氯通道抑制剂缓解栽培大豆盐伤害的离子特征

浙江农业学报 ›› 2022, Vol. 34 ›› Issue (10): 2095-2104.

氯通道抑制剂缓解栽培大豆盐伤害的离子特征

孟娜¹(), 薛辉², 魏明¹, 魏胜华¹

1.安徽工程大学生物与食品工程学院,安徽芜湖 241000
2.安徽珩戍林业规划设计有限公司,安徽芜湖 241000

收稿日期:2021-03-29 出版日期:2022-10-25 发布日期:2022-10-26
作者简介:孟娜(1977—),女,安徽宿州人,博士研究生,高级实验师,主要从事植物逆境生理与分子生物学研究。E-mail: dreammn@126.com
基金资助:
国家级大学生创新创业训练计划(202010363052);安徽工程大学国家自然科学基金预研项目(2018yyzr13)

Ion characteristics on chloride channel blocker ameliorating salt injury to Glycine max

MENG Na¹(), XUE Hui², WEI Ming¹, WEI Shenghua¹

1. College of Biological and Food Engineering, Anhui Polytechnic University, Wuhu 241000, Anhui, China
2. Anhui Hengshu Forestry Planning and Design Co., Ltd., Wuhu 241000, Anhui, China

Received:2021-03-29 Online:2022-10-25 Published:2022-10-26

摘要/Abstract

摘要：

为了从离子响应的视角探究氯通道抑制剂缓解栽培大豆幼苗盐伤害的作用机理,以大豆栽培品种绥农35为试验材料,采用大豆幼苗期盐胁迫外加氯通道抑制剂的方法,比较不同处理幼苗的相关生理指标、解剖结构和离子组的差异。结果显示:外加不同氯离子抑制剂对盐胁迫作用效果不同,分别表现为ZnCl₂能缓解栽培大豆幼苗的盐伤害作用,但尼氟灭酸(NFA)和蒽-9-羧酸(9-AC)则相反,加重了盐害作用;外加Zn²⁺,根部次生导管孔径回升,皮层细胞厚度减少,有利于降低根部总消耗,缩短水分和矿质离子横向运输的距离;外加Zn²⁺,叶部叶绿素含量、叶绿素荧光参数PSⅡ最大光化学效率(F_v/F_m)值,N $O 3 -$ 以及微量元素(Fe, Mn, Cu和Zn)含量均回升,而Cl^-含量回落介于对照组与盐处理组之间。综上表明,盐胁迫下外加Zn²⁺可重建幼苗叶部的离子稳态,降低叶部Cl^-毒害,提高幼苗抗氧化能力和光合能力,有效缓解大豆幼苗受到的伤害。

关键词: 盐胁迫, 大豆, 氯通道抑制剂, 离子稳态

Abstract:

To elucidate the mechanism on chloride channel inhibitor Z $n 2 +$ in ameliorating salt injury to soybean from the perspective of ion response, in this study, Glycine max cultivar Suinong 35 was used as the experimental material. The differences in physiological indexes, anatomical structure and ionomics of seedlings under different treatments were compared by the method of NaCl stress plus chloride channel inhibitors. The results showed that under NaCl stress plus additional three chloride channel inhibitors, ZnCl₂ could ameliorate salt injury in soybean to a certain extent, but nifluoric acid (NFA) and anthracene-9-carboxylic acid (9-AC) showed the reverse. Under NaCl stress plus Zn²⁺, the diameter of secondary vessel in root was markedly recovered, while cortex thickness in roots was decreased, which was helpful to reduce the metabolic cost and shorten the distance of water absorption from root to vascular cylinder. Under NaCl stress plus Zn²⁺, total chlorophyll content, F_v/F_m, N $O 3 -$ and microelement contents (Fe, Mn, Cu and Zn) rose again, while leaf Cl^- content was reduced between the control and NaCl-stressed seedlings. In summary, NaCl stress plus Zn²⁺ can effectively ameliorate salt injury to soybean seedlings by reestablishing ion homeostasis, which reduces Cl^- toxicity in leaves and improves antioxidant capacity and photosynthetic capacity.

Key words: salt stress, Glycine max, chloride channel blocker, ion homeostasis

中图分类号:

S565.1

孟娜, 薛辉, 魏明, 魏胜华. 氯通道抑制剂缓解栽培大豆盐伤害的离子特征[J]. 浙江农业学报, 2022, 34(10): 2095-2104.

MENG Na, XUE Hui, WEI Ming, WEI Shenghua. Ion characteristics on chloride channel blocker ameliorating salt injury to Glycine max[J]. Acta Agriculturae Zhejiangensis, 2022, 34(10): 2095-2104.

图/表 7

图1 不同盐浓度(A)和氯离子通道抑制剂(B)栽培大豆绥农35植株表型比较从左到右依次是A ( Control, 100、140和200 mmol·L-1NaCl ), B (Control, NaCl, Zn2+, 9-AC和NFA)。

Fig.1 The appearance of Glycine max cultivar Suinong 35 under different concentrations of NaCl treatments (A) and different chloride channel inhibitors (B) From left to right, A (Control, 100, 140 and 200 mmol·L-1NaCl), B (Control, NaCl, Zn2+, 9-AC and NFA).

表1 不同处理下栽培大豆品种绥农35植株的形态参数比较

Table 1 Comparison of morphological parameters in Glycine max cultivar Suinong 35 under different treatments

处理 Treatment	叶片长度 Leaf length/cm	叶片宽度 Leaf width/cm	叶片面积 Leaf area/cm²	叶片周长 Leaf perimeter/cm	根长 Root length/cm	株高 Plant height/cm
对照组Control	7.20±0.04 a	3.64±0.009 a	18.47±0.04 a	17.48±0.65 a	16.0±0.67 a	62.0±0.67 a
盐处理组NaCl	4.52±0.016 c	2.41±0.007 c	8.09±0.14 c	8.93±0.27 c	12.33±0.44 b	38.0±1.33 c
盐+Zn²⁺处理组	5.32±0.018 b	2.91±0.007 b	11.70±0.23 b	12.88±0.43 b	17.0±1.33 a	45.0±0.67 b
NaCl+ Zn²⁺

图2 不同处理下栽培品种绥农35植株根部解剖结构和结构参数特征比较从左到右依次是:Control、NaCl、NaCl+ Zn2+。A,根部解剖结构特征;B,根部表皮细胞。px,初生木质部;sx,次生木质部;sp,次生韧皮部;st,中柱;c,皮层;ec,表皮细胞。

Fig.2 Comparison of anatomical structures and parameters of roots in Glycine max cultivar Suinong 35 under different treatments From left to right: Control, NaCl, NaCl+ Zn2+, respectively. A, Cross-sections of roots; B, Epidermis cell of roots. px, Primary xylem; sx, Second xylem; sp, Second phloem; st, Stele; c, Cortex; ec, Epidermis cell.

表2 不同处理下栽培大豆品种绥农35根部结构参数

Table 2 Root structure parameters of Glycine max cultivar Suinong 35 under different treatments

处理 Treatment	根直径 Root diameter/μm	皮层厚度 Cortex thickness/μm	皮层/根直径 Cortex/Root diameter/%	次生导管直径 Secondary vessel diameter/μm	中柱直径 Stele diameter/μm	中柱/根直径 Stele/Root diameter/%
对照组Control	2 836.70±78.42 a	592.98±42.50 a	20.90±1.41 b	47.99±5.15 a	1 471.86±57.02 a	51.89±3.09 a
盐处理组NaCl	1 636.90±6.32 c	451.01±9.09 b	27.56±0.65 a	38.74±3.36 b	703.12±9.32 c	42.95±0.60 b
盐+Zn²⁺处理组	2 274.27±20.08 b	603.07±33.50 a	26.52±1.40 a	37.36±2.92 b	1 000.96±12.56 b	44.01±0.74 b
NaCl+ Zn²⁺

图3 不同处理下栽培大豆绥农35植株叶部总叶绿素含量(A)和Fv/Fm (B)值数据均3个重复平均值,不同字母表示处理间差异显著(P<0.05)。下同。

Fig.3 Total chlorophyll contents (A) and Fv/Fm value (B) of Glycine max cultivar Suinong 35 under different treatments Statistical data were expressed as x -±s of three replicates. Means in bars with different letters indicated significant differences (P<0.05) among treatments according to Duncan’s multiple-range test.The same as below.

图4 不同处理下测定栽培大豆绥农35植株根、茎和叶部Cl-(A)和 NO 3 -(B)含量

Fig.4 Contents of Cl-(A) and NO 3 -(B) of roots, stems and leaves in Glycine max cultivar Suinong 35 under different treatments

表3 不同处理下栽培大豆绥农35叶部元素含量

Table 3 Element contents in leaves of Glycine max cultivar Suinong 35 under different treatments

处理 Treatments	K/(g· kg^-1)	Na/(g· kg^-1)	Ca/(g· kg^-1)	Mg/(g· kg^-1)	Fe/(mg· kg^-1)	Mn/(mg· kg^-1)	Cu/(mg· kg^-1)	Zn/(mg· kg^-1)	Mo/(mg· kg^-1)
对照组 Control	43.181± 0.691 b	1.115± 0.088 b	10.108± 0.800 a	3.448± 0.273 a	309.248± 4.087 b	189.023± 2.593 a	12.878± 0.666 b	87.122± 3.273 c	2.812± 0.193 a
盐处理组 NaCl	64.190± 2.653 a	21.671± 0.738 a	10.565± 1.126 a	3.308± 0.174 a	241.392± 8.566 c	111.332± 5.287 c	13.753± 0.907 b	102.387± 1.947 b	1.586± 0.280 b
盐+Zn²⁺处理组 NaCl+ Zn²⁺	62.221± 1.280 a	21.565± 1.647 a	9.333± 0.280 a	3.714± 0.186 a	495.529± 9.173 a	148.938± 2.387 b	19.747± 0.745 a	147.867± 7.561 a	0.737± 0.113 c

参考文献 38

[1]	MUNNS R, TESTER M. Mechanisms of salinity tolerance[J]. Annual Review of Plant Biology, 2008, 59(1): 651-681. DOI URL
[2]	武维华. 植物生理学[M]. 2版. 北京: 科学出版社, 2008: 520.
[3]	LUO Q Y, YU B J, LIU Y L. Differential sensitivity to chloride and sodium ions in seedlings of Glycine max and G.soja under NaCl stress[J]. Journal of Plant Physiology, 2005, 162(9): 1003-1012. DOI URL
[4]	TEAKLE N L, TYERMAN S D. Mechanisms of Cl^- transport contributing to salt tolerance[J]. Plant, Cell & Environment, 2010, 33(4): 566-589.
[5]	PARIDA A K, DAS A B. Salt tolerance and salinity effects on plants: a review[J]. Ecotoxicology and Environmental Safety, 2005, 60(3): 324-349. PMID
[6]	朱晓林, 魏小红, 王宝强, 等. c-GMP诱导对盐胁迫下番茄的转录组分析[J]. 浙江农业学报, 2020, 32(10): 1788-1797. DOI
	ZHU X L, WEI X H, WANG B Q, et al. Transcriptome analysis of tomato under salt stress induced by c-GMP[J]. Acta Agriculturae Zhejiangensis, 2020, 32(10): 1788-1797. (in Chinese with English abstract) DOI
[7]	OSAKABE Y, YAMAGUCHI-SHINOZAKI K, SHINOZAKI K, et al. ABA control of plant macroelement membrane transport systems in response to water deficit and high salinity[J]. New Phytologist, 2014, 202(1): 35-49. DOI PMID
[8]	LAHNER B, GONG J M, MAHMOUDIAN M, et al. Genomic scale profiling of nutrient and trace elements in Arabidopsis thaliana[J]. Nature Biotechnology, 2003, 21(10): 1215-1221. DOI URL
[9]	丁广大, 刘佳, 石磊, 等. 植物离子组学: 植物营养研究的新方向[J]. 植物营养与肥料学报, 2010, 16(2): 479-484.
	DING G D, LIU J, SHI L, et al. Plant inomics: a new field in plant nutrition[J]. Plant Nutrition and Fertilizer Science, 2010, 16(2): 479-484. (in Chinese with English abstract)
[10]	NGUYEN C T, AGORIO A, JOSSIER M, et al. Characterization of the chloride channel-like, AtCLCg, involved in chloride tolerance in Arabidopsis thaliana[J]. Plant and Cell Physiology, 2016, 57(4): 764-775. DOI URL
[11]	WONG T H, LI M W, YAO X Q, et al. The GmCLC1 protein from soybean functions as a chloride ion transporter[J]. Journal of Plant Physiology, 2013, 170(1): 101-104. DOI URL
[12]	WHITE P. Chloride in soils and its uptake and movement within the plant: a review[J]. Annals of Botany, 2001, 88(6): 967-988. DOI URL
[13]	屈娅娜, 於丙军. 氯离子通道抑制剂对盐胁迫下野生和栽培大豆幼苗离子含量等生理指标的影响[J]. 南京农业大学学报, 2008, 31(2): 17-21.
	QU Y N, YU B J. Effects of chloride channel blockers on ion contents and other physiological indexes of Glycine soja and Glycine max seedlings under NaCl stress[J]. Journal of Nanjing Agricultural University, 2008, 31(2): 17-21. (in Chinese with English abstract)
[14]	商静, 许嘉阳, 范艺宽, 等. 高氯土壤条件下烤烟对Cl^-通道抑制剂的生理响应[J]. 植物营养与肥料学报, 2017, 23(2): 460-467.
	SHANG J, XU J Y, FAN Y K, et al. Physiological responses of flue-cured tobacco under the high chloride to chloride channel inhibitors[J]. Journal of Plant Nutrition and Fertilizer, 2017, 23(2): 460-467. (in Chinese with English abstract)
[15]	王灵燕. 钠盐和氯盐胁迫对甘薯幼苗生长及光合作用的效应[D]. 济南: 山东师范大学, 2012.
	WANG L Y. Effect of sodium and chloride salt stress on the growth and photosynthesis of sweet potato seedlings[D]. Jinan: Shandong Normal University, 2012. (in Chinese with English abstract)
[16]	付春旭, 姜成喜, 付亚书, 等. 高产、优质大豆新品种绥农35的选育与示范推广[J]. 大豆科技, 2013(3): 31-33.
	FU C X, JIANG C X, FU Y S, et al. Breeding, demonstration and popularization of a new soybean cultivar Suinong 35 with high yield and good quality[J]. Soybean Science & Technology, 2013(3): 31-33. (in Chinese)
[17]	孙启高, 宋书银, 王宇飞, 等. 介绍双子叶植物叶结构分类术语[J]. 植物分类学报, 1997, 35(3): 275-288.
	SUN Q G, SONG S Y, WANG Y F, et al. Introduction to terminology of classification of dicotyledonous leaf architecture[J]. Acta Phytotaxonomica Sinica, 1997, 35(3): 275-288. (in Chinese with English abstract)
[18]	孟娜, 黄嘉宏, 贾瑞, 等. 盐逆境下氯离子通道抑制剂对栽培大豆离子吸收、转运和含量的影响[J]. 西北农业学报, 2020, 29(12): 1814-1821.
	MENG N, HUANG J H, JIA R, et al. Effect of chloride channel blockers on ion absorption, transport and content of Glycine max seedlings under NaCl induced stress[J]. Acta Agriculturae Boreali-Occidentalis Sinica, 2020, 29(12): 1814-1821. (in Chinese with English abstract)
[19]	周强, 李萍, 曹金花, 等. 测定植物体内氯离子含量的滴定法和分光光度法比较[J]. 植物生理学通讯, 2007, 43(6): 1163-1166.
	ZHOU Q, LI P, CAO J H, et al. Comparison on titration and spectrophotometric methods for determination of chloride content in plants[J]. Plant Physiology Communications, 2007, 43(6): 1163-1166. (in Chinese)
[20]	王学奎. 植物生理生化实验原理和技术[M]. 2版. 北京: 高等教育出版社, 2006.
[21]	孟娜, 徐航, 魏明, 等. 叶面喷施烯效唑对盐胁迫下大豆幼苗生理及解剖结构的影响[J]. 西北植物学报, 2017, 37(10): 1988-1995.
	MENG N, XU H, WEI M, et al. Effect of foliar uniconazole spraying under salt stress on physiological and anatomical characteristics in Glycine max[J]. Acta Botanica Boreali-Occidentalia Sinica, 2017, 37(10): 1988-1995. (in Chinese with English abstract)
[22]	MENG N, YU B J, GUO J S. Ameliorative effects of inoculation with Bradyrhizobium japonicum on Glycine max and Glycine soja seedlings under salt stress[J]. Plant Growth Regulation, 2016, 80(2): 137-147. DOI URL
[23]	ARDINI F, SOGGIA F, ABELMOSCHI M L, et al. Effect of heat stress on the ionomic profile of Nicotiana langsdorffii wild-type and mutant genotypes[J]. International Journal of Environmental Analytical Chemistry, 2016, 96(5): 460-473. DOI URL
[24]	CHIMUNGU J G, BROWN K M, LYNCH J P. Reduced root cortical cell file number improves drought tolerance in maize[J]. Plant Physiology, 2014, 166(4): 1943-1955. DOI PMID
[25]	CHIMUNGU J G, BROWN K M, LYNCH J P. Reduced root cortical cell file number improves drought tolerance in maize[J]. Plant Physiology, 2014, 166(4): 1943-1955. DOI PMID
[26]	王继安, 宁海龙, 罗秋香, 等. 大豆品种间叶绿素含量、RUBP活性、希尔反应活力及其与产量间的关系[J]. 东北农业大学学报, 2004, 35(2): 129-134.
	WANG J A, NING H L, LUO Q X, et al. The content of chlorophyll, the activity of RUBP and Hill and their correlations with yield[J]. Journal of Northeast Agricultural University, 2004, 35(2): 129-134. (in Chinese with English abstract)
[27]	MARWOOD C A, SOLOMON K R, GREENBERG B M. Chlorophyll fluorescence as a bioindicator of effects on growth in aquatic macrophytes from mixtures of polycyclic aromatic hydrocarbons[J]. Environmental Toxicology and Chemistry, 2001, 20(4): 890-898. PMID
[28]	BETHKE P C, DREW M C. Stomatal and nonstomatal components to inhibition of photosynthesis in leaves of Capsicum annuum during progressive exposure to NaCl salinity[J]. Plant Physiology, 1992, 99(1): 219-226. DOI URL
[29]	TAVAKKOLI E, FATEHI F, COVENTRY S, et al. Additive effects of Na⁺ and Cl^- ions on barley growth under salinity stress[J]. Journal of Experimental Botany, 2011, 62(6): 2189-2203. DOI URL
[30]	El-FOULY M M, MOBARAK Z M, SALAMA Z A. Micronutrients (Fe, Mn, Zn) foliar spray for increasing salinity tolerance in wheat Triticum aestivum L[J]. African Journal of Plant Science, 2011, 5(5), 314-322.
[31]	IQBAL M N, RASHEED R, ASHRAF M Y, et al. Exogenously applied zinc and copper mitigate salinity effect in maize (Zea mays L.) by improving key physiological and biochemical attributes[J]. Environmental Science and Pollution Research, 2018, 25(24): 23883-23896. DOI URL
[32]	安振锋, 方正. 植物锰营养研究进展[J]. 河北农业科学, 2002, 6(4): 35-41.
	AN Z F, FANG Z. The advance of manganese nutrition in plant[J]. Journal of Hebei Agricultural Sciences, 2002, 6(4): 35-41. (in Chinese with English abstract)
[33]	ZHU J K. Plant salt tolerance[J]. Trends in Plant Science, 2001, 6(2): 66-71. PMID
[34]	陆安桥, 张峰举, 王学琴, 等. 盐胁迫对苗期湖南稷子K⁺、Na⁺含量与分布的影响[J]. 浙江农业学报, 2021, 33(3): 396-403. DOI
	LU A Q, ZHANG F J, WANG X Q, et al. Effects of NaCl and Na₂SO₄stress on content and distribution of K⁺and Na⁺of Echinochloa frumentacea seedlings[J]. Acta Agriculturae Zhejiangensis, 2021, 33(3): 396-403. (in Chinese with English abstract)
[35]	汪洪, 汪立刚, 周卫, 等. 干旱条件下土壤中锌的有效性及与植物水分利用的关系[J]. 植物营养与肥料学报, 2007, 13(6): 1178-1184.
	WANG H, WANG L G, ZHOU W, et al. Soil zinc availability under water stress condition and its relationship with plant water utilization: a review[J]. Plant Nutrition and Fertilizer Science, 2007, 13(6): 1178-1184. (in Chinese with English abstract)
[36]	王小玲, 高柱, 黄益宗, 等. 铜胁迫对3种草本植物生长和重金属积累的影响[J]. 生态毒理学报, 2014, 9(4): 699-706.
	WANG X L, GAO Z, HUANG Y Z, et al. Effects of copper stress on three kinds of herbaceous plants growth and heavy metal accumulation[J]. Asian Journal of Ecotoxicology, 2014, 9(4): 699-706. (in Chinese with English abstract)
[37]	刘鹏. 钼胁迫对植物的影响及钼与其它元素相互作用的研究进展[J]. 农业环境保护, 2002, 21(3): 276-278.
	LIU P. Effects of stress of molybdenum on plants and interaction between molybdenum and other elements[J]. Agro-Environmental Protection, 2002, 21(3): 276-278. (in Chinese with English abstract)
[38]	SCHMUTZ J, CANNON S B, SCHLUETER J, et al. Genome sequence of the palaeopolyploid soybean[J]. Nature, 2010, 463(7278): 178-183. DOI URL