植物耐盐机制研究进展及展望

doi:10.7668/hbnxb.20195207

华北农学报 ›› 2024, Vol. 39 ›› Issue (5): 80-92. doi: 10.7668/hbnxb.20195207

所属专题： 盐碱胁迫；热点论文；栽培生理；

• 耕作栽培·生理生化 • 上一篇下一篇

植物耐盐机制研究进展及展望

王大江¹^,⁴, 刘昭¹^,², 路翔¹^,², 高源¹^,⁴, 孙思邈¹, 郭含欣¹, 田雯¹^,², 王霖¹, 李子琛¹, 李连文¹, 王昆¹^,⁴, 刘继红²^,³

¹ 中国农业科学院果树研究所,农业农村部园艺作物种质资源利用重点实验室,辽宁兴城 125100
² 石河子大学农学院,特色果树栽培生理与种质资源利用兵团重点实验室,新疆石河子 832000
³ 华中农业大学园林学院,果蔬园艺作物种质创新与利用全国重点实验室,湖北武汉 430070
⁴ 国家盐碱地综合利用技术创新中心,山东东营 257000

收稿日期:2024-05-26 出版日期:2024-10-28
通讯作者:
王昆(1971-),男,辽宁兴城人,研究员,硕士,主要从事果树种质资源研究。
刘继红(1971-),男,湖北武汉人,教授,博士,主要从事果树逆境分子生理研究。
作者简介:
王大江(1985-),男,河南南阳人,副研究员,博士,主要从事果树种质资源研究。
基金资助:
中国农业科学院创新工程项目(CAAS-ASTIP-2021-RIP-02)

Advances and Prospect on Mechanism of Salt Tolerance in Plants

WANG Dajiang¹^,⁴, LIU Zhao¹^,², LU Xiang¹^,², GAO Yuan¹^,⁴, SUN Simiao¹, GUO Hanxin¹, TIAN Wen¹^,², WANG Lin¹, LI Zichen¹, LI Lianwen¹, WANG Kun¹^,⁴, LIU Jihong²^,³

¹ Research Institute of Pomology,Chinese Academy of Agricultural Sciences,Key Laboratory of Horticultural Crops Germplasm Resources Utilization,Ministry of Agriculture and Rural Affairs of the People's Republic of China,Xingcheng 125100,China
² Agricultural College,Shihezi University,Xinjiang Production and Construction Corps Key Laboratory of Special Fruits and Vegetables Cultivation Physiology and Germplasm Resources Utilization,Shihezi 832000,China
³ College of Horticulture and Forestry Sciences,Huazhong Agricultural University,National Key Laboratory for Germplasm Innovation & Utilization of Horticultural Crops,Wuhan 430070,China
⁴ National Center of Technology Innovation for Comprehensive Utilization of Saline-Alkali Land,Dongying 257000,China

Received:2024-05-26 Published:2024-10-28

摘要/Abstract

摘要：

植物的生长和生产面临各种生物和非生物胁迫,其中盐胁迫严重影响植物的正常生长发育、品质和产量形成。植物在长期的演化过程中,进化出了适应盐胁迫的形态结构、生理生化反应和遗传基础。在形态结构上,耐盐植物的叶片具有蜡质层、气孔密度低于盐敏感植物,另外具有泌盐或者阻断功能的盐腺、毛状体、盐囊泡、凯氏带等结构;在生理活动调节方面,一方面耐盐植物具有高的酶促和非酶促抗氧化物质,如SOD、CAT、酚类物质等,另一方面耐盐植物具有较高的渗透调节物质含量,或者在盐胁迫下可以合成渗透调节物质,包括有机物质可溶性蛋白和糖以及无机盐离子等;在分子机理方面,SOS途径是研究得最为清晰的离子调控途径,通过SOS1、SOS2和SOS3协同作用维持细胞内Na⁺/K⁺平衡;此外,植物激素及碳代谢途径在植物耐盐过程中亦具有重要作用。本研究通过综述植物耐盐研究进展,探讨耐盐植物在形态结构、生理基础、遗传分子基础和转基因手段在响应盐胁迫的潜在研究重点和方向,有助于研究人员快速找到切入点,逐步完善植物的耐盐机理体系,加快耐盐植物的高效利用。

关键词: 植物, 耐盐, 抗氧化, 渗透调控, 离子毒害

Abstract:

Plant growth and production are faced with various biological and abiotic stresses,among which salt stress seriously affects the normal growth and development,quality and yield formation of plants.Plants have evolved morphological structure,physiological and biochemical reactions and genetic basis to adapt to salt stress during the long process of evolution.In terms of morphological structure,the leaves of salt-tolerant plants have waxy layer and lower stomatal density than those of salt-sensitive plants,and salt glands,microhairs,salt vesicles,and casparian strip have salt secretion or blocking functions.In terms of physiological activity regulation,on the one hand,salt-tolerant plants have high enzymatic and non-enzymatic antioxidant substances,such as SOD,CAT,phenolic substances,on the other hand,salt-tolerant plants have a high content of osmoregulatory substances,or can synthesize osmoregulatory substances under salt stress,including soluble proteins and sugars of organic substances and inorganic ions.In terms of molecular mechanism,SOS pathway is the most clearly studied ion regulation pathway,which maintains intracellular Na⁺/K⁺ balance through the synergistic action of SOS1,SOS2 and SOS3.In addition,plant hormones and carbon metabolism pathways also play an important role in the process of plant salt tolerance.This paper summarizes the research progress of salt-tolerant plants,and discusses the potential research focus and direction of salt-tolerant plants in terms of morphological structure,physiological basis,genetic molecular basis and transgenic methods in response to salt stress,which will help researchers quickly find the breakthrough point,gradually improve the mechanism system of salt-tolerant plants,and accelerate the efficient utilization of salt-tolerant plants.

Key words: Plant, Salt tolerance, Antioxidant, Osmotic regulation, Ion toxicity

王大江, 刘昭, 路翔, 高源, 孙思邈, 郭含欣, 田雯, 王霖, 李子琛, 李连文, 王昆, 刘继红. 植物耐盐机制研究进展及展望[J]. 华北农学报, 2024, 39(5): 80-92. doi: 10.7668/hbnxb.20195207.

WANG Dajiang, LIU Zhao, LU Xiang, GAO Yuan, SUN Simiao, GUO Hanxin, TIAN Wen, WANG Lin, LI Zichen, LI Lianwen, WANG Kun, LIU Jihong. Advances and Prospect on Mechanism of Salt Tolerance in Plants[J]. Acta Agriculturae Boreali-Sinica, 2024, 39(5): 80-92. doi: 10.7668/hbnxb.20195207.

http://www.hbnxb.net/CN/Y2024/V39/I5/80

参考文献 173

[1]	Munns R, Tester M. Mechanisms of salinity tolerance[J]. Annual Review of Plant Biology, 2008, 59:651-681.doi:10.1146/annurev.arplant.59.032607.092911. pmid: 18444910
[2]	Wang D J, Gao Y, Sun S M, Lu X, Li Q S, Li L W, Wang K, Liu J H. Effects of salt stress on the antioxidant activity and malondialdehyde, solution protein, proline, and chlorophyll contents of three Malus species[J]. Life, 2022, 12(11):1929.doi:10.3390/life12111929.
[3]	Jamil A, Riaz S, Ashraf M, Foolad M R. Gene expression profiling of plants under salt stress[J]. Critical Reviews in Plant Sciences, 2011, 30(5):435-458.doi:10.1080/07352689.2011.605739.
[4]	Alasvandyari F, Mahdavi B, Madah H. Glycine betaine affects the antioxidant system and ion accumulation and reduces salinity-induced damage in safflower seedlings[J]. Archives of Biological Sciences, 2017, 69(1):139-147.doi:10.2298/abs160216089a.
[5]	Chen X, Cheng X W, Zhu H, Bañuelos G, Shutes B, Wu H T. Influence of salt stress on propagation,growth and nutrient uptake of typical aquatic plant species[J]. Nordic Journal of Botany, 2019, 37(12):e02411.doi:10.1111/njb.02411.
[6]	Soltabayeva A, Ongaltay A, Omondi J O, Srivastava S. Morphological,physiological and molecular markers for salt-stressed plants[J]. Plants, 2021, 10(2):243.doi:10.3390/plants10020243.
[7]	于沛玉. 盐胁迫下K⁺对珠美海棠幼苗生理特性的影响[D]. 天津: 天津农学院, 2014.
	Yu P Y. Effects potassium ion on physiological property of Malus zumi seedlings under salt stress[D]. Tianjin:Tianjin Agricultural University, 2014.
[8]	Mohammad M, Shibli R, Ajlouni M, Nimri L. Tomato root and shoot responses to salt stress under different levels of phosphorus nutrition[J]. Journal of Plant Nutrition, 1998, 21(8):1667-1680.doi:10.1080/01904169809365512.
[9]	Meloni D A, Oliva M A, Ruiz H A, Martinez C A. Contribution of proline and inorganic solutes to osmotic adjustment in cotton under salt stress[J]. Journal of Plant Nutrition, 2001, 24(3):599-612.doi:10.1081/PLN-100104983.
[10]	戴伟民, 蔡润, 潘俊松, 何欢乐. 盐胁迫对番茄幼苗生长发育的影响[J]. 上海农业学报, 2002, 18(1):58-62.doi:10.3969/j.issn.1000-3924.2002.01.014.
	Dai W M, Cai R, Pan J S, He H L. Effects of salt stress on growth development of tomato[J]. Acta Agriculturae Shanghai, 2002, 18(1):58-62.
[11]	高玉坤, 杨溥原, 项晓冬, 魏世林, 任根增, 殷丛培, 梁红凯, 崔江慧, 常金华. 不同耐盐高粱品种全生育期对盐胁迫的响应[J]. 华北农学报, 2020, 35(6):113-121.doi:10.7668/hbnxb.20191411.
	Gao Y K, Yang P Y, Xiang X D, Wei S L, Ren G Z, Yin C P, Liang H K, Cui J H, Chang J H. Response of different salt tolerant sorghum varieties to salt stress in the whole growth period[J]. Acta Agriculturae Boreali-Sinica, 2020, 35(6):113-121. doi: 10.7668/hbnxb.20191411
[12]	El-Hendawy S E, Hu Y C, Schmidhalter U. Growth,ion content,gas exchange,and water relations of wheat genotypes differing in salt tolerances[J]. Australian Journal of Agricultural Research, 2005, 56(2):123.doi:10.1071/ar04019.
[13]	Feng Y N, Cui J Q, Zhou T, Liu Y, Yue C P, Huang J Y, Hua Y P. Comprehensive dissection into morpho-physiologic responses,ionomic homeostasis,and transcriptomic profiling reveals the systematic resistance of allotetraploid rapeseed to salinity[J]. BMC Plant Biology, 2020, 20(1):534.doi:10.1186/s12870-020-02734-4.
[14]	Tao R R, Ding J F, Li C Y, Zhu X K, Guo W S, Zhu M. Evaluating and screening of agro-physiological indices for salinity stress tolerance in wheat at the seedling stage[J]. Frontiers in Plant Science, 2021, 12:646175.doi:10.3389/fpls.2021.646175.
[15]	王庆彪, 王艳萍, 令狐波, 钱慧慧, 赵秋菊, 张丽. 盐胁迫对萝卜幼苗生长及RsCAT和RsSOD基因表达的影响[J]. 华北农学报, 2021, 36(6):1-6.doi:10.7668/hbnxb.20192155.
	Wang Q B, Wang Y P, Linghu B, Qian H H, Zhao Q J, Zhang L. Effect of salt stress on seedling growth and transcription of RsCAT and RsSOD genes in radish[J]. Acta Agriculturae Boreali-Sinica, 2021, 36(6):1-6.
[16]	Tester M, Davenport R. Na⁺ tolerance and Na⁺ transport in higher plants[J]. Annals of Botany, 2003, 91(5):503-527.doi:10.1093/aob/mcg058. pmid: 12646496
[17]	Munns R. Comparative physiology of salt and water stress[J]. Plant,Cell & Environment, 2002, 25(2):239-250.doi:10.1046/j.0016-8025.2001.00808.x.
[18]	Hedrich R, Shabala S. Stomata in a saline world[J]. Current Opinion in Plant Biology, 2018, 46:87-95.doi:10.1016/j.pbi.2018.07.015. pmid: 30138845
[19]	Badawi G H, Yamauchi Y, Shimada E, Sasaki R, Kawano N, Tanaka K, Tanaka K. Enhanced tolerance to salt stress and water deficit by overexpressing superoxide dismutase in tobacco (Nicotiana tabacum) chloroplasts[J]. Plant Science, 2004, 166(4):919-928.doi:10.1016/j.plantsci.2003.12.007.
[20]	Allakhverdiev S I, Murata N. Salt stress inhibits photosystems Ⅱ and Ⅰ in cyanobacteria[J]. Photosynthesis Research, 2008, 98(1):529-539.doi:10.1007/s11120-008-9334-x.
[21]	Munns R, Day D A, Fricke W, Watt M, Arsova B, Barkla B J, Bose J, Byrt C S, Chen Z H, Foster K J, Gilliham M, Henderson S W, Jenkins C L D, Kronzucker H J, Miklavcic S J, Plett D, Roy S J, Shabala S, Shelden M C, Soole K L, Taylor N L, Tester M, Wege S, Wegner L H, Tyerman S D. Energy costs of salt tolerance in crop plants[J]. New Phytologist, 2020, 225(3):1072-1090.doi:10.1111/nph.15864. pmid: 31004496
[22]	Turgut Yiğit A, Yilmaz O, UzĪlday B, Özgür Uzİday R, Türkan İ. Plant response to salinity:an analysis of ROS formation,signaling,and antioxidant defense[J]. Turkish Journal of Botany, 2020, 44(1):1-13.doi:10.3906/bot-1911-15.
[23]	Waszczak C, Carmody M, Kangasjärvi J. Reactive oxygen species in plant signaling[J]. Annual Review of Plant Biology, 2018, 69:209-236.doi:10.1146/annurev-arplant-042817-040322. pmid: 29489394
[24]	Smirnoff N, Arnaud D. Hydrogen peroxide metabolism and functions in plants[J]. The New Phytologist, 2019, 221(3):1197-1214.doi:10.1111/nph.15488.
[25]	Sies H, Jones D P. Reactive oxygen species (ROS) as pleiotropic physiological signaling agents[J]. Nature Reviews Molecular Cell Biology, 2020, 21(7):363-383.doi:10.1038/s41580-020-0230-3.
[26]	Pang C H, Wang B S. Oxidative stress and salt tolerance in plants[J]. Progress in Botany, 2008, 69:231-245.doi:10.1007/978-3-540-72954-9_9.
[27]	Tewari R K, Kumar P, Sharma P N. Antioxidant responses to enhanced generation of superoxide anion radical and hydrogen peroxide in the copper-stressed mulberry plants[J]. Planta, 2006, 223(6):1145-1153.doi:10.1007/s00425-005-0160-5. pmid: 16292566
[28]	王瀚祥, 李广存, 徐建飞, 王万兴, 金黎平. 植物耐盐机理研究进展[J]. 作物杂志, 2022(5):1-12.doi:10.16035/j.issn.1001-7283.2022.05.001.
	Wang H X, Li G C, Xu J F, Wang W X, Jin L P. Advances in research on salt tolerance mechanism of plants[J]. Crops, 2022(5):1-12.
[29]	赵龙. 盐生植物碱地肤耐盐生理及分子机制研究[D]. 长春: 东北师范大学, 2018.
	Zhao L. Pysiological and molecular mechanisms underlying salt tolerance in halophyte Kochia sieversiana[D]. Changchun: Northeast Normal University, 2018.
[30]	van Zelm E, Zhang Y X, Testerink C. Salt tolerance mechanisms of plants[J]. Annual Review of Plant Biology, 2020, 71:403-433.doi:10.1146/annurev-arplant-050718-100005. pmid: 32167791
[31]	薛琼琼, 赵露露, 王云霞, 赵猛. 盐生植物耐盐性研究进展[J]. 中国野生植物资源, 2021, 40(5):60-65.doi:10.3969/j.issn.1006-9690.2021.05.011.
	Xue Q Q, Zhao L L, Wang Y X, Zhao M. Research progress on salt tolerance of halophytes[J]. Chinese Wild Plant Resources, 2021, 40(5):60-65.
[32]	Zhu J K. Regulation of ion homeostasis under salt stress[J]. Current Opinion in Plant Biology, 2003, 6(5):441-445.doi:10.1016/s1369-5266(03)00085-2. pmid: 12972044
[33]	刘晓, 刘晓红, 宋姝, 吕婉嘉, 杨新豪, 马云霞, 杨永青. 盐碱胁迫下植物体内离子平衡调控的机制[J]. 植物生理学报, 2023, 59(4):715-726.doi:10.13592/j.cnki.ppj.300091.
	Liu X, Liu X H, Song S, Lü W J, Yang X H, Ma Y X, Yang Y Q. Regulation of ion homeostasis for salinity tolerance in plants[J]. Plant Physilogy Journal, 2023, 59(4):715-726.
[34]	Tomeo N J, Rosenthal D M. Variable mesophyll conductance among soybean cultivars sets a tradeoff between photosynthesis and water-use-efficiency[J]. Plant Physiology, 2017, 174(1):241-257.doi:10.1104/pp.16.01940. pmid: 28270627
[35]	Zuo Z Y, Ye F, Wang Z S, Li S X, Li H, Guo J H, Mao H P, Zhu X C, Li X N. Salt acclimation induced salt tolerance in wild-type and chlorophyl b-deficient mutant wheat[J]. Plant,Soil and Environment, 2021, 67(1):26-32.doi:10.17221/429/2020-pse.
[36]	赵澍, 廖里平, 张宏武, 马媛, 李钢铁. 盐胁迫对酸枣幼苗光合生理特性的影响[J]. 干旱区资源与环境, 2018, 32(5):149-153.doi:10.13448/j.cnki.jalre.2018.155.
	Zhao S, Liao L P, Zhang H W, Ma Y, Li G T. Impacts of salt stress on characteristics of photosynthetic of Zizyphus jujube seedlings[J]. Journal of Arid Land Resources and Environment, 2018, 32(5):149-153.
[37]	付乃鑫, 贺明荣, 诸葛玉平, 代兴龙, 胡国庆, 董元杰. 外源SA对盐胁迫下冬小麦幼苗生长的缓解效应及其机理[J]. 中国农业大学学报, 2019, 24(3):10-17.doi:10.11841/j.issn.1007-4333.2019.03.02.
	Fu N X, He M R, Zhuge Y P, Dai X L, Hu G Q, Dong Y J. Effects and mechanisms of exogenous SA alleviating the growth of winter wheat seedlings under salt stress[J]. Journal of China Agricultural University, 2019, 24(3):10-17.
[38]	尹宝丝. 盐胁迫对黑枸杞幼苗生长及光合特性的影响[D]. 北京: 北京林业大学, 2019.
	Yin B S. Effects of salt treatments on characteristics of growth and photosynthesis of Lycium ruthenicum Murr.[D]. Beijing: Beijing Forestry University, 2019.
[39]	宋倩云. 冬青对盐胁迫的生理响应及耐盐筛选[D]. 南京: 南京林业大学, 2020.
	Song Q Y. Physiological response of hollies to salt stress and the selection of salt-tolerant species[D]. Nanjing: Nanjing Forestry University, 2020.
[40]	赵可夫. 植物对盐渍逆境的适应[J]. 生物学通报, 2002, 37(6):7-10.doi:10.3969/j.issn.0006-3193.2002.06.003.
	Zhao K F. Adaptation to saline stress in plant[J]. Bulletin of Biology, 2002, 37(6):7-10.
[41]	Beacham A M, Hand P, Pink D A, Monaghan J M. Analysis of Brassica oleracea early stage abiotic stress responses reveals tolerance in multiple crop types and for multiple sources of stress[J]. Journal of the Science of Food and Agriculture, 2017, 97(15):5271-5277.doi:10.1002/jsfa.8411. pmid: 28474472
[42]	张风娟. 盐生植物耐盐结构的研究现状[J]. 河北职业技术师范学院学报, 2003, 17(4):75-78.doi:10.3969/j.issn.1672.7983.2003.04.020.
	Zhang F J. The study status quo of halophyte's salt tolerant structure[J]. Journal of Hebei Vocation-Technical Teachers College, 2003, 17(4):75-78.
[43]	梁爽, 王广野. 盐生植物耐盐结构研究进展[J]. 长春师范大学学报, 2015, 34(3):71-74.
	Liang S, Wang G Y. Research advance on salt-resistant structures of halophytes[J]. Journal of Changchun Normal University, 2015, 34(3):71-74.
[44]	Munns R, Passioura J B, Colmer T D, Byrt C S. Osmotic adjustment and energy limitations to plant growth in saline soil[J]. New Phytologist, 2020, 225(3):1091-1096.doi:10.1111/nph.15862. pmid: 31006123
[45]	Yuan F, Leng B Y, Wang B S. Progress in studying salt secretion from the salt glands in recretohalophytes:how do plants secrete salt?[J]. Frontiers in Plant Science, 2016, 7:977.doi:10.3389/fpls.2016.00977.
[46]	Yuan F, Wang B S. Handbook of Halophytes:adaptation of recretohalophytes to salinity[M]. Cambridge: Springer Press, 2020.doi:10.1007/978-3-030-17854-3_32-1.
[47]	Pollak G, Waisel Y. Salt secretion in Aeluropus litoralis (Willd.) parl[J]. Annals of Botany, 1970, 34(4):879-888.doi:10.1093/oxfordjournals.aob.a084419.
[48]	陆静梅, 李建东, 胡阿林, 历锡亮. 二色补血草叶片泌盐结构的扫描电镜观察[J]. 应用生态学报, 1995, 6(4):355-358.doi:10.13287/j.1001-9332.1995.0072.
	Lu J M, Li J D, Hu A L, Li X L. ESM observation of secretory salt structure in Limonium bicolor leaf[J]. Chinese Journal of Applied Ecology, 1995, 6(4):879-888.
[49]	Zhang M J, Chen Z, Yuan F, Wang B S, Chen M. Integrative transcriptome and proteome analyses provide deep insights into the molecular mechanism of salt tolerance in Limonium bicolor[J]. Plant Molecular Biology, 2022, 108(1/2):127-143.doi:10.1007/s11103-021-01230-z.
[50]	Yuan F, Wang X, Zhao B Q, et al. The genome of the recretohalophyte Limonium bicolor provides insights into salt gland development and salinity adaptation during terrestrial evolution[J]. Molecular Plant, 2022, 15(6):1024-1044.doi:10.1016/j.molp.2022.04.011.
[51]	Ramadan T, Flowers T J. Effects of salinity and benzyl adenine on development and function of microhairs of Zea mays L.[J]. Planta, 2004, 219(4):639-648.doi:10.1007/s00425-004-1269-7. pmid: 15098124
[52]	Jarvis D E, Ho Y S, Lightfoot D J, et al. The genome of Chenopodium quinoa[J]. Nature, 2017, 542:307-312.doi:10.1038/nature21370.
[53]	Mangelson H, Jarvis D E, Mollinedo P, Rollano-Penaloza O M, Palma-Encinas V D, Gomez-Pando L R, Jellen E N, Maughan P J. The genome of Chenopodium pallidicaule:an emerging Andean super grain[J]. Applications in Plant Sciences, 2019, 7(11):e11300.doi:10.1002/aps3.11300.
[54]	Chen J, Xiao Q, Wu F H, Dong X J, He J X, Pei Z M, Zheng H L, Näsholm T. Nitric oxide enhances salt secretion and Na⁺ sequestration in a mangrove plant,Avicennia marina,through increasing the expression of H⁺-ATPase and Na⁺/H⁺ antiporter under high salinity[J]. Tree Physiology, 2010, 30(12):1570-1585.doi:10.1093/treephys/tpq086.
[55]	杨洪兵. 苹果属植物抗盐机理的研究[D]. 北京: 中国农业大学, 2004.doi:10.7666/d.Y658934.
	Yang H B. Study on salt resistance mechanism of Malus plants[D]. Beijing: China Agricultural University, 2004.
[56]	Tan W K, Lin Q S, Lim T M, Kumar P, Loh C S. Dynamic secretion changes in the salt glands of the mangrove tree species Avicennia officinalis in response to a changing saline environment[J]. Plant,Cell & Environment, 2013, 36(8):1410-1422.doi:10.1111/pce.12068.
[57]	Duan L N, Dietrich D, Ng C H, Chan P M Y, Bhalerao R, Bennett M J, Dinneny J R. Endodermal ABA signaling promotes lateral root quiescence during salt stress in Arabidopsis seedlings[J]. The Plant Cell, 2013, 25(1):324-341.doi:10.1105/tpc.112.107227.
[58]	陆静梅, 李建东. 三种双子叶耐盐碱植物根的解剖研究[J]. 东北师大学报(自然科学版), 1994, 26(3):96-99.doi:10.16163/j.cnki.22-1123/n.1994.03.027.
	Lu J M, Li J D. The anatomical study of three dicotyledons roots in saline-alkali soil[J]. Journal of Northeast Normal Universtiy, 1994, 26(3):96-99.
[59]	Reinhardt D H, Rost T L. Salinity accelerates endodermal development and induces an exodermis in cotton seedling roots[J]. Environmental and Experimental Botany, 1995, 35(4):563-574.doi:10.1016/0098-8472(95)00015-1.
[60]	Karahara I, Ikeda A, Kondo T, Uetake Y. Development of the Casparian strip in primary roots of maize under salt stress[J]. Planta, 2004, 219(1):41-47.doi:10.1007/s00425-004-1208-7. pmid: 14986139
[61]	Barberon M, Vermeer J E M, De Bellis D, Wang P, Naseer S, Andersen T G, Humbel B M, Nawrath C, Takano J, Salt D E, Geldner N. Adaptation of root function by nutrient-induced plasticity of endodermal differentiation[J]. Cell, 2016, 164(3):447-459.doi:10.1016/j.cell.2015.12.021. pmid: 26777403
[62]	Wang Y Y, Cao Y B, Liang X Y, Zhuang J H, Wang X F, Qin F, Jiang C F. A dirigent family protein confers variation of Casparian strip thickness and salt tolerance in maize[J]. Nature Communications, 2022, 13(1):2222.doi:10.1038/s41467-022-29809-0.
[63]	Galvan-Ampudia C S, Julkowska M M, Darwish E, Gandullo J, Korver R A, Brunoud G, Haring M A, Munnik T, Vernoux T, Testerink C. Halotropism is a response of plant roots to avoid a saline environment[J]. Current Biology, 2013, 23(20):2044-2050.doi:10.1016/j.cub.2013.08.042. pmid: 24094855
[64]	Korver R A, van den Berg T, Meyer A J, Galvan-Ampudia C S, Ten Tusscher K H W J, Testerink C. Halotropism requires phospholipase Dζ1-mediated modulation of cellular polarity of auxin transport carriers[J]. Plant,Cell & Environment, 2020, 43(1):143-158.doi:10.1111/pce.13646.
[65]	Yu B, Zheng W N, Xing L, Zhu J K, Persson S, Zhao Y. Root twisting drives halotropism via stress-induced microtubule reorientation[J]. Developmental Cell, 2022, 57(20):2412-2425.e6.doi:10.1016/j.devcel.2022.09.012. pmid: 36243013
[66]	Sun H J, Sun X W, Wang H, Ma X L. Advances in salt tolerance molecular mechanism in tobacco plants[J]. Hereditas, 2020, 157(1):5.doi:10.1186/s41065-020-00118-0. pmid: 32093781
[67]	Qi Y C, Liu W Q, Qiu L Y, Zhang S M, Ma L, Zhang H. Overexpression of glutathione S-transferase gene increases salt tolerance of Arabidopsis[J]. Russian Journal of Plant Physiology, 2010, 57(2):233-240.doi:10.1134/S102144371002010X.
[68]	Pang C H, Li K, Wang B S. Overexpression of SsCHLAPXs confers protection against oxidative stress induced by high light in transgenic Arabidopsis thaliana[J]. Physiologia Plantarum, 2011, 143(4):355-366.doi:10.1111/j.1399-3054.2011.01515.x.
[69]	Li K, Pang C H, Ding F, Sui N, Feng Z T, Wang B S. Overexpression of Suaeda salsa stroma ascorbate peroxidase in Arabidopsis chloroplasts enhances salt tolerance of plants[J]. South African Journal of Botany, 2012, 78:235-245.doi:10.1016/j.sajb.2011.09.006.
[70]	Negi N P, Shrivastava D C, Sharma V, Sarin N B. Overexpression of CuZnSOD from Arachis hypogaea alleviates salinity and drought stress in tobacco[J]. Plant Cell Reports, 2015, 34(7):1109-1126.doi:10.1007/s00299-015-1770-4.
[71]	Mittler R. Oxidative stress,antioxidants and stress tolerance[J]. Trends in Plant Science, 2002, 7(9):405-410.doi:10.1016/s1360-1385(02)02312-9. pmid: 12234732
[72]	Arias-Moreno D M, Jiménez-Bremont J F, Maruri-López I, Delgado-Sánchez P. Effects of catalase on chloroplast arrangement in Opuntia streptacantha chlorenchyma cells under salt stress[J]. Scientific Reports, 2017, 7(1):8656.doi:10.1038/s41598-017-08744-x. pmid: 28819160
[73]	Sharma P, Dubey R S. Ascorbate peroxidase from rice seedlings:properties of enzyme isoforms,effects of stresses and protective roles of osmolytes[J]. Plant Science, 2004, 167(3):541-550.doi:10.1016/j.plantsci.2004.04.028.
[74]	Singh N, Mishra A, Jha B. Over-expression of the peroxisomal ascorbate peroxidase (SbpAPX) gene cloned from halophyte Salicornia brachiata confers salt and drought stress tolerance in transgenic tobacco[J]. Marine Biotechnology, 2014, 16(3):321-332.doi:10.1007/s10126-013-9548-6.
[75]	Huang D J, Ou B X, Prior R L. The chemistry behind antioxidant capacity assays[J]. Journal of Agricultural and Food Chemistry, 2005, 53(6):1841-1856.doi:10.1021/jf030723c. pmid: 15769103
[76]	Ksouri R, Megdiche W, Debez A, Falleh H, Grignon C, Abdelly C. Salinity effects on polyphenol content and antioxidant activities in leaves of the halophyte Cakile maritima[J]. Plant Physiology and Biochemistry, 2007, 45(3/4):244-249.doi:10.1016/j.plaphy.2007.02.001.
[77]	Falleh H, Jalleli I, Ksouri R, Boulaaba M, Guyot S, Magné C, Abdelly C. Effect of salt treatment on phenolic compounds and antioxidant activity of two Mesembryanthemum edule provenances[J]. Plant Physiology and Biochemistry, 2012, 52:1-8.doi:10.1016/j.plaphy.2011.11.001. pmid: 22305062
[78]	Valifard M, Mohsenzadeh S, Kholdebarin B, Rowshan V. Effects of salt stress on volatile compounds,total phenolic content and antioxidant activities of Salvia mirzayanii[J]. South African Journal of Botany, 2014, 93:92-97.doi:10.1016/j.sajb.2014.04.002.
[79]	Shao H B, Liang Z S, Shao M A, Sun Q. Dynamic changes of anti-oxidative enzymes of 10 wheat genotypes at soil water deficits[J]. Colloids and Surfaces B,Biointerfaces, 2005, 42(3/4):187-195.doi:10.1016/j.colsurfb.2005.02.007.
[80]	杜玉玲. 外源水杨酸对盐胁迫唐古特白刺ROS代谢以及AsA-GSH循环的影响[D]. 哈尔滨: 东北农业大学, 2017.
	Du Y L. Effects of exogenous salicylic acid on ROS metabolism and AsA-GSH cycle in Nitraria Tangutorum Bor.under salt stress.[D]. Harbin:Northeast Agricultural University, 2017.
[81]	陈沁, 刘友良. 谷胱甘肽对盐胁迫大麦叶片活性氧清除系统的保护作用[J]. 作物学报, 2000, 26(3):365.doi:10.3321/j.issn:0496-3490.2000.03.019.
	Chen Q, Liu Y L. Effect of glutathione on active oxygen scavenging system in leaves of barley seedlings under salt stress[J]. Acta Agronomica Sinica, 2000, 26(3):365.
[82]	Boestfleisch C, Papenbrock J. Changes in secondary metabolites in the halophytic putative crop species Crithmum maritimum L.,Triglochin maritima L.and Halimione portulacoides (L.) Aellen as reaction to mild salinity[J]. PLoS One, 2017, 12(4):e0176303.doi:10.1371/journal.pone.0176303.
[83]	He J, Ng O W J, Qin L. Salinity and salt-priming impact on growth,photosynthetic performance,and nutritional quality of edible Mesembryanthemum crystallinum L.[J]. Plants, 2022, 11(3):332.doi:10.3390/plants11030332.
[84]	Verbruggen N, Hermans C. Proline accumulation in plants:a review[J]. Amino Acids, 2008, 35(4):753-759.doi:10.1007/s00726-008-0061-6. pmid: 18379856
[85]	杨洪兵. 外源Mg²⁺对荞麦耐盐性的影响[J]. 华北农学报, 2012, 27(6):130-133.doi:10.3969/j.issn.1000-7091.2012.06.026.
	Yang H B. Effect of exogenous Mg²⁺ on salt tolerance of buckwheat[J]. Acta Agriculturae Boreali-Sinica, 2012, 27(6):130-133.
[86]	Shabala S, Shabala L. Ion transport and osmotic adjustment in plants and bacteria[J]. Biomolecular Concepts, 2011, 2(5):407-419.doi:10.1515/BMC.2011.032. pmid: 25962045
[87]	Zou C L, Liu D, Wu P R, Wang Y B, Gai Z J, Liu L, Yang F F, Li C F, Guo G H. Transcriptome analysis of sugar beet (Beta vulgaris L.) in response to alkaline stress[J]. Plant Molecular Biology, 2020, 102(6):645-657.doi:10.1007/s11103-020-00971-7.
[88]	Teh C Y, Shaharuddin N A, Ho C L, Mahmood M. Exogenous proline significantly affects the plant growth and nitrogen assimilation enzymes activities in rice (Oryza sativa) under salt stress[J]. Acta Physiologiae Plantarum, 2016, 38(6):151.doi:10.1007/s11738-016-2163-1.
[89]	Butt M, Sattar A, Abbas T, Sher A, Ijaz M, Ul-Allah S, Shaheen M R, Kaleem F. Foliage applied proline induces salt tolerance in chili genotypes by regulating photosynthetic attributes,ionic homeostasis,and antioxidant defense mechanisms[J]. Horticulture,Environment,and Biotechnology, 2020, 61(4):693-702.doi:10.1007/s13580-020-00236-8.
[90]	Konstantinova T, Parvanova D, Atanassov A, Djilianov D. Freezing tolerant tobacco,transformed to accumulate osmoprotectants[J]. Plant Science, 2002, 163(1):157-164.doi:10.1016/s0168-9452(02)00090-0.
[91]	Szabados L, Savouré A. Proline:a multifunctional amino acid[J]. Trends in Plant Science, 2010, 15(2):89-97.doi:10.1016/j.tplants.2009.11.009. pmid: 20036181
[92]	刘晓蕊, 尚丽霞, 蔡勤安, 于志晶, 马瑞. 耐盐转基因植物研究进展[J]. 东北农业科学, 2021, 46(4):27-33.doi:10.16423/j.cnki.1003-8701.2021.04.007.
	Liu X R, Shang L X, Cai Q A, Yu Z J, Ma R. Research progress of salt tolerant transgenic plants[J]. Journal of Northeast Agricultural Sciences, 2021, 46(4):27-33.
[93]	李班, 吕莹, 杨明煊, 宋婷, 于放, 刘志文. 盐碱胁迫对甘蓝型油菜生理及分子机制的影响[J]. 华北农学报, 2022, 37(3):86-93.doi:10.7668/hbnxb.20192719.
	Li B, Lü Y, Yang M X, Song T, Yu F, Liu Z W. Effects of saline-alkali stress on physiology and molecular mechanism of Brassica napus L.[J]. Acta Agriculturae Boreali-Sinica, 2022, 37(3):86-93.
[94]	De la Torre-González A, Montesinos-Pereira D, Blasco B, Ruiz J M. Influence of the proline metabolism and glycine betaine on tolerance to salt stress in tomato (Solanum lycopersicum L.) commercial genotypes[J]. Journal of Plant Physiology, 2018, 231:329-336.doi:10.1016/j.jplph.2018.10.013. pmid: 30388672
[95]	Li Q L, Gao X R, Yu X H, Wang X Z, An L J. Molecular cloning and characterization of betaine aldehyde dehydrogenase gene from Suaeda liaotungensis and its use in improved tolerance to salinity in transgenic tobacco[J]. Biotechnology Letters, 2003, 25(17):1431-1436.doi:10.1023/A:1025003628446.
[96]	Chen T H H, Murata N. Glycinebetaine protects plants against abiotic stress:mechanisms and biotechnological applications[J]. Plant,Cell & Environment, 2011, 34(1):1-20.doi:10.1111/j.1365-3040.2010.02232.x.
[97]	Xu Z J, Sun M L, Jiang X F, Sun H P, Dang X M, Cong H Q, Qiao F. Glycinebetaine biosynthesis in response to osmotic stress depends on jasmonate signaling in watermelon suspension cells[J]. Frontiers in Plant Science, 2018, 9:1469.doi:10.3389/fpls.2018.01469. pmid: 30369936
[98]	Abebe T, Guenzi A C, Martin B, Cushman J C. Tolerance of mannitol-accumulating transgenic wheat to water stress and salinity[J]. Plant Physiology, 2003, 131(4):1748-1755.doi:10.1104/pp.102.003616. pmid: 12692333
[99]	Penna S. Building stress tolerance through over-producing trehalose in transgenic plants[J]. Trends in Plant Science, 2003, 8(8):355-357.doi:10.1016/S1360-1385(03)00159-6. pmid: 12927963
[100]	da Silva J M, Arrabça M C. Contributions of soluble carbohydrates to the osmotic adjustment in the C4 grass Setaria sphacelata:a comparison between rapidly and slowly imposed water stress[J]. Journal of Plant Physiology, 2004, 161(5):551-555.doi:10.1078/0176-1617-01109.
[101]	Fujii H, Verslues P E, Zhu J K. Arabidopsis decuple mutant reveals the importance of SnRK2 kinases in osmotic stress responses in vivo[J]. Proceedings of the National Academy of Sciences of the United States of America, 2011, 108(4):1717-1722.doi:10.1073/pnas.1018367108.
[102]	Mustilli A C, Merlot S, Vavasseur A, Fenzi F, Giraudat J. Arabidopsis OST1 protein kinase mediates the regulation of stomatal aperture by abscisic acid and acts upstream of reactive oxygen species production[J]. The Plant Cell, 2002, 14(12):3089-3099.doi:10.1105/tpc.007906.
[103]	Yoshida R, Hobo T, Ichimura K, Mizoguchi T, Takahashi F, Aronso J, Ecker J R, Shinozaki K. ABA-activated SnRK2 protein kinase is required for dehydration stress signaling in Arabidopsis[J]. Plant & Cell Physiology, 2002, 43(12):1473-1483.doi:10.1093/pcp/pcf188.
[104]	杨升, 张华新, 陈秋夏, 杨秀艳. 沙枣幼苗根尖离子流对NaCl胁迫的响应[J]. 植物生态学报, 2017, 41(4):489-496.doi:10.17521/cjpe.2016.0091.
	Yang S, Zhang H X, Chen Q X, Yang X Y. Responses of apical ion fluxes to NaCl stress in Elaeagnus angustifolia seedlings[J]. Chinese Journal of Plant Ecology, 2017, 41(4):489-496.
[105]	Faisal S, Mujtaba S M,Asma, Mahboob W. Polyethylene glycol mediated osmotic stress impacts on growth and biochemical aspects of wheat (Triticum aestivum L.)[J]. Journal of Crop Science and Biotechnology, 2019, 22(3):213-223.doi:10.1007/s12892-018-0166-0.
[106]	Jacob P T, Siddiqui S A, Rathore M S. Seed germination,seedling growth and seedling development associated physiochemical changes in Salicornia brachiata (Roxb.) under salinity and osmotic stress[J]. Aquatic Botany, 2020, 166:103272.doi:10.1016/j.aquabot.2020.103272.
[107]	董宏图, 解超杰, 侯佩臣, 李爱学, 王晓冬. 高盐胁迫下小麦幼苗离子吸收动态及耐盐性筛选[J]. 中国生态农业学报, 2021, 29(4):762-770.doi:10.13930/j.cnki.cjea.200664.
	Dong H T, Xie C J, Hou P C, Li A X, Wang X D. Dynamic of ionic absorption and salt tolerance screening in wheat seedling under salt stress[J]. Chinese Journal of Eco-Agriculture, 2021, 29(4):762-770.
[108]	张华宁, 李孟军, 郭秀林, 张艳敏, 刘子会. 盐胁迫下不同K⁺吸收抑制剂对小麦根系K⁺/Na⁺比和质膜相关蛋白活性的影响[J]. 华北农学报, 2017, 32(5):154-162.doi:10.7668/hbnxb.2017.05.024.
	Zhang H N, Li M J, Guo X L, Zhang Y M, Liu Z H. Effects of K⁺ absorption inhibitors on K⁺/Na⁺ ratios and activities of plasma membrane proteins under salt stress in Triticum aestivum L.[J]. Acta Agriculturae Boreali-Sinica, 2017, 32(5):154-162.
[109]	Mei X D, Dai T, Shen Y B. Adaptive strategy of Nitraria sibirica to transient salt,alkali and osmotic stresses via the alteration of Na⁺/K⁺ fluxes around root tips[J]. Journal of Forestry Research, 2023, 34(2):425-432.doi:10.1007/s11676-022-01486-1.
[110]	张倩, 李笑佳, 张淑英. 硅对盐胁迫下棉花幼苗生长和渗透调节系统的影响[J]. 华北农学报, 2019, 34(6):110-117.doi:10.7668/hbnxb.201751589.
	Zhang Q, Li X J, Zhang S Y. Effects of silicon on growth and osmotic regulation of cotton seedlings under salt stress[J]. Acta Agriculturae Boreali-Sinica, 2019, 34(6):110-117. doi: 10.7668/hbnxb.201751589
[111]	Chérel I, Lefoulon C, Boeglin M, Sentenac H. Molecular mechanisms involved in plant adaptation to low K⁺ availability[J]. Journal of Experimental Botany, 2014, 65(3):833-848.doi:10.1093/jxb/ert402.
[112]	Anschütz U, Becker D, Shabala S. Going beyond nutrition:regulation of potassium homoeostasis as a common denominator of plant adaptive responses to environment[J]. Journal of Plant Physiology, 2014, 171(9):670-687.doi:10.1016/j.jplph.2014.01.009. pmid: 24635902
[113]	Genc Y, Oldach K, Taylor J, Lyons G H. Uncoupling of sodium and chloride to assist breeding for salinity tolerance in crops[J]. New Phytologist, 2016, 210(1):145-156.doi:10.1111/nph.13757. pmid: 26607560
[114]	Oyiga B C, Sharma R C, Baum M, Ogbonnaya F C, Léon J, Ballvora A. Allelic variations and differential expressions detected at quantitative trait loci for salt stress tolerance in wheat[J]. Plant,Cell & Environment, 2018, 41(5):919-935.doi:10.1111/pce.12898.
[115]	Zhao S S, Zhang Q K, Liu M Y, Zhou H P, Ma C L, Wang P P. Regulation of plant responses to salt stress[J]. International Journal of Molecular Sciences, 2021, 22(9):4609.doi:10.3390/ijms22094609.
[116]	Azhar N, Su N N, Shabala L, Shabala S. Exogenously applied 24-epibrassinolide (EBL) ameliorates detrimental effects of salinity by reducing K⁺ efflux via depolarization-activated K⁺ channels[J]. Plant & Cell Physiology, 2017, 58(4):802-810.doi:10.1093/pcp/pcx026.
[117]	Su Y, Luo W G, Lin W H, Ma L Y, Kabir M H. Model of cation transportation mediated by high-affinity potassium transporters (HKTs) in higher plants[J]. Biological Procedures Online, 2015, 17:1.doi:10.1186/s12575-014-0013-3. pmid: 25698907
[118]	Hamamoto S, Horie T, Hauser F, Deinlein U, Schroeder J I, Uozumi N. HKT transporters mediate salt stress resistance in plants:from structure and function to the field[J]. Current Opinion in Biotechnology, 2015, 32:113-120.doi:10.1016/j.copbio.2014.11.025. pmid: 25528276
[119]	Buschmann P H, Vaidyanathan R, Gassmann W, Schroeder J I. Enhancement of Na⁺ uptake currents,time-dependent inward-rectifying K⁺ channel currents,and K⁺channel transcripts by K⁺ starvation in wheat root cells[J]. Plant Physiology, 2000, 122(4):1387-1398.doi:10.1104/pp.122.4.1387. pmid: 10759535
[120]	Ahmad P, Abdel Latef A A, Abd Allah E F, Hashem A, Sarwat M, Anjum N A, Gucel S. Calcium and potassium supplementation enhanced growth,osmolyte secondary metabolite production,and enzymatic antioxidant machinery in cadmium-exposed chickpea (Cicer arietinum L.)[J]. Frontiers in Plant Science, 2016, 7:513.doi:10.3389/fpls.2016.00513.
[121]	Zhu J K. Cell signaling under salt,water and cold stresses[J]. Current Opinion in Plant Biology, 2001, 4(5):401-406.doi:10.1016/s1369-5266(00)00192-8. pmid: 11597497
[122]	Zhu J K. Salt and drought stress signal transduction in plants[J]. Annual Review of Plant Biology, 2002, 53:247-273.doi:10.1146/annurev.arplant.53.091401.143329.
[123]	Gong D M, Guo Y, Schumaker K S, Zhu J K. The SOS3 family of calcium sensors and SOS2 family of protein kinases in Arabidopsis[J]. Plant Physiology, 2004, 134(3):919-926.doi:10.1104/pp.103.037440.
[124]	Du W M, Lin H X, Chen S, Wu Y S, Zhang J, Fuglsang A T, Palmgren M G, Wu W H, Guo Y. Phosphorylation of SOS3-like calcium-binding proteins by their interacting SOS2-like protein kinases is a common regulatory mechanism in Arabidopsis[J]. Plant Physiology, 2011, 156(4):2235-2243.doi:10.1104/pp.111.173377.
[125]	Bassil E, Coku A, Blumwald E. Cellular ion homeostasis:emerging roles of intracellular NHX Na⁺/H⁺ antiporters in plant growth and development[J]. Journal of Experimental Botany, 2012, 63(16):5727-5740.doi:10.1093/jxb/ers250. pmid: 22991159
[126]	Katschnig D, Bliek T, Rozema J, Schat H. Constitutive high-level SOS1 expression and absence of HKT1;1 expression in the salt-accumulating halophyte Salicornia dolichostachya[J]. Plant Science, 2015, 234:144-154.doi:10.1016/j.plantsci.2015.02.011. pmid: 25804817
[127]	Zhu J K. Abiotic stress signaling and responses in plants[J]. Cell, 2016, 167(2):313-324.doi:10.1016/j.cell.2016.08.029.
[128]	Dragwidge J M, Scholl S, Schumacher K, Gendall A R. NHX-type Na⁺ (K⁺)/H⁺ antiporters are required for TGN/EE trafficking and endosomal ion homeostasis in Arabidopsis thaliana[J]. Journal of Cell Science, 2019, 132(7):226472 doi:10.1242/jcs.226472.
[129]	Achard P, Cheng H, De Grauwe L, Decat J, Schoutteten H, Moritz T, Van Der Straeten D, Peng J R, Harberd N P. Integration of plant responses to environmentally activated phytohormonal signals[J]. Science, 2006, 311(5757):91-94.doi:10.1126/science.1118642. pmid: 16400150
[130]	Orman-Ligeza B, Morris E C, Parizot B, et al. The xerobranching response represses lateral root formation when roots are not in contact with water[J]. Current Biology, 2018, 28(19):3165-3173.e5.doi:10.1016/j.cub.2018.07.074. pmid: 30270188
[131]	Irving H R, Gehring C A, Parish R W. Changes in cytosolic pH and calcium of guard cells precede stomatal movements[J]. Proceedings of the National Academy of Sciences of the United States of America, 1992, 89(5):1790-1794.doi:10.1073/pnas.89.5.1790. pmid: 11607281
[132]	Brandt B, Munemasa S, Wang C, et al. Calcium specificity signaling mechanisms in abscisic acid signal transduction in Arabidopsis guard cells[J]. eLife, 2015, 4:e03599.doi:10.7554/eLife.03599.
[133]	Gomez-Roldan V, Fermas S, Brewer P B, et al. Strigolactone inhibition of shoot branching[J]. Nature, 2008, 455(7210):189-194.doi:10.1038/nature07271.
[134]	Umehara M, Hanada A, Yoshida S, Akiyama K, Arite T, Takeda-Kamiya N, Magome H, Kamiya Y, Shirasu K, Yoneyama K, Kyozuka J, Yamaguchi S. Inhibition of shoot branching by new terpenoid plant hormones[J]. Nature, 2008, 455(7210):195-200.doi:10.1038/nature07272.
[135]	Van Ha C, Leyva-González M A, Osakabe Y, et al. Positive regulatory role of strigolactone in plant responses to drought and salt stress[J]. Proceedings of the National Academy of Sciences of the United States of America, 2014, 111(2):851-856.doi:10.1073/pnas.1322135111. pmid: 24379380
[136]	Qiao Y H, Lu W X, Wang R, Nisa Z U, Yu Y, Jin X X, Yu L J, Chen C. Identification and expression analysis of strigolactone biosynthetic and signaling genes in response to salt and alkaline stresses in soybean(Glycine max)[J]. DNA and Cell Biology, 2020, 39(10):1850-1861.doi:10.1089/dna.2020.5637.
[137]	Bleecker A B, Kende H. Ethylene:a gaseous signal molecule in plants[J]. Annual Review of Cell and Developmental Biology, 2000, 16:1-18.doi:10.1146/annurev.cellbio.16.1.1. pmid: 11031228
[138]	Cao W H, Liu J, He X J, Mu R L, Zhou H L, Chen S Y, Zhang J S. Modulation of ethylene responses affects plant salt-stress responses[J]. Plant Physiology, 2007, 143(2):707-719.doi:10.1104/pp.106.094292.
[139]	Zhang L X, Li Z F, Quan R D, Li G J, Wang R G, Huang R F. An AP2 domain-containing gene,ESE1,targeted by the ethylene signaling component EIN3 is important for the salt response in Arabidopsis[J]. Plant Physiology, 2011, 157(2):854-865.doi:10.1104/pp.111.179028.
[140]	Peng J Y, Li Z H, Wen X, Li W Y, Shi H, Yang L S, Zhu H Q, Guo H W. Salt-induced stabilization of EIN3/EIL1 confers salinity tolerance by deterring ROS accumulation in Arabidopsis[J]. PLoS Genetics, 2014, 10(10):e1004664.doi:10.1371/journal.pgen.1004664.
[141]	Dou L R, He K K, Higaki T, Wang X F, Mao T L. Ethylene signaling modulates cortical microtubule reassembly in response to salt stress[J]. Plant Physiology, 2018, 176(3):2071-2081.doi:10.1104/pp.17.01124. pmid: 29431630
[142]	Yu Y W, Wang J, Shi H, Gu J T, Dong J G, Deng X W, Huang R F. Salt stress and ethylene antagonistically regulate nucleocytoplasmic partitioning of COP1 to control seed germination[J]. Plant Physiology, 2016, 170(4):2340-2350.doi:10.1104/pp.15.01724. pmid: 26850275
[143]	Schachtman D P, Goodger J Q D. Chemical root to shoot signaling under drought[J]. Trends in Plant Science, 2008, 13(6):281-287.doi:10.1016/j.tplants.2008.04.003. pmid: 18467158
[144]	Nishiyama R, Watanabe Y, Fujita Y, et al. Analysis of cytokinin mutants and regulation of cytokinin metabolic genes reveals important regulatory roles of cytokinins in drought,salt and abscisic acid responses,and abscisic acid biosynthesis[J]. The Plant Cell, 2011, 23(6):2169-2183.doi:10.1105/tpc.111.087395. pmid: 21719693
[145]	Wang L, Li Q T, Lei Q, Feng C, Gao Y N, Zheng X D, Zhao Y, Wang Z, Kong J. MzPIP2;1:an aquaporin involved in radial water movement in both water uptake and transportation,altered the drought and salt tolerance of transgenic Arabidopsis[J]. PLoS One, 2015, 10(11):e0142446.doi:10.1371/journal.pone.0142446.
[146]	Allegra M, Reiter R J, Tan D X, Gentile C, Tesoriere L, Livrea M A. The chemistry of melatonin’s interaction with reactive species[J]. Journal of Pineal Research, 2003, 34(1):1-10.doi:10.1034/j.1600-079x.2003.02112.x. pmid: 12485365
[147]	陈楠, 张维, 张晓明. 褪黑素调控辣椒幼苗响应盐胁迫下的叶片生理特性[J]. 中国瓜菜, 2023, 36(5):84-90.doi:10.3969/j.issn.1673-2871.2023.05.012.
	Chen N, Zhang W, Zhang X M. Melatonin regulates physiological characteristics of pepper seedlings in response to salt stress[J]. China Cucurbits and Vegetables, 2023, 36(5):84-90.
[148]	郭远航, 王洪博, 白宝伟, 张磊, 赵丰年, 吕东雪, 贾婷, 王兴鹏. 外源褪黑素对大豆幼苗盐胁迫的缓解效应[J]. 华北农学报, 2024, 39(2):116-125.doi:10.7668/hbnxb.20194665.
	Guo Y H, Wang H B, Bai B W, Zhang L, Zhao F N, Lü D X, Jia T, Wang X P. Effects of exogenous melatonin on salt stress reduction in soybean seedlings[J]. Acta Agriculturae Boreali-Sinica, 2024, 39(2):116-125. doi: 10.7668/hbnxb.20194665
[149]	王彪, 宋世佳, 李东晓, 董伟欣, 张月辰. 褪黑素通过H₂O₂调控盐胁迫下小豆Na⁺/K⁺平衡机制[J]. 华北农学报, 2023, 38(6):62-71.doi:10.7668/hbnxb.20194433.
	Wang B, Song S J, Li D X, Dong W X, Zhang Y C. The mechanism of melatonin regulating Na⁺/K⁺ balance by mediating H₂O₂ in adzuki bean under salt stress[J]. Acta Agriculturae Boreali-Sinica, 2023, 38(6):62-71.
[150]	Savaldi-Goldstein S, Peto C, Chory J. The epidermis both drives and restricts plant shoot growth[J]. Nature, 2007, 446(7132):199-202.doi:10.1038/nature05618.
[151]	Ubeda-Tomás S, Swarup R, Coates J, Swarup K, Laplaze L, Beemster G T S, Hedden P, Bhalerao R, Bennett M J. Root growth in Arabidopsis requires gibberellin/DELLA signalling in the endodermis[J]. Nature Cell Biology, 2008, 10(5):625-628.doi:10.1038/ncb1726. pmid: 18425113
[152]	Ubeda-Tomás S, Federici F, Casimiro I, Beemster G T S, Bhalerao R, Swarup R, Doerner P, Haseloff J, Bennett M J. Gibberellin signaling in the endodermis controls Arabidopsis root meristem size[J]. Current Biology, 2009, 19(14):1194-1199.doi:10.1016/j.cub.2009.06.023. pmid: 19576770
[153]	Hacham Y, Holland N, Butterfield C, Ubeda-Tomas S, Bennett M J, Chory J, Savaldi-Goldstein S. Brassinosteroid perception in the epidermis controls root meristem size[J]. Development, 2011, 138(5):839-848.doi:10.1242/dev.061804. pmid: 21270053
[154]	González-García M P, Vilarrasa-Blasi J, Zhiponova M, Divol F, Mora-García S, Russinova E, Caño-Delgado A I. Brassinosteroids control meristem size by promoting cell cycle progression in Arabidopsis roots[J]. Development, 2011, 138(5):849-859.doi:10.1242/dev.057331. pmid: 21270057
[155]	Zhao Y, Dong W, Zhang N B, Ai X H, Wang M C, Huang Z G, Xiao L T, Xia G M. A wheat allene oxide cyclase gene enhances salinity tolerance via jasmonate signaling[J]. Plant Physiology, 2014, 164(2):1068-1076.doi:10.1104/pp.113.227595. pmid: 24326670
[156]	Knight H, Trewavas A J, Knight M R. Calcium signalling in Arabidopsis thaliana responding to drought and salinity[J]. The Plant Journal, 1997, 12(5):1067-1078.doi:10.1046/j.1365-313x.1997.12051067.x.
[157]	Stephan A B, Kunz H H, Yang E, Schroeder J I. Rapid hyperosmotic-induced Ca²⁺ responses in Arabidopsis thaliana exhibit sensory potentiation and involvement of plastidial KEA transporters[J]. Proceedings of the National Academy of Sciences of the United States of America, 2016, 113(35):E5242-E5249.doi:10.1073/pnas.1519555113.
[158]	Han N, Lan W J, He X, Shao Q, Wang B S, Zhao X J. Expression of a Suaeda salsa vacuolar H⁺/Ca²⁺ transporter gene in Arabidopsis contributes to physiological changes in salinity[J]. Plant Molecular Biology Reporter, 2012, 30(2):470-477.doi:10.1007/s11105-011-0353-y.
[159]	Vivek P J, Tuteja N, Soniya E V. CDPK1 from ginger promotes salinity and drought stress tolerance without yield penalty by improving growth and photosynthesis in Nicotiana tabacum[J]. PLoS One, 2013, 8(10):e76392.doi:10.1371/journal.pone.0076392.
[160]	Zhang S Q, Klessig D F. MAPK cascades in plant defense signaling[J]. Trends in Plant Science, 2001, 6(11):520-527.doi:10.1016/s1360-1385(01)02103-3. pmid: 11701380
[161]	Jonak C, Nakagami H, Hirt H. Heavy metal stress.Activation of distinct mitogen-activated protein kinase pathways by copper and cadmium[J]. Plant Physiology, 2004, 136(2):3276-3283.doi:10.1104/pp.104.045724.
[162]	Yang C J, Wang R N, Gou L Z, Si Y C, Guan Q J. Overexpression of Populus trichocarpa mitogen-activated protein kinase Kinase4 enhances salt tolerance in tobacco[J]. International Journal of Molecular Sciences, 2017, 18(10):2090.doi:10.3390/ijms18102090.
[163]	Sunagawa H, Cushman J, Agarie S. Crassulacean acid metabolism may alleviate production of reactive oxygen species in a facultative CAM plant,the common ice plant Mesembryanthemum crystallinum L.[J]. Plant Production Science, 2010, 13(3):256-260.doi:10.1626/pps.13.256.
[164]	陆静梅, 李建东, 景德章, 杨凤清, 张洪芹. 星星草Puccinellia tenuiflora (Turcz.) Scribn.et Merr.解剖研究[J]. 东北师大学报(自然科学版), 1994(1):63-66.doi:10.16163/j.cnki,22-1123/n.1994.01.016.
	Lu J M, Li J D, Jing D Z, Yang F Q, Zhang H Q. The anatomical study of Puccinella tenuiflora (Turcz.) Scribn.et Merr.[J]. Journal of Northeast Normal University, 1994(1):63-66.
[165]	陆静梅, 张常钟, 张洪芹, 杨凤清. 单子叶植物耐盐碱的形态解剖特征与生理适应的相关性研究[J]. 东北师大学报(自然科学版), 1994(2):79-82.doi:10.16163/j.cnki,22-1123/n.1994.02.021.
	Lu J M, Zhang C Z, Zhang H Q, Yang F Q. The character of morphology anatomy of monocotyledons in saline-alkali soil of resistant and the study physiological adaptability interrelation[J]. Journal of Northeast Normal University, 1994(2):79-82.
[166]	Liu Y M, Xiong Y, Bassham D C. Autophagy is required for tolerance of drought and salt stress in plants[J]. Autophagy, 2009, 5(7):954-963.doi:10.4161/auto.5.7.9290. pmid: 19587533
[167]	Yang X C, Srivastava R, Howell S H, Bassham D C. Activation of autophagy by unfolded proteins during endoplasmic reticulum stress[J]. The Plant Journal, 2016, 85(1):83-95.doi:10.1111/tpj.13091. pmid: 26616142
[168]	万雪, 荆文旭, 魏磊, 邢旭明, 史树德. 甜菜幼苗对盐胁迫的生理响应及其自噬现象[J]. 华北农学报, 2021, 36(4):90-95.doi:10.7668/hbnxb.20191924.
	Wan X, Jing W X, Wei L, Xing X M, Shi S D. Physiological characteristics and autophagy of sugar beet seedlings in response to salt stress[J]. Acta Agriculturae Boreali-Sinica, 2021, 36(4):90-95. doi: 10.7668/hbnxb.20191924
[169]	Mittler R, Zandalinas S I, Fichman Y, Van Breusegem F. Reactive oxygen species signaling in plant stress responses[J]. Nature Reviews Molecular Cell Biology, 2022, 23(10):663-679.doi:10.1038/s41580-022-00499-2.
[170]	梁栋, 杨于杰, 耿彪, 景盼盼, 苏献存, 武林锐, 渠云芳, 黄晋玲. 棉花远缘杂交新种质耐盐性鉴定及生理特性分析[J]. 华北农学报, 2024, 39(1):95-103.doi:10.7668/hbnxb.20194209.
	Liang D, Yang Y J, Geng B, Jing P P, Su X C, Wu L R, Qu Y F, Huang J L. Identification of salt tolerance and physiological characteristics of new germplasm of distant hybridization of cotton[J]. Acta Agriculturae Boreali-Sinica, 2024, 39(1):95-103. doi: 10.7668/hbnxb.20194209
[171]	Zhang A N, Liu Y, Wang F M, Li T F, Chen Z H, Kong D Y, Bi J G, Zhang F Y, Luo X X, Wang J H, Tang J J, Yu X Q, Liu G L, Luo L J. Enhanced rice salinity tolerance via CRISPR/Cas9-targeted mutagenesis of the OsRR22 gene[J]. Molecular Breeding, 2019, 39(3):47.doi:10.1007/s11032-019-0954-y.
[172]	莫天宇, 徐善斌, 邹德堂, 王敬国, 刘化龙, 孙健, 贾琰, 赵宏伟, 郑洪亮. 利用CRISPR/Cas9技术敲除OsEIL1和OsEIL2基因改良水稻耐盐性[J]. 华北农学报, 2021, 36(1):71-80.doi:10.7668/hbnxb.20190798.
	Mo T Y, Xu S B, Zou D T, Wang J G, Liu H L, Sun J, Jia Y, Zhao H W, Zheng H L. Enhancing salt tolerance of rice by knocking out OsEIL1 and OsEIL2 via CRISPR/Cas9 system[J]. Acta Agriculturae Boreali-Sinica, 2021, 36(1):71-80.
[173]	程利华. 转ScALDH21基因棉花的获得及耐盐性鉴定[D]. 乌鲁木齐: 新疆农业大学, 2022.
	Cheng L H. Acquisition of transgenic ScALDH21 cotton and identification of transgenic cotton on salt tolerance[D]. Urumqi: Xinjiang Agricultural University, 2022.

选择文件类型/文献管理软件名称

选择包含的内容

植物耐盐机制研究进展及展望

Advances and Prospect on Mechanism of Salt Tolerance in Plants

RichHTML

PDF

可视化

被引次数

摘要/Abstract

引用本文

使用本文

参考文献 173

相关文章 15

编辑推荐

Metrics

本文评价

[1]	肖陈耀东, 刘涛, 刘仕志, 张淑英. 外源H₂O₂对盐胁迫下棉花幼苗生理特性的影响[J]. 华北农学报, 2024, 39(5): 93-101.
[2]	尹明达, 罗蕊, 任文静, 王志妍, 苏志敏, 李茹鑫, 陈圳, 李玉玲, 王艳, 黄凤兰. *蓖麻PIP5K2基因克隆及功能预测*[J]. 华北农学报, 2024, 39(4): 72-79.
[3]	郭远航, 王洪博, 白宝伟, 张磊, 赵丰年, 吕东雪, 贾婷, 王兴鹏. 外源褪黑素对大豆幼苗盐胁迫的缓解效应[J]. 华北农学报, 2024, 39(2): 116-125.
[4]	沈巧玲, 刘光敏, 王亚钦, 赵子璇, 魏蕾, 何洪巨, 刘海河. 壳寡糖对羽衣甘蓝芽苗硫代葡萄糖苷及抗氧化活性的影响[J]. 华北农学报, 2024, 39(1): 104-112.
[5]	梁栋, 杨于杰, 耿彪, 景盼盼, 苏献存, 武林锐, 渠云芳, 黄晋玲. 棉花远缘杂交新种质耐盐性鉴定及生理特性分析[J]. 华北农学报, 2024, 39(1): 95-103.
[6]	闫锋, 董扬, 赵富阳, 侯晓敏, 李清泉, 王立达, 胡继芳, 范国权, 刘凯. 植物生长调节剂对谷子生长发育的调控效应[J]. 华北农学报, 2023, 38(S1): 112-118.
[7]	邓森文, 张德朝, 方建林, 王博, 杨帆, 罗斐晖, 黎婷, 张大为. 大田条件下油菜镉积累特征[J]. 华北农学报, 2023, 38(S1): 125-130.
[8]	马晓蕾, 胡朋举, 郭颂, 刘翠丽, 张冉冉, 李玉荣, 陶佩君, 王瑾, 杨永庆. 花生耐盐碱品种的筛选及评价[J]. 华北农学报, 2023, 38(S1): 145-153.
[9]	史建硕, 潘丽佳, 蒋龙刚, 贾舟, 任燕利, 侯升林, 王丽英. 外源γ-氨基丁酸对高温胁迫下番茄幼苗生长、抗氧化酶活性及光合特性的影响[J]. 华北农学报, 2023, 38(S1): 188-193.
[10]	杨宇, 刘雨晨, 周美春, 程璧瑄, 宋建国, 罗乐, 于超. 低温沙藏下密刺蔷薇种子内含物变化研究[J]. 华北农学报, 2023, 38(S1): 253-260.
[11]	彭多姿, 黄颢源, 范占煌, 戴悦, 元妙新, 张振乾. 不同杂交类型甘蓝型油菜在重金属污染土中的适应性研究[J]. 华北农学报, 2023, 38(4): 181-189.
[12]	王亚丽, 魏琦超, 李成伟. *玉米PLATZ基因GRMZM2G006585启动子的克隆及表达分析*[J]. 华北农学报, 2023, 38(2): 21-30.
[13]	孙德慧, 王晓宇, 王莹, 霍红雁, 刘海臣, 徐惠, 张继星. *蓖麻类钙调蛋白基因RcCML42克隆与表达载体构建*[J]. 华北农学报, 2023, 38(1): 94-101.
[14]	杜强, 韩玲玲, 肖秀文, 李锦程, 沈梦宇, 王志龙, 陈秋红. 水稻DUF760基因家族的全基因组鉴定与表达谱分析[J]. 华北农学报, 2023, 38(1): 1-8.
[15]	程利华, 杨红兰, 张大伟, 马清倩, 张道远. *转ScALDH21基因棉花的耐盐性分析*[J]. 华北农学报, 2022, 37(S1): 1-7.

模态框（Modal）标题

选择文件类型/文献管理软件名称

选择包含的内容

植物耐盐机制研究进展及展望

Advances and Prospect on Mechanism of Salt Tolerance in Plants

RichHTML

PDF

可视化

被引次数

摘要/Abstract

引用本文

使用本文

参考文献 173

相关文章 15

编辑推荐

Metrics

本文评价