Comprehensive Evaluation of Drought Tolerance and Physiological Regulatory Mechanism of Wheat Seedlings

doi:10.15933/j.cnki.1004-3268.2026.07.001

Abstract

Abstract: With 120 wheat germplasms as materials，PEG6000 solution was used to simulate drought stress at seedling stage.Plant height，root length，fresh weight of shoot，dry weight of shoot，fresh weight of root，dry weight of root，root⁃shoot ratio and SPAD value were determined，and their drought tolerance coefficients were calculated.Principal component analysis（PCA）was performed based on their drought tolerance coefficients，cluster analysis was conducted using the comprehensive drought tolerance value（D），and comprehensive evaluation of drought tolerance of wheat germplasms was carried out.Highly drought⁃tolerant and drought⁃sensitive germplasms were selected to determine H₂O₂，malondialdehyde（MDA），proline（Pro），soluble sugar（SS）contents and catalase（CAT）activity，so as to analyze the physiological regulatory mechanism of drought tolerance.The pre⁃experiment confirmed that 25% was the optimal PEG6000 concentration，all phenotypic traits showed extremely significant difference compared with the control under this concentration. Drought stress obviously decreased fresh weight of shoot，dry weight of shoot，and SPAD value，but increased root length and root⁃shoot ratio.The correlations among phenotypic traits changed obviously under drought stress compared with those under normal water condition. Under drought stress，the correlation between root length and dry weight of shoot disappeared，SPAD value was significantly positively correlated with plant height and fresh weight.Genotype，drought and their interaction had extremely significant effects on all eight phenotypic traits.PCA showed that the first four principal components accounted for 86.67% of the total variation.The drought tolerance coefficients of all traits were extremely significantly positively correlated with D value，among which the correlation coefficients between drought tolerance coefficients of fresh weight of shoot，dry weight of shoot，fresh weight of root，dry weight of root and D value were higher， which were core indexes for drought tolerance identification. Cluster analysis based on D value showed that the wheat germplasms were divided into five groups，and 15 highly drought⁃tolerant germplasms were screened out，including Xinong 109，Luohan 22，Shannong 116，Xinmai 26，Yu’an X208，Bainong 207，Luyuan 502，Bainong 4199，Xinmai 60，Xinong 519，Wenmai 801，Saidemai 8，Yunong 186，Xinong 20，and Fengdecun 20.Compared with highly drought⁃sensitive germplasms， highly drought⁃tolerant germplasms had significantly lower H₂O₂ and MDA contents，and significantly higher CAT activity，Pro and SS contents，and CAT activity，Pro and SS contents were significantly or extremely significantly positively correlated with D value，indicating that highly drought⁃tolerant germplasms alleviate drought damage by enhancing CAT activity，accumulating Pro and SS，and reducing H_₂O_₂ and MDA accumulation.

Key words: Wheat, Germplasm resources, Seedling stage, Drought tolerance, Comprehensive evaluation, Physiological regulation

摘要： 以120份小麦种质为试验材料，利用 PEG6000溶液模拟干旱胁迫，测定株高、根长、地上部鲜质量、根鲜质量、地上部干质量、根干质量、根冠比、SPAD值，计算其耐旱系数，并基于表型性状耐旱系数进行主成分分析，然后通过耐旱性综合评价值（D）进行聚类分析，综合评价小麦种质苗期耐旱性，最后选取高耐旱种质和高敏旱种质测定其干旱胁迫条件下的过氧化氢（H₂O₂）、丙二醛（MDA）、脯氨酸（Pro）、蔗糖（SS）含量及过氧化氢酶（CAT）活性，分析其耐旱生理调控机制。结果表明，通过预试验确定小麦苗期耐旱性鉴定的最佳PEG6000溶液质量分数为25%，该质量分数下各表型性状与未干旱处理间差异均达到极显著水平。干旱胁迫处理后，小麦种质地上部鲜质量、地上部干质量及SPAD值均大幅下降，根长、根冠比均明显升高。相关性与方差分析显示，干旱胁迫条件下表型性状间的相关性较正常水分条件下发生明显改变，其中，根长与地上部干质量的相关性消失，SPAD 值与株高、根鲜质量均呈极显著正相关；基因型、干旱及二者互作对8个表型性状均产生极显著影响。基于表型性状耐旱系数的主成分分析表明，前4个主成分的累计贡献率达86.67%。8个表型性状的耐旱系数与D值均呈极显著正相关，其中，地上部鲜质量、根鲜质量、地上部干质量、根干质量的耐旱系数与D值的相关系数较高，可作为小麦苗期耐旱性鉴定的核心指标。基于D值的聚类分析表明，供试材料可划分为5个组，筛选出高耐旱种质15份，包括西农109、洛旱22、山农116、新麦26、豫安X208、百农207、鲁原502、百农4199、新麦60、西农519、温麦801、赛德麦8、豫农186、西农20、丰德存20。高耐旱种质的H₂O₂、MDA含量均显著低于高敏旱种质，CAT活性、Pro含量、SS含量均显著高于高敏旱种质，且CAT活性、Pro含量和SS含量与D值均呈显著或极显著正相关，表明高耐旱种质通过提升CAT活性、积累Pro和SS，降低H_₂O_₂和MDA含量，减轻干旱伤害。

关键词: 小麦, 种质资源, 苗期, 耐旱性, 综合评价, 生理调控

CLC Number:

S512

Liu Yuhang, Shi Yuhan, Yang Yiting, Li Xueting, Jiang Mengxin, Li Jiayi, Li Junhua, Sang Shifei, Feng Liuchun, Wang Li. Comprehensive Evaluation of Drought Tolerance and Physiological Regulatory Mechanism of Wheat Seedlings[J]. Journal of Henan Agricultural Sciences, 2026, 55(7): 1-10.

刘宇航, 史钰涵, 杨义婷, 李雪婷, 蒋梦鑫, 李佳怡, 李俊华, 桑世飞, 丰柳春, 王丽. 小麦种质资源苗期耐旱性综合评价及其生理调控机制[J]. 河南农业科学, 2026, 55(7): 1-10.

Figures/Tables 10

References

［1］Gupta P K，Balyan H S，Gahlaut V，et al. Hybrid wheat：past，present and future［J］. Theoretical and Applied Genetics，2019，132（9）：2463⁃2483.

［2］Mwadzingeni L，Shimelis H，Dube E，et al. Breeding wheat for drought tolerance：progress and technologies ［J］. Journal of Integrative Agriculture，2016，15（5）：935⁃943.

［3］Hickey L T，N Hafeez A，Robinson H，et al. Breeding crops to feed 10 billion［J］. Nature Biotechnology，2019，37（7）：744⁃754.

［4］Mao H D，Li S M，Wang Z X，et al. Regulatory changes in TaSNAC8‑6A are associated with drought tolerance in wheat seedlings［J］. Plant Biotechnology Journal，2020，18（4）：1078⁃1092.

［5］傅晓艺，王红光，刘志连，等.水分胁迫对不同小麦幼苗期生长的影响及抗旱品种筛选［J］. 作物杂志，2023（4）：224⁃229.

Fu X Y，Wang H G，Liu Z L，et al. Effects of water stress on growth of different wheat varieties at seedling stage and selection of drought resistant varieties［J］.Crops，2023（4）：224⁃229.

［6］Zhang H，Zhao Y，Zhu J K. Thriving under stress：how plants balance growth and the stress response［J］.Developmental Cell，2020，55（5）：529⁃543.

［7］Ahmed H G M，Zeng Y W，Shah A N，et al. Conferring of drought tolerance in wheat（Triticum aestivum L.）genotypes using seedling indices［J］. Frontiers in Plant Science，2022，13：961049.

［8］李瑞雪，孙任洁，汪泰初，等. 植物抗旱性鉴定评价方法及抗旱机制研究进展［J］. 生物技术通报，2017，33（7）：40⁃48.

Li R X，Sun R J，Wang T C，et al. Research progress on identification and evaluation methods，and mechanism of drought resistance in plants［J］. Biotechnology Bulletin，2017，33（7）：40⁃48.

［9］周全，路秋梅，赵张晨，等. 244份春小麦苗期抗旱性的鉴定［J］. 中国农业科学，2024，57（9）：1646⁃1657.

Zhou Q，Lu Q M，Zhao Z C，et al. Identification of drought resistance of 244 spring wheat varieties at seedling stage［J］. Scientia Agricultura Sinica，2024，57（9）：1646⁃1657.

［10］Run Y H，Cheng X Y，Dou W，et al. Wheat potassium transporter TaHAK13 mediates K+ absorption and maintains potassium homeostasis under low potassium stress［J］. Frontiers in Plant Science，2022，13：1103235.

［11］Muhammad I，Yang L，Ahmad S，et al.Melatonin⁃priming enhances maize seedling drought tolerance by regulating the antioxidant defense system［J］. Plant Physiology，2023，191（4）：2301⁃2315.

［12］Han S C，Wang Y L，Li Y X，et al. The OsNAC41⁃RoLe1⁃OsAGAP module promotes root development and drought resistance in upland rice［J］.Molecular Plant，2024，17（10）：1573⁃1593.

［13］Zhou Y X，Wang D Z，Wang H，et al. Integrative omics of the genetic basis for wheat WUE and drought resilience reveal the function of TaMYB7⁃A1［J］.Nature Communications，2025，16：8622.

［14］关睿捷，温嘉茂，张恬畅，等. 114份春小麦苗期耐旱性综合评价与筛选［J］. 种子，2025，44（10）：28⁃36.
Guan R J，Wen J M，Zhang T C，et al. Comprehensive evaluation and screening of drought tolerance in 114 spring wheat seedlings［J］. Seed，2025，44（10）：28⁃36.

［15］胡润慧，汪军成，司二静，等. 小麦苗期耐旱耐盐种质筛选及抗旱耐盐综合评价［J］. 作物学报，2025，51（9）：2371⁃2386.
Hu R H，Wang J C，Si E J，et al. Screening of drought and salt tolerant germplasm during wheat seedling stage and comprehensive evaluation of drought and salt tolerance［J］. Acta Agronomica Sinica，2025，51 （9）：2371⁃2386.

［16］Ahmed K，Shabbir G，Ahmed M. Exploring drought tolerance for germination traits of diverse wheat genotypes at seedling stage：a multivariate analysis approach［J］. BMC Plant Biology，2025，25（1）：390.

［17］Tardieu F，Simonneau T，Muller B. The physiological basis of drought tolerance in crop plants：a scenario⁃dependent probabilistic approach［J］. Annual Review of Plant Biology，2018，69：733⁃759.

［18］Bapela T，Shimelis H，Tsilo T J，et al. Genetic improvement of wheat for drought tolerance：progress，challenges and opportunities［J］. Plants，2022，11（10）：1331.

［19］Eziz A，Yan Z B，Tian D，et al. Drought effect on plant biomass allocation：a meta⁃analysis［J］.Ecology and Evolution，2017，7（24）：11002⁃11010.

［20］Laxa M，Liebthal M，Telman W，et al. The role of the plant antioxidant system in drought tolerance［J］.Antioxidants，2019，8（4）：94.

［21］Pecka J，Kraus K，Zelený M，et al. Exogenous proline modulates physiological responses and induces stress memory in wheat under repeated and delayed drought stress［J］. Agronomy，2025，15（6）：1370.

［22］彭超军，华夏，王松峰，等. 干旱胁迫对不同小麦品种灌浆期旗叶光合及生理特性的影响［J］. 河南农业科学，2025，54（2）：40⁃47.
Peng C J，Hua X，Wang S F，et al. Effect of drought stress on photosynthetic and physiological characteristics of flag leaves of different wheat varieties at filling stage［J］. Journal of Henan Agricultural Sciences，2025，54（2）：40⁃47.

［23］黄钰，翁雪莲，Indrila Dey Traye，等. 基于萌发期和苗期主要形态指标的耐旱小麦种质筛选与评价［J］. 江苏农业科学，2025，53（14）：106⁃116.
Huang Y，Weng X L，Traye I D，et al. Screening and evaluation of drought⁃tolerant wheat germplasms based on main morphological indices at germination and seedling stages［J］. Jiangsu Agricultural Sciences，2025，53（14）：106⁃116.

［24］Yan S Y，Weng B S，Jing L S，et al. Effects of drought stress on water content and biomass distribution in summer maize（Zea mays L.）［J］. Frontiers in Plant Science，2023，14：1118131.

［25］周敏姑，闫云才，高文，等. 基于多光谱遥感和CNN的玉米地上生物量估算模型［J］. 农业机械学报，2024，55（9）：238⁃248.
Zhou M G，Yan Y C，Gao W，et al. Estimating aboveground biomass of maize based on multispectral remote sensing and convolutional neural network［J］.Transactions of the Chinese Society for Agricultural Machinery，2024，55（9）：238⁃248.

［26］Narayanan S，Mohan A，Gill K S，et al. Variability of root traits in spring wheat germplasm［J］. PLoS One，2014，9（6）：e100317.

［27］Wu Y，Wang H Z，Yang X W，et al. Soil water effect on root activity，root weight density，and grain yield in winter whea［t J］. Crop Science，2017，57（1）：437⁃443.

［28］Fasoula V A，Fasoula D A. Principles underlying genetic improvement for high and stable crop yield potential［J］. Field Crops Research，2002，75（2/3）：191⁃209.

［29］Haghpanah M，Hashemipetroudi S，Arzani A，et al.Drought tolerance in plants：physiological and molecular responses［J］. Plants，2024，13（21）：2962.

［30］Tian G，Wang S B，Wu J H，et al. Allelic variation of TaWD40‑4B.1 contributes to drought tolerance by modulating catalase activity in wheat［J］. Nature Communications，2023，14（1）：1200.

［31］Zhu C G，Jing B Y，Lin T，et al. Phosphorylation of sugar transporter TST2 by protein kinase CPK27 enhances drought tolerance in tomato［J］. Plant Physiology，2024，195（2）：1005⁃1024.

［32］王安泽，张阿里，窦松岩，等. 不同小麦品种苗期抗旱性鉴定及根际微生物功能研究［J］. 干旱地区农业研究，2024，42（6）：8⁃18.
Wang A Z，Zhang A L，Dou S Y，et al. Identification of drought resistance of different wheat varieties at seedling stage and functional analysis of rhizosphere microorganisms［J］. Agricultural Research in the Arid Areas，2024，42（6）：8⁃18.

［33］Gao C X. Genome engineering for crop improvement and future agriculture［J］. Cell，2021，184（6）：1621⁃1635.

［34］Liang Y M，Liu H J，Yan J B，et al. Natural variation in crops：realized understanding，continuing promise［J］.Annual Review of Plant Biology，2021，72：357⁃385.

[1]	Li Fangjie, Shi Dakun, Wei Xiaoyi, Wei Feng, Liu Junheng, Mei Yongxian. Comprehensive Evaluation of Lodging Resistance of 78 Maize Inbred Lines and Screening of Core Indexes [J]. Journal of Henan Agricultural Sciences, 2026, 55(7): 11-19.
[2]	ZHOU Jinghang, WANG Zenghao, ZHANG Lei, ZHANG Xia, LIU Jiajia, HU Tiezhu, ZHANG Dazhong. Effects of Addition Amounts，Treatment Durations and Pretreatment Durations of Different Chalcogens on Cadmium Absorption and Transport of Wheat Seedlings under Cadmium Stress [J]. Journal of Henan Agricultural Sciences, 2026, 55(6): 1-10.
[3]	WANG Yubo, WANG Yixing, DING Lin, XU Hao, ZHAO Wei, LI Na, JIE Yize, TIAN Zhicheng, MENG Panpan, TANG Jianwei, DONG Chunhao, LI Qiaoyun, YIN Guihong, HUANG Zhenpu. Identification and Agronomic Trait Analysis of Pre⁃harvest Sprouting⁃Resistant Mutants of Zhoumai 22 [J]. Journal of Henan Agricultural Sciences, 2026, 55(6): 11-22.
[4]	WANG Junmei, LIU Lulu, LIU Kai, LI Yahong, FENG Chaohong, SHI Ruijie, HAN Zihang, DU Xijun, XU Fei. Molecular Response of Wheat Serine/Threonine Kinase to Blumeria graminis f.sp.tritici and Plant Hormones [J]. Journal of Henan Agricultural Sciences, 2026, 55(6): 81-91.
[5]	LIANG Lu, YIN Kun, ZHANG Weijie, ZHUANG Qiuli, XU Bohan, TIAN Kaiyue, ZHANG Dongjie, ZHANG Donghui, HUANG Yubo. Impact of Vermicompost Application Rate for Winter Wheat on Dry Matter Accumulation，Transportation，and Yield of Subsequent Maize in Lime Concretion Black Soil [J]. Journal of Henan Agricultural Sciences, 2026, 55(5): 43-52.
[6]	TIAN Luyao, CHEN Fabo, SUN Chun, YAN Qingjiu. Evaluation of Drought Resistance of Radish under Combined Stress of High Temperature and Drought Based on Multiple Indicators [J]. Journal of Henan Agricultural Sciences, 2026, 55(5): 116-127.
[7]	MA Xuhui, ZHANG Fuyan, ZHAO Wan, WANG Jiahuan, CHENG Zhongjie, ZHANG Zhongzhou, LI Liulong, DING Jiacun, ZHAO Chi, WANG Hanfeng, CHEN Xiaojie. Analysis of High and Stable Yield Performance and Genetic Characteristics of New Wheat Cultivar Yutong 112 [J]. Journal of Henan Agricultural Sciences, 2026, 55(4): 23-32.
[8]	ZHANG Yunhong, YANG Yonghui, ZHANG Sensen, ZHANG Zeli, ZHANG Shuiqing, GAO Cuimi, HE Fang, HAN Weifeng, ZHANG Huan, PAN Xiaoying, WANG Bohan. Effects of Different Soil Conditioners on the Quality of Acidified Sandy Ginger Black Soil，and the Growth and Nutrient Absorption of Wheat [J]. Journal of Henan Agricultural Sciences, 2026, 55(4): 79-89.
[9]	TAO Hua, DING Jingli, REN Zhijie, HE Ailing, LIU Gaoyuan, DU Jun. Effects of Nitrogen Reduction Combined with Different Types of Fertilizers on Yield and Nitrogen Use of Winter Wheat [J]. Journal of Henan Agricultural Sciences, 2026, 55(2): 1-11.
[10]	WEI Xiaoyi, LIU Zhicheng, ZHANG Zhanhui, SHI Dakun, LI Fangjie, HONG Defeng, SUN Pei, LI Zhi, WEI Feng. Preliminary Study on Molecular Mechanism of Drought Tolerance of Maize Inbred Line Xin 4095 Based on miRNA‐mRNA Integration Analysis Technology [J]. Journal of Henan Agricultural Sciences, 2026, 55(1): 40-51.
[11]	NI Tianhong, ZHANG He, XU Hui, GONG Shoufu, ZHANG Yan, CHEN Liwen. Effect of Storage Temperature on Seed Quality of Camellia oleifera Abel [J]. Journal of Henan Agricultural Sciences, 2026, 55(1): 173-180.
[12]	ZHANG Suyu, LIU Hongjie, REN Dechao, YANG Mingda, GE Jun, ZHAO Jingling, ZHU Peipei, ZHENG Dongfang. Effects of Exogenous Regulators on Superior and Inferior Grain Filling and Yield of Wheat [J]. Journal of Henan Agricultural Sciences, 2025, 54(9): 23-33.
[13]	WANG Haodong, LU Yifang, YANG Shan, LIU Fengying, WANG Gang. Effect of the gacS Gene in Pseudomonas chlororaphis on the Biocontrol Efficacy against Wheat Take‑all Disease [J]. Journal of Henan Agricultural Sciences, 2025, 54(9): 112-118.
[14]	MA Shangying, XIA Tingting, HAN Pengbin, ZHANG Mengjiao, MAO Yingjie, WANG Zhiqiang, XIN Zeyu, LIN Tongbao, LIAN Yanhao, REN Yongzhe. Effects of Application Rate and Method of Nitrogen on Winter Wheat Nitrogen Uptake and Utilization，Yield and Soil Nitrate Nitrogen Residue under Wide‑Narrow Row Planting [J]. Journal of Henan Agricultural Sciences, 2025, 54(8): 38-50.
[15]	JIN Jianghua, XIA Bing, LI Xu, YU Dapeng, GUO Wei, CHEN Tao, WU Wenxin, LI Sijun, WANG Xinyue, HUANG Jie, DENG Xiaohua. Fuzzy Comprehensive Evaluation of Leaf Quality and Economic Traits of Paddy‑Stubble Tobacco with Root Zone Application of Microbial Fertilizer [J]. Journal of Henan Agricultural Sciences, 2025, 54(8): 82-91.