黄瓜嫁接愈合进程及相关影响因素研究进展

doi:10.16420/j.issn.0513-353x.2024-0688

摘要/Abstract

摘要：

嫁接可有效提升植株抗逆能力、提高产量及果实品质，已广泛应用于黄瓜生产栽培。嫁接成活率与嫁接苗品质受到嫁接技术及愈合环境等多因素共同影响，明确黄瓜嫁接愈合进程相关机理与影响因素、探究无损检测技术在黄瓜嫁接愈合中的应用方式对工厂化嫁接育苗体系优化有一定指导意义。本文对黄瓜嫁接愈合进程的组织学、细胞学、生理生化、分子、表型组学机理进行了总结，综合分析了砧木—接穗亲和性、嫁接方法与苗龄、激素、养分、环境因子对嫁接愈合的影响，并从黄瓜嫁接愈合无损检测模型建立、外源物质施用方式探究、嫁接方法对嫁接愈合的内在影响机理解析、嫁接愈合关键基因的功能验证与环境因素对嫁接植株内源物质合成转运的影响五个方面对黄瓜嫁接愈合机理及生产应用进行展望。旨在为后续生产过程中黄瓜嫁接苗无损化检测、嫁接技术优化与愈合环境精准化调控提供参考。

关键词: 黄瓜, 嫁接, 愈合进程, 嫁接愈合环境

Abstract:

Grafting has been extensively utilized in cucumber production and cultivation，significantly boosting plant stress resistance，yield，and fruit quality. The success rate of grafting and the quality of grafted seedlings are jointly influenced by various factors，including grafting techniques and healing conditions. Clarifying the mechanisms and influencing factors associated with the healing process of cucumber grafting，as well as exploring the application of non-destructive testing techniques in this context，holds considerable significance for optimizing industrialized grafting seedling production systems. Consequently，this paper summarizes the histological，cytological，physiological and biochemical，molecular，and phenotypic omics mechanisms underlying the cucumber grafting healing process. It also reviews the impacts of stock-scion compatibility，grafting methods，seedling age，hormones，nutrients，and environmental factors on the healing process. Additionally，the paper provides insights into cucumber grafting healing mechanisms and production applications from five perspectives：the establishment of non-destructive testing models for cucumber grafting healing，the exploration of exogenous substance application methods，the analysis of the internal mechanisms by which grafting methods affect healing，the functional validation of key genes related to grafting healing，and the influence of environmental factors on the synthesis and transport of endogenous substances in grafted plants. The ultimate goal is to offer a reference for non-destructive testing of grafted cucumber seedlings，optimization of grafting techniques，and precise regulation of healing environments in subsequent production processes.

Key words: cucumber, graft, healing process, graft healing environment

林黄昉, 程赛川, 林碧英, 陈琰臻, 叶惠澜, 邵卿卿, 林鸿辉, 翟挺楷, 王锴, 申宝营. 黄瓜嫁接愈合进程及相关影响因素研究进展[J]. 园艺学报, 2025, 52(10): 2799-2815.

LIN Huangfang, CHENG Saichuan, LIN Biying, CHEN Yanzhen, YE Huilan, SHAO Qingqing, LIN Honghui, ZHAI Tingkai, WANG Kai, SHEN Baoying. Research Progress on the Healing Process of Cucumber Grafting and Related Influencing Factors[J]. Acta Horticulturae Sinica, 2025, 52(10): 2799-2815.

图/表 2

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功能类别 Functional category	具体分类 Specific classification	相关基因 Related gene	主要功能 Key feature	参考文献 Reference
激素响应 Hormone response	生长素 Auxin	IAA、ARF、ALF4	与JAs、ANAC069、ANAC071互作，促进愈伤组织与维管束形成，加快嫁接愈合 Interact with JAs，ANAC069，and ANAC071 to promote the formation of callus and vascular bundles，and accelerate graft healing	Matsuoka et al.,2021
	乙烯 Ethylene	RAP2.6L、ERF114、ERF115	通过上调ANAC071、ERF114、ERF115，抑制RAP2.6L，加快嫁接愈合 Up-regulate ANAC071，ERF114 and ERF115，inhibit RAP2.6L and accelerating graft healing	马庆等,2013
	细胞分裂素 Cytokinin	-	诱导PIN蛋白合成，促进生长素物质运输，加快嫁接愈合 Induce PIN protein synthesis，promote auxin material transport，and accelerate graft healing	谢露露等,2020
	脱落酸 Abscisic acid	-	与生长素共同调控CmoANT，加快嫁接愈合 Co-regulate CmoANT with auxin to speed up graft healing	谢露露等,2020
	褪黑素 Melatonin	-	上调CAD、HCT表达水平，促进木质素合成，加快嫁接愈合 Up-regulate CAD and HCT，promote lignin synthesis，and accelerating graft healing	Zhu et al.,2024a
组织重连 Organizational reconnection	细胞壁代谢基因 Cell wall metabolism gene	PMEI5	抑制PME合成，提高下胚轴细胞硬度，加快嫁接愈合 Inhibit PME synthesis，improve hypocotyl cell stiffness，and accelerate graft healing	Zhang et al.,2021
	细胞壁代谢基因 Cell wall metabolism gene	GH9B3	正向调控激素信号转导、MAPK途径与蔗糖合成途径，促进木质素积累，加快嫁接愈合 Positively regulate hormone signal transduction, MAPK pathway and sucrose synthesis pathway，promote lignin accumulation，and accelerate graft healing	Zhu et al.,2023,2024b
	转录因子 Transcription factor	ANAC069、ANAC071	促进韧皮部重连，加快嫁接愈合 Promote phloem reconnection and speed up graft healing	Matsuoka et al.,2021
		WOX3、WOX4、WOX13、WOX14	调控PXY、EXP、PLL、GH9B3等基因，促进形成层细胞增殖分化与木质素积累，加快嫁接愈合 Regulate genes such as PXY，EXP，PLL，GH9B3，promote the proliferation and differentiation of cambium cells and the accumulation of lignin，and accelerate graft healing	Etchells et al.,2013；Melnyk,2016,2017；Cui et al.,2021；Thomas et al.,2021；Ikeuchi et al.,2022
		DOF6、DOF2.1、HCA2、TMO6	促进细胞壁修复，其中TMO6可与纤维素CEL3结合，共同加快嫁接愈合 Promotes cell wall repair，in which TMO6 can bind to cellulose CEL3 to work together to speed up graft healing	Zhang et al.,2021
环境因子响应 Environmental factor response	光 Light	PAT1	促进愈伤组织形成，加快嫁接愈合 Promote callus formation and speed up graft healing	Feng et al.,2024
环境因子响应 Environmental factor response	温度 Temperature	PHYB、PIF4	形成PHYB-PIF4-生长素模块响应高温，加快嫁接愈合 The formation of PHYB-PIF4-auxin module responds to high temperature and accelerates graft healing	Meng et al.,2025

功能类别 Functional category	具体分类 Specific classification	相关基因 Related gene	主要功能 Key feature	参考文献 Reference
激素响应 Hormone response	生长素 Auxin	IAA、ARF、ALF4	与JAs、ANAC069、ANAC071互作，促进愈伤组织与维管束形成，加快嫁接愈合 Interact with JAs，ANAC069，and ANAC071 to promote the formation of callus and vascular bundles，and accelerate graft healing	Matsuoka et al.,2021
	乙烯 Ethylene	RAP2.6L、ERF114、ERF115	通过上调ANAC071、ERF114、ERF115，抑制RAP2.6L，加快嫁接愈合 Up-regulate ANAC071，ERF114 and ERF115，inhibit RAP2.6L and accelerating graft healing	马庆等,2013
	细胞分裂素 Cytokinin	-	诱导PIN蛋白合成，促进生长素物质运输，加快嫁接愈合 Induce PIN protein synthesis，promote auxin material transport，and accelerate graft healing	谢露露等,2020
	脱落酸 Abscisic acid	-	与生长素共同调控CmoANT，加快嫁接愈合 Co-regulate CmoANT with auxin to speed up graft healing	谢露露等,2020
	褪黑素 Melatonin	-	上调CAD、HCT表达水平，促进木质素合成，加快嫁接愈合 Up-regulate CAD and HCT，promote lignin synthesis，and accelerating graft healing	Zhu et al.,2024a
组织重连 Organizational reconnection	细胞壁代谢基因 Cell wall metabolism gene	PMEI5	抑制PME合成，提高下胚轴细胞硬度，加快嫁接愈合 Inhibit PME synthesis，improve hypocotyl cell stiffness，and accelerate graft healing	Zhang et al.,2021
	细胞壁代谢基因 Cell wall metabolism gene	GH9B3	正向调控激素信号转导、MAPK途径与蔗糖合成途径，促进木质素积累，加快嫁接愈合 Positively regulate hormone signal transduction, MAPK pathway and sucrose synthesis pathway，promote lignin accumulation，and accelerate graft healing	Zhu et al.,2023,2024b
	转录因子 Transcription factor	ANAC069、ANAC071	促进韧皮部重连，加快嫁接愈合 Promote phloem reconnection and speed up graft healing	Matsuoka et al.,2021
		WOX3、WOX4、WOX13、WOX14	调控PXY、EXP、PLL、GH9B3等基因，促进形成层细胞增殖分化与木质素积累，加快嫁接愈合 Regulate genes such as PXY，EXP，PLL，GH9B3，promote the proliferation and differentiation of cambium cells and the accumulation of lignin，and accelerate graft healing	Etchells et al.,2013；Melnyk,2016,2017；Cui et al.,2021；Thomas et al.,2021；Ikeuchi et al.,2022
		DOF6、DOF2.1、HCA2、TMO6	促进细胞壁修复，其中TMO6可与纤维素CEL3结合，共同加快嫁接愈合 Promotes cell wall repair，in which TMO6 can bind to cellulose CEL3 to work together to speed up graft healing	Zhang et al.,2021
环境因子响应 Environmental factor response	光 Light	PAT1	促进愈伤组织形成，加快嫁接愈合 Promote callus formation and speed up graft healing	Feng et al.,2024
环境因子响应 Environmental factor response	温度 Temperature	PHYB、PIF4	形成PHYB-PIF4-生长素模块响应高温，加快嫁接愈合 The formation of PHYB-PIF4-auxin module responds to high temperature and accelerates graft healing	Meng et al.,2025

砧木品类 Rootstock category	砧木品种 Rootstock cultivar	接穗品种 Scion	嫁接方法 Grafting method	成活率/% Survival rate	主要性状 Main characters			参考文献 References
砧木品类 Rootstock category	砧木品种 Rootstock cultivar	接穗品种 Scion	嫁接方法 Grafting method	成活率/% Survival rate	果实产量/ （kg · hm^-2） Fruit yield	果实品质 Fruit quality	植株抗逆能力 Plant stress resistance	参考文献 References
南瓜 Cucurbita L. .	京欣砧5号 Jingxin- zhen 5	津优35 Jinyou 35	插接法 Peg grafting	97.00	277.47	果皮无蜡粉；果肉深绿 Peel without wax powder；Fruit flesh dark green	耐低温；抗猝倒病、枯萎病、霜霉病 Low temperature resistance；Resistance to Damping off disease，Fusarium Wilt and downy mildew	张帆等,2011
	金妈妈1989 Kim Mama 1989	北农佳秀 Beinong Jiaxiu	贴接法 Patch grafting	94.00	581.67	果皮无蜡粉；果肉浅绿，甜脆，营养价值丰富Peel without wax powder；Fruit flesh light green；sweet and crispy, and has rich in nutritional value	耐高温 High temperature resistance	王书娟等,2022
	亮丽王 Liangli- wang	津优1号 Jinyou 1	插接法 Peg grafting	92.86	381.02	果皮无蜡粉；果肉营养价值丰富Peel without wax powder；Fruit flesh rich in nutritional value	抗枯萎病 Resistance to Fusarium Wilt	田守胜等,2018
	火凤凰 Fire Phoenix	津优1号 Jinyou 1	插接法 Peg grafting	96.19	326.22	果皮无蜡粉 Peel without wax powder	抗根结线虫 Root knot nematode resistance	田守胜等,2018；邱艳华,2018
	日本青秀 Japanese Aohide	丰田全能 Toyota Versatile	插接法 Peg grafting	100.00	507.73	果肉脆甜，营养价值丰富 Fruit flesh sweet and crispy，and rich in nutritional value	抗枯萎病 Resistance to Fusarium Wilt	潘玲华等,2014
瓠瓜 Lagenaria Ser.	瓠子瓜 Gourds	夏满田 Xia Mantian	插接法 Peg grafting	93.90	452.28	-	抗病性强 Strong disease resistance	梁祖珍等,2014
瓠瓜 Lagenaria Ser.	华瓠杂3号Hua Gourd Miscellan-eous 3	津研4号 Jinyan 4	插接法 Peg grafting	97.60	116.06	早熟，营养价值丰富 Premature and rich in nutritional value	-	刘芬,2009
丝瓜 Luffa Mill.	江蔬1号 Jiangshu 1	津春4号 Jinchun 4	劈接法 Cleft grafting	94.79	-	营养价值丰富 Rich in nutritional value	抗逆性强 Strong stress resistance	陶美奇等,2018
	五叶香 Wuye- xiang	春秋王2号Chun- qiuwang 2	劈接法 Cleft grafting	91.33	100.24	果肉脆甜，营养价值丰富Fruit flesh sweet and crispy，and rich in nutritional value	耐高温 High temperature resistance	李思思等,2017；郝婷等,2014
甜瓜 Cucumis melo L.	东方脆甜/威盛1号 Oriental crisp sweet/Wei-sheng 1	津优35 Jinyou 35	中间砧嫁接法Intermediate rootstock grafting	88.37	343.24	果形优；营养价值丰富 Excellent fruit shape；Rich in nutritional value	具抗逆性 Stress resistance	王立霞等,2019
甜瓜 Cucumis melo L.	伊丽莎白/威盛1号 Elizabeth/Wei- sheng 1	津优35 Jinyou 35	中间砧嫁接法Intermediate rootstock grafting	85.82	332.27	果形优；风味佳；营养价值丰富Excellent fruit shape and flavor；Rich in nutritional value	具抗逆性 Stress resistance	王立霞等,2019
苦瓜 Momordica charantia L.	傲美 Aomei	春秋王2号Chunqiuwang 2	劈接法 Cleft grafting	54.00	-	营养价值较丰富 Relatively rich in nutritional value	较耐高温 Relatively high temperature resistant	郝婷等,2014；李思思等,2017,2018
苦瓜 Momordica charantia L.	长绿 Changlü	津优35 Jinyou 35	劈接法 Cleft grafting	-	-	营养价值丰富 Rich in nutritional value	耐高温 High temperature resistance	魏莹,2019；陶美奇,2020

砧木品类 Rootstock category	砧木品种 Rootstock cultivar	接穗品种 Scion	嫁接方法 Grafting method	成活率/% Survival rate	主要性状 Main characters			参考文献 References
砧木品类 Rootstock category	砧木品种 Rootstock cultivar	接穗品种 Scion	嫁接方法 Grafting method	成活率/% Survival rate	果实产量/ （kg · hm^-2） Fruit yield	果实品质 Fruit quality	植株抗逆能力 Plant stress resistance	参考文献 References
南瓜 Cucurbita L. .	京欣砧5号 Jingxin- zhen 5	津优35 Jinyou 35	插接法 Peg grafting	97.00	277.47	果皮无蜡粉；果肉深绿 Peel without wax powder；Fruit flesh dark green	耐低温；抗猝倒病、枯萎病、霜霉病 Low temperature resistance；Resistance to Damping off disease，Fusarium Wilt and downy mildew	张帆等,2011
	金妈妈1989 Kim Mama 1989	北农佳秀 Beinong Jiaxiu	贴接法 Patch grafting	94.00	581.67	果皮无蜡粉；果肉浅绿，甜脆，营养价值丰富Peel without wax powder；Fruit flesh light green；sweet and crispy, and has rich in nutritional value	耐高温 High temperature resistance	王书娟等,2022
	亮丽王 Liangli- wang	津优1号 Jinyou 1	插接法 Peg grafting	92.86	381.02	果皮无蜡粉；果肉营养价值丰富Peel without wax powder；Fruit flesh rich in nutritional value	抗枯萎病 Resistance to Fusarium Wilt	田守胜等,2018
	火凤凰 Fire Phoenix	津优1号 Jinyou 1	插接法 Peg grafting	96.19	326.22	果皮无蜡粉 Peel without wax powder	抗根结线虫 Root knot nematode resistance	田守胜等,2018；邱艳华,2018
	日本青秀 Japanese Aohide	丰田全能 Toyota Versatile	插接法 Peg grafting	100.00	507.73	果肉脆甜，营养价值丰富 Fruit flesh sweet and crispy，and rich in nutritional value	抗枯萎病 Resistance to Fusarium Wilt	潘玲华等,2014
瓠瓜 Lagenaria Ser.	瓠子瓜 Gourds	夏满田 Xia Mantian	插接法 Peg grafting	93.90	452.28	-	抗病性强 Strong disease resistance	梁祖珍等,2014
瓠瓜 Lagenaria Ser.	华瓠杂3号Hua Gourd Miscellan-eous 3	津研4号 Jinyan 4	插接法 Peg grafting	97.60	116.06	早熟，营养价值丰富 Premature and rich in nutritional value	-	刘芬,2009
丝瓜 Luffa Mill.	江蔬1号 Jiangshu 1	津春4号 Jinchun 4	劈接法 Cleft grafting	94.79	-	营养价值丰富 Rich in nutritional value	抗逆性强 Strong stress resistance	陶美奇等,2018
	五叶香 Wuye- xiang	春秋王2号Chun- qiuwang 2	劈接法 Cleft grafting	91.33	100.24	果肉脆甜，营养价值丰富Fruit flesh sweet and crispy，and rich in nutritional value	耐高温 High temperature resistance	李思思等,2017；郝婷等,2014
甜瓜 Cucumis melo L.	东方脆甜/威盛1号 Oriental crisp sweet/Wei-sheng 1	津优35 Jinyou 35	中间砧嫁接法Intermediate rootstock grafting	88.37	343.24	果形优；营养价值丰富 Excellent fruit shape；Rich in nutritional value	具抗逆性 Stress resistance	王立霞等,2019
甜瓜 Cucumis melo L.	伊丽莎白/威盛1号 Elizabeth/Wei- sheng 1	津优35 Jinyou 35	中间砧嫁接法Intermediate rootstock grafting	85.82	332.27	果形优；风味佳；营养价值丰富Excellent fruit shape and flavor；Rich in nutritional value	具抗逆性 Stress resistance	王立霞等,2019
苦瓜 Momordica charantia L.	傲美 Aomei	春秋王2号Chunqiuwang 2	劈接法 Cleft grafting	54.00	-	营养价值较丰富 Relatively rich in nutritional value	较耐高温 Relatively high temperature resistant	郝婷等,2014；李思思等,2017,2018
苦瓜 Momordica charantia L.	长绿 Changlü	津优35 Jinyou 35	劈接法 Cleft grafting	-	-	营养价值丰富 Rich in nutritional value	耐高温 High temperature resistance	魏莹,2019；陶美奇,2020