梨响应干旱胁迫的研究进展

doi:10.16420/j.issn.0513-353x.2026-0453

摘要/Abstract

摘要：

干旱是限制全球梨产业发展的重要非生物胁迫，对梨树生长发育和果实品质形成产生重要影响。本文中从渗透调节、抗氧化防御、水分平衡等方面概述了梨响应干旱胁迫的生理机制，阐述了干旱信号感知、转录调控途径、翻译后修饰等分子调控机制，结合品种、资源筛选和新品种选育总结了梨抗旱育种进展，列举当前梨抗旱调控措施，剖析了现有研究存在的问题，展望未来研究热点，以期为梨抗旱栽培与育种提供理论依据和指导。

关键词: 梨, 干旱, 胁迫, 响应机制

Abstract:

Drought is an important abiotic stress that limits the development of the global pear industry and has an important impact on the growth and development of pear trees and the formation of fruit quality. In this paper，the physiological mechanism of pear in response to drought stress was summarized from the aspects of osmotic regulation，antioxidant defense and water balance，and the molecular regulation mechanisms such as drought signal perception，transcriptional regulation pathway and post-translational modification were expounded. Combined with the screening of cultivar resources and the breeding of new cultivars，the progress of drought-resistant breeding of pear was summarized. The current drought-resistant regulation measures of pear were listed，the existing problems in the existing research were analyzed，and the future research hotspots were prospected，in order to provide theoretical basis and guidance for drought-resistant cultivation and breeding of pear.

Key words: pear, drought, stress, responding mechanism

刘晓敏, 徐亮亮, 刘亚玲, 吴俊. 梨响应干旱胁迫的研究进展[J]. 园艺学报, 2026, 53(7): 1955-1976.

LIU Xiaomin, XU Liangliang, LIU Yaling, WU Jun. Research Progress of Drought Stress Response in Pear[J]. Acta Horticulturae Sinica, 2026, 53(7): 1955-1976.

图/表 5

参考文献 93

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来源物种 Source species	基因名称 Gene name	抗旱机制 Mechanism of drought resistance	参考文献 Reference
杜梨 Pyrus. betulifolia	PbATG5	自噬基因，正调控抗旱性；过表达拟南芥抗旱性增强，VIGS沉默降低抗旱性 Autophagy gene，positively regulates drought tolerance；overexpression in Arabidopsis enhances drought tolerance，VIGS silencing reduces drought tolerance	贾昕等,2026
	rolB	提高根冠比、色素含量及抗氧化酶活性，增强抗旱性 Increases root/shoot ratio，pigment content，and antioxidant enzyme activities，enhancing drought tolerance	王靖冉等,2024
	PbDRM	干旱响应性移动mRNA，从砧木运输至接穗，正调控抗旱 Drought-responsive mobile mRNA，transported from rootstock to scion，positively regulates drought tolerance	Hao et al.,2020
	PbHsfC1a	热激转录因子，靶向PbPIP1;4和PbNCED4，调控ABA和H₂O₂信号通路 Heat shock transcription factor，targets PbPIP1;4 and PbNCED4，regulates ABA and H₂O₂ signaling pathways	Zhang et al.,2024
	PbERF3	AP2/ERF转录因子，与PbHsfC1a互作，共同激活PbPIP1;4和PbNCED4，正调控抗旱 AP2/ERF transcription factor，interacts with PbHsfC1a，co-activates PbPIP1;4 and PbNCED4，positively regulates drought tolerance	Zhang et al.,2024
	PbeNAC1	NAC转录因子，与PbeDREB1/2A互作，过表达增强抗旱耐寒 NAC transcription factor，interacts with PbeDREB1/2A；overexpression enhances drought and cold tolerance	Jin et al.,2017
	PbrMYB21	R2R3-MYB，调控精氨酸脱羧酶影响多胺合成，正调控抗旱 R2R3-MYB，regulates arginine decarboxylase affecting polyamine synthesis，positively regulates drought tolerance	Li et al.,2017
	PbrWRKY53	WRKY转录因子，结合PbrNCED1启动子，促进维生素C积累，正调控抗旱 WRKY transcription factor，binds PbrNCED1 promoter，promotes vitamin C accumulation，positively regulates drought tolerance	Liu et al.,2019
	PbbZIP88	与PbSRK2E互作，激活PbATL18表达，促进气孔关闭，增强抗旱 Interacts with PbSRK2E，activates PbATL18 expression，promotes stomatal closure，enhances drought tolerance	Lin et al.,2025b
白梨 P. bretschneideri	PbrPLD2	磷脂酶D，过表达拟南芥和梨增强抗旱，促进气孔关闭 Phospholipase D；overexpression in Arabidopsis and pear enhances drought tolerance and promotes stomatal closure	Lin et al.,2025a
	PbrPUB18	U-box E3泛素连接酶，过表达拟南芥增强抗旱 U-box E3 ubiquitin ligase；overexpression in Arabidopsis enhances drought tolerance	Wang et al.,2021
	PbBPC5	BPC转录因子，VIGS沉默降低抗旱性，影响ROS清除 BPC transcription factor；VIGS silencing reduces drought tolerance and affects ROS scavenging	Jia et al.,2026
砂梨 P. pyrifolia	PpyCBF2/3	CBF转录因子，过表达拟南芥增强抗旱、耐盐、耐寒 CBF transcription factors；overexpression in Arabidopsis enhances drought，salt，and cold tolerance	Ahmad et al.,2019
新疆梨 P. sinkiangensis	PsHAT5	HD-Zip蛋白；过表达番茄增强抗旱、耐盐，但降低低温耐受性 HD-Zip protein；overexpression in tomato enhances drought and salt tolerance，but reduces cold tolerance	Liu et al.,2022
新疆梨 P. sinkiangensis	PsHB7	HD-Zip I转录因子，过表达番茄增强抗旱性 HD-Zip I transcription factor；overexpression in tomato enhances drought tolerance	刘晓燕等,2025

来源物种 Source species	基因名称 Gene name	抗旱机制 Mechanism of drought resistance	参考文献 Reference
杜梨 Pyrus. betulifolia	PbATG5	自噬基因，正调控抗旱性；过表达拟南芥抗旱性增强，VIGS沉默降低抗旱性 Autophagy gene，positively regulates drought tolerance；overexpression in Arabidopsis enhances drought tolerance，VIGS silencing reduces drought tolerance	贾昕等,2026
	rolB	提高根冠比、色素含量及抗氧化酶活性，增强抗旱性 Increases root/shoot ratio，pigment content，and antioxidant enzyme activities，enhancing drought tolerance	王靖冉等,2024
	PbDRM	干旱响应性移动mRNA，从砧木运输至接穗，正调控抗旱 Drought-responsive mobile mRNA，transported from rootstock to scion，positively regulates drought tolerance	Hao et al.,2020
	PbHsfC1a	热激转录因子，靶向PbPIP1;4和PbNCED4，调控ABA和H₂O₂信号通路 Heat shock transcription factor，targets PbPIP1;4 and PbNCED4，regulates ABA and H₂O₂ signaling pathways	Zhang et al.,2024
	PbERF3	AP2/ERF转录因子，与PbHsfC1a互作，共同激活PbPIP1;4和PbNCED4，正调控抗旱 AP2/ERF transcription factor，interacts with PbHsfC1a，co-activates PbPIP1;4 and PbNCED4，positively regulates drought tolerance	Zhang et al.,2024
	PbeNAC1	NAC转录因子，与PbeDREB1/2A互作，过表达增强抗旱耐寒 NAC transcription factor，interacts with PbeDREB1/2A；overexpression enhances drought and cold tolerance	Jin et al.,2017
	PbrMYB21	R2R3-MYB，调控精氨酸脱羧酶影响多胺合成，正调控抗旱 R2R3-MYB，regulates arginine decarboxylase affecting polyamine synthesis，positively regulates drought tolerance	Li et al.,2017
	PbrWRKY53	WRKY转录因子，结合PbrNCED1启动子，促进维生素C积累，正调控抗旱 WRKY transcription factor，binds PbrNCED1 promoter，promotes vitamin C accumulation，positively regulates drought tolerance	Liu et al.,2019
	PbbZIP88	与PbSRK2E互作，激活PbATL18表达，促进气孔关闭，增强抗旱 Interacts with PbSRK2E，activates PbATL18 expression，promotes stomatal closure，enhances drought tolerance	Lin et al.,2025b
白梨 P. bretschneideri	PbrPLD2	磷脂酶D，过表达拟南芥和梨增强抗旱，促进气孔关闭 Phospholipase D；overexpression in Arabidopsis and pear enhances drought tolerance and promotes stomatal closure	Lin et al.,2025a
	PbrPUB18	U-box E3泛素连接酶，过表达拟南芥增强抗旱 U-box E3 ubiquitin ligase；overexpression in Arabidopsis enhances drought tolerance	Wang et al.,2021
	PbBPC5	BPC转录因子，VIGS沉默降低抗旱性，影响ROS清除 BPC transcription factor；VIGS silencing reduces drought tolerance and affects ROS scavenging	Jia et al.,2026
砂梨 P. pyrifolia	PpyCBF2/3	CBF转录因子，过表达拟南芥增强抗旱、耐盐、耐寒 CBF transcription factors；overexpression in Arabidopsis enhances drought，salt，and cold tolerance	Ahmad et al.,2019
新疆梨 P. sinkiangensis	PsHAT5	HD-Zip蛋白；过表达番茄增强抗旱、耐盐，但降低低温耐受性 HD-Zip protein；overexpression in tomato enhances drought and salt tolerance，but reduces cold tolerance	Liu et al.,2022
新疆梨 P. sinkiangensis	PsHB7	HD-Zip I转录因子，过表达番茄增强抗旱性 HD-Zip I transcription factor；overexpression in tomato enhances drought tolerance	刘晓燕等,2025

选育方法 Breeding method	品种 Cultivar	亲本 Parent	选育单位 Breeding organization	参考文献Reference
野生变异选育 Wild variant selection	金珠沙梨 Jinzhu Shali	野生砂梨变异 Wild Pyrus pyrifolia variant	河南省豫西山区（洛宁县） Western Henan mountainous area（Luoning County）	王红波,2014
实生选育 Seedling selection	新苹梨 Xinpingli	苹果梨实生选育 Seedling of Pingguoli	辽宁省果树科学研究所 Liaoning Institute of Pomology	王家珍,2004
杂交育种 Cross breeding	寒红梨 Hanhong Li	南果梨 × 晋酥梨 Nanguoli × Jinsuli	吉林省农业科学院果树研究所 Pomology Institute，Jilin Academy of Agricultural Sciences	张茂君等,2004
	八月酥 Bayuesu	栖霞大香水梨 × 郑州鹅梨 Qixia Daxiangshuili × Zhengzhou Eli	中国农业科学院郑州果树研究所 Zhengzhou Fruit Research Institute，CAAS	魏闻东,1994
	甘梨2号 Ganli 2	四百目梨 × 早酥 Sibaimu × Zaosu	甘肃省农业科学院林果花卉研究所 Institute of Fruit and Floriculture，Gansu Academy of Agricultural Sciences	李红旭,2022
	华梨2号 Huali 2	二宫白 × 菊水 Ninomiyahakuri × Kikusui	华中农业大学 Huazhong Agricultural University	罗正荣,2003
	锦丰梨 Jinfeng Li	苹果梨 × 慈梨 Pingguoli × Cili	中国农业科学院果树研究所 Institute of Pomology，CAAS	喻菊芳和苏东岩,1983
	新梨7号 Xinli 7	库尔勒香梨 × 早酥 Korla Pear × Zaosu	新疆塔里木农垦大学 Tarim Agricultural Reclamation University，Xinjiang	刘建萍,2001
	八月红 August Red	早巴 × 早酥 Zaoba × Zaosu	陕西省果树研究所 Shaanxi Fruit Research Institute	冯月秀等,1995
芽变选育 Bud sport selection	红星梨（红茄梨） Red Star Pear（Red Clapps Favorite）	茄梨芽变 Sport of Clapps’ Favorite	美国 USA	葛武伟和李兴敏,2003
	水晶梨（新高芽变） Suisho（Niitaka sport）	新高芽变 Sport of Niitaka	韩国 Korea	张培芳等,2008
	红茄梨 Red Clapps Favorite	茄梨芽变 Sport of Clapps Favorite	美国 USA	孙振华,2006
杂交育种 Cross breeding	早生黄金 Early Gold	新高 × 新兴 Niitaka × Shinko	韩国 Korea	陈修会等,2002
	黄金梨 Whangkeumbae	新高 × 二十世纪 Niitaka × Nijisseiki	韩国 Korea	张培芳等,2008
	新高梨（大果水晶） Niitaka（Daguo Shuijing）	天之川 × 今村秋 Yanogawa × Imamuraaki	日本 Japan	刘乐昌等,2001

选育方法 Breeding method	品种 Cultivar	亲本 Parent	选育单位 Breeding organization	参考文献Reference
野生变异选育 Wild variant selection	金珠沙梨 Jinzhu Shali	野生砂梨变异 Wild Pyrus pyrifolia variant	河南省豫西山区（洛宁县） Western Henan mountainous area（Luoning County）	王红波,2014
实生选育 Seedling selection	新苹梨 Xinpingli	苹果梨实生选育 Seedling of Pingguoli	辽宁省果树科学研究所 Liaoning Institute of Pomology	王家珍,2004
杂交育种 Cross breeding	寒红梨 Hanhong Li	南果梨 × 晋酥梨 Nanguoli × Jinsuli	吉林省农业科学院果树研究所 Pomology Institute，Jilin Academy of Agricultural Sciences	张茂君等,2004
	八月酥 Bayuesu	栖霞大香水梨 × 郑州鹅梨 Qixia Daxiangshuili × Zhengzhou Eli	中国农业科学院郑州果树研究所 Zhengzhou Fruit Research Institute，CAAS	魏闻东,1994
	甘梨2号 Ganli 2	四百目梨 × 早酥 Sibaimu × Zaosu	甘肃省农业科学院林果花卉研究所 Institute of Fruit and Floriculture，Gansu Academy of Agricultural Sciences	李红旭,2022
	华梨2号 Huali 2	二宫白 × 菊水 Ninomiyahakuri × Kikusui	华中农业大学 Huazhong Agricultural University	罗正荣,2003
	锦丰梨 Jinfeng Li	苹果梨 × 慈梨 Pingguoli × Cili	中国农业科学院果树研究所 Institute of Pomology，CAAS	喻菊芳和苏东岩,1983
	新梨7号 Xinli 7	库尔勒香梨 × 早酥 Korla Pear × Zaosu	新疆塔里木农垦大学 Tarim Agricultural Reclamation University，Xinjiang	刘建萍,2001
	八月红 August Red	早巴 × 早酥 Zaoba × Zaosu	陕西省果树研究所 Shaanxi Fruit Research Institute	冯月秀等,1995
芽变选育 Bud sport selection	红星梨（红茄梨） Red Star Pear（Red Clapps Favorite）	茄梨芽变 Sport of Clapps’ Favorite	美国 USA	葛武伟和李兴敏,2003
	水晶梨（新高芽变） Suisho（Niitaka sport）	新高芽变 Sport of Niitaka	韩国 Korea	张培芳等,2008
	红茄梨 Red Clapps Favorite	茄梨芽变 Sport of Clapps Favorite	美国 USA	孙振华,2006
杂交育种 Cross breeding	早生黄金 Early Gold	新高 × 新兴 Niitaka × Shinko	韩国 Korea	陈修会等,2002
	黄金梨 Whangkeumbae	新高 × 二十世纪 Niitaka × Nijisseiki	韩国 Korea	张培芳等,2008
	新高梨（大果水晶） Niitaka（Daguo Shuijing）	天之川 × 今村秋 Yanogawa × Imamuraaki	日本 Japan	刘乐昌等,2001

调控措施 Regulation measure	具体措施 Specific measure	作用 Effect	参考文献 Reference
水分与土壤管理措施 Water and soil management	黑地膜覆盖 Black plastic film mulching	保墒；提升表层土壤pH，加速有机质分解 Moisture conservation；increasing topsoil pH，accelerating organic matter decomposition	曹刚等,2022
	玉米秸秆覆盖 Maize straw mulching	保墒；降低土壤pH，提升表层有机质、速效钾和速效铁含量 Moisture conservation；reducing soil pH，increasing topsoil organic matter，available K and Fe	曹刚等,2022
	覆膜滴灌、适度灌水次数、渗水深度 Film-mulched drip irrigation，appropriate frequency，infiltration depth	提高叶片结构紧密度，促进ABA合成，提高抗旱基因PbDREB表达量 Increasing leaf structural tightness，promoting ABA synthesis，upregulating drought-resistant gene PbDREB	黄海娜等,2018
	调亏灌溉 Regulated deficit irrigation	建立枝水势与土壤含水量正相关关系，为智能灌溉提供依据 Establishing positive correlation between branch water potential and soil water content，providing basis for smart irrigation	孙桂丽等,2016
外源物质施用 Exogenous substance application	外源ABA Exogenous ABA	增强光合系统、ABA生物合成与信号转导、类黄酮积累、糖代谢、活性氧清除 Enhancing photosynthetic system，ABA biosynthesis and signaling，flavonoid accumulation，sugar metabolism，and ROS scavenging	Guo et al.,2025
	外源ABA Exogenous ABA	增加甜菜碱醛脱氢酶活性与丰度，提高甜菜碱浓度，参与渗透调节 Increasing betaine aldehyde dehydrogenase activity/abundance，elevating betaine concentration，participating in osmotic adjustment	Gao et al.,2004
	外源甜菜碱 Exogenous betaine	降低MDA和相对电导率，提高SOD活性；维持细胞膨压、保护生物大分子 Reducing MDA and relative conductivity，increasing SOD activity；maintaining cell turgor，protecting macromolecules	魏树伟等,2012
	外源水杨酸 Exogenous salicylic acid	提高脯氨酸积累，减缓相对含水量下降，提高SOD活性，降低MDA和质膜透性 Increasing proline accumulation，relative water content decline，enhancing SOD activity，reducing MDA and plasma membrane permeability	魏树伟等,2013
	接种丛枝菌根真菌 Inoculation with arbuscular mycorrhizal fungi	降低MDA，提高抗氧化酶活性和脯氨酸含量；增强抗氧化和渗透调节 Reducing MDA，increasing antioxidant enzyme activities and proline content；enhancing antioxidant and osmotic adjustment	栗晗等,2024
	施用生物炭 Biochar application	缓解根系呼吸代谢抑制，维持糖酵解和三羧酸循环贡献率，提高磷酸戊糖途径酶活性 Alleviating inhibition of root respiratory metabolism，maintaining contribution of glycolysis and TCA cycle，increasing pentose phosphate pathway enzyme activity	杜国栋等,2016
	施用保水剂 Water-retaining agent application	促进生长，延长干旱下生存期 Promoting growth，prolonging survival period under drought	杨青松等,2009
农艺措施 Agricultural strategies	干旱期疏果 Fruit thinning during drought	促进气孔调节和土壤水分保存，增加根系碳有效性 Promoting stomatal regulation and soil water conservation，increasing carbon availability for roots	Marsal et al.,2008
	软儿梨间作甜菜 Intercropping soft pear with sugar beet	提高成活率，获得经济效益 Increasing survival rate，obtaining economic benefits	关淑琳和张正鹏,2018
	综合夏剪（拉枝、环刻、晚剪、扭梢、环剥） Integrated summer pruning: branch bending，girdling，late pruning，twisting，ring barking	提高总枝量、萌芽率和成花率；改善树体结构和光照条件 Increasing total branch number，budding rate and flowering rate；improving tree structure and light condition	魏晓东等,2008