外源EBR调控大豆幼苗抗旱的根系响应

doi:10.11924/j.issn.1000-6850.casb2025-0512

摘要/Abstract

摘要：

干旱胁迫严重抑制大豆根系生长与活力，为探明外源2,4-表油菜素内酯（EBR）调控大豆幼苗抗旱的根系生理机制，以'辽鲜一号'为材料，采用盆栽控水法，设置0.5、1.0、1.5、2.0 mg/L 4个EBR浓度，研究其对干旱胁迫下大豆幼苗根系形态、保护酶活性、活性氧积累、伤流强度及内源激素（ABA、CTK）的影响。结果表明：与CK相比，干旱胁迫极显著降低了大豆幼苗的主根长、根表面积、根体积、主根直径及根干重(P<0.01)，提高活性氧积累并降低根系活力；叶面喷施1.0~1.5 mg/L EBR能显著缓解干旱胁迫对大豆幼苗根系生长的抑制，其中，1.0 mg/L EBR处理效果最优，可使大豆幼苗主根长、根表面积、根体积、主根直径、根干重等分别较单纯干旱处理分别提高46.6%、30.6%、34.9%、43.8%、40.0%，且上述各指标均显著高于单纯干旱胁迫处理。叶面喷施0.5~1.5 mg/LEBR后极显著提高了干旱胁迫下大豆幼苗根尖的超氧化物歧化酶(SOD)、过氧化物酶(POD)和过氧化氢酶(CAT)等保护酶活性(P<0.01)，极显著降低了超氧阴离子自由基(O₂^-·)产生速率和过氧化氢(H₂O₂)含量(P<0.01)，不同程度地降低了根系伤流液中ABA含量，提高了CTK含量和根系伤流强度，以1.0 mg/L EBR处理大豆幼苗根尖保护酶活性、伤流强度和CTK含量最高，O₂^-·产生速率、H₂O₂含量以及ABA含量最低。1.0 mg/L EBR处理大豆幼苗根尖SOD活性、POD活性和CAT活性分别较单纯干旱处理提高了14.1%、17.2%和50.6%，O₂^-·产生速率和H₂O₂含量分别较单纯干旱处理降低了33.0%和17.8%。因此，外源EBR是通过增强干旱胁迫下大豆幼苗根系伤流强度和根尖保护酶活性、提高CTK含量、降低ABA含量以及抑制活性氧积累来缓解干旱胁迫。以1.0 mg/L EBR处理对大豆幼苗抗旱性诱导效果最好。本研究为大豆抗旱化控栽培提供理论依据，未来可进一步开展花期/结荚期调控与产量形成机制研究。

关键词: 大豆, 根系生长, 保护酶活性, 活性氧积累, 伤流强度, 激素含量

Abstract:

To explore the effects of exogenous 2,4-Epibrassinolide (EBR) on root morphology and physiology of soybean seedlings under the drought stress, using ' Liaoxian No.1' as the material, the effects of 0.5-2.0 mg/L EBR treatment on root morphology, protective enzyme activity, reactive oxygen species accumulation, bleeding intensity and endogenous hormones (ABA, CTK ) of soybean seedlings under drought stress were studied by using pot water control method. The results showed that compared with CK, the drought stress treatment extremely significantly reduced the main root length, root surface area, root volume, main root diameter and root dry weight of soybean seedlings (P<0.01). Foliar spraying of 1.0-1.5 mg/L EBR could significantly alleviate the inhibition of root growth of soybean seedlings by drought stress. Among them, the main root length, root surface area, root volume, main root diameter and root dry weight of soybean seedlings treated with 1.0 mg/L EBR increased by 46.6%, 30.6%, 34.9%, 43.8% and 40.0% respectively. And all the above indicators were significantly higher than those of simple drought stress treatment. Foliar spraying 0.5-1.5 mg/L EBR significantly increased the activities of protective enzymes such as superoxidedismutase (SOD), peroxidase (POD), and catalase (CAT) in the root tips of soybean seedlings under drought stress (P<0.01), and significantly reduced the superoxide anion radical (O₂^-·) production rate and hydrogen peroxide (H₂O₂) content in the root tips of soybean seedlings under drought stress (P<0.01), reduced the ABA content in the root wound fluid to varying degrees, and increased the CTK content and root bleeding intensity. Soybean seedlings treated with 1.0 mg/L EBR had the highest protective enzyme activity in the root tip, bleeding intensity and CTK content, while the O₂^-· production rate, H₂O₂ content and ABA content were the lowest. Compared with the simple drought treatment, the SOD activity, POD activity and CAT activity in the root tips of soybean seedlings treated with 1.0 mg/L EBR increased by 14.0%, 17.2% and 50.6% respectively, the O₂^-· roduction rate and H₂O₂ content decreased by 33.0% and 17.8% respectively. Therefore, exogenous EBR alleviated the soybean drought stress by enhancing the root bleeding intensity and root tip protection enzyme activities of soybean seedlings under drought stress, increasing the CTK content, reducing the ABA content and inhibiting the active oxygen accumulation. Treatment with 1.0 mg/L EBR had the best effects on inducing drought resistance in soybean seedlings.

Key words: soybean (Glycine max L.), root growth, protective enzyme activity, reactive oxygen species accumulation, bleeding intensity, hormone content

中图分类号:

S311作物生理学

徐芬芬, 韩金多. 外源EBR调控大豆幼苗抗旱的根系响应[J]. 中国农学通报, 2026, 42(8): 53-59.

XU Fenfen, HAN Jinduo. Root Response Mechanism of Exogenous EBR Regulating Drought Resistance in Soybean Seedlings[J]. Chinese Agricultural Science Bulletin, 2026, 42(8): 53-59.

图/表 4

参考文献 33

[1]	邓辉茗, 龙聪颖, 蔡仕珍, 等. PEG-6000模拟干旱协迫对大蓟叶片生理特性的影响[J]. 西北植物学报, 2017, 37(5):959-964.
[2]	胡国玉, 夏英萍, 杜祥备, 等. 安徽省夏大豆产量及限制因素分析[J]. 中国油料作物学报, 2023, 45(4):793-802. doi: 10.19802/j.issn.1007-9084.2022145
[3]	LE D T, NISHIYAMA R, WATANABE Y, et al. Differential gene expression in soybean leaf tissues at late developmental stages under drought stress revealed by genome-wide transcriptome analysis[J]. Plos one, 2012, 7(11):49522. doi: 10.1371/journal.pone.0049522 pmid: 23189148
[4]	耿孝宇, 张翔, 刘洋, 等. 江苏省滨海盐碱地籼粳杂交稻产量优势形成特征[J]. 作物学报, 2024(5):1253-1270. doi: 10.3724/SP.J.1006.2024.32034
[5]	CRAMER G R, LUCHLI A, EPSTEIN E. Effects of NaCl and CaCl₂ on ion activities in complex nutrient solutions and root growth of cotton[J]. Plant physiology, 1986, 81(3):792-797. doi: 10.1104/pp.81.3.792 URL
[6]	JIA Y B, YANG X E, FENG Y, et al. Differential response of root morphology to potassium deficient stress among rice genotypes varying in potassium efficiency[J]. Journal of Zhejiang University science b, 2008, 9(5):427-434. doi: 10.1631/jzus.B0710636 URL
[7]	SRINIVASARAO C H, BENZIONI A, ESHEL A, et al. Effects of salinity on root morphorlogy and nutrient acquisition by Faba beans (Vicia faba L.)[J]. Journal of the Indian society of soil science, 2004, 52(2):184-191.
[8]	徐芬芬, 郭定生, 王楠. NaCl预处理诱导大豆幼苗的抗旱性[J]. 分子植物育种, 2018, 16(10):3317-3320.
[9]	梁晓艳. 烯效唑对干旱胁迫下苗期大豆根系的调控[D]. 大庆: 黑龙江八一农垦大学, 2019.
[10]	卢洁春. 生物源植物生长物质对大豆生长发育及产量的影响[D]. 大庆: 黑龙江八一农垦大学, 2019.
[11]	张秀玲, 孙颖, 孟岩, 等. 干旱胁迫下外源水杨酸对野生大豆生理特性的影响[J]. 中国野生植物资源, 2022, 41(1):9-12,19
[12]	邹京南. 外源褪黑素对干旱胁迫下大豆光合及生长的影响[D]. 大庆: 黑龙江八一农垦大学, 2019.
[13]	王玺越. 甲哌鎓对大豆苗期干旱胁迫的调控效应[D]. 哈尔滨: 东北农业大学, 2023.
[14]	郑崇兰. 三唑酮对大豆花期长期干旱胁迫的缓解效应[D]. 南京: 南京农业大学, 2016.
[15]	马野, 原换换, 裴新涌, 等. 干旱胁迫下外源油菜素内酯对玉米幼苗光合作用和D1蛋白的调控效应[J]. 华北农学报, 2017, 32(3):118-124. doi: 10.7668/hbnxb.2017.03.018
[16]	吴杨, 高慧纯, 张必弦. 24-表油菜素内酯对盐碱胁迫下大豆生育、生理及细胞超微结构的影响[J]. 中国农业科学, 2017, 50(5):811-821. doi: 10.3864/j.issn.0578-1752.2017.05.004
[17]	殷欣, 张海玲, 吴杨. 24-表油菜素内酯对镉胁迫下大豆苗期生理特性的影响[J]. 核农学报, 2016, 30(2):0364-0371.
[18]	魏鑫, 倪虹, 张会慧, 等. 外源脱落酸和油菜素内酯对干旱胁迫下大豆幼苗抗旱性的影响[J]. 中国油料作物学报, 2016, 38(5):605-610. doi: 10.7505/j.issn.1007-9084.2016.05.010
[19]	罗晓峰, 代宇佳. 三种植物生长调节剂对大豆生长发育及产量的影响[J]. 核农学报, 2021, 35(4):0980-0988.
[20]	李合生. 植物生理生化实验原理和技术[M]. 北京: 高等教育出版,2000:164-169,258-260.
[21]	王爱国, 罗广华. 植物的超氧物自由基与羟胺反应的定量关系, 植物生理学通讯, 1990(6):55-57.
[22]	刘俊, 吕波, 徐朗莱. 植物叶片中过氧化氢含量测定方法的改进, 生物化学与生物物理进展, 2000, 27(5):548-551.
[23]	OSMONT K S, SIBOUT R, HARDTKE C S. Hidden branches: developments in root system architecture[J]. Annual review of plant biology, 2007, 58(1):93-113. doi: 10.1146/arplant.2007.58.issue-1 URL
[24]	毋翔, 张义凯, 张鹏, 等. 2,4-表油菜素内酯对生物炭基质育秧水稻秧苗根系生长及生理特性的影响[J]. 中国水稻科学, 2024, 38(6):685-694. doi: 10.16819/j.1001-7216.2024.240503
[25]	MORILLON R, CATTEROU M, SANGWAN R S, et al. Brassinolide may control aquaporin activities in Arabidopsis thaliana[J]. Planta, 2001, 212(2):199-204. doi: 10.1007/s004250000379 URL
[26]	TANVEER M, SHAHZAD B, SHARMA A, et al. 2,4-Epibrassinolide application in plants: an implication for improving drought stress tolerance in plants[J]. Plant physiology and biochemistry, 2019, 135:295-303. doi: 10.1016/j.plaphy.2018.12.013 URL
[27]	CHOI M H, SHIM S M, KIM G H. Protective effect of black raspberry seed containing anthocyanins against oxidative damage to DNA, protein, and lipid[J]. Journal of food science & technology, 2016, 53(2):1214-1221.
[28]	HASHEMINASAB H, ASSAD M T, ALIAKBARI A, et al. Influence of drought stress on oxidative damage and antioxidant defense systems in tolerant and susceptible wheat genotypes[J]. Journal of agricultural science, 2012, 4(8):20-30.
[29]	MORITA S, OKAMOTO M, ABE J, et al. Bleeding rate of field-grown maize with reference to root system development[J]. Japanese journal of crop science, 2000, 69(1):80-85. doi: 10.1626/jcs.69.80 URL
[30]	毛紫琳. 干旱胁迫对杂交稻与常规稻干物质分配与产量形成的影响[D]. 武汉: 华中农业大学, 2023.
[31]	李春艳, 李思忠, 楚光红. 不同滴水量处理下大豆根系生长与花荚形成的关系[J]. 南京农业大学学报, 2016, 39(2):198-204.
[32]	刘丹, 郭广昊, 刘磊, 等. 外源喷施油菜素内酯对碱胁迫下甜菜生长的影响[J]. 西北农业学报, 2018, 27(10):1461-1469.
[33]	刘亚西, 毛芮, 杨梦含, 等. 外源激素对干旱胁迫下黑麦草生理特性的影响及抗旱性评价[J]. 草业科学, 2024, 41(2):425-436.

处理	主根长/cm	根表面积/(cm²/株)	根体积/(cm³/株)	主根直径/mm	根干重/(g/株)
CK	13.6±1.2 aA	165.30±15.22 aA	1.20±0.20 aA	0.53±0.04 aA	0.68±0.06 aA
干旱	8.8±0.7 dC	119.86±10.17 cB	0.83±0.05 bB	0.32±0.02 cdCD	0.45±0.05 eC
干旱+0.5 mg/L EBR	10.8±0.8 bcBC	130.64±12.29 bcB	0.96±0.08 bAB	0.42±0.03 bcBC	0.56±0.04 cB
干旱+1.0 mg/L EBR	12.9±1.0 aAB	156.54±17.57 aA	1.12±0.09 aAB	0.46±0.03 bAB	0.63±0.03 abAB
干旱+1.5 mg/L EBR	11.0±0.9 bBC	136.75±10.28 bB	0.93±0.72 bB	0.38±0.03 cdBCD	0.58±0.04 bcB
干旱+2.0 mg/L EBR	9.0±0.7 cdC	125.37±11.56 bcB	0.89±0.62 bB	0.33±0.02 cdCD	0.48±0.05 deC

处理	主根长/cm	根表面积/(cm²/株)	根体积/(cm³/株)	主根直径/mm	根干重/(g/株)
CK	13.6±1.2 aA	165.30±15.22 aA	1.20±0.20 aA	0.53±0.04 aA	0.68±0.06 aA
干旱	8.8±0.7 dC	119.86±10.17 cB	0.83±0.05 bB	0.32±0.02 cdCD	0.45±0.05 eC
干旱+0.5 mg/L EBR	10.8±0.8 bcBC	130.64±12.29 bcB	0.96±0.08 bAB	0.42±0.03 bcBC	0.56±0.04 cB
干旱+1.0 mg/L EBR	12.9±1.0 aAB	156.54±17.57 aA	1.12±0.09 aAB	0.46±0.03 bAB	0.63±0.03 abAB
干旱+1.5 mg/L EBR	11.0±0.9 bBC	136.75±10.28 bB	0.93±0.72 bB	0.38±0.03 cdBCD	0.58±0.04 bcB
干旱+2.0 mg/L EBR	9.0±0.7 cdC	125.37±11.56 bcB	0.89±0.62 bB	0.33±0.02 cdCD	0.48±0.05 deC

处理	SOD活性/(U/gFW)	POD活性/(U/gFW)	CAT活性/(U/gFW)	O₂^-·产生速率/[mmol/(min·mg)]	H₂O₂含量/(μmol/gFW)
CK	132.4±12.2 cC	253.5±20.8 cC	35.7±2.2 cC	12.6±1.2 dD	23.8±1.8 dB
干旱	156.3±13.4 bB	266.4±24.6 cBC	43.3±3.6 cBC	20.3±1.8 aA	32.1±2.1 aA
干旱+0.5 mg/L EBR	168.7±15.6 abA	285.3±22.7 bB	55.4±5.2 bA	17.6±1.6 bcBC	29.2±1.7 bA
干旱+1.0 mg/L EBR	178.3±15.2 aA	312.3±13.3 aA	65.2±4.4 aA	13.6±0.8 dD	26.4±1.5 cdB
干旱+1.5 mg/L EBR	157.8±15.1 abA	283.6±15.6 bB	52.3±4.2 bB	16.3±1.5 cC	28.4±2.6 bcAB
干旱+2.0 mg/L EBR	157.0±13.5 bA	275.6±20.2 bcB	56.7±3.7 bA	18.9±1.3 abAB	30.2±2.5 abA

处理	SOD活性/(U/gFW)	POD活性/(U/gFW)	CAT活性/(U/gFW)	O₂^-·产生速率/[mmol/(min·mg)]	H₂O₂含量/(μmol/gFW)
CK	132.4±12.2 cC	253.5±20.8 cC	35.7±2.2 cC	12.6±1.2 dD	23.8±1.8 dB
干旱	156.3±13.4 bB	266.4±24.6 cBC	43.3±3.6 cBC	20.3±1.8 aA	32.1±2.1 aA
干旱+0.5 mg/L EBR	168.7±15.6 abA	285.3±22.7 bB	55.4±5.2 bA	17.6±1.6 bcBC	29.2±1.7 bA
干旱+1.0 mg/L EBR	178.3±15.2 aA	312.3±13.3 aA	65.2±4.4 aA	13.6±0.8 dD	26.4±1.5 cdB
干旱+1.5 mg/L EBR	157.8±15.1 abA	283.6±15.6 bB	52.3±4.2 bB	16.3±1.5 cC	28.4±2.6 bcAB
干旱+2.0 mg/L EBR	157.0±13.5 bA	275.6±20.2 bcB	56.7±3.7 bA	18.9±1.3 abAB	30.2±2.5 abA

激素种类	测定时间/ h	CK	干旱	干旱+0.5 mg/L EBR	干旱+1.0 mg/L EBR	干旱+1.5 mg/L EBR	干旱+2.0 mg/L EBR
ABA含量/(ng/g)	24	1.32±0.12 dC	1.53±0.15 aA	1.47±0.11 abAB	1.38±0.10 cdBC	1.42±0.12 bcBC	1.46±0.13 abAB
	48	1.35±0.09 eC	2.64±0.21 aA	2.55±0.22 bABC	2.32±0.18 dC	2.34±0.13 dC	2.43±0.19 cBC
	72	1.41±0.08 dD	1.87±0.15 aA	1.75±0.13 bABC	1.65±0.14 cC	1.69±0.15 bBC	1.76±0.16 abAB
CTK含量/(ng/g)	24	0.53±0.03 dD	0.65±0.05 cCD	0.88±0.08 aAB	0.95±0.07 aA	0.85±0.06 abAB	0.76±0.05 bBC
	48	0.55±0.02 dD	0.86±0.06 cC	1.21±0.10 bB	1.43±0.13 aA	1.25±0.11 bB	1.22±0.08 bB
	72	0.61±0.05 dD	0.93±0.06 cC	1.32 ±0.11 bB	1.53±0.15 aA	1.46±0.10 bAB	1.33±0.13 bB