作物对干旱胁迫的响应机制及提高作物抗旱能力的调控措施研究进展

doi:10.11924/j.issn.1000-6850.casb2021-1042

中国农学通报 ›› 2022, Vol. 38 ›› Issue (29): 31-44.doi: 10.11924/j.issn.1000-6850.casb2021-1042

所属专题：生物技术

作物对干旱胁迫的响应机制及提高作物抗旱能力的调控措施研究进展

王硕(), 贾潇倩, 何璐, 李浩然, 王红光, 何建宁, 李东晓, 房琴(), 李瑞奇

河北农业大学农学院/省部共建华北作物改良与调控国家重点实验室/河北省作物生长调控实验室,河北保定 071001

收稿日期:2021-11-03 修回日期:2022-01-27 出版日期:2022-10-15 发布日期:2022-10-14
通讯作者: 房琴
作者简介:王硕,女,1997年出生,河北石家庄人,硕士研究生,主要从事作物栽培生理生态研究。通信地址：071001 河北省保定市河北农业大学农学院,E-mail： 1342229002@qq.com。
基金资助:
“河北农业大学引进人才科研专项”(YJ201851);“国家小麦产业技术体系项目”(CARS-03-05);“保定市科技计划”(1911ZN010)

Response Mechanism of Crops to Drought Stress and Measures for Improving Drought Resistance of Crops: Research Progress

WANG Shuo(), JIA Xiaoqian, HE Lu, LI Haoran, WANG Hongguang, HE Jianning, LI Dongxiao, FANG Qin(), LI Ruiqi

College of Agronomy, Hebei Agricultural University/State Key Laboratory of North China Crop Improvement and Regulation/Key Laboratory of Crop Growth Regulation of Hebei Province, Baoding, Hebei 071001

Received:2021-11-03 Revised:2022-01-27 Online:2022-10-15 Published:2022-10-14
Contact: FANG Qin

摘要/Abstract

摘要：

干旱是影响作物生长发育和产量的最主要非生物胁迫之一,在气候变化背景下,作物遭受干旱胁迫的风险越来越大。为了应对干旱,作物表现出一系列的抵御机制,包括形态特征和生理生化（抗氧化酶、渗透调节物质、内源激素）特性改变。本研究从上述2个方面总结了作物对干旱胁迫的响应机制,并对提高作物抗旱能力的调控措施进行了论述,主要包括：（1）筛选抗旱性品种,促进对深层土壤贮水的吸收利用;（2）地面覆盖,有利于降低土壤蒸发,增加土壤含水量;（3）节水灌溉技术,如微喷灌、滴灌等灌溉方式能实现少量多次灌溉,根区局部灌溉有利于调节气孔关闭,减少奢侈蒸腾,降低土壤蒸发;（4）抗蒸腾剂,在作物枝干及叶面表层形成超薄透光的保护膜,抑制作物水分过度蒸腾;（5）植物生长调节剂,调控植物生理代谢,增强抗旱性;（6）纳米肥料,改变作物生理生化反应,促进植株生长发育;（7）生物炭,有利于土壤通气保水,改善土壤的物理性质和土壤的持水能力。本研究系统地对以上7种措施提高作物抗旱能力的作用机理、应用前景及存在问题进行了论述,以期为应对干旱胁迫提供理论依据和技术参考。

关键词: 干旱, 形态特征, 生理生化, 抗旱能力, 调控措施

Abstract:

Drought is one of the most important abiotic stresses affecting the growth and yield of crops. The risk of crops suffering from drought stress is increasing under climate change. In order to cope with drought, crops show a series of resistance mechanisms, including the changes of morphological characteristics, and physiological and biochemical characteristics (antioxidant enzyme, osmotic regulation substances, and endogenous hormone), which play an important role in improving the resistance of crops under drought conditions. This paper summarized the response mechanism of crops to drought stress and introduced the measures to improve the drought resistance ability of crops, including: (1) selecting drought-resistance cultivars to promote the absorption of deep soil water; (2) soil mulching, which could reduce soil evaporation and increase soil water content; (3) water-saving irrigation techniques, such as micro-sprinkler irrigation and drip irrigation, which could precisely control the irrigation application and increase irrigation frequency; in addition, partial root-zone irrigation could reduce luxurious transpiration and soil evaporation by regulating stomatal closure; (4) anti-transpirants, which could inhibit excessive transpiration of water by forming ultrathin transparent protective film on the surface of crop branches and leaves; (5) plant growth regulators, which could enhance crop drought resistance by regulating plant physiological metabolism; (6) nano-fertilizers, which could promote plant growth and development by changing physiological and biochemical reactions of crops and enhance drought resistance of crops; (7) biochar, which is beneficial to soil aeration and water retention, and could improve soil physical properties and soil water retention capacity. In order to provide a theoretical basis and technical reference for coping with drought stress, this paper systematically discussed the action mechanism, application prospect and existing problems of the above 7 measures.

Key words: drought, morphological characteristics, physiological and biochemical characteristics, drought resistance, regulation measures

中图分类号:

王硕, 贾潇倩, 何璐, 李浩然, 王红光, 何建宁, 李东晓, 房琴, 李瑞奇. 作物对干旱胁迫的响应机制及提高作物抗旱能力的调控措施研究进展[J]. 中国农学通报, 2022, 38(29): 31-44.

WANG Shuo, JIA Xiaoqian, HE Lu, LI Haoran, WANG Hongguang, HE Jianning, LI Dongxiao, FANG Qin, LI Ruiqi. Response Mechanism of Crops to Drought Stress and Measures for Improving Drought Resistance of Crops: Research Progress[J]. Chinese Agricultural Science Bulletin, 2022, 38(29): 31-44.

图/表 2

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外源生长调节剂类型	试验环境		施用方法			作物		作用	参考文献
褪黑素	温室		棉花第一朵白花出现后,喷施100、 200 μmol褪黑素100 mL, 和干旱胁迫同时进行			棉花 (Gossypium hirsutum L.)		调节花粉碳水化合物代谢, 改善干旱条件下棉花花粉活力	[132]
褪黑素	防雨棚		幼苗3叶期每天加入100 μmol褪黑素溶液200 mL,共7天, 之后进行干旱胁迫处理			油菜 (Brassica napus L.)		促进主根和侧根的生长和气孔开放, 改善了光合能力,提高CAT活性	[133]
ABA	大田环境		干旱处理后,开花期每天17点喷施 20 mg/L的ABA溶液（每平方米施用 500 mL）,共4天			小麦 (Triticum aestivum L.)		增加了叶片脯氨酸的积累,提高了叶片抗氧化酶(SOD、POD、CAT和APX)的活性,降低了玉米素/赤霉素、水杨酸、脱落酸、和精胺/亚精胺的比值, 延长了旗叶的功能期	[124]
BRs	大田环境		在开花期或灌浆期水分胁迫前10天, 喷施0、0.05、0.1 mg/L的BR溶液2 L			小麦 (Triticum aestivum L.)		增加了脯氨酸的含量,降低了丙二醛的积累,减少了细胞膜脂质过氧化。0.1 mg/L的BR溶液效果最佳	[134]
BRs	室内控制环境		对种子灭菌后用0.4 μmol 24-表油菜素内酯(EBR)溶液浸泡3 h,种子萌发后3天,将幼苗移植到0.4 μmol EBR溶液中处理24 h,之后用5%的甘露醇模拟干旱胁迫			小麦 (Triticum aestivum L.)		增加了ABA,降低了IAA和CK的含量, 促进了脱氢蛋白(DHN)的积累	[135]
IBA	室内控制环境		浓度为10^-7mol IBA、10^-8mol IBA、 10^-11mol的IBA溶液培养幼苗			玉米 (Zea mays L.)		低浓度(10^-11mol)的IBA可以刺激根的生长,但不会影响细胞壁的发育;高浓度(10^-7mol)的IBA诱发了离根尖较近的质外屏障（凯氏带和木质部）的发育, 提高了根部木质素含量	[126]
多巴胺	室内控制环境		含有100 μmol的多巴胺营养液灌溉1 m高的苹果树苗,处理10天, 同时进行干旱胁迫			苹果 (Malus domestica Borkh)		调控氮、次生化合物和氨基酸代谢相关基因的表达, 激活Ca²⁺通路,提高苹果的耐旱性	[129]
亚精胺(SPD)	室内控制环境		浓度为0.05 mmol和0.1 mmol的Spd溶液培养幼苗2天,之后15% PEG-6000 模拟干旱胁迫			玉米 (Zea mays L.)		外源施用SPD后,玉米幼苗叶绿体超微结构排列更加有序,增加了内源多胺、吲哚乙酸、玉米苷、赤霉素含量,降低了水杨酸含量。外源SPD可以缓解光合速率下降,减轻干旱对细胞结构与功能的损害。 0.1 mmol效果最佳	[136]
腐胺(PUT)	室内控制环境		浓度为0.5 mmol的腐胺溶液中培养7天,同时用PEG模拟干旱胁迫			小麦 (Triticum aestivum L.)		腐胺提高了叶片中游离水杨酸的含量,改变了类囊体膜中的脂肪酸组成,增加了植株地上部鲜重和净光合速率。同时脱氢酶,乙烯合成酶等一系列蛋白酶的基因表达上升	[137]
硝普钠	室内控制环境		10% PEG溶液中加入50、100、250、 500 μmol的硝普钠溶液			甜叶菊 (Stevia rebaudiana (Bertoni) Hemsl.)		加入硝普钠后减轻活性氧的危害,增加植株分枝数和分枝长度,有利于维持细胞内的离子平衡, 稳定渗透功能。50~100 μmol效果最好	[138]
脯氨酸	大田环境		植株移栽后的60天和75天各施用1次0、1、2 mmol的脯氨酸, 同时进行干旱胁迫			洋葱 (Allium cepa L.)		外源脯氨酸增加了叶片相对含水量、光合效率和渗透保护剂的含量,1~2 mmol脯氨酸效果最好	[127]
SA		温室		种子发芽一周后喷洒0.5 mmol的水杨酸溶液,分蘖至开花期进行干旱胁迫	小麦 (Triticum aestivum L.)		SA能够缓解膜电解质渗漏和膜脂过氧化, 提高抗氧化酶的活性,维持细胞膜的稳定性		[139]
SA		温室		第1次于发芽后1周喷施0.5 mmol的SA溶液,第2、3次分别于干旱胁迫前24 h和后24 h喷施0.5 mmol的SA溶液	玉米 (Zea mays L.)		喷施SA后增加了株高、地上部鲜重、叶面积、地上部干物质积累量、多胺含量、可溶性糖和碳水化合物的含量,提高了抗氧化酶的活性		[140]
茉莉酸甲酯 (MeJA)		室内控制环境		浓度为0.1 μmol的MeJA处理3日龄的幼苗,培养24 h,之后用甘露醇模拟干旱胁迫	小麦 (Triticum aestivum L.)		外源喷施MeJA增强了小麦TADHN基因的表达和脱氢蛋白酶的积累		[141]
MeJA		室内控制环境		种子在0、2.5、5 mmol MeJA溶液中浸泡24 h,发芽后在光照培养箱中培养14天,之后用PEG模拟干旱胁迫	水稻 (Oryza sativa L.)		MeJA提高了叶片的水势,WUE和相对含水量, 降低了电导率,改变植株根茎组织中多重代谢产物的浓度,从而影响水稻种子的萌发和幼苗的生长		[142]
冠菌素(COR)		大田环境		灌水前1周,喷施0.1 mol的COR溶液	小麦 (Triticum aestivum L.)		COR提高了叶片的光合参数和SPAD, COR还可以诱导根发育,增强冬小麦抗旱能力		[131]

外源生长调节剂类型	试验环境		施用方法			作物		作用	参考文献
褪黑素	温室		棉花第一朵白花出现后,喷施100、 200 μmol褪黑素100 mL, 和干旱胁迫同时进行			棉花 (Gossypium hirsutum L.)		调节花粉碳水化合物代谢, 改善干旱条件下棉花花粉活力	[132]
褪黑素	防雨棚		幼苗3叶期每天加入100 μmol褪黑素溶液200 mL,共7天, 之后进行干旱胁迫处理			油菜 (Brassica napus L.)		促进主根和侧根的生长和气孔开放, 改善了光合能力,提高CAT活性	[133]
ABA	大田环境		干旱处理后,开花期每天17点喷施 20 mg/L的ABA溶液（每平方米施用 500 mL）,共4天			小麦 (Triticum aestivum L.)		增加了叶片脯氨酸的积累,提高了叶片抗氧化酶(SOD、POD、CAT和APX)的活性,降低了玉米素/赤霉素、水杨酸、脱落酸、和精胺/亚精胺的比值, 延长了旗叶的功能期	[124]
BRs	大田环境		在开花期或灌浆期水分胁迫前10天, 喷施0、0.05、0.1 mg/L的BR溶液2 L			小麦 (Triticum aestivum L.)		增加了脯氨酸的含量,降低了丙二醛的积累,减少了细胞膜脂质过氧化。0.1 mg/L的BR溶液效果最佳	[134]
BRs	室内控制环境		对种子灭菌后用0.4 μmol 24-表油菜素内酯(EBR)溶液浸泡3 h,种子萌发后3天,将幼苗移植到0.4 μmol EBR溶液中处理24 h,之后用5%的甘露醇模拟干旱胁迫			小麦 (Triticum aestivum L.)		增加了ABA,降低了IAA和CK的含量, 促进了脱氢蛋白(DHN)的积累	[135]
IBA	室内控制环境		浓度为10^-7mol IBA、10^-8mol IBA、 10^-11mol的IBA溶液培养幼苗			玉米 (Zea mays L.)		低浓度(10^-11mol)的IBA可以刺激根的生长,但不会影响细胞壁的发育;高浓度(10^-7mol)的IBA诱发了离根尖较近的质外屏障（凯氏带和木质部）的发育, 提高了根部木质素含量	[126]
多巴胺	室内控制环境		含有100 μmol的多巴胺营养液灌溉1 m高的苹果树苗,处理10天, 同时进行干旱胁迫			苹果 (Malus domestica Borkh)		调控氮、次生化合物和氨基酸代谢相关基因的表达, 激活Ca²⁺通路,提高苹果的耐旱性	[129]
亚精胺(SPD)	室内控制环境		浓度为0.05 mmol和0.1 mmol的Spd溶液培养幼苗2天,之后15% PEG-6000 模拟干旱胁迫			玉米 (Zea mays L.)		外源施用SPD后,玉米幼苗叶绿体超微结构排列更加有序,增加了内源多胺、吲哚乙酸、玉米苷、赤霉素含量,降低了水杨酸含量。外源SPD可以缓解光合速率下降,减轻干旱对细胞结构与功能的损害。 0.1 mmol效果最佳	[136]
腐胺(PUT)	室内控制环境		浓度为0.5 mmol的腐胺溶液中培养7天,同时用PEG模拟干旱胁迫			小麦 (Triticum aestivum L.)		腐胺提高了叶片中游离水杨酸的含量,改变了类囊体膜中的脂肪酸组成,增加了植株地上部鲜重和净光合速率。同时脱氢酶,乙烯合成酶等一系列蛋白酶的基因表达上升	[137]
硝普钠	室内控制环境		10% PEG溶液中加入50、100、250、 500 μmol的硝普钠溶液			甜叶菊 (Stevia rebaudiana (Bertoni) Hemsl.)		加入硝普钠后减轻活性氧的危害,增加植株分枝数和分枝长度,有利于维持细胞内的离子平衡, 稳定渗透功能。50~100 μmol效果最好	[138]
脯氨酸	大田环境		植株移栽后的60天和75天各施用1次0、1、2 mmol的脯氨酸, 同时进行干旱胁迫			洋葱 (Allium cepa L.)		外源脯氨酸增加了叶片相对含水量、光合效率和渗透保护剂的含量,1~2 mmol脯氨酸效果最好	[127]
SA		温室		种子发芽一周后喷洒0.5 mmol的水杨酸溶液,分蘖至开花期进行干旱胁迫	小麦 (Triticum aestivum L.)		SA能够缓解膜电解质渗漏和膜脂过氧化, 提高抗氧化酶的活性,维持细胞膜的稳定性		[139]
SA		温室		第1次于发芽后1周喷施0.5 mmol的SA溶液,第2、3次分别于干旱胁迫前24 h和后24 h喷施0.5 mmol的SA溶液	玉米 (Zea mays L.)		喷施SA后增加了株高、地上部鲜重、叶面积、地上部干物质积累量、多胺含量、可溶性糖和碳水化合物的含量,提高了抗氧化酶的活性		[140]
茉莉酸甲酯 (MeJA)		室内控制环境		浓度为0.1 μmol的MeJA处理3日龄的幼苗,培养24 h,之后用甘露醇模拟干旱胁迫	小麦 (Triticum aestivum L.)		外源喷施MeJA增强了小麦TADHN基因的表达和脱氢蛋白酶的积累		[141]
MeJA		室内控制环境		种子在0、2.5、5 mmol MeJA溶液中浸泡24 h,发芽后在光照培养箱中培养14天,之后用PEG模拟干旱胁迫	水稻 (Oryza sativa L.)		MeJA提高了叶片的水势,WUE和相对含水量, 降低了电导率,改变植株根茎组织中多重代谢产物的浓度,从而影响水稻种子的萌发和幼苗的生长		[142]
冠菌素(COR)		大田环境		灌水前1周,喷施0.1 mol的COR溶液	小麦 (Triticum aestivum L.)		COR提高了叶片的光合参数和SPAD, COR还可以诱导根发育,增强冬小麦抗旱能力		[131]

肥料类型	试验环境	施用方法	作物	作用	参考文献
纳米硼肥 (80~90 nm)	大田环境	分蘖期叶面喷施2 g/L纳米硼肥溶液,每隔15天喷施1次	小麦 (Triticum aestivum L.)	提高了籽粒蛋白质含量、SOD和POD 活性、产量和产量构成	[149]
纳米锌肥 (18 nm)	温室	种子萌发后2周,试验盆内加入 2.17 mg/kg纳米氧化锌肥料	小麦 (Triticum aestivum L.)	小麦抽穗期提前	[146]
纳米锌肥 (18 nm)	室内控制环境	纳米锌肥和土壤混匀,3个浓度： 1、3、5 mg/kg	高粱 (Sorghum bicolor L.)	促进了植株生长,增加了干物质积累量和N、K素积累量,5 mg/kg效果最好	[150]
纳米锌肥 (20~30 nm)	室内控制环境	发芽2周后,用100 mg/L纳米锌肥溶液叶片喷施,每隔1周喷施1次,总共4次,总用量为2 L	小麦 (Triticum aestivum L.)	促进了小麦生长,提高了叶片叶绿素含量、叶片POD和SOD活性	[151]
纳米铁肥 (10~18nm)	室内控制环境	浓度为5、10、15、20、50、90、120 mg/L处理种子7天	谷子 (Setaria italica L.)	增加了脯氨酸、叶绿素和可溶性糖含量, 并随浓度增加而增加	[152]
纳米铁肥 (15~34 nm) 纳米凝胶 (51~94 nm)	室内控制环境	肥料和土壤混匀,浓度为0、25、50、 100 mg/kg	水稻 (Oryza sativa L.)	增加了生物量、抗氧化酶活性、光合作用效率、叶绿素含量,降低了H₂O₂含量。 100 mg/kg效果最好	[153]
纳米铁肥 (50~100 nm)	室内控制环境	把适量浓度的0、25、50、100 mg/kg的纳米铁肥溶液均匀倒入试验盆内	小麦 (Triticum aestivum L.)	增加了光合速率、降低了H₂O₂和MDA 含量,提高了SOD和POD活性。 100 mg/kg效果最好	[154]
纳米硅肥	室内控制环境	在玉米6~8叶期喷施浓度为0、100和200 ppm的溶液	玉米 (Zea mays L.)	100 ppm增加了籽粒中的N含量;200 ppm增加了地上部N、K、Cu、Mg、Si的含量	[155]
壳聚糖纳米肥料	温室	干旱处理后第3天,叶面喷施	甘蔗 (Saccharum spp. L)	提高光合速率、增加了根干重和根冠比	[156]

肥料类型	试验环境	施用方法	作物	作用	参考文献
纳米硼肥 (80~90 nm)	大田环境	分蘖期叶面喷施2 g/L纳米硼肥溶液,每隔15天喷施1次	小麦 (Triticum aestivum L.)	提高了籽粒蛋白质含量、SOD和POD 活性、产量和产量构成	[149]
纳米锌肥 (18 nm)	温室	种子萌发后2周,试验盆内加入 2.17 mg/kg纳米氧化锌肥料	小麦 (Triticum aestivum L.)	小麦抽穗期提前	[146]
纳米锌肥 (18 nm)	室内控制环境	纳米锌肥和土壤混匀,3个浓度： 1、3、5 mg/kg	高粱 (Sorghum bicolor L.)	促进了植株生长,增加了干物质积累量和N、K素积累量,5 mg/kg效果最好	[150]
纳米锌肥 (20~30 nm)	室内控制环境	发芽2周后,用100 mg/L纳米锌肥溶液叶片喷施,每隔1周喷施1次,总共4次,总用量为2 L	小麦 (Triticum aestivum L.)	促进了小麦生长,提高了叶片叶绿素含量、叶片POD和SOD活性	[151]
纳米铁肥 (10~18nm)	室内控制环境	浓度为5、10、15、20、50、90、120 mg/L处理种子7天	谷子 (Setaria italica L.)	增加了脯氨酸、叶绿素和可溶性糖含量, 并随浓度增加而增加	[152]
纳米铁肥 (15~34 nm) 纳米凝胶 (51~94 nm)	室内控制环境	肥料和土壤混匀,浓度为0、25、50、 100 mg/kg	水稻 (Oryza sativa L.)	增加了生物量、抗氧化酶活性、光合作用效率、叶绿素含量,降低了H₂O₂含量。 100 mg/kg效果最好	[153]
纳米铁肥 (50~100 nm)	室内控制环境	把适量浓度的0、25、50、100 mg/kg的纳米铁肥溶液均匀倒入试验盆内	小麦 (Triticum aestivum L.)	增加了光合速率、降低了H₂O₂和MDA 含量,提高了SOD和POD活性。 100 mg/kg效果最好	[154]
纳米硅肥	室内控制环境	在玉米6~8叶期喷施浓度为0、100和200 ppm的溶液	玉米 (Zea mays L.)	100 ppm增加了籽粒中的N含量;200 ppm增加了地上部N、K、Cu、Mg、Si的含量	[155]
壳聚糖纳米肥料	温室	干旱处理后第3天,叶面喷施	甘蔗 (Saccharum spp. L)	提高光合速率、增加了根干重和根冠比	[156]

作物对干旱胁迫的响应机制及提高作物抗旱能力的调控措施研究进展

Response Mechanism of Crops to Drought Stress and Measures for Improving Drought Resistance of Crops: Research Progress

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