Study on the Relationship between pH and Organic Acid in the Root of Agropyron cristatum Tawukumu under Alkaline Stress

doi:10.6048/j.issn.1001-4330.2021.10.021

Abstract

Abstract:

【Objective】 To study the mechanism of alkali resistance of Agropyron cristatum 'Tawukumu'and the relationship between organic acids secreted by roots and pH value in regulating alkali resistance of the plants. 【Methods】 Two kinds of alkaline salts Na₂CO₃ and NaHCO₃ were mixed at a molar ratio of 9∶1. Alkali stress was simulated by artificial simulation under natural alkaline habitat conditions of 10, 30, 60, 90 and 120 mmol/L. The pH value and organic acid species and content of the seedlings were determined to study their physiological response to stress. 【Results】 Under alkaline stress, the radicle and embryo of Agropyron tawukumensis grew slowly, and the radicle growth rate was only 0.40 times that of the control, which was more significantly inhibited. With the increase of the concentration of treatment solution, the roots of seedlings secreted organic acids mainly malic acid and citric acid, which interacted with each other to resist the damage of high pH to plants kept the pH in the roots between 6.65 and 6.68. Although the organic acids secreted by the roots of Agropyricacrassifera increased significantly in the process of alkali stress, they could not prevent the damage of high pH to the plant seedlings. 【Conclusion】 When the concentration of treatment solution is 60 mmol/L, that is, the pH value is 9.8, the organic acid secretion was the most appropriate concentration to regulate the pH value of Agropyricacrassifera root.

Key words: Agropyron cristatum Tawukumu; alkaline stress; pH; organic acids

摘要：

【目的】研究塔乌库姆冰草耐碱性机理,分析根系分泌的有机酸与pH值在调控植物抗碱性之间的关系。【方法】将2种碱性盐Na₂CO₃、NaHCO₃按摩尔比9∶1混合,人工模拟10、30、60、90、120 mmol/L天然碱性生境条件对塔乌库姆冰草幼苗进行碱胁迫,测定pH值与有机酸种类及含量,分析其对胁迫的生理响应。【结果】碱胁迫导致塔乌库姆冰草胚根、胚芽生长缓慢,其中胚根生长率仅为对照的0.40倍,受到明显抑制;随处理液浓度的增加,幼苗根系分泌以苹果酸、柠檬酸为主的有机酸,使植物根系内pH保持在6.65~6.68。冰草根系分泌的有机酸在碱胁迫过程中虽然明显增加,但不能阻止高浓度的pH对植物幼苗的伤害。【结论】碱处理液浓度为60 mmol/L,即pH为9.8时,有机酸分泌量为调控冰草根系pH最适宜浓度。

关键词: 塔乌库姆冰草, 碱胁迫, pH值, 有机酸

CLC Number:

S543

Mei KE, Yurong HOU, Yigong ZHANG. Study on the Relationship between pH and Organic Acid in the Root of Agropyron cristatum Tawukumu under Alkaline Stress[J]. Xinjiang Agricultural Sciences, 2021, 58(10): 1929-1937.

柯梅, 侯钰荣, 张一弓. 碱胁迫下冰草根系pH值与有机酸含量变化[J]. 新疆农业科学, 2021, 58(10): 1929-1937.

Figures/Tables 6

References 46

[1]	Rozema J, Flowers T. Ecology. Crops for a salinized world[J]. Science, 2008, 322(5907):1478-1480. DOI PMID
[2]	杨真, 王宝山. 中国盐渍土资源现状及改良利用对策[J]. 山东农业科学, 2015, 47(4):125-130.
	YANG Zhen, WANG Baoshan. Present Status of Saline Soil Resources and Countermeasures for Improvement and Utilization in China[J]. Shandong Agricultural Sciences, 2015, 47(4):125-130.
[3]	杨劲松. 中国盐渍土研究的发展历程与展望[J]. 土壤学报, 2008,(5):837-845.
	YANG Jingsong. Development and Prospect of the research on salt affected soils in China[J]. Acta Pedologica Sinica, 2008,(5):837-845.
[4]	王佳丽, 黄贤金, 钟太洋, 等. 盐碱地可持续利用研究综述[J]. 地理学报, 2011, 66(5):673-684.
	WANG Jiali, HUANG Xianjin, ZHONG Taiyang, et al. Review on Sustainable Utilization of Salt-affected Land[J]. Acta Pedologica Sinica, 2011, 66(5):673-684.
[5]	曾华. 植物耐盐碱机制研究进展[J]. 北方水稻, 2017, 47(2):58-61.
	ZENG Hua. Research progress in plant salt-alkali resistance mechanism[J]. North Rice, 2017, 47(2):58-61.
[6]	赵海新, 黄晓群, 陈书强, 等. 碱胁迫下不同水稻品种微观结构响应解析[J]. 东北农业大学学报, 2020, 51(11):11-22.
	ZHAO Haixin, HUANG Xiaoqun, CHEN Shuqiang, et al. Response analysis of microstructure of different rice varieties under alkali stress[J]. Journal of Northeast Agricultural University, 2020, 51(11):11-22.
[7]	徐晓雪. 高粱幼苗对碱胁迫的生理响应及代谢组学研究[D]. 沈阳:沈阳农业大学, 2020.
	XU Xiaoxue. Physiological responses and metabonomics of sorghum seedlings to alkali stress[D]. Shenyang: Shenyang Agricultural University, 2020.
[8]	易善军, 孙振元, 韩蕾, 等. 植物耐碱机理及相关基因研究进展[J]. 世界林业研究, 2011, 24(1):28-32.
	YI Shanjun, SUN Zhenyuan, HAN Lei, et al. Research Advance on Alkali-resistant Mechanism of Plants and related Genes[J]. World Forestry Research, 2011, 24(1):28-32.
[9]	杨国会. 碱胁迫诱导小冰麦有机酸积累和分泌的研究[J]. 西北农林科技大学学报(自然科学版), 2010, 38(7):77-84.
	YANG Guohui. Study on organic acids accumulation and secretion of alkali stress induced in wheat-wheatgrass[J]. Journal of Northwest A&F University(Nat. Sci. Ed), 2010, 38(7):77-84.
[10]	崔亚茹. 苦荞根系分泌有机酸对土壤低氮胁迫的响应[D]. 临汾: 山西师范大学, 2019.
	CUI Yaru. Response of Organic Acids Secreted by Roots of Tartary Buckwheat under Low Nitrogen Stress from Pot Experiment[D]. Linfen: Shanxi Normal University, 2019.
[11]	崔翠, 蔡靖, 张硕新. 核桃根系分泌物化感物质的分离与鉴定[J]. 林业科学, 2013, 49(2):54-60.
	CUI Cui, CAI Jing, ZHANG Shuoxin. Isolation and Identification of the Allelochemicals in Walnut (Juglansregia) Root Exudates[J]. Scientia Silvae Sinicae, 2013, 49(2):54-60.
[12]	贺永华, 沈东升, 朱荫湄. 根系分泌物及其根际效应[J]. 科技通报, 2006,(6):761-766.
	HE Yonghua, SHEN Dongsheng, ZHU Yinmei. Root Exudates and Their Rhizospheric Effects[J]. Bulletin of Science and Technology, 2006, (6):761-766.
[13]	张锡洲, 李廷轩, 王永东. 植物生长环境与根系分泌物的关系[J]. 土壤通报, 2007,(4):785-789.
	ZHANG Xizhou, LI Tingxuan, WANG Yongdong. Relationship between Growth Environment and Root Exudates of Plants: A Review[J]. Chinese Journal of soil Science, 2007,(4):785-789.
[14]	Wanchao Chen, and Pengjuan Cui, and Huaying Sun, and Weiqing Guo, and Chunwu Yang, and Hao Jin, and Bin Fang, and Decheng Shi. Comparative effects of salt and alkali stresses on organic acid accumulation and ionic balance of seabuckthorn (Hippophae rhamnoides L.)[J]. Industrial Crops and Products, 2009.
[15]	杨春武, 李长有, 尹红娟, 等. 小冰麦(Triticumaestivum-Agropyron intermedium)对盐胁迫和碱胁迫的生理响应[J]. 作物学报, 2007, 33(8):1255-1261.
	YANG Chunwu, LI Changyou, YIN Hongjuan, et al. Physiological Response of Xiaobingmai(Triticumaestivum-Agropyron intermedium)to Salt-Stress and Alkali Stress[J]. Acta Agronomica Sinica, 2007, 33(8):1255-1261.
[16]	贾渊. 荒漠草原植物根分泌物及其有机酸组分对土壤中微生物及养分的影响[D]. 呼和浩特:内蒙古师范大学, 2019.
	JIA Yuan. Effects of plant root exudates and their organic acids on soil microbes and nutrients in desert steppe[D]. Hohhot: Inner Mongolia Normal University, 2019.
[17]	吕家强, 李长有, 杨春武, 等. 天然盐碱土壤对虎尾草茎叶有机酸积累影响及胁迫因子分析[J]. 草业学报, 2015, 24(4):95-103.
	LÜ Jiaqiang, LI Changyou, YANG Chunwu, et al. Effect of natural saline soil on organic acid accumulation in the stem and leaf of Chloris virgata and analysis of stress factors[J]. Acta Prataculturae Sinica, 2015, 24(4):95-103.
[18]	贾纳提, 李学森, 米克什, 等. 塔乌库姆冰草特征分析及适应性的研究[J]. 草食家畜, 2012,(1):91-92.
	JIA Nati, LI Xuesen, MI Keshi, et al. Study of Feature Analysis and Adaptability on AgropyronCristatum (Linn.) Gaertn. cv.Tawukumu[J]. Grass-Feeding Livestock, 2012, (1):91-92.
[19]	新疆维吾尔自治区地方标准[J]. 草食家畜, 2015,(3):72-76.
	Local standards of Xinjiang Uygur Autonomous Region[J]. Grass-Feeding Livestock, 2015,(3):72-76.
[20]	张一弓, 贾纳提, 李学森, 等. 碱胁迫下塔乌库姆冰草的萌发及生理响应[J]. 草地学报, 2014, 22(4):783-788.
	ZHANG Yigong, JIA Nati, LI Xueseng, et al. Seed Germination and Physiological Responses of Agropyroncristatum?? Tawukumu? under Alkali Stress[J]. Acta Agrestia Sinica, 2014, 22(4):783-788.
[21]	李秀峰, 张欣欣, 柳参奎, 等. 水稻苹果酸酶基因在逆境下的表达特性研究[J]. 基因组学与应学, 2012, 31(4):327-332.
	LI Xiufeng, ZHANG Xinxin, LIU Chankui, et al. Expression Characteristics of Rice (Oryza sativa L.) Malic Enzyme (OsNA-DP-ME3) Gene under Environmental Stress[J]. Genomics and Applied Biology, 2012, 31(4):327-332.
[22]	李德华, 向春雷, 姜益泉, 等. 低磷胁迫下水稻不同品种根系有机酸分泌的差异[J]. 中国农学通报, 2005,(11):186-188, 201.
	LI Dehua, XIANG Chunlei, JIANG Yiquan, et al. Difference of Organic Acid Secretion from Roots of Various Rice Varieties under the Stress of Low Phosphorus[J]. Chinese Agricultural Science Bulletin, 2005, (11):186-188,201.
[23]	杨春武. 虎尾草和水稻抗碱机制研究[D]. 吉林:东北师范大学, 2010.
	YANG Chunwu. Mechanisms of alkali tolerance in Chloris virgata and rice(Oryza Sativa)[D]. Jilin: Northeast Normal University, 2010.
[24]	韩萌. 混合盐碱胁迫对7种禾种子萌发及生理特性的影响[D]. 哈尔滨:哈尔滨师范大学, 2013.
	HAN Meng. Effects of saline-alkaloid stress on the seed germination and physiologial characteristics of seven grass seeds[D]. Harbin: Harbin Normal University, 2003.
[25]	Hussain M K, Rehman O U. Breeding sunflower for salt tolerance: Association of shoot growth and mature plant traits for salt tolerance in cultivated sunflower (Helianthus annuus L.)[J]. Helia, 1995, 18:69-76.
[26]	庞丙亮, 曹帮华, 张玉娟, 等. 不同盐碱胁迫对紫菀生理生化的影响[J]. 林业科技发, 2011, 25(3):44-46.
	PANG Bingliang, CAO Banghua, ZHANG Yujuan, et al. Effect saline-alkali stresses on physiological and biochemical characteristics of Aster tongolensis[J]. Journal of Forestry Engineering, 2011, 25(3):44-46.
[27]	Lynch J P,. Deikman J. Phosphorus in Plant Biology:Regulatory Roles in Molecular,Cellular,Organismic and Ecosystem Processes[J]. American Society of Plant Physiologists Rockville,Maryland USA. 1998: 157.
[28]	Cao L M, Pan X H. A preliminary study on the tolerant mechanism office[J]. Acta Agron Sin, 2002, 28(2):260-264.
[29]	Noordwijk M, Willigen P D, Ehlert P, et al. A simple model of P uptake by crops as a possible basis for P fertilizer recommendations[J]. Netherlands Journal of Agricultural Science, 1990, 38(3A):317-332. DOI URL
[30]	洪常青, 聂艳丽. 根系分泌物及其在植物营养中的作用[J]. 生态环境, 2003,(4):508-511.
	HONG Changqing, NIE Yanli. Effects of root exudates on plant nutrition[J]. Ecology and Environment, 2003, (4):508-511.
[31]	沈宏, 严小龙. 根分泌物研究现状及其在农业与环境领域的应用[J]. 农村生态环境, 2000,(3):51-54.
	SHEN Hong, YAN Xiaolong. Status of the Study on Root Exudates andIts Application to Agriculture and Environment[J]. Rural Eco-Environment, 2000,(3):51-54.
[32]	López-Bucio J, Nieto-Jacobo M F, Ramírez-Rodríguez V, et al. Organic acid metabolism in plants: from adaptive physiology to transgenic varieties for cultivation in extreme soils[J]. Plant Sci, 2000, 160:1-13. PMID
[33]	SH I D C, YIN S J, YANG G H, et al. Citric acid accumulation in an alkali tolerant plant Puccinelliatenu flora under alkaline stress[J]. Act a Bot Sin, 2002, 44(5) : 537-540.
[34]	SHI D C, SH ENG Y M. Effect of various salt- alkaline mixed stress conditions on sunflower seedling and analysis of their stress factors[J]. Environ J Exp Bot, 2005, 54:8-21. DOI URL
[35]	吴彩霞, 傅华. 根系分泌物的作用及影响因素[J]. 草业科学, 2009, 26(9):24-29.
	WU Caixia, FU Hua. Effects and roles of root exudates[J]. Pratacultural Science, 2009, 26(9):24-29.
[36]	黄文斌, 马瑞, 杨迪, 等. 土壤逆境下植物根系分泌的有机酸及其对植物生态适应性的影响(英文)[J]. 安徽农业科学与技术, 2014, 15(7):1167-1173.
	HUANG Wenbin, MA Rui, YANG Di, et al. Organic Acids Secreted from Plant Roots under Soil Stress and Their Effects on Plant Ecological Adaptability[J]. Agricultural Science & Technology, 2014, 15(7):1167-1173.
[37]	王惠君, 薛卫杰, 张昕, 等. 叶面喷施苹果酸对水稻Cd积累特性的影响[J]. 农业环境科学学报,2020, 2021, 40(2):269-278.
	WANG Huijun, XUE Weijie, ZHANG Xin, et al. Effects of foliar application with malic acid on the accumulation of cadmium in rice[J]. Journal of Agro-Environment Science, 2021, 40(2):269-278.
[38]	Tao P, Guo W P, Li B Y, et al. Genome-wide identification, classification, and analysis of NADP-ME family members from 12 crucifer species[J]. Molecular Genetics and Genomics, 2016, 291(3):1167-1180. DOI URL
[39]	Sun X, Han G L, Meng Z, et al. Roles of malic enzymes in plant development and stress responses[J]. Plant Signaling and Behavior, 2019, 14(10):1-8.
[40]	Bela K, Horváth E, Gallé ?, et al. Plant glutathione peroxidases: Emerging role of the antioxidant enzymes in plant development and stress responses[J]. Journal of Plant Physiology, 2015, 176:192-201. DOI URL
[41]	董秋丽, 夏方山, 董宽虎. NaCl胁迫对芨芨草苗期脯氨酸代谢的影响[J]. 草业学报, 2010, 19(5):71-76.
	DONG Qiuli, XIA Fangshan, DONG Kuanhu Effects of NaCl stress on proline metabolism of Achnatherumsplendens seedling[J]. Acta Prataculturae Sinica, 2010, 19(5):71-76.
[42]	杨春武, 李长有, 张美丽, 等. 盐、碱胁迫下小冰麦体内的pH及离子平衡[J]. 应用生态学报, 2008,(5):1000-1005. PMID
	YANG Chunwu, LI Changyou, ZHANG Meili, et al. pH and ion balance in wheat-wheatgrass under salt- or alkali stress[J]. Chinese Journal of Applied Ecology, 2008,(5):1000-1005. PMID
[43]	Alhendawi R A, Mheld V, Kirkby E A, et al. Influence of increasing bicarbonate concentrations on plant growth, organic acid accumulation in roots and iron uptake by barley, sorghum, and maize[J]. Journal of Plant Nutrition, 1997, 20(12):1731-1753. DOI URL
[44]	Timpa J D, Burke J J, Quisenberry J E, et al. Effects of Water Stress on the Organic Acid and Carbohydrate Compositions of Cotton Plants[J]. Plant Physiology, 1986, 82(3):724-728. PMID
[45]	石德成, 殷立娟. 盐(NaCl)与碱(Na₂CO₃)对星星草胁迫作用的差异[J]. Journal of Integrative Plant Biology, 1993,(2):144-149.
	SHI Decheng, YIN Lijuan. Difference between salt(NaCl)and Alkaline(Na₂CO₃)stresses on PuccineuiaTenuiflora(Griseb.)scribn.ET Merr. plants[J]. Journal of Integrative Plant Biology, 1993, (2):144-149.
[46]	Francoise F, Daniel L R, John G. Effects of salt stress on amino acid,organic acid and carbohydrate composition of root,bactericide and cymosely of alfalfa[J]. Plant Physiol, 1991, 96:1228-1236. DOI URL

分类 Classification	浓度 Concentration (mmol/L)	电导率 Electric conductivity (μS/cm)	水势 Water potential (MPa)	pH值	相对含水量 Relative water content (%)
对照(CK)	0	2.17	-0.05	6.10	87.82
碱处理 Alkali treatment	10	3.23	-0.14	9.17	86.02
	30	4.85	-0.23	9.46	85.65
	60	5.38	-0.30	9.76	84.75
	90	6.25	-0.35	9.92	83.80
	120	7.14	-0.39	10.00	82.24
	150	8.01	-0.51	10.08	80.82

分类 Classification	浓度 Concentration (mmol/L)	电导率 Electric conductivity (μS/cm)	水势 Water potential (MPa)	pH值	相对含水量 Relative water content (%)
对照(CK)	0	2.17	-0.05	6.10	87.82
碱处理 Alkali treatment	10	3.23	-0.14	9.17	86.02
	30	4.85	-0.23	9.46	85.65
	60	5.38	-0.30	9.76	84.75
	90	6.25	-0.35	9.92	83.80
	120	7.14	-0.39	10.00	82.24
	150	8.01	-0.51	10.08	80.82

名称 Tag	浓度 Concentration(mmol/L)	处理前 Treatment front 根系外pH值 Root environment pH	处理后 After treatment
名称 Tag	浓度 Concentration(mmol/L)	处理前 Treatment front 根系外pH值 Root environment pH	根系外pH值 Root environment pH	根系组织pH值 Root tissue pH
对照(CK)	0	6.097±0.032^f	6.497±0.020^d	6.610±0.017^b
碱处理 Alkali treatment	10	9.167±0.038^e	8.487±0.097^c	6.647±0.009^ab
	30	9.463±0.084^d	8.580±0.042^c	6.667±0.020^ab
	60	9.757±0.038^c	8.960±0.036^b	6.662±0.029^ab
	90	9.917±0.027^b	9.007±0.033^b	6.680±0.023^a
	120	10.000±0.015^ab	9.093±0.018^ab	6.653±0.009^ab
	150	10.077±0.023^a	9.160±0.025^a	6.669±0.010^ab

名称 Tag	浓度 Concentration(mmol/L)	处理前 Treatment front 根系外pH值 Root environment pH	处理后 After treatment
名称 Tag	浓度 Concentration(mmol/L)	处理前 Treatment front 根系外pH值 Root environment pH	根系外pH值 Root environment pH	根系组织pH值 Root tissue pH
对照(CK)	0	6.097±0.032^f	6.497±0.020^d	6.610±0.017^b
碱处理 Alkali treatment	10	9.167±0.038^e	8.487±0.097^c	6.647±0.009^ab
	30	9.463±0.084^d	8.580±0.042^c	6.667±0.020^ab
	60	9.757±0.038^c	8.960±0.036^b	6.662±0.029^ab
	90	9.917±0.027^b	9.007±0.033^b	6.680±0.023^a
	120	10.000±0.015^ab	9.093±0.018^ab	6.653±0.009^ab
	150	10.077±0.023^a	9.160±0.025^a	6.669±0.010^ab