种子处理对模拟干旱胁迫玉米苗期生长与养分吸收的调控效应

doi:10.3864/j.issn.0578-1752.2026.12.005

摘要/Abstract

摘要：

【目的】种子处理可增强植物对非生物胁迫的耐受力，探讨种子处理对干旱胁迫下玉米苗期生长状况及养分吸收的调控效应，拟为玉米高产高效抗逆栽培提供依据。【方法】采用盆栽试验，设置对照（CK）、超声（US）、超声+多糖（US+OS）、超声+微生物（US+B）和综合技术（超声+多糖+微生物，IPM）5个种子处理，解析模拟干旱胁迫下玉米苗期农艺性状、光合生理、抗氧化酶活性、养分吸收等参数变化特征，结合Mantel检验与随机森林，分析指标间的潜在关系及影响玉米养分吸收的关键参数。【结果】种子处理可显著改善玉米株高与叶面积，玉米根长、根表面积、根体积和根直径均有不同程度地增加。与对照相比，轻度干旱、中度干旱和重度干旱条件下玉米根系干重分别提高69.23%— 171.15%、59.09%—218.18%和93.84%—264.52%，差异达显著水平（P<0.05）。种子处理显著调节了干旱胁迫下玉米叶片叶绿素相对含量和气体交换参数，增加了叶片与根系的抗氧化酶活性，改善了光合物质生产和生理代谢能力，叶片与根系养分吸收有所上升，其中超声+微生物处理增加效果最好，干旱胁迫下玉米叶片和根系的氮吸收提高31.59%— 59.21%和39.79%—82.35%，磷吸收提高33.70%—69.94%和51.66%—88.69%，钾吸收提高47.71%—78.17%和57.59%— 108.08%。Mantel检验与随机森林结果表明，种子处理通过改善多个生长参数，综合提升玉米抗旱能力，其中生物量、叶面积和株高等形态特征是影响玉米氮、磷、钾等养分吸收的关键参数。【结论】种子处理通过调节干旱胁迫下玉米苗期地上农艺性状和根系形态特征，提高植株光合生理代谢和抗氧化酶活性，促进养分吸收与物质积累，其中超声+微生物处理适合北方旱地农田玉米生产。

关键词: 种子处理, 干旱胁迫, 玉米根系, 抗氧化酶, 养分吸收

Abstract:

【Objective】Seed treatments can enhance the tolerance of plants to abiotic stress. This study explored the effects of different seed treatment methods under drought stress on the growth and nutrient uptake of maize at the seedling stage, to provide a theoretical reference for maize cultivation with high yield, high efficiency and strong ecological adaptability.【Method】A pot experiment was conducted, with five seed treatments, including control (CK), ultrasound (US), ultrasound+polysaccharide (US+OS), ultrasound+microorganism (US+B), and integrated technology (US+polysaccharide+microorganism, IPM). The agronomic traits, photosynthetic physiology, antioxidant enzyme activities, and nutrient uptake of maize at the seedling stage under different simulated drought stress conditions were investigated. Combining with the Mantel test and random forest, the potential relationships among the indicators affecting the nutrient uptake of maize were analyzed.【Result】Compared with the CK, seed treatment significantly improved the plant height, leaf area, root length, root surface area, root volume and root diameter. These stimulative features were conducive to increasing the root dry weight by 69.23%-171.15% under light drought, 59.09%-218.18% under moderate drought, and 93.84%-264.52% under severe drought. Seed treatment significantly regulated the SPAD values and gas exchange parameters under drought stress, and increased the antioxidant enzyme activities, thereby improving the production of photosynthetic substances and the ability of osmotic regulation. The changes in physiological parameters directly promoted the functional metabolism of the plants, which was conducive to optimizing nutrient uptake of the leaves and roots. The US+B treatment achieved the greatest increase, with the nitrogen, phosphorus and potassium uptake of maize under drought stress increased by 31.59%-59.21% for leaves and 39.79%-82.35% for roots, 33.70%-69.94% for leaves and 51.66%-88.69% for roots, and 47.71%-78.17% for leaves and 57.59%- 108.08% for roots, respectively. The Mantel test and the random forest analysis revealed that seed treatment obviously enhanced multiple growth parameters, thereby comprehensively improving the drought resistance of maize. The plant biomass, leaf area, and plant height were the key parameters affecting nutrient uptake.【Conclusion】Seed treatment could promote nutrient uptake and dry matter accumulation by optimizing the aboveground agronomic traits and root morphological characteristics of maize at the seedling stage under drought stress, and improving the photosynthetic physiological metabolism and antioxidant enzyme activities of leaves. The US+B treatment was expected to be a potential practice for enhancing the maize productivity in the dryland farmlands of the North China.

Key words: seed treatment, drought stress, maize roots, antioxidant enzyme, nutrient uptake

杨静雯, 常力匀, 卢伟岸, 王海华, 姜英, 张雯, 宫香伟, 吕国华. 种子处理对模拟干旱胁迫玉米苗期生长与养分吸收的调控效应[J]. 中国农业科学, 2026, 59(12): 2591-2605.

YANG JingWen, CHANG LiYun, LU WeiAn, WANG HaiHua, JIANG Ying, ZHANG Wen, GONG XiangWei, LÜ GuoHua. Effects of Seed Treatment on Growth and Nutrient Uptake of Maize at the Seedling Stage Under Simulated Drought Stress[J]. Scientia Agricultura Sinica, 2026, 59(12): 2591-2605.

图/表 10

图1

图2

图3

图4

图5

图6

表1

干旱胁迫下种子处理对玉米苗期叶片光合特性的影响"

干旱胁迫 Drought stress	种子处理 Seed treatment	干旱胁迫20 d Drought stress for 20 d				干旱胁迫27 d Drought stress for 27 d
干旱胁迫 Drought stress	种子处理 Seed treatment	净光合速率 P_n (μmol·m^-2·s^-1)	气孔导度 G_s (μmol·m^-2·s^-1)	胞间CO₂ 浓度C_i (μmol·mol^-1）	蒸腾速率 T_r (mmol·m^-2·s^-1)	净光合速率 P_n (μmol·m^-2·s^-1)	气孔导度 G_s (μmol·m^-2·s^-1)	胞间CO₂ 浓度C_i (μmol·mol^-1)	蒸腾速率 T_r (mmol·m^-2·s^-1)
常规处理 Normal treatment	CK	20.53±0.59c	0.08±0.01c	21.80±1.75c	2.73±0.40c	35.60±0.53c	0.36±0.04a	150.33±9.81a	10.38±0.45ab
	US	26.43±2.03b	0.30±0.03a	25.70±2.76ab	4.47±0.60b	41.70±0.89a	0.36±0.03a	155.00±2.00a	11.28±0.68a
	US+OS	30.37±2.58b	0.15±0.01b	24.50±1.16ab	5.62±0.14a	38.83±1.01b	0.30±0.04a	123.67±1.15b	8.66±0.94c
	US+B	36.57±1.79a	0.18±0.01b	25.20±1.82a	4.50±0.40b	37.70±1.31b	0.29±0.02a	125.00±1.15b	8.83±0.42c
	IPM	35.07±1.37a	0.18±0.01b	22.80±1.78bc	6.62±0.75a	39.00±0.56b	0.33±0.08a	150.33±1.53a	9.95±0.07b
轻度干旱 Light drought	CK	30.77±0.90c	0.16±0.01b	44.00±3.57d	3.96±0.43c	19.77±0.85c	0.15±0.01b	99.13±9.42c	6.44±0.90b
	US	37.60±2.01b	0.21±0.02ab	51.77±1.25c	5.44±0.41b	26.30±1.97a	0.20±0.01a	122.27±4.56b	7.83±0.42a
	US+OS	39.90±1.64ab	0.23±0.02a	58.57±1.61ab	5.66±0.38ab	23.30±1.15b	0.20±0.02a	146.67±11.15a	8.00±0.81a
	US+B	42.20±0.61a	0.23±0.01a	62.30±2.34a	6.17±0.26a	24.33±1.36ab	0.16±0.01b	99.07±3.00c	6.91±0.15ab
	IPM	39.37±1.25b	0.22±0.01a	54.97±1.08bc	5.62±0.2ab	22.23±1.40bc	0.15±0.01b	106.17±9.25c	6.45±0.55b
中度干旱 Moderate drought	CK	27.40±1.71c	0.17±0.01bc	22.63±1.07d	3.36±0.23c	20.07±0.25d	0.14±0.01ab	67.03±3.80ab	5.99±0.83a
	US	36.83±1.29b	0.16±0.02c	30.43±0.71c	5.07±0.29a	23.63±0.42bc	0.14±0.01ab	47.03±3.03d	5.94±0.39a
	US+OS	34.40±1.10b	0.16±0.02c	46.63±1.99a	4.34±0.08b	22.47±1.27c	0.13±0.01b	75.03±3.87a	5.70±0.39c
	US+B	36.47±2.02b	0.21±0.03b	42.60±2.10b	4.95±0.18a	25.73±1.27a	0.15±0.01a	55.30±7.38cd	6.36±0.42a
	IPM	39.63±0.29a	0.24±0.02a	42.10±1.59b	5.09±0.10a	24.83±1.25ab	0.14±0.01ab	62.80±7.26bc	5.72±0.55a
重度干旱 Severe drought	CK	25.83±1.60c	0.17±0.01c	38.20±0.62d	5.44±0.17c	19.13±0.80c	0.13±0.05b	69.77±0.55ab	4.49±0.83c
	US	34.57±2.12b	0.19±0.01c	44.00±2.43c	6.46±0.49b	29.33±0.75a	0.14±0.02ab	65.10±2.82c	5.34±0.36bc
	US+OS	41.53±2.06a	0.34±0.03a	44.70±1.73c	7.51±0.30a	24.70±0.26b	0.15±0.01ab	85.43±4.06a	5.83±0.48ab
	US+B	41.33±0.21a	0.27±0.01b	55.43±1.84a	7.92±0.58a	24.40±0.66b	0.15±0.02ab	72.50±4.8ab	6.06±0.43ab
	IPM	33.93±1.91b	0.20±0.02c	50.93±1.76b	5.34±0.11c	28.57±0.01a	0.18±0.01a	73.60±4.49ab	6.71±0.55a
ANOVA
干旱胁迫Drought stress		**	**	**	**	**	**	**	**
种子处理Seed treatment		**	**	**	**	**	ns	**	*
干旱胁迫×种子处理 Drought stress ×Seed treatment		**	**	**	**	**	*	**	**

表1

图7

图8

图9

参考文献 37

[1]	XUE Z Y, CHEN Y, YIN Y X, CHEN W L, JIAO Y, DENG P, DAI S B. Spatio-temporal characteristics and driving factors of flash drought in northern China from 1978 to 2020. Global and Planetary Change, 2024, 232: 104326. doi: 10.1016/j.gloplacha.2023.104326
[2]	ELLOUZI H, BEN SLIMENE DEBEZ I, AMRAOUI S, RABHI M, HANANA M, ALYAMI N M, DEBEZ A, ABDELLY C, ZORRIG W. Effect of seed priming with auxin on ROS detoxification and carbohydrate metabolism and their relationship with germination and early seedling establishment in salt stressed maize. BMC Plant Biology, 2024, 24(1): 704. doi: 10.1186/s12870-024-05413-w pmid: 39054427
[3]	ZHUANG D W, LI H B, WANG Y Q, ZHOU D M, ZHAO L J. Nanoparticle-elicited eustress intensifies cucumber plant adaptation to water deficit. Environmental Science & Technology, 2025, 59(7): 3613-3623. doi: 10.1021/acs.est.4c13531
[4]	JIANG Y J, LI Q H, MAO W Q, TANG W T, WHITE J F Jr, LI H Y. Endophytic bacterial community of Stellera chamaejasme L. and its role in improving host plants' competitiveness in grasslands. Environmental Microbiology, 2022, 24(8): 3322-3333. doi: 10.1111/emi.v24.8
[5]	PARMAR S, SHARMA V K, LI T, TANG W T, LI H Y. Fungal seed endophyte FZT214 improves Dysphania ambrosioides Cd tolerance throughout different developmental stages. Frontiers in Microbiology, 2022, 12: 783475. doi: 10.3389/fmicb.2021.783475
[6]	姜雅文, 解文艳, 何久兴, 龚民, 霍秋燕, 杨浠, 韩伟, 吕国华. 微生物种子包衣促进干旱条件下小麦种子萌发及幼苗生长. 中国农业气象, 2025, 46(5): 609-618.
	JIANG Y W, XIE W Y, HE J X, GONG M, HUO Q Y, YANG X, HAN W, LÜ G H. Microbial seed coating promote wheat seed germination and seedling growth under drought stress condition. Chinese Journal of Agrometeorology, 2025, 46(5): 609-618. (in Chinese) doi: 10.3969/j.issn.1000-6362.2025.05.002
[7]	AIN N U, NAVEED M, HUSSAIN A, MUMTAZ M Z, RAFIQUE M, BASHIR M A, ALAMRI S, SIDDIQUI M H. Impact of coating of urea with Bacillus-augmented zinc oxide on wheat grown under salinity stress. Plants, 2020, 9(10): 1375. doi: 10.3390/plants9101375
[8]	赵艳军, 王宝军, 刘婧, 魏盼, 秦海峰, 梁镇海. 超声对小麦种子萌发特性的生物学效应研究. 种子, 2012, 31(9): 112-114.
	ZHAO Y J, WANG B J, LIU J, WEI P, QIN H F, LIANG Z H. The biological effect of ultrasound on wheat seeds germination. Seed, 2012, 31(9): 112-114. (in Chinese)
[9]	郭丹钊, 王斌, 朱彬. 干旱胁迫条件下细菌多糖对玉米苗期抗旱性的影响. 安徽农业科学, 2007, 35(34): 11050-11051.
	GUO D Z, WANG B, ZHU B. Effect of bacterial polysaccharides on maize drought resistance in the seedling stage under drought stress. Journal of Anhui Agricultural Sciences, 2007, 35(34): 11050-11051. (in Chinese)
[10]	REHMAN A, FAROOQ M, NAVEED M, NAWAZ A, SHAHZAD B. Seed priming of Zn with endophytic bacteria improves the productivity and grain biofortification of bread wheat. European Journal of Agronomy, 2018, 94: 98-107. doi: 10.1016/j.eja.2018.01.017
[11]	AMUTHA M. Establishment of Beauveria bassiana (balsamo) vuillemin as an endophytein cotton. International Journal of Current Microbiology and Applied Sciences, 2017, 6(6): 2506-2513.
[12]	WANG J, LIU W B, YIN D Q. Impacts of integrated meteorological and agricultural drought on global maize yields. Agricultural Water Management, 2025, 318: 109727. doi: 10.1016/j.agwat.2025.109727
[13]	VADEZ V, GRONDIN A, CHENU K, HENRY A, LAPLAZE L, MILLET E J, CARMINATI A. Crop traits and production under drought. Nature Reviews Earth & Environment, 2024, 5(3): 211-225.
[14]	韩雪, 高馨竹, 龚伟军, 宋金钊, 赵良洲, 李海燕. 微生物种子包衣的应用与研究进展. 微生物学通报, 2023, 50(12): 5534-5547.
	HAN X, GAO X Z, GONG W J, SONG J Z, ZHAO L Z, LI H Y. The microbial seed coating and its application. Microbiology China, 2023, 50(12): 5534-5547. (in Chinese)
[15]	罗晟, 赵泽文, 任新宇, 魏宏宇, 马雅静, 潘起涛, 李荣同, 龚国胜, 程新. 屎肠球菌胞外多糖对镉胁迫下水稻种子萌发及幼苗生长的影响. 农业环境科学学报, 2020, 39(9): 1888-1899.
	LUO S, ZHAO Z W, REN X Y, WEI H Y, MA Y J, PAN Q T, LI R T, GONG G S, CHENG X. Effect of exopolysaccharide from Enterococcus faecium on rice seed germination and seedling growth under cadmium stress. Journal of Agro-Environment Science, 2020, 39(9): 1888-1899. (in Chinese)
[16]	王荣荣, 陈天鹏, 尹豪杰, 蒋桂英. 不同抗旱性春小麦根系生长对干旱胁迫的响应及滴灌复水补偿效应. 中国农业科学, 2023, 56(24): 4826-4841. doi:10.3864/j.issn.0578-1752.2023.24.003.
	WANG R R, CHEN T P, YIN H J, JIANG G Y. Response and drip irrigation re-watering compensation effect of spring wheat roots to drought stress with different drought tolerance varieties. Scientia Agricultura Sinica, 2023, 56(24): 4826-4841. doi:10.3864/j.issn.0578-1752.2023.24.003. (in Chinese)
[17]	ZVINAVASHE A T, LIM E, SUN H, MARELLI B. A bioinspired approach to engineer seed microenvironment to boost germination and mitigate soil salinity. PNAS, 2019, 116(51): 25555-25561. doi: 10.1073/pnas.1915902116 pmid: 31776251
[18]	柳嘉怡, Diao Zhuo, 魏嘉琳, 王海华. 浒苔多糖铁对盐胁迫条件下小麦生长的影响. 现代农业科技, 2021(3): 1-3.
	LIU J Y, DIAO Z, WEI J L, WANG H H. Effect of Enteromorpha polysaccharide iron on wheat growth under salt-stress. Modern Agricultural Science and Technology, 2021(3): 1-3. (in Chinese)
[19]	闫振华, 刘东尧, 贾绪存, 杨琴, 陈艺博, 董朋飞, 王群. 花期高温干旱对玉米雄穗发育、生理特性和产量影响. 中国农业科学, 2021, 54(17): 3592-3608. doi:10.3864/j.issn.0578-1752.2021.17.004.
	YAN Z H, LIU D Y, JIA X C, YANG Q, CHEN Y B, DONG P F, WANG Q. Maize tassel development, physiological traits and yield under heat and drought stress during flowering stage. Scientia Agricultura Sinica, 2021, 54(17): 3592-3608. doi:10.3864/j.issn.0578-1752.2021.17.004. (in Chinese)
[20]	ZHANG W, LONG A R, JI X J, SUN Z X, TIAN P, JIN C C, GONG X W, JIANG Y, QI H, YU H Q. Tillage combined with straw return increases maize yield and water use by regulating root morphological distribution and nitrogen metabolism in Northeast China. Soil and Tillage Research, 2026, 256: 106876. doi: 10.1016/j.still.2025.106876
[21]	PIRI R, MORADI A, BALOUCHI H, SALEHI A. Improvement of cumin (Cuminum cyminum) seed performance under drought stress by seed coating and biopriming. Scientia Horticulturae, 2019, 257: 108667. doi: 10.1016/j.scienta.2019.108667
[22]	张思聪, 孔雷蕾, 唐湘如. 超声波预处理对作物种子及幼苗的影响综述. 安徽农业科学, 2017, 45(21): 11-12, 43.
	ZHANG S C, KONG L L, TANG X R. Effects of ultrasonic pretreatment on seed and seedling. Journal of Anhui Agricultural Sciences, 2017, 45(21): 11-12, 43. (in Chinese)
[23]	VOOTHULURU P, WU Y J, SHARP R E. Not so hidden anymore: Advances and challenges in understanding root growth under water deficits. The Plant Cell, 2024, 36(5): 1377-1409. doi: 10.1093/plcell/koae055 pmid: 38382086
[24]	MAYEGOWDA S B, GADILINGAPPA M N. Microbial siderophores: a new insight on healthcare applications. BME Frontiers, 2025, 6: 112. doi: 10.34133/bmef.0112
[25]	GUO J L, SUN Y Y, RAO Z W, CHEN Y S, LIU Z J, JIN H, HE X, YANG Z P, ZHANG Q. Drought stress at different growth stages decrease dry weight and active constituent yield accumulation by inhibiting root morphology in Scutellaria baicalensis Georgi. Industrial Crops and Products, 2025, 223: 120280. doi: 10.1016/j.indcrop.2024.120280
[26]	ZOU J, HU W, LI Y X, ZHU H H, HE J Q, WANG Y H, MENG Y L, CHEN B L, ZHAO W Q, WANG S S, ZHOU Z G. Leaf anatomical alterations reduce cotton’s mesophyll conductance under dynamic drought stress conditions. The Plant Journal, 2022, 111(2): 391-405. doi: 10.1111/tpj.v111.2
[27]	宫香伟, 刘春娟, 冯乃杰, 郑殿峰, 王畅. S3307和DTA-6对大豆不同冠层叶片光合特性及产量的影响. 植物生理学报, 2017, 53(10): 1867-1876.
	GONG X W, LIU C J, FENG N J, ZHENG D F, WANG C. Effects of plant growth regulators S₃₃₀₇ and DTA-6 on photosynthetic characteristics and yield in soybean canopy. Plant Physiology Journal, 2017, 53(10): 1867-1876. (in Chinese)
[28]	齐先科, 李淼, 李彩虹, 渠琛玲. 介质阻挡放电低温等离子体处理对小麦种子活力及幼苗生理的影响. 农业工程学报, 2024, 40(1): 301-309.
	QI X K, LI M, LI C H, QU C L. Wheat seed vigor and seedling physiology using dielectric barrier discharge plasmas. Transactions of the Chinese Society of Agricultural Engineering, 2024, 40(1): 301-309. (in Chinese)
[29]	ZHANG K K, HAN X M, FU Y F, KHAN Z, ZHANG B J, BI J G, HU L Y, LUO L J. Biochar coating promoted rice growth under drought stress through modulating photosynthetic apparatus, chloroplast ultrastructure, stomatal traits and ROS homeostasis. Plant Physiology and Biochemistry, 2024, 216: 109145. doi: 10.1016/j.plaphy.2024.109145
[30]	王慧, 王冬梅, 张泽洲, 任怀新, 黄薇, 谢正丰. 外源褪黑素对干旱胁迫下黑麦草和苜蓿抗氧化能力及养分吸收的影响. 应用生态学报, 2022, 33(5): 1311-1319. doi: 10.13287/j.1001-9332.202205.004
	WANG H, WANG D M, ZHANG Z Z, REN H X, HUANG W, XIE Z F. Effects of exogenous melatonin on antioxidant capacity and nutrient uptake of Lolium perenne and Medicago sativa under drought stress. Chinese Journal of Applied Ecology, 2022, 33(5): 1311-1319. (in Chinese) doi: 10.13287/j.1001-9332.202205.004
[31]	CHEN S, LIU H L, YANGZONG Z, GARDEA-TORRESDEY J L, WHITE J C, ZHAO L J. Seed priming with reactive oxygen species-generating nanoparticles enhanced maize tolerance to multiple abiotic stresses. Environmental Science & Technology, 2023, 57(48): 19932-19941. doi: 10.1021/acs.est.3c07339
[32]	栗晗, 江尚焘, 彭海英, 李培根, 顾长宜, 张金莲, 陈廷速, 徐阳春, 沈其荣, 董彩霞. 接种土著和外源AM真菌对杜梨抗旱性的影响及其适应机制. 中国农业科学, 2024, 57(1): 159-172. doi:10.3864/j.issn.0578-1752.2024.01.011.
	LI H, JIANG S T, PENG H Y, LI P G, GU C Y, ZHANG J L, CHEN T S, XU Y C, SHEN Q R, DONG C X. Effects of inoculation with indigenous and exogenous arbuscular mycorrhizal fungi on drought resistance of Pyrus betulaefolia and its adaptation mechanism. Scientia Agricultura Sinica, 2024, 57(1): 159-172. doi:10.3864/j.issn.0578-1752.2024.01.011. (in Chinese)
[33]	PANTIGOSO H A, OSSOWICKI A, STRINGLIS I A, CARRIÓN V J. Hub metabolites at the root-microbiome interface: Unlocking plant drought resilience. Trends in Plant Science, 2025, 30(9): 1046-1059. doi: 10.1016/j.tplants.2025.04.007
[34]	FAN X Y, GE A H, QI S S, GUAN Y F, WANG R, YU N, WANG E T. Root exudates and microbial metabolites: signals and nutrients in plant-microbe interactions. Science China Life Sciences, 2025, 68(8): 2290-2302. doi: 10.1007/s11427-024-2876-0
[35]	QIAO Z N, SHI Y L, YI J J, ZHU J Q, KANG Q Z, QU L B, YANG R, LU J K, ZHAO C C. Low frequency ultrasound enhanced the antioxidant activity and isoflavones accumulation of soybean sprouts by inducing oxidant stress. Food Bioscience, 2024, 60: 104360. doi: 10.1016/j.fbio.2024.104360
[36]	WU K J, HU C X, LIAO P Y, HU Y L, SUN X C, TAN Q L, PAN Z Y, XU S J, DONG Z H, WU S W. Potassium stimulates fruit sugar accumulation by increasing carbon flow in Citrus sinensis. Horticulture Research, 2024, 11(11): uhae240.
[37]	ZHANG M L, HU Y Y, HAN W, CHEN J, LAI J S, WANG Y. Potassium nutrition of maize: Uptake, transport, utilization, and role in stress tolerance. The Crop Journal, 2023, 11(4): 1048-1058. doi: 10.1016/j.cj.2023.02.009