Advances in the Research on Salt Tolerance of Malus Plants

doi:10.16420/j.issn.0513-353x.2025-0738

Acta Horticulturae Sinica ›› 2026, Vol. 53 ›› Issue (7): 1929-1954.doi: 10.16420/j.issn.0513-353x.2025-0738

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Advances in the Research on Salt Tolerance of Malus Plants

LIU Zhao¹^,², GAO Yuan¹, SUN Simiao¹, WANG Kun¹, GUO Hanxin¹, SHANG Wei¹, WANG Lin¹, TIAN Wen¹^,², LI Zichen¹, SUN Yanming¹, LI Lianwen¹, WANG Dajiang¹^,³^,^*()

¹ Research Institute of Pomology， Chinese Academy of Agricultural Sciences; Key Laboratory of Horticultural Crops Germplasm Resources Utilization，Ministry of Agriculture and Rural Affairs，Xingcheng， Liaoning 125100， China
² College of Agriculture， Shihezi University;The Key Laboratory of Special Fruits and Vegetables Cultivation Physiology and Germplasm Resources Utilization Xinjiang Production and Construction Corps，Shihezi， Xinjiang 832000， China
³ National Center Technology Innovation for Comprehensive Utilization of Saline-Alkali Land， Dongying， Shandong 257000， China

Received:2026-02-12 Revised:2026-05-28 Online:2026-07-23 Published:2026-07-23
Contact: WANG Dajiang

Abstract

Abstract:

In-depth research on the response mechanisms and defense strategies of Malus plants to salt stress is of great theoretical significance for the development of salt tolerant cultivation techniques for apples and the breeding of salt tolerant varieties. This review comprehensively analyzed the advanced results on the effects of salt stress in Malus plants，the identification，evaluation and screening of Malus plants under salt stress，the physiological processes and molecular mechanisms of salt tolerance，and the identification of salt-tolerant functional genes，aiming to obtain a comprehensive understanding of salt tolerance in Malus plants and identify the current situation and existing problems in the research，and propose the main research directions. It will provide selection and reference for the practical application of salt-tolerant Malus and the selection and breeding of salt-tolerant apple rootstocks.

Key words: Malus, salt stress, salt-tolerant mechanism

LIU Zhao, GAO Yuan, SUN Simiao, WANG Kun, GUO Hanxin, SHANG Wei, WANG Lin, TIAN Wen, LI Zichen, SUN Yanming, LI Lianwen, WANG Dajiang. Advances in the Research on Salt Tolerance of Malus Plants[J]. Acta Horticulturae Sinica, 2026, 53(7): 1929-1954.

Figures/Tables 5

References 138

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抗盐能力 Salt tolerance	种质 Germplasm	参考文献 Reference	抗盐能力 Salt tolerance	种质 Germplasm	参考文献 Reference
高High	沙果 Malus asiatica	石桂英和刘捍中，1990	中Moderate	M2（M. × domestica）	矫彩娣等,2025
	珠美海棠 M. zumi	马继龙等，1996		M9（M. × domestica）	巴桑,1990
	楸子 M. prunifolia	尹蓉，2010		M16（M. × domestica）	巴桑,1990
	东北黄海棠 M. prunifolia	尹蓉，2010		M26（M. × domestica）	巴桑,1990
	新疆黄海棠 M. prunifolia	石桂英和刘捍中，1990		MM106（M. × domestica）	巴桑,1990
	‘难咽’M.micromalus‘Nanyan’	石桂英和刘捍中，1990	低Low	丽江山荆子 M. rockii	杜中军等,2001
	Luo-2（M. × domestica）	杜中军等，2001		山荆子 M. baccata	杜中军等,2001
	Luo-1（M. × domestica）	杜中军等，2001		毛山荆子 M. manshurica	潘增光等,1999
	花叶海棠 M. transitoria	李晓林等，1996		小克莱 M. pumila	查霞娟等,1986
	大果楸子M. prunifolia	李晓林等，1996		林芝海棠 M. rockii	赵进春等,1995
	小金海棠 M. sikkimensis	尹蓉，2010		雅江变叶 M. toringoides	石桂英和刘捍中,1990
	八棱海棠 M. robusta	张志晓等，2017		红三叶海棠 M. sieboldii	石桂英和刘捍中,1990
	平顶海棠 M. robusta	张志晓等，2017		巴东海棠 M. hupehensis	石桂英和刘捍中,1990
	富平楸子 M. prunifolia	李春容，2023		河南海棠 M. honanensis	石桂英和刘捍中,1990
	草原海棠 M. ioensis	林冰冰等，2016		湖北海棠 M. hupehensis	杜明辉,2023
	平邑甜茶 M. hupehensis	傅明洋，2013		理县楸子 M. prunifolia	李晓林等,1996
	新疆野苹果 M. sieversii	傅明洋，2013		三叶海棠 M. sieboldii	李晓林等,1996
	崂山柰子 M. prunifolia	傅明洋，2013		马尔康海棠 M. maerkangensis	李晓林等,1996
	G935（M. × domestica）	李春容，2023		西府海棠 M. micromalus	李晓林等,1996
中Moderate	平邑甜茶 M. hupehensis	赵进春等，1995		宿萼花叶海棠 M. transitoria	李晓林等,1996
	八棱海棠 M. robusta	宗鹏鹏等，2013		卢氏红果 M. sieboldii	Yin et al.,2010
	新疆野苹果 M. sieversii	林冰冰等，2016		樱叶海棠 M. cerasifolia	尹蓉，2010
	倒挂珍珠 M. halliana	石桂英和刘捍中，1990		德钦海棠 M. prunifolia	杜中军等,2001
	垂丝海棠 M. halliana	张德等，2021		SH15（M. × domestica）	杜中军等,2001
	富平楸子 M. prunifolia	Yin et al.，2010		M4（M. × domestica）	巴桑,1990
	平顶海棠 M. robusta	林冰冰等，2016		M11（M. × domestica）	巴桑,1990
	崂山柰子 M. prunifolia	林冰冰等，2016		M27（M. × domestica）	巴桑,1990
	K15（M. × domestica）	矫彩娣等，2025

抗盐能力 Salt tolerance	种质 Germplasm	参考文献 Reference	抗盐能力 Salt tolerance	种质 Germplasm	参考文献 Reference
高High	沙果 Malus asiatica	石桂英和刘捍中，1990	中Moderate	M2（M. × domestica）	矫彩娣等,2025
	珠美海棠 M. zumi	马继龙等，1996		M9（M. × domestica）	巴桑,1990
	楸子 M. prunifolia	尹蓉，2010		M16（M. × domestica）	巴桑,1990
	东北黄海棠 M. prunifolia	尹蓉，2010		M26（M. × domestica）	巴桑,1990
	新疆黄海棠 M. prunifolia	石桂英和刘捍中，1990		MM106（M. × domestica）	巴桑,1990
	‘难咽’M.micromalus‘Nanyan’	石桂英和刘捍中，1990	低Low	丽江山荆子 M. rockii	杜中军等,2001
	Luo-2（M. × domestica）	杜中军等，2001		山荆子 M. baccata	杜中军等,2001
	Luo-1（M. × domestica）	杜中军等，2001		毛山荆子 M. manshurica	潘增光等,1999
	花叶海棠 M. transitoria	李晓林等，1996		小克莱 M. pumila	查霞娟等,1986
	大果楸子M. prunifolia	李晓林等，1996		林芝海棠 M. rockii	赵进春等,1995
	小金海棠 M. sikkimensis	尹蓉，2010		雅江变叶 M. toringoides	石桂英和刘捍中,1990
	八棱海棠 M. robusta	张志晓等，2017		红三叶海棠 M. sieboldii	石桂英和刘捍中,1990
	平顶海棠 M. robusta	张志晓等，2017		巴东海棠 M. hupehensis	石桂英和刘捍中,1990
	富平楸子 M. prunifolia	李春容，2023		河南海棠 M. honanensis	石桂英和刘捍中,1990
	草原海棠 M. ioensis	林冰冰等，2016		湖北海棠 M. hupehensis	杜明辉,2023
	平邑甜茶 M. hupehensis	傅明洋，2013		理县楸子 M. prunifolia	李晓林等,1996
	新疆野苹果 M. sieversii	傅明洋，2013		三叶海棠 M. sieboldii	李晓林等,1996
	崂山柰子 M. prunifolia	傅明洋，2013		马尔康海棠 M. maerkangensis	李晓林等,1996
	G935（M. × domestica）	李春容，2023		西府海棠 M. micromalus	李晓林等,1996
中Moderate	平邑甜茶 M. hupehensis	赵进春等，1995		宿萼花叶海棠 M. transitoria	李晓林等,1996
	八棱海棠 M. robusta	宗鹏鹏等，2013		卢氏红果 M. sieboldii	Yin et al.,2010
	新疆野苹果 M. sieversii	林冰冰等，2016		樱叶海棠 M. cerasifolia	尹蓉，2010
	倒挂珍珠 M. halliana	石桂英和刘捍中，1990		德钦海棠 M. prunifolia	杜中军等,2001
	垂丝海棠 M. halliana	张德等，2021		SH15（M. × domestica）	杜中军等,2001
	富平楸子 M. prunifolia	Yin et al.，2010		M4（M. × domestica）	巴桑,1990
	平顶海棠 M. robusta	林冰冰等，2016		M11（M. × domestica）	巴桑,1990
	崂山柰子 M. prunifolia	林冰冰等，2016		M27（M. × domestica）	巴桑,1990
	K15（M. × domestica）	矫彩娣等，2025

响应盐胁迫的类型 Types of responses to salt stress	基因名称 Gene name	基因功能 Gene function	参考文献 Reference
离子稳态 Ion homeostasis	MdNHX2 MdNHX5	受盐诱导上调表达，正向调控苹果耐盐性 Salt-induced up-regulated expression and positively regulated salt tolerance of apple	黄鑫,2023
	MdNHX1	过表达MdNHX1可提高M26的耐盐耐旱性 Overexpression of MdNHX1 enhanced the salt and drought tolerance of M26	Li et al.,2013
	MdHKT1	受盐诱导上调表达，调控离子平衡 Salt-induced up-regulated expression and regulated of ion balance	李超,2016
	MdAKT1	减少Na⁺积累，维持Na⁺/K⁺稳态 Reduce the accumulation of Na⁺ and maintain Na⁺/K⁺ homeostasis	郭晶,2024
	MdHAK1	MdWRKY17结合 MdHAK1的启动子并激活其表达 MdWRKY17 binds to the promoter of MdHAK1 and activates its expression	Wang et al.,2025a
	MdCCX2	MdCCX2抑制盐胁迫下的Na⁺积累 MdCCX2 inhibited the accumulation of Na⁺ under salt stress	Yang et al.,2021
渗透调节 Osmotic regulation	MdPFPK5	过表达MdPFPK5增加苹果愈伤组织中可溶性糖的积累 Overexpression of MdPFPK5 increased the accumulation of soluble sugars in apple callus	Zhang et al.,2024c
	MdTyDC	过表达MdTyDC增加了苹果愈伤组织中脯氨酸的积累 Overexpression of MdTyDC increased the accumulation of proline in apple callus	Wang et al.,2020
	MdALMT14	促进苹果酸的积累提高耐盐性 Promote the accumulation of malic acid to enhance salt tolerance	路静,2019
	MdSWEET1	促进可溶性糖的积累Promote the accumulation of soluble sugar	路静等,2019
	MdSUT2.2	盐诱导MdSUT2.2发生磷酸化，促进蔗糖的积累 Salt induced phosphorylation of MdSUT2.2 and promoted the accumulation of sucrose	Ma et al.,2018
	MdSTP1	MdSTP1促进葡萄糖和蔗糖的积累 MdSTP1 promotes the accumulation of glucose and sucrose	董元花,2020
信号转导 Signal transduction	MdMAPK6	过表达MdMAPK6增强苹果愈伤组织的耐盐性 Overexpression of MdMAPK6 enhanced the salt tolerance of apple callus	Li et al.,2025c
	MdCNGC15A	沉默MdCNGC15A增强苹果植株耐盐性 Silencing MdCNGC15A enhanced the salt tolerance of apple plants	Qiu et al.,2024
	MhABF	过表达MhABF增强苹果愈伤组织的耐盐性 Overexpression of MhABF enhanced the salt tolerance of apple callus	王双成,2022
信号转导 Signal transduction	MhPYL	负向调控苹果愈伤组织耐盐性Negatively regulates the salt tolerance of apple callus	王双成,2022
信号转导 Signal transduction	MpSnRK2.10	过表达MpSnRK2.10增强ABA信号的输出 Overexpression of MpSnRK2.10 enhanced the output of ABA signaling	Shao et al.,2019
	MdPP2CA	MdPP2CA负向调控苹果耐盐性 MdPP2CA negatively regulated the salt tolerance of apple	陈萃,2024
	MdPYL9	过表达MdPYL9提高组培苹果苗的耐盐性 Overexpression of MdPYL9 enhanced the salt tolerance of apple seedlings in tissue culture	刘铭潇,2023
	MdSOS2	过表达MdSOS2增强苹果愈伤组织的耐盐性 Overexpression of MdSOS2 enhanced the salt tolerance of apple callus	Hu et al.,2012
	MdSOS3	过表达MdSOS3增加SOD，POD活性，降低ROS含量 Overexpression of MdSOS3 increased the activities of SOD and POD and decreased the content of ROS	郭晶,2024
	MdCIPK13	正向调控MdSUT2.2，提高苹果植株耐盐性 Positively regulated MdSUT2.2 to enhance the salt tolerance of apple plants	马齐军,2018
	MdCBL10.1	过表达MdCBL10.1 提高苹果愈伤组织耐盐性 Overexpression of MdCBL10.1 enhanced the salt tolerance in apple callus	Chen et al.,2021
	MdMAPK3	MdMAPK3磷酸化MdWRKY17，增强其转录激活MdHAK1的能力 MdMAPK3 phosphorylates MdWRKY17 and enhances its ability to transcriptively activate MdHAK1	Wang et al.,2025a
抗氧化系统 Antioxidant system	MhLAR	过表达MhLAR增加苹果愈伤组织中SOD，POD和CAT活性 Overexpression of MhLAR increased the activities of SOD，POD and CAT in apple callus	Li et al.,2025b
	MdTPR16	过表达MdTPR16减少MDA和ROS积累 Overexpression of MdTPR16 reduced the accumulation of MDA and ROS	Wang et al.,2025c
	MdHSP70-38	过表达MdHSP70-38减少MDA，H₂O₂和超氧阴离子积累 Overexpression of MDHSP70-38 reduced the accumulation of MDA，H₂O₂ and superoxide anion	Han et al.,2025
	MdXERICO	过表达MdXERICO减少ACS含量和SOD，POD和CAT活性 Overexpression of MdXERICO reduced the content of ACS and the activities of SOD，POD and CAT	杨超,2024
	MdATL16	促进ROS清除来增强苹果耐盐性 Enhanced apple salt tolerance by promoting ROS scavenging	Yuan et al.,2024
	MhZDS	过表达MhZDS增加Pro含量和SOD，POD和CAT活性 Overexpression of MhZDS increased the content of Pro and the activities of SOD，POD and CAT	Wang et al.,2023
	MdUGT83L3	过表达MdUGT83L3促进花青素积累和ROS清除 Overexpression of MdUGT83L3 promotes the accumulation of anthocyanin and ROS scavenging	李茹,2024
	MdZAT17	过表达MdZAT17促进花青素积累，减少ROS积累 Overexpression of MdZAT17 promoted anthocyanin accumulationand reduced ROS accumulation	王大儒,2023
激素调节 Hormone regulation	MdIAA29	MdIAA29制抑MdARF4蛋白对MdGH3.9和MdNCED3的调控，增加IAA和ABA含量以增强耐盐性 MdIAA29 inhibits the regulation of MdGH3.9 and MdNCED3 by MdARF4 protein，and increases the content of IAA and ABA to enhance salt tolerance	Lei et al.,2024
	MdGH3-2/12	MdGH3-2/12基因沉默后苹果植株耐盐性下降 Salt tolerance of apple decreased after MdGH3-2/12 gene silencing	马梦楠,2021
	MdARF2	MdARF2降低苹果对ABA信号通路的敏感性，并增强其耐盐性 MdARF2 reduced apple sensitivity to the ABA signaling pathway and enhanced its salt tolerance	Wang et al.,2022
	MdCRF6	过表达MdCRF6抑制苹果愈伤组织中花青素积累，降低耐盐性 Overexpression of MdCRF6 inhibited the accumulation of anthocyanin and reduced the salt tolerance in apple callus	安建平等,2017
特定基因或蛋白 Specific genes or proteins	MdTLP7	Tubby-like蛋白MdTLP7与MdNAC72L互作促进MdHsp70-8和MdAGP的表达，增强苹果对盐的耐受性 The Tubby-like protein MdTLP7 interacted with MdNAC72L to promote the expression of MdHsp70-8 and MdAGP，and enhanced the salt tolerance of apples	Wang et al.,2025b
	MdLBD37	LBD蛋白MdLBD37正向调控苹果对干旱和盐的适应性 The LBD protein MdLBD37 positively regulated the adaptation of apple to drought and salt	Li et al.,2025a
	MbACD11	MbACD11缓解由ROS积累引起的氧化应激，增强苹果耐盐性 MbACD11 alleviated oxidative stress caused by ROS accumulation and enhances the salt tolerance of apples	Liu et al.,2024b
	MdAGO1.2	MdAGO1.2提高抗氧化酶活性、抑制ROS积累 MdAGO1.2 enhanced the activity of antioxidant enzymes and inhibited the accumulation of ROS	王梦琦,2023
	MdCP37	MdCP37负调控苹果耐盐性MdCP37 negatively regulated the salt tolerance of apple	巩星遥等,2022
	MdFER	异位表达MdFER提高抗氧化酶活性降低ROS积累Ectopic expression of MdFER enhanced the activity of antioxidant enzymes and reduced the accumulation of ROS	谢寅焕,2022
	MdVQ37	MdVQ37负调控苹果耐盐性MdVQ37 negatively regulated the salt tolerance of apple	Dong et al.,2022
	MdANK2B	过表达MdANK2B增强苹果抗氧化酶活性 Overexpression of MdANK2B enhanced the activities of antioxidant enzymes in apple	张富军,2021
	MdPIP1;1， MdPIP1;2	过表达MdPIP1;1、MdPIP1;2提高苹果愈伤组织耐盐性 Overexpression of MdPIP1;1 and MdPIP1;2 enhanced the salt tolerance of apple callus	杨蕾蕾,2020

响应盐胁迫的类型 Types of responses to salt stress	基因名称 Gene name	基因功能 Gene function	参考文献 Reference
离子稳态 Ion homeostasis	MdNHX2 MdNHX5	受盐诱导上调表达，正向调控苹果耐盐性 Salt-induced up-regulated expression and positively regulated salt tolerance of apple	黄鑫,2023
	MdNHX1	过表达MdNHX1可提高M26的耐盐耐旱性 Overexpression of MdNHX1 enhanced the salt and drought tolerance of M26	Li et al.,2013
	MdHKT1	受盐诱导上调表达，调控离子平衡 Salt-induced up-regulated expression and regulated of ion balance	李超,2016
	MdAKT1	减少Na⁺积累，维持Na⁺/K⁺稳态 Reduce the accumulation of Na⁺ and maintain Na⁺/K⁺ homeostasis	郭晶,2024
	MdHAK1	MdWRKY17结合 MdHAK1的启动子并激活其表达 MdWRKY17 binds to the promoter of MdHAK1 and activates its expression	Wang et al.,2025a
	MdCCX2	MdCCX2抑制盐胁迫下的Na⁺积累 MdCCX2 inhibited the accumulation of Na⁺ under salt stress	Yang et al.,2021
渗透调节 Osmotic regulation	MdPFPK5	过表达MdPFPK5增加苹果愈伤组织中可溶性糖的积累 Overexpression of MdPFPK5 increased the accumulation of soluble sugars in apple callus	Zhang et al.,2024c
	MdTyDC	过表达MdTyDC增加了苹果愈伤组织中脯氨酸的积累 Overexpression of MdTyDC increased the accumulation of proline in apple callus	Wang et al.,2020
	MdALMT14	促进苹果酸的积累提高耐盐性 Promote the accumulation of malic acid to enhance salt tolerance	路静,2019
	MdSWEET1	促进可溶性糖的积累Promote the accumulation of soluble sugar	路静等,2019
	MdSUT2.2	盐诱导MdSUT2.2发生磷酸化，促进蔗糖的积累 Salt induced phosphorylation of MdSUT2.2 and promoted the accumulation of sucrose	Ma et al.,2018
	MdSTP1	MdSTP1促进葡萄糖和蔗糖的积累 MdSTP1 promotes the accumulation of glucose and sucrose	董元花,2020
信号转导 Signal transduction	MdMAPK6	过表达MdMAPK6增强苹果愈伤组织的耐盐性 Overexpression of MdMAPK6 enhanced the salt tolerance of apple callus	Li et al.,2025c
	MdCNGC15A	沉默MdCNGC15A增强苹果植株耐盐性 Silencing MdCNGC15A enhanced the salt tolerance of apple plants	Qiu et al.,2024
	MhABF	过表达MhABF增强苹果愈伤组织的耐盐性 Overexpression of MhABF enhanced the salt tolerance of apple callus	王双成,2022
信号转导 Signal transduction	MhPYL	负向调控苹果愈伤组织耐盐性Negatively regulates the salt tolerance of apple callus	王双成,2022
信号转导 Signal transduction	MpSnRK2.10	过表达MpSnRK2.10增强ABA信号的输出 Overexpression of MpSnRK2.10 enhanced the output of ABA signaling	Shao et al.,2019
	MdPP2CA	MdPP2CA负向调控苹果耐盐性 MdPP2CA negatively regulated the salt tolerance of apple	陈萃,2024
	MdPYL9	过表达MdPYL9提高组培苹果苗的耐盐性 Overexpression of MdPYL9 enhanced the salt tolerance of apple seedlings in tissue culture	刘铭潇,2023
	MdSOS2	过表达MdSOS2增强苹果愈伤组织的耐盐性 Overexpression of MdSOS2 enhanced the salt tolerance of apple callus	Hu et al.,2012
	MdSOS3	过表达MdSOS3增加SOD，POD活性，降低ROS含量 Overexpression of MdSOS3 increased the activities of SOD and POD and decreased the content of ROS	郭晶,2024
	MdCIPK13	正向调控MdSUT2.2，提高苹果植株耐盐性 Positively regulated MdSUT2.2 to enhance the salt tolerance of apple plants	马齐军,2018
	MdCBL10.1	过表达MdCBL10.1 提高苹果愈伤组织耐盐性 Overexpression of MdCBL10.1 enhanced the salt tolerance in apple callus	Chen et al.,2021
	MdMAPK3	MdMAPK3磷酸化MdWRKY17，增强其转录激活MdHAK1的能力 MdMAPK3 phosphorylates MdWRKY17 and enhances its ability to transcriptively activate MdHAK1	Wang et al.,2025a
抗氧化系统 Antioxidant system	MhLAR	过表达MhLAR增加苹果愈伤组织中SOD，POD和CAT活性 Overexpression of MhLAR increased the activities of SOD，POD and CAT in apple callus	Li et al.,2025b
	MdTPR16	过表达MdTPR16减少MDA和ROS积累 Overexpression of MdTPR16 reduced the accumulation of MDA and ROS	Wang et al.,2025c
	MdHSP70-38	过表达MdHSP70-38减少MDA，H₂O₂和超氧阴离子积累 Overexpression of MDHSP70-38 reduced the accumulation of MDA，H₂O₂ and superoxide anion	Han et al.,2025
	MdXERICO	过表达MdXERICO减少ACS含量和SOD，POD和CAT活性 Overexpression of MdXERICO reduced the content of ACS and the activities of SOD，POD and CAT	杨超,2024
	MdATL16	促进ROS清除来增强苹果耐盐性 Enhanced apple salt tolerance by promoting ROS scavenging	Yuan et al.,2024
	MhZDS	过表达MhZDS增加Pro含量和SOD，POD和CAT活性 Overexpression of MhZDS increased the content of Pro and the activities of SOD，POD and CAT	Wang et al.,2023
	MdUGT83L3	过表达MdUGT83L3促进花青素积累和ROS清除 Overexpression of MdUGT83L3 promotes the accumulation of anthocyanin and ROS scavenging	李茹,2024
	MdZAT17	过表达MdZAT17促进花青素积累，减少ROS积累 Overexpression of MdZAT17 promoted anthocyanin accumulationand reduced ROS accumulation	王大儒,2023
激素调节 Hormone regulation	MdIAA29	MdIAA29制抑MdARF4蛋白对MdGH3.9和MdNCED3的调控，增加IAA和ABA含量以增强耐盐性 MdIAA29 inhibits the regulation of MdGH3.9 and MdNCED3 by MdARF4 protein，and increases the content of IAA and ABA to enhance salt tolerance	Lei et al.,2024
	MdGH3-2/12	MdGH3-2/12基因沉默后苹果植株耐盐性下降 Salt tolerance of apple decreased after MdGH3-2/12 gene silencing	马梦楠,2021
	MdARF2	MdARF2降低苹果对ABA信号通路的敏感性，并增强其耐盐性 MdARF2 reduced apple sensitivity to the ABA signaling pathway and enhanced its salt tolerance	Wang et al.,2022
	MdCRF6	过表达MdCRF6抑制苹果愈伤组织中花青素积累，降低耐盐性 Overexpression of MdCRF6 inhibited the accumulation of anthocyanin and reduced the salt tolerance in apple callus	安建平等,2017
特定基因或蛋白 Specific genes or proteins	MdTLP7	Tubby-like蛋白MdTLP7与MdNAC72L互作促进MdHsp70-8和MdAGP的表达，增强苹果对盐的耐受性 The Tubby-like protein MdTLP7 interacted with MdNAC72L to promote the expression of MdHsp70-8 and MdAGP，and enhanced the salt tolerance of apples	Wang et al.,2025b
	MdLBD37	LBD蛋白MdLBD37正向调控苹果对干旱和盐的适应性 The LBD protein MdLBD37 positively regulated the adaptation of apple to drought and salt	Li et al.,2025a
	MbACD11	MbACD11缓解由ROS积累引起的氧化应激，增强苹果耐盐性 MbACD11 alleviated oxidative stress caused by ROS accumulation and enhances the salt tolerance of apples	Liu et al.,2024b
	MdAGO1.2	MdAGO1.2提高抗氧化酶活性、抑制ROS积累 MdAGO1.2 enhanced the activity of antioxidant enzymes and inhibited the accumulation of ROS	王梦琦,2023
	MdCP37	MdCP37负调控苹果耐盐性MdCP37 negatively regulated the salt tolerance of apple	巩星遥等,2022
	MdFER	异位表达MdFER提高抗氧化酶活性降低ROS积累Ectopic expression of MdFER enhanced the activity of antioxidant enzymes and reduced the accumulation of ROS	谢寅焕,2022
	MdVQ37	MdVQ37负调控苹果耐盐性MdVQ37 negatively regulated the salt tolerance of apple	Dong et al.,2022
	MdANK2B	过表达MdANK2B增强苹果抗氧化酶活性 Overexpression of MdANK2B enhanced the activities of antioxidant enzymes in apple	张富军,2021
	MdPIP1;1， MdPIP1;2	过表达MdPIP1;1、MdPIP1;2提高苹果愈伤组织耐盐性 Overexpression of MdPIP1;1 and MdPIP1;2 enhanced the salt tolerance of apple callus	杨蕾蕾,2020

转录因子 Transcription factor	响应盐胁迫的成员 Members response to salt stress	响应盐胁迫的表现 Performance in response to salt stress	参考文献 Reference
WRKY	MdWRKY9	过表达MdWRKY9诱导MdNHX1与MdSOS2的上调表达 Overexpression of MdWRKY9 induced up-regulated expression of MdNHX1 and MdSOS2	Zhao et al.,2025
	MdWRKY75	MdWRKY75能够结合MdSOS3的启动子并激活其转录 MdWRKY75 can bind to the promoter of MdSOS3 and activate its transcription	郭晶,
	MdWRKY18	MdWRKY18与MdGRF8相互作用，增强苹果植株耐盐性 The interaction between MdWRKY18 and MdGRF8 enhanced the salt tolerance of apple plants	Fan et al.,2023
	MdWRKY55	MdWRKY55与MdNAC17-L互作通过激活MdNHX1表达提高苹果耐盐性 The interaction between MdWRKY55 and MdNAC17-L enhanced the salt tolerance of apples by activating the expression of MdNHX1	Su et al.,2022
	MdWRKY30	过表达MdWRKY30增强苹果愈伤组织耐盐性 Overexpression of MdWRKY30 enhanced salt tolerance in apple callus	Dong et al.,2020
	MxWRKY55，MxWRKY64	异源过表达MxWRKY55和MxWRKY64增强拟南芥对盐胁迫的耐受力 Heterologous overexpression of MxWRKY55 and MxWRKY64 enhanced the tolerance of Arabidopsis to salt stress	张兆源,2019
	MxWRKY106	异源过表达MxWRKY106增强拟南芥的耐盐性 Heterologous overexpression of MxWRKY106 enhances the salt tolerance of Arabidopsis	柴丽静,2019
	MdWRKY18，MdWRKY40	MdWRKY18和MdWRKY40增强王林愈伤对盐胁迫的耐受性 MdWRKY18 and MdWRKY40 enhanced the tolerance of‘Orin’to salt stress	许海峰等,2018
NAC	MbNAC22	异源过表达MbNAC22的拟南芥对盐和干旱的耐受性更高 Heterologous overexpression of MbNAC22 enhances the salt and drought tolerance of Arabidopsis	Jiao et al.,2023
	MdNAC1	MdNAC1负调控苹果植株根系生长发育，进而减弱其耐盐性 MdNAC1 negatively regulates the growth and development of apple root system，and then weakens its salt tolerance	岳宏,2022
	MbNAC25，MbNAC29	异源过表达MbNAC25和MbNAC29增强拟南芥的耐盐性 Heterologous overexpression of MbNAC25 and MbNAC29 enhances the salt tolerance of Arabidopsis thaliana	丁海滨,2019
	MdNAC29	MdNAC29与脱水响应基因启动子结合并抑制其表达，负调控苹果的耐盐性MdNAC29 negatively regulated salt tolerance of apple by binding to the promoters of dehydration-responsive genes and inhibiting their expression	郭运娜,2018
	MdNAC143	过表达MdNAC143增强苹果愈伤组织的耐盐性 Overexpression of MdNAC143 enhanced the salt tolerance of apple callus	张全艳等,2017
MYB	MdMYB44-like	MdMYB44-like通过MdPYL8-MdPP2CA模块调控ABA介导的苹果耐盐性 MdMYB44-like regulated ABA-mediated salt tolerance in apple through the MdPYL8-MdPP2CA module	陈萃,2024
MYB	MdMYB108L	MdMYB108L正调控苹果耐盐基因MdNHX1的转录，提高其耐盐性 MdMYB108L positively regulated the transcription of salt tolerance gene MdNHX1 to improve the salt tolerance of apple	Du et al.,2022
	MdMYB46	MdMYB46激活胁迫响应信号来增强苹果对盐和渗透胁迫的耐受性 MdMYB46 activates the stress response signal to enhance the tolerance of apples to salt and osmotic stress	Chen et al.,2019
	MdMYB2	过表达MdMYB2增强苹果愈伤组织耐盐性 Overexpression of MdMYB2 enhanced the tolerance of apple callus to salt stress	曲丰佳等,2017
	MdMYB4	过表达MdMYB4增强苹果愈伤组织耐盐性 Overexpression of MdMYB4 enhanced the tolerance of apple callus to salt stress	曲常志,2017
AP2/ERF	MdERF023	抑制ABA信号传导和Na⁺/H⁺稳态来降低苹果耐盐性 Inhibition of ABA signaling and Na⁺/H⁺ homeostasis to reduce apple salt tolerance	Chen et al.,2024
	MdERF114	MdERF114通过调控离子平衡，降低ROS水平，增强苹果根系耐盐性 MdERF114 enhanced salt tolerance of apple roots by regulating ion balance and reducing ROS levels	刘宇松,2022
	MdERF72	MdERF72在响应高盐胁迫过程中起到正调控作用 MdERF72 plays a positive regulatory role in response to high salt stress	王佳慧等, 2019
	MdERF014	过表达MdERF014增强苹果愈伤组织耐盐性 Overexpression of MdERF014 enhanced the tolerance of apple callus to salt stress	齐晨辉等,2019
	MdERF4	过表达MdERF4负调控耐盐性和乙烯响应 Overexpression of MdERF4 negatively regulated salt tolerance and ethylene response	An et al.,2018
bZIP	MhbZIP23	异源过表达MhbZIP23增强拟南芥对盐碱的耐受性 Heterologous overexpression of MhbZIP23 enhanced the tolerance of Arabidopsis thaliana to salt and alkali stresses	Liu et al.,2024a
bZIP	MdbZIP26	异源表达MdbZIP26提高拟南芥对高盐和干旱胁迫的耐受性 Heterologous expression of MdbZIP26 enhanced the tolerance of Arabidopsis thaliana to high salt and drought stresses	Wan et al.,2022
bHLH	MhbHLH122	过表达MhbHLH122能增强苹果愈伤组织耐盐性 Overexpression of MhbHLH122 enhanced the tolerance of apple callus to salt stress	朱晓悦等,2024
	MdbHLH100	MdbHLH100与MdZAT10a-like互作负调控苹果耐盐性 The interaction between MdbHLH100 and MdZAT10a-like negatively regulated salt tolerance of apple	Yu et al.,2022
	MxbHLH18，MxbHLH30	MxbHLH18和MxbHLH30通过调控ABA合成和信号转导提高耐盐性 MxbHLH18 and MxbHLH30 enhanced salt tolerance by regulating ABA synthesis and signal transduction	徐天龙,2022
	MdbHLH137-like	干扰MdbHLH137-like降低转基因愈伤组织对盐胁迫的耐受能力 Interference with MdbHLH137-like reduced the tolerance of transgenic calli to salt stress	刘媛,2022
	MdbHLH106L	MdbHLH106L通过激活MdNHX1启动子和与MdZAT10互作介导耐盐性 MdbHLH106L mediates salt tolerance by activating the MdNHX1 promoter and interacting with MdZAT10	Zou et al.,2021
SPL	MdSPL13	过表达MdSPL13增强了苹果耐盐性 Overexpression of MdSPL13 enhanced the salt tolerance of apple	Ma et al.,2020
SPL	MdSPL13	miR156-MdSPL3通过调控MdWRKY33的表达响应苹果盐胁迫 miR156-MdSPL3 responds to apple salt stress by regulating the expression of MdWRKY33	薛浩,2019
HD-Zip	MdHB7-like	MdHB7-like促进MdCCX1表达增加了Na⁺外排和苹果耐盐性 The promotion of MdCCX1 expression by MdHB7-like increased Na⁺ efflux and salt tolerance of apple	Yang et al.,2023
HD-Zip	MdHB-7	MdHB-7减少ROS和Na⁺的积累以及促进脯氨酸的积累，增强其耐盐性 MdHB-7 reduced the accumulation of ROS and Na⁺ and promoted the accumulation of proline to enhance its salt tolerance	Zhao et al.,2021
NF-Y	MdNF-YA8	过表达MdNF-YA8增强苹果愈伤组织的耐盐性 Overexpression of MdNF-YA8 enhanced the tolerance of apple callus to salt stress	王梦瑜,2022

Advances in the Research on Salt Tolerance of Malus Plants

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