related enzymes and increase lignin content	Peng et al.,2024
	较低浓度Cd/Pb Low concentration of Cd/Pb		大豆 G. max（L.）Merr.	相关基因转录水平提高，增加木质素含量 The transcription levels of related genes are increased, thereby enhancing lignin content	Pawlak et al.,2011
	较低浓度Cd Low concentration of Cd		羽扇豆 Lupinus micranthus Guss.	相关基因转录水平下降，降低木质素含量 The transcription levels of related genes are decreased，thereby reducing lignin content	Pawlak et al.,2011
	较低浓度Pb Low concentration of Pb		羽扇豆 L. micranthus Guss.	相关基因转录水平提高，增加木质素含量 The transcription levels of related genes are increased，thereby enhancing lignin content	Pawlak et al.,2011
	较高浓度Cd/Pb Low concentration of Cd/Pb		大豆/羽扇豆 G. max（L.）Merr./ L. micranthus Guss.	激活相关酶活性，增加木质素含量 Activate the activity of related enzymes and increase lignin content	Pawlak et al.,2011
其他化合物 Other compounds	植酸 Phytic acid		苹果 M. pumila Mill.	抑制相关酶活性，降低木质素含量Inhibit the activity of related enzymes and reduce lignin content	Fang et al.,2022
	DPI		茭白Z. latifolia（Griseb.） Turcz. ex Stapf	下调相关基因，降低木质素含量 Down-regulate related genes and reduce lignin content	Qian et al.,2023
	H₂O₂		茭白Z. latifolia（Griseb.） Turcz. ex Stapf	上调相关基因，增加木质素含量 Up-regulate the activity of related enzymes and the expression of related genes, and increase lignin content	Qian et al.,2023
	H₂O₂		甜瓜C. melo L.	上调相关基因，增加木质素含量 Up-regulate the activity of related enzymes and the expression of related genes, and increase lignin content	Liu et al.,2021a
	TKL		秋葵A. esculentus（L.）Moench	激活相关酶活性，增加木质素含量Activate the activity of related enzymes and increase lignin content	Zhang et al.,2023
	CO₂		胡萝卜 D. carota subsp. sativa （Hoffm.）Arcang.	增加木质素含量 Increase lignin content	Wang et al.,2018
	蛋氨酸处理 Methionine treatment		枣 Ziziphus jujuba Mill.	上调相关酶活性和基因表达，增加木质素含量 Up-regulate the activity of related enzymes and the expression of related genes, and increase lignin content	Liu et al.,2022
	琼脂低聚糖处理 Agar oligosaccharide treatment		桃 P. persica（L.）Batsch	上调相关酶活性和基因表达，增加木质素含量 Up-regulate the activity of related enzymes and the expression of related genes, and increase lignin content	Li et al.,2024
机械损伤 Mechanical damage	机械损伤 Mechanical damage		甜瓜 C. melo L.	激活相关酶活性，增加木质素含量Activate the activity of related enzymes and increase lignin content	Liu et al.,2018
病害 Plant disease	核盘菌感染 Sclerotinia sclerotiorum infection		甘蓝型油菜 B. napus L.	上调相关基因，增加木质素含量 Up-regulate related genes and increase lignin content	Kerstin et al.,2021
	炭疽病菌接种Colletotrichum inoculation		柿 Diospyros kaki L.f.	激活DkWRKY8和DkWRKY10转录因子，上调相关基因表达，增加木质素含量 Activate transcription factors DkWRKY8 and DkWRKY10, up-regulate the expression of related genes, and increase lignin content	Fan et al.,2024
	胡萝卜欧文氏菌感染 Erwinia carotovora infection		大白菜 B. rapa subsp. pekinensis	上调相关基因，增加木质素含量 Up-regulate related genes and increase lignin content	Zhang et al.,2007

环境因素 Environmental factor	处理 Treatment	物种 Species		对木质素含量的影响 Effect on lignin content	参考文献 Reference
温度 Temperature	低温 Low temperature	茭白Zizania latifoli（Griseb.）Turcz. ex Stapf		上调相关基因，增加木质素含量 Up-regulate related genes and increase lignin content	Qian et al.,2023
		血橙Citrus × sinensis‘Blood Orange’		上调相关基因，增加木质素含量 Up-regulate related genes and increase lignin content	Xie et al.,2022
		香蕉 Musa acuminate‘（AAA）’		上调相关基因，增加木质素含量 Up-regulate related genes and increase lignin content	Xiao et al.,2023
		大白菜 B. rapa subsp. pekinensis		上调相关基因，增加木质素含量 Up-regulate related genes and increase lignin content	Dai et al.,2022
		茭白Z. latifolia （Griseb.）Turcz. ex Stapf		抑制相关酶活性，降低木质素含量 Inhibit the activity of related enzymes and reduce lignin content	Yang et al.,2022a
		甘蔗Saccharum officinarum L.		下调相关基因，降低木质素含量 Down-regulate related genes and reduce lignin content	Santos et al.,2015
		甘蔗S. officinarum L.		上调相关基因，增加木质素含量 Up-regulate related genes and increase lignin content	Santos et al.,2015
	高温 Hight temperature	木瓜 Carica papaya L.		上调相关基因，增加木质素含量 Up-regulate related genes and increase lignin content	Rahman et al.,2024
	高温 Hight temperature	柑橘C. reticulata Blanco		增加木质素含量 Increase lignin content	Zandalinas et al.,2017
肥料 Fertilizer	Si	大豆 Glycine max（L.）Merr.		激活相关酶活性，增加木质素含量 Activate the activity of related enzymes and increase lignin content	Sajad et al.,2021
	Si	番茄 Solanum lycopersicum L.		上调相关基因，增加木质素含量 Up-regulate related genes and increase lignin content	Xie et al.,2024
	Si	甘蓝型油菜 B. napus L.		上调相关酶活性和基因表达，增加木质素含量 Up-regulate the activity of related enzymes and the expression of related genes, and increase lignin content	Hu et al.,2022
	Se	辣椒Capsicum annuum L.		上调相关基因，增加木质素含量 Up-regulate related genes and increase lignin content	Li et al.,2021
	Mn	黄瓜C. sativus L.		增加木质素含量Increase lignin content	Eskandari & Sharifnabi,2019
盐 Salt	不同浓度 Different concentrations		马齿苋 Portulaca oleracea L.	相关基因差异表达，调节木质素含量 Differential expression of related genes regulates lignin content	Li et al.,2023
	盐处理 Salt treatment		苹果 M. × domestica Borkh.	诱导MdSND1激活MdMYB46/83的转录，增加木质素含量 Induce MdSND1 to activate the transcription of MdMYB46/83, thereby increasing lignin content	Chen et al.,2020
	盐胁迫 Salt stress		胡萝卜 D. carota L.	上调相关基因，增加木质素含量 Up-regulate related genes and increase lignin content	陈皓炜等,2021
	盐胁迫 Salt stress		甜樱桃 P. avium L.	上调PaLectinL7表达，激活相关酶活性，增加木质素含量 Up-regulate the expression of PaLectinL7，activate the activity of related enzymes，and increase lignin content	Wu et al.,2023
	高盐环境 High-salt environment		葫芦 L. siceraria	激活相关酶活性，增加木质素含量 Activate the activity of related enzymes and increase lignin content	Liu et al.,2023a
	盐度增加Increase in salinity		叶用莴苣 Lactuca sativa	木质素含量随着盐度的增加而增加 Lignin content increased with the increase in salinity	Garrido et al.,2014
	盐胁迫 Salt stress		甜瓜 C. melo L.	激活相关酶活性，增加木质素含量 Activate the activity of related enzymes and increase lignin content	Liu et al.,2018
水 Water	干旱Drought		鹰嘴豆 Cicer arietinum L.	调控相关基因表达，最终导致增加木质素含量 Regulate the expression of related genes, ultimately leading to an increase in lignin content	Sharma et al.,2020
			甜瓜 C. melo L.	上调相关基因，增加木质素含量 Up-regulate related genes and increase lignin content	Liu et al.,2021b
			苹果 M. × domestica Borkh.	MdMRLK2受体样激酶激活相关酶活性，增加木质素含量 The receptor-like kinase MdMRLK2 activates the activity of related enzymes，thereby increasing lignin content	Jing et al.,2024
			甜瓜 C. melo L.	激活相关酶活性，增加木质素含量 Activate the activity of related enzymes and increase lignin content	Liu et al.,2021b
			猕猴桃 A. chinensis Planch.	激活相关酶活性，增加木质素含量 Activate the activity of related enzymes and increase lignin content	Wang et al.,2024c
	涝害Water logging injury		茄子 S. melongena L.	调控 miRNA，增加木质素含量 Regulate miRNAs and increase lignin content	Jiang et al.,2023
	涝害Water logging injury		玉米 Zea mays L.	降低木质素含量 Reduce lignin content	王倩等,2024
光 Light	遮荫 Shading		大豆 G. max（L.）Merr.	下调相关基因，降低木质素含量 Down-regulate related genes and reduce lignin content	Liu et al.,2018
	遮荫 Shading		芦笋 Asparagus officinalis L.	下调相关基因，降低木质素含量 Down-regulate related genes and reduce lignin content	Ma et al.,2023
	黑暗处理 Dark treatment		芹菜 Apium graveolens	下调相关基因，降低木质素含量 Down-regulate related genes and reduce lignin content	Zhu et al.,2024
	蓝光 Blue light		梨 Pyrus L.	上调相关基因，增加木质素含量 Up-regulate related genes and increase lignin content	Wang et al.,2022
	蓝光（低光强） Blue light (low light intensity)		大豆 G. max（L.）Merr.	上调相关基因，增加木质素含量 Up-regulate related genes and increase lignin content	He et al.,2024
	蓝光与红光（3:1） Blue light and red light (3:1)		甜瓜 C. melo L.	上调相关基因，增加木质素含量 Up-regulate related genes and increase lignin content	Jing et al.,2018
	红蓝光（3:1） Red and blue light (3:1)		叶用莴苣 L. sativa L.	下调相关基因，降低木质素含量 Down-regulate related genes and reduce lignin content	Sng et al.,2021
重金属 Heavy metals	Cu		芥菜 B. juncea（L.）Czern. et Coss.	增加木质素含量 Increase lignin content	Rather et al.,2022
	Cu		辣椒 C. annuum L.	抑制相关酶活性，降低木质素含量 Inhibit the activity of related enzymes and reduce lignin content	Dı́az et al.,2001
重金属 Heavy metal	Cd		叶用莴苣 L. sativa L.	增加木质素含量 Increase lignin content	Gao et al.,2022
	Cd		芥菜 B. juncea（L.） Czern. et Coss.	激活相关酶活性，增加木质素含量Activate the activity of related enzymes and increase lignin content	Peng et al.,2024
	较低浓度Cd/Pb Low concentration of Cd/Pb		大豆 G. max（L.）Merr.	相关基因转录水平提高，增加木质素含量 The transcription levels of related genes are increased, thereby enhancing lignin content	Pawlak et al.,2011
	较低浓度Cd Low concentration of Cd		羽扇豆 Lupinus micranthus Guss.	相关基因转录水平下降，降低木质素含量 The transcription levels of related genes are decreased，thereby reducing lignin content	Pawlak et al.,2011
	较低浓度Pb Low concentration of Pb		羽扇豆 L. micranthus Guss.	相关基因转录水平提高，增加木质素含量 The transcription levels of related genes are increased，thereby enhancing lignin content	Pawlak et al.,2011
	较高浓度Cd/Pb Low concentration of Cd/Pb		大豆/羽扇豆 G. max（L.）Merr./ L. micranthus Guss.	激活相关酶活性，增加木质素含量 Activate the activity of related enzymes and increase lignin content	Pawlak et al.,2011
其他化合物 Other compounds	植酸 Phytic acid		苹果 M. pumila Mill.	抑制相关酶活性，降低木质素含量Inhibit the activity of related enzymes and reduce lignin content	Fang et al.,2022
	DPI		茭白Z. latifolia（Griseb.） Turcz. ex Stapf	下调相关基因，降低木质素含量 Down-regulate related genes and reduce lignin content	Qian et al.,2023
	H₂O₂		茭白Z. latifolia（Griseb.） Turcz. ex Stapf	上调相关基因，增加木质素含量 Up-regulate the activity of related enzymes and the expression of related genes, and increase lignin content	Qian et al.,2023
	H₂O₂		甜瓜C. melo L.	上调相关基因，增加木质素含量 Up-regulate the activity of related enzymes and the expression of related genes, and increase lignin content	Liu et al.,2021a
	TKL		秋葵A. esculentus（L.）Moench	激活相关酶活性，增加木质素含量Activate the activity of related enzymes and increase lignin content	Zhang et al.,2023
	CO₂		胡萝卜 D. carota subsp. sativa （Hoffm.）Arcang.	增加木质素含量 Increase lignin content	Wang et al.,2018
	蛋氨酸处理 Methionine treatment		枣 Ziziphus jujuba Mill.	上调相关酶活性和基因表达，增加木质素含量 Up-regulate the activity of related enzymes and the expression of related genes, and increase lignin content	Liu et al.,2022
	琼脂低聚糖处理 Agar oligosaccharide treatment		桃 P. persica（L.）Batsch	上调相关酶活性和基因表达，增加木质素含量 Up-regulate the activity of related enzymes and the expression of related genes, and increase lignin content	Li et al.,2024
机械损伤 Mechanical damage	机械损伤 Mechanical damage		甜瓜 C. melo L.	激活相关酶活性，增加木质素含量Activate the activity of related enzymes and increase lignin content	Liu et al.,2018
病害 Plant disease	核盘菌感染 Sclerotinia sclerotiorum infection		甘蓝型油菜 B. napus L.	上调相关基因，增加木质素含量 Up-regulate related genes and increase lignin content	Kerstin et al.,2021
	炭疽病菌接种Colletotrichum inoculation		柿 Diospyros kaki L.f.	激活DkWRKY8和DkWRKY10转录因子，上调相关基因表达，增加木质素含量 Activate transcription factors DkWRKY8 and DkWRKY10, up-regulate the expression of related genes, and increase lignin content	Fan et al.,2024
	胡萝卜欧文氏菌感染 Erwinia carotovora infection		大白菜 B. rapa subsp. pekinensis	上调相关基因，增加木质素含量 Up-regulate related genes and increase lignin content	Zhang et al.,2007