燕麦对碱胁迫的阳离子响应机制

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萨如拉, 刘景辉, 刘伟, 白健慧, 王占海. 燕麦对碱胁迫的阳离子响应机制. 作物学报, 2014, 40(2): 362-368[SA Ru-La, LIU Jing-Hui, LIU Wei, BAI Jian-Hui, WANG Zhan-Hai. Cation-Responsive Mechanisms of Oats to Alkali Stress. 作物学报, 2014, 40(2): 362-368] 复制到剪切板

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燕麦对碱胁迫的阳离子响应机制

萨如拉¹, 刘景辉^1,^*, 刘伟¹, 白健慧¹, 王占海²

¹内蒙古农业大学农学院, 内蒙古呼和浩特 010019

²呼伦贝尔市农业科学研究所, 内蒙古呼伦贝尔 162650

^* 通讯作者(Corresponding author): 刘景辉, E-mail:cauljh@aliyun.com, Tel: 0471-87281507

收稿日期:2013-11-24

基金:本研究由国家自然科学基金项目(31060174, 30660084), 内蒙古自然科学基金项目(2010Zd07, 200607010301), 国家现代农业产业技术体系建设专项(CARS-08-B-5)和内蒙古农业大学科技创新团队项目(NDTD2010-8)资助。

摘要

以耐碱性燕麦品种Vao-9和碱敏感性品种白燕5号为试验材料, 采用盆栽法, 用25、50、75和100 mmol L^-1碱浓度(Na₂CO₃和NaHCO₃按摩尔比1∶1混合)进行短期(14 d)和长期(28 d)胁迫处理, 观测两品种根、茎、叶中Na⁺、K⁺、Ca²⁺、Mg²⁺吸收及分配特点, 并从离子平衡吸收与分配角度, 探讨燕麦对碱胁迫的生理适应机制。胁迫处理14 d后, 燕麦体内Na⁺增加, K⁺下降, Ca²⁺和Mg²⁺变化不大, 且两品种间各器官中4种离子的分配比例差异不显著。胁迫处理28 d后, 两品种各器官中Na⁺增幅较大, K⁺、Ca²⁺和Mg²⁺降幅较大。Vao-9植株体内Na⁺、Ca²⁺含量大于白燕5号, 但K⁺、Mg²⁺含量与白燕5号无显著差异, 但两品种间各器官中4种离子的分配特点不同; 当胁迫浓度达到100 mmol L^-1时, 与白燕5号相比, Vao-9叶片中少分配5.9个百分点Na⁺, 多分配13.5个百分点K⁺、28.9个百分点Ca²⁺、10.9个百分点Mg²⁺, 茎中多分配5.4个百分点Na⁺, 少分配9.8个百分点K⁺, 根中少分配28.9个百分点Ca²⁺、10.9个百分点Mg²⁺, 因而Vao-9叶片中Na⁺ /K⁺、Na⁺/Ca²⁺、Na⁺/Mg²⁺值较白燕5号低。可见, 燕麦通过提高阳离子选择吸收及器官分配能力以适应碱胁迫。

关键词: 碱胁迫; 燕麦; 阳离子平衡

Cation-Responsive Mechanisms of Oats to Alkali Stress

SA Ru-La¹, LIU Jing-Hui^1,^*, LIU Wei¹, BAI Jian-Hui¹, WANG Zhan-Hai²

¹Agricultural College, Inner Mongolia Agricultural University, Hohhot 010019, China

²Institute of Agricultural Sciences, Hulun Buir 162650, China

第一作者联系方式: E-mail:sarula_0922@126.com, Tel: 15848101483

Abstract

In a pot experiment, an alkali-tolerant oat variety, Vao-9, and an alkali-sensitive variety, Baiyan 5, were exposed to 25, 50, 75, and 100 mmol L^-1 of alkali stress (molar ratio of Na₂CO₃:NaHCO₃ = 1:1). The contents of Na⁺, K⁺, Ca²⁺, and Mg²⁺ absorbed by oat seedling and their distribution characteristics in root, stem, and leaf were measured after a short-term (14 d) or a long-term (28 d) stress treatment. The objective was to understand the physiological adaptation to alkali stress in oat in the view of ion balance absorption and distribution. After short-term stress, the cation contents in oat plants showed the variations of increased Na⁺, decreased K⁺, and minor changes in Ca²⁺ and Mg²⁺. Besides, the distribution proportions of the four ions in various organs were not significantly different between the two varieties. Compared to short-term stress, long-term stress resulted in larger increase of Na⁺ content and larger decreases of K⁺, Ca²⁺, and Mg²⁺ contents in all organs of both varieties. Vao-9 absorbed more Na⁺and Ca²⁺ than Baiyan 5, and the absorptions of K⁺ and Mg²⁺ were not significantly different between varieties, but the distribution characteristics of the four ions in various organs were different between the two varieties. Under 100 mmol L^-1 alkali stress, 5.9 percent Na⁺ in leaf were lower distributed and 13.5 percent K⁺, 28.9 Ca²⁺, 10.9 Mg²⁺ more distributed in Vao-9 than in Baiyan 5, 5.4 percent Na⁺ in stem higher and 9.8 K⁺lower in Vao-9 than in Baiyan 5, 28.9 Ca²⁺, 10.9 Mg²⁺ in root were lower in Vao-9 than in Baiyan 5, As a result, the Na⁺ content and the ratios of Na⁺/K⁺, Na⁺/Ca²⁺, and Na⁺/Mg²⁺ in leaf were lower in Vao-9 than in Baiyan 5. Clearly, oat plant has the mechanism of selective absorption and distribution of various cations in different organs in response to alkali stress.

Keyword: Alkali stress; Oat; Cation iquilibrium

Show Figures

盐渍化土壤严重影响作物生长和产量^{[ 1]}。Na⁺是造成植物盐害及产生盐渍生境的主要离子。由于Na⁺和K⁺有相似的离子半径和水合能^{[ 2]}, 高浓度Na⁺竞争K⁺, 使植物体内K⁺高亲和力AKT1等离子通道及HKT1等转运蛋白结合位的K⁺吸收及转运受到抑制^{[ 3]}。高浓度Na⁺可置换质膜和细胞内膜上结合的Ca²⁺, 破坏膜系统完整性, 增加膜透性, 扰乱离子的选择运输能力。在盐碱胁迫下, 由于阳离子竞争效应, 植物对Mg²⁺的吸收和向地上部运输受Na⁺、K⁺、Ca²⁺抑制^{[ 4, 5]}, 而这4种离子的平衡吸收选择与分配能力与其耐盐碱性有关^{[ 6]}。

燕麦( Arena sativa)作为粮饲兼用的一年生草本植物, 具有抗寒、耐贫瘠和耐盐碱等特性, 被称为盐碱地改良的先锋作物^{[ 7]}。研究表明, 燕麦在盐碱胁迫下合成和积累有机溶质(脯氨酸和可溶性糖)以及吸收和积累无机离子(Na⁺)来进行渗透调节, 以适应盐碱胁迫, 且不同时期、不同燕麦品种间存在差异^{[ 7, 8, 9]}。盐碱胁迫下, 植物对Na⁺、K⁺、Ca²⁺、Mg²⁺吸收选择及分配能力不仅与胁迫时间和胁迫程度有关, 且与品种类型有关^{[ 10, 11, 12, 13]}。目前, 不同燕麦品种不同器官中Ca²⁺、Mg²⁺对燕麦耐碱性影响的研究报道较少。本试验选用耐碱性不同的2个燕麦品种, 通过不同浓度碱溶液处理, 观测短期和长期胁迫后不同器官中Na⁺、K⁺、Ca²⁺、Mg²⁺平衡吸收与分配规律差异, 为揭示燕麦对碱胁迫的阳离子响应机制提供依据。

1 材料与方法

1.1 试验材料与处理

耐碱性燕麦品种Vao-9由加拿大东部谷物与油料作物研究中心提供, 碱敏感性品种白燕5号由吉林省白城市农业科学院提供。

2012年4月至7月, 在内蒙古农业大学燕麦产业研究中心日光温室内进行盆栽试验。将种子播于直径22 cm、高30 cm的塑料盆内, 每盆装7.5 kg细沙, 定苗20株。每隔2 d浇一次Hogland营养液, 每次500 mL。待燕麦长至二叶一心时进行短期(14 d)和长期(28 d)胁迫处理。

按摩尔比1∶1混合2种碱性盐(Na₂CO₃、NaHCO₃), 配成25、50、75和100 mmol L^-1浓度碱液, 分别加入Hogland营养液(共500 mL), 作为胁迫处理液; 对照只加Hogland营养液500 mL。3次重复。为避免盐激反应, 各处理每天递增预定浓度的1/4, 每隔2 d加一次碱溶液, 使各处理同一天达到预定浓度, 此时作为胁迫第1天, 分别于处理后第14天和第28天取样。

1.2 阳离子测定方法

取出每盆中所有植株, 用自来水冲洗干净, 再用蒸馏水冲洗, 并用吸水纸吸去表面附着的水分。将根、茎、叶分开, 105℃杀青后80℃烘干至恒量。样品磨碎后准确称取干样0.3000 g, 放入50 mL的塑料管中, 加1 mol L^-1盐酸25.0 mL, 盖紧管口, 振荡2 h后用定量滤纸过滤。用火焰光度计测定K⁺(直接用滤液)和Na⁺(滤液稀释5倍后)含量。

取0.5000 g干样放入150 mL三角瓶, 加HNO₃-HClO₄ (4∶1)溶液20 mL, 放通风厨里在电热板上加热至140~150℃, 消煮过夜, 至冒白烟为止。把消煮液倒在 25 mL的比色管里定容, 在原子吸收光谱仪上测Ca²⁺和Mg²⁺含量。

1.3 数据分析

采用Microsoft Excel绘制柱形图和SPSS 17. 0软件进行试验数据的方差分析(one-way ANOVA)。

2 结果与分析

2.1 碱胁迫对燕麦不同器官中阳离子吸收及分配的影响

两品种的Na⁺含量均随碱浓度的升高而增加, 且处理间有极显著差异( P<0.01)。短期胁迫后, Vao-9 Na⁺总量大于白燕5号, 但Na⁺在各器官的分布品种间无显著差异(图1-A1)。长期胁迫后, 两品种Na⁺总量差异不大, 分别为67.7 mg g^-1 DW和65.6 mg g^-1 DW, 但Na⁺在不同器官中的分布品种间差异明显, Vao-9主要在茎中积累, 而白燕5号在叶片中有大量积累(图1-B1)。在100 mmol L^-1长期胁迫下, Vao-9叶片中Na⁺分配百分比较白燕5号低5.9个百分点, 而根和茎中的分配百分比分别较白燕5号高0.5和5.4个百分点(图1-C1)。

K⁺主要分布在茎和叶中, 根中含量很低。随碱胁迫浓度增加和时间的推移, K⁺含量在各器官中均呈下降趋势, 且两品种表现一致(图1-A2, B2)。各器官中不同碱浓度处理的K⁺含量有极显著差异( P<0.01)。在短期胁迫下, Vao-9吸收的K⁺总量小于白燕5号, 但两品种不同器官中分布差异不显著(图1-A2); 长期胁迫下, Vao-9的K⁺总量为118.1 mg g^-1 DW, 白燕5号为118.2 mg g^-1 DW, 但K⁺在茎、叶中的分布两品种间不同, 尤其是75 mmol L^-1和100 mmol L^-1处理(图1-B2), 其中100 mmol L^-1处理叶片中K⁺分配百分比, Vao-9较白燕5号高13.5个百分点, 而茎中则少9.8个百分点(图1-C2)。

短期胁迫下, 不同处理Vao-9各器官的Ca²⁺含量始终大于白燕5号, 且处理间变化幅度较小(图1-A3)。与短期胁迫相比, 长期胁迫使两品种根和叶中的Ca²⁺骤增, 而茎中减少。在长期胁迫下, 随碱浓度增加, 两品种茎、叶中Ca²⁺含量降低; 而根中Ca²⁺含量, Vao-9中先降低再升高, 白燕5号表现持续增加, 且各处理间差异显著(图1-B3)。Vao-9的Ca²⁺总量(36.5 mg g^-1 DW)大于白燕5号(29.6 mg g^-1 DW)。4种浓度处理的叶片Ca²⁺含量均低于对照, 但Vao-9的下降百分率(10.3%~22.2%)明显低于白燕5号(42.8%~79.6%)。100 mmol L^-1长期胁迫下, Ca²⁺在茎、叶片中分配比例, Vao-9分别比白燕5号高1.5和28.9个百分点, 而根中比例则低30.4个百分点(图1-C3)。

短期胁迫下, 除Vao-9叶片中Mg²⁺含量随碱浓度的增加表现先上升后下降外, 其他均呈逐渐降低趋势, 但Vao-9的降幅小于白燕5号(图1-A4)。长期胁迫下, 除白燕5号根中Mg²⁺含量升高外, 其他均呈降低趋势; 两品种的Mg²⁺总量无显著差异, 但Mg²⁺在不同器官中的分布特点不同。两品种叶片中Mg²⁺含量随碱浓度的升高而下降, Vao-9较对照下降0~20.6%, 低于白燕5号的11.7%~36.7% (图1-B4)。与Ca²⁺相似, 100 mmol L^-1长期胁迫下, Vao-9的茎和叶中比白燕5号分配了较高比例的Mg²⁺, 而根中Mg²⁺分配比例则低于白燕5号(图1-C4)。

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图1 不同浓度碱胁迫下燕麦根、茎、叶中4种阳离子含量与分配百分率短期(14 d)和长期(28 d)胁迫下, 各处理的碱浓度依次为0 (CK)、25 (T1)、50 (T2)、75 (T3)和100 mmol L^-1 (T4)。Fig. 1 Contents and distribution ratios of four cations in root, stem, and leaf of oat under different concentrations of alkali stressAlkali concentrations in different treatments were 0 (CK), 25 (T1), 50 (T2), 75 (T3), and 100 mmol L^-1 (T4) under both short-term (14 d) and long-term (28 d) stress.

2.2 碱胁迫对燕麦不同器官中离子含量比值的影响

两品种的Na⁺/K⁺值随着胁迫浓度的增加而增加, 在根中最大, 这主要是根中K⁺含量低所致。短期胁迫下, Vao-9茎、叶Na⁺/K⁺值大于白燕5号, 根Na⁺/K⁺值在胁迫浓度达到50 mmol L^-1时大于白燕5号; 长期胁迫下, Vao-9叶片Na⁺/K⁺值较白燕5号小, 且白燕5号各处理间均达极显著差异, 但根和茎的Na⁺/K⁺值较白燕5号大, 其中100 mmol L^-1处理的根、茎、叶Na⁺/K⁺值, Vao-9分别为白燕5号的144.4%、140.8%和51.2% (表1)。

Na⁺/Ca²⁺值表现为茎>叶>根; 两品种茎和叶的Na⁺/Ca²⁺值呈现随着胁迫浓度增加而增加的趋势, 且白燕5号处理间存在极显著差异, 但短期胁迫下75 mmol L^-1处理的茎Na⁺/Ca²⁺值高于100 mmol L^-1处理, Vao-9叶Na⁺/Ca²⁺值在浓度达50 mmol L^-1时基本稳定。无论短期还是长期胁迫, 白燕5号的茎、叶Na⁺/Ca²⁺值均高于Vao-9, 长期胁迫下的差异更为明显, 尤其是长期胁迫的100 mmol L^-1处理, Vao-9的茎、叶Na⁺/Ca²⁺值分别只有白燕5号的60.4%和17.7% (表1)。

表1 短期和长期碱胁迫对燕麦不同器官中Na⁺/K⁺、Na⁺/Ca²⁺和Na⁺/Mg²⁺值的影响 Table 1 Effects of short-term and long-term alkali stresses on Na⁺/K⁺, Na⁺/Ca²⁺, and Na⁺/Mg²⁺ ratios in different organs of oat

离子比类型 Type of ion ratio	处理 Treatment	白燕5号Baiyan 5			Vao-9
离子比类型 Type of ion ratio	处理 Treatment	根 Root	茎 Stem	叶 Leaf	根 Root	茎 Stem	叶 Leaf
短期碱胁迫 Short-term alkali stress
Na⁺/K⁺	CK	0.32 D	0.11 D	0.16 B	0.28 D	0.13 D	0.14 C
	T1	1.11 A	0.35 C	0.22 AB	0.69 C	0.45 C	0.34 B
	T2	0.77 B	0.44 BC	0.30 AB	0.88 B	0.57 B	0.39 AB
	T3	0.69 C	0.52 AB	0.36 A	1.19 A	0.61 B	0.45 A
	T4	1.14 A	0.60 A	0.40 A	1.20 A	0.71 A	0.46 A
Na⁺/Ca²⁺	CK	1.01 D	2.22 E	2.18 E	0.69 C	1.49 E	0.89 C
	T1	2.19 B	5.85 D	3.18 D	1.81 B	4.03 D	1.60 B
	T2	2.08 C	8.31 C	4.06 C	2.11 A	4.74 C	2.46 A
	T3	2.32 A	11.16 A	5.31 B	1.86 B	7.54 A	2.17 A
	T4	2.03 C	8.71 B	7.06 A	1.85 B	6.81 B	2.43 A
Na⁺/Mg²⁺	CK	16.73 D	29.72 E	36.14 E	17.17 E	32.38 E	30.81 E
	T1	32.19 C	84.73 D	52.59 D	36.15 D	83.37 D	58.41 D
	T2	32.54 C	131.70 C	74.83 C	47.41 B	124.17 C	72.70 B
	T3	37.50 B	153.24 B	86.42 B	53.92 A	129.66 B	70.14 C
	T4	42.00 A	165.91 A	91.07 A	45.53 C	133.48 A	78.53 A
长期碱胁迫 Long-term alkali stress
Na⁺/K⁺	CK	0.54 E	0.15 E	0.12 E	0.44 D	0.19 D	0.16 C
	T1	0.80 D	0.40 D	0.39 D	0.90 C	0.45 C	0.34 B
	T2	0.87 C	0.54 C	0.56 C	0.94 C	0.80 B	0.68 A
	T3	1.11 B	0.65 B	0.94 B	1.26 B	0.87 AB	0.65 A
	T4	1.17 A	0.71 A	1.27 A	1.69 A	1.00 A	0.65 A
Na⁺/Ca²⁺	CK	0.60 AB	1.40 E	0.43 E	0.50 D	1.65 E	0.53 D
	T1	0.63 AB	16.54 C	2.03 D	0.89 A	7.49 D	1.18 C
	T2	0.70 A	15.48 D	4.55 C	0.66 BC	13.02 C	1.92 B
	T3	0.57 B	20.12 B	5.37 B	0.59 CD	31.20 A	2.09 A
	T4	0.62 AB	39.18 A	12.07 A	0.74 B	23.68 B	2.14 A
Na⁺/Mg²⁺	CK	29.43 C	48.49 E	26.99 E	22.70 D	55.33 E	32.73 E
	T1	29.36 C	118.54 D	82.56 D	40.18 C	118.89 D	65.23 D
	T2	33.24 B	145.91 C	117.61 C	40.93 C	194.75 C	111.37 C
	T3	33.30 B	176.63 B	183.82 B	44.46 B	204.18 B	119.71 B
	T4	38.31 A	217.20 A	243.11 A	55.77 A	238.66 A	129.20 A

短期(14 d)和长期(28 d)胁迫下, 各处理的碱浓度依次为0 (CK)、25 (T1)、50 (T2)、75 (T3)和100 mmol L^-1 (T4)。数据后不同字母表示处理间达极显著差异( P< 0.01)。

Alkali concentrations in different treatments were 0 (CK), 25 (T1), 50 (T2), 75 (T3), and 100 mmol L^-1 (T4) under both short-term (14 d) and long-term (28 d) stress. Values followed by different letters are significantly different among treatments at P< 0.01.

Na⁺/Mg²⁺值也表现为茎>叶>根。随着胁迫浓度提高, Na⁺/Mg²⁺值在各器官中也呈增加趋势, 但短期胁迫下, Vao-9根Na⁺/Mg²⁺值大于白燕5号, 茎、叶Na⁺/Mg²⁺值小于白燕5号。长期胁迫下两品种的茎、叶Na⁺/Mg²⁺值均大于短期胁迫的Na⁺/Mg²⁺值。长期胁迫下, Vao-9的叶Na⁺/Mg²⁺值远小于白燕5号, 且处理间差异极显著, 而根、茎Na⁺/Mg²⁺值则表现为Vao-9大于白燕5号; 当100 mmol L^-1长期胁迫时, Vao-9根、茎、叶中Na⁺/Mg²⁺值分别为白燕5号的145.6%、109.9%和53.1% (表1)。

3 讨论

渗透调节在作物耐盐碱机制中有重要作用^{[ 14, 15, 16]}。参与渗透调节的阳离子主要是Na⁺、K⁺和Ca²⁺, 其在植物器官中的分配比例因物种、品种和生育时期而异^{[ 17, 18, 19]}。通过调节阳离子在不同器官、组织或细胞内的区域(隔)化的分布, 有助于减轻离子毒害^{[ 20]}。本研究中, 长期和短期胁迫后, 两品种的Na⁺含量均显著增加, 且向茎、叶中大量积累; 两种胁迫强度下, Vao-9的Na⁺总量总是大于白燕5号, 长期胁迫下, Na⁺在Vao-9茎中的积累及分配较高, 而在叶片中的积累相对较低。这一方面是因为Vao-9通过地上部积累大量离子, 增大地上部和根部的渗透势差, 促进水分从根部向地上部运输, 改善地上部的水分状况, 促进生长^{[ 21, 22]}; 另一方面是通过把吸收的大部分Na⁺分配在茎部, 从而减少叶片受伤害^{[ 23, 24]}。

盐碱土壤中高浓度Na⁺经常阻止高亲和力K⁺转运蛋白导致作物体内Na⁺内流增加、K⁺内流减少, 使植物细胞膜去极化而减少净K⁺的吸收, 因此盐渍环境中植物叶片中K⁺减少、Na⁺/K⁺值升高。在胁迫下植物细胞质保持高K⁺/Na⁺的能力是植物适应盐碱胁迫的决定因子之一。本试验中, 长期胁迫下, 两品种吸收的K⁺总量比短期胁迫的降幅, Vao-9小于白燕5号, 且长期胁迫下两品种不同器官中K⁺分配差异显著, Vao-9叶片中积累及分配较高K⁺含量, 尤其是75 mmol L^-1处理下叶片中积累K⁺能力更强, 使叶片中保持较低Na⁺/K⁺值。在长期胁迫下, 虽然与Vao-9吸收的K⁺总量无差异, 但白燕5号积累的大部分K⁺分配在茎部, 使胁迫下叶片中Na⁺/K⁺值较高。这与赵旭等^{[ 25]}在小麦耐盐性研究中的结果一致。

Ca²⁺不仅对于保持植物细胞膜的完整性和稳定性有重要作用, 而且与植物的耐盐碱能力有重要关联^{[ 26]}, 由于胁迫下高活性Na⁺置换质膜上的Ca²⁺, 细胞中Ca²⁺浓度降低, 导致质膜上Ca²⁺失去平衡, 失去保护细胞的作用, 使细胞膜的稳定性和选择性下降而破坏作物体内离子平衡。本研究中, 碱胁迫降低了两品种的Ca²⁺含量, 但随着胁迫时间的延长, Ca²⁺总量增加, 这与范远等^{[ 11]}在白燕2号上试验结果不一致, 可能不同基因型品种间存在差异导致适应胁迫的机制有所不同。碱胁迫条件下, Vao-9积累的Ca²⁺总量大于白燕5号, 且长期胁迫各处理下Vao-9根、叶中积累及分配较多Ca²⁺, 而白燕5号叶片中分配的Ca²⁺远少于根中, 使胁迫下Vao-9可以保持叶片中较低的Na⁺/Ca²⁺值。这有利于降低叶片细胞膜透性, 保持叶片中离子平衡和细胞正常代谢。同时与长期胁迫相比, 短期胁迫的各处理中, 两品种Ca²⁺含量始终保持较低水平和较小变幅, 说明碱胁迫下燕麦吸收及分配的Ca²⁺除基因型差异外还受胁迫时间的影响。

Mg²⁺是叶绿素分子的重要组成部分, 而盐碱胁迫下作物Na⁺增加Ca²⁺降低进一步限制了K⁺吸收, Ca²⁺吸收的降低和Na⁺的大量积累进一步限制Mg²⁺吸收, 由于K⁺与Mg²⁺间存在拮抗作用, 且主要发生在由根系向地上部的运输过程中^{[ 4, 5]}, 所以在盐碱胁迫下作物叶片中保持较低Na⁺/Mg²⁺值是衡量植物耐盐碱的重要指标之一^{[ 27, 28]}。本研究中, 碱胁迫降低燕麦Mg²⁺含量, 且随着胁迫时间的延长而减少, 但Vao-9中降幅小于白燕5号。长期胁迫下两品种各器官中分布差异显著, Vao-9在叶中积累较多Mg²⁺, 而白燕5号则在根中积累较多Mg²⁺, 因而Vao-9叶片中保持较低的Na⁺/Mg²⁺值。这证实了不同刺槐品种存在Mg²⁺吸收选择能力差异的结果^{[ 29]}。在棉花、碱地肤上的研究表明, 在盐、碱胁迫下, 作物对Na⁺的区域化分配, K⁺、Ca²⁺、Mg²⁺的吸收, 与叶片细胞质膜H⁺-ATPase、液泡膜H⁺-ATPase、H⁺-PPase和Na⁺/H⁺逆向运输蛋白等活性有关^{[ 30, 31]}。燕麦的耐碱机制是否与上述研究结果一致, 需进一步探讨。

The authors have declared that no competing interests exist.

作者已声明无竞争性利益关系。The authors have declared that no competing interests exist.

参考文献

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1	Evelin H, Kapoor R, Giri B. Arbuscular mycorrhizal fungi in alleviation of salt stress: a review. Ann Bot, 2009, 104: 1263-1281 [本文引用:1] [JCR: 0.657]
2	Parida A K, Das A B, Mittra. Effects of salt on growth, ion accumulation photosynthesis and leaf anatomy of the mangrove, Bruguiera parviflora. Trees-Struct Func, 2004, 18: 167-174 [本文引用:1] [JCR: 1.925]
3	Borsani O, Valpuesta V, Botella M A. Developing salt tolerant plants in a newcentury: a molecular biologyapproach. Plant Cell, Tissue Organ Cult, 2003, 73: 101-115 [本文引用:1] [JCR: 3.633]
4	Lauchli A. Calcium, salinity and the plasma membrane. In: Leonard R T, Hepler P K, eds. Calcium in plant growth and development. Rockville, Mary-land , USA: American Society of Plant Physiologists, 1990. pp26-35 [本文引用:2]
5	李娟. 植物钾、钙、镁素营养的研究进展. 福建稻麦科技, 2007, 25(1): 39-42 Li J. Research progress in potassium, calcium, magnesium nutrition of plant. Fujian Sci Tech Rice Wheat, 2007, 25(1): 39-42 (in Chinese) [本文引用:2] [CJCR: 0.3109]
6	鄂志国, 张丽靖. 水稻盐胁迫应答的分子机制. 杂交水稻, 2010, 25(2): 1-5 E Z G, Zhang L J. Molecular mechanism of rice responses to salt stress. Hybrid Rice, 2010, 25(2): 1-5 (in Chinese with English abstract) [本文引用:1] [CJCR: 0.605]
7	范远, 任长忠, 李品芳, 任图生. 盐碱胁迫下燕麦生长及阳离子吸收特征. 应用生态学报, 2011, 22: 2875-2882 Fan Y, Ren C Z, Li P F, Ren T S. Oat growth and cation absorption characteristics under salt and alkali stress. Chin J Appl Ecol, 2011, 22: 2875-2882 (in Chinese with English abstract) [本文引用:2] [CJCR: 1.742]
8	王波, 宋凤斌. 燕麦对盐碱胁迫的反应和适应性. 生态环境, 2006, 15: 625-629 Wang B, Song F B. Physiological responses and adaptive capacity of oats to saline-alkali stress. Ecol Environ, 2006, 15: 625-629 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.2453]
9	毛明艳, 方正. NaHCO₃胁迫对燕麦幼苗生长及相关生理指标的影响. 安徽农业科学, 2009, 37: 4468-4470 Mao M Y, Fang Z. Effects of NaHCO₃ stress on the growth and some physiological indexes in of oat seedlings. J Anhui Agric Sci, 2009, 37: 4468-4470 (in Chinese with English abstract) [本文引用:1] [CJCR: 0.687]
10	Mühling K H, Lauchli A. Effect of salt stress on growth and cation compartmentation in leaves of two plant species differing in salt tolerance. J Plant Physiol, 2002, 59: 137-146 [本文引用:1] [JCR: 2.699]
11	商学芳, 董树亭, 郑世英, 王丽燕. 玉米种子萌发过程中Na^{+、K^{+和Ca²+含量变化与耐盐性关系}}. 作物学报, 2008, 34: 333-336 Shang X F, Dong S T, Zheng S Y, Wang L Y. Relationship between changes of Na^{+, K^{+, and Ca²+ contents during seed germination and salt tolerance in maize}}. Acta Agron Sin, 2008, 34: 333-336 (in Chinese with English abstract) [本文引用:2] [CJCR: 1.667]
12	王学征, 李秋红, 吴凤芝. NaCl胁迫下栽培型番茄Na^{+、K^{+吸收、分配和转运特性}}. 中国农业科学, 2010, 43: 1423-1432 Wang X Z, Li Q H, Wu F Z. Study on the Characteristics of absorption, distribution and selective transport of Na^{+ and K^{+ in tomato plants under salt stress}}. Sci Agric Sin, 2010, 43: 1423-1432 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.889]
13	任志彬, 王志刚, 聂庆娟, 黄大庄, 马向超. 盐胁迫对锦带花幼苗生长及不同部位 Na^{+、K^{+、Ca²+、Mg²+离子质量分数的影响}}. 东北林业大学学报, 2011, 39(5): 24-27 Ren Z B, Wang Z G, Nie Q J, Huang D Z, Ma X C. Effect of salt stress on growth and ion contents in Weigela florida cuttings. J North For Univ, 2011, 39(5): 24-27 (in Chinese with English abstract) [本文引用:1]
14	Talukder N M, Dutta R K. Salinity effect on mineral nutrient distribution along roots and shoots of rice (Oryza sativa L. ) genotypes differing in salt tolerance. Arch Agron Soil Sci, 2011, 57: 33-45 [本文引用:1]
15	王晓冬, 王成, 马智宏, 侯瑞锋, 高权, 陈泉. 短期NaCl胁迫对不同小麦品种幼苗K^{+吸收和Na^{+、K^{+积累的影响}}}. 生态学报, 2011, 31: 2822-2830 Wang X D, Wang C, Ma Z H, Hou R F, Gao Q, Chen Q. Effect of short term salt stress on the absorption of K^{+ and accumulation of Na^{+, K^{+ in seedlings of different wheat varieties}}}. Acta Ecol Sin, 2011, 31: 2822-2830 (in Chinese with English abstract) [本文引用:1]
16	於丙军, 罗庆云, 刘友良. 盐胁迫对盐生野大豆生长和离子分布的影响. 作物学报, 2001, 27: 776-780 Yu B J, Luo Q Y, Liu Y L. Effects of salt stress on growth and ionic distribution of salt-bornGlycine soja. Acta Agron Sin, 2001, 27: 776-780 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.667]
17	马建华, 郑海雷, 赵中秋, 张春光. 植物抗盐机理研究进展. 生命科学研究, 2001, 5: 175-179 Ma J H, Zheng H L, Zhao Z Q, Zhang C G. Progress in mechanisms of plant resistance to salt stress. Life Sci Res, 2001, 5: 175-179 (in Chinese with English abstract) [本文引用:1] [CJCR: 0.59]
18	郭伟, 王庆祥, 于立河. 盐碱混合胁迫对小麦幼苗阳离子吸收和分配的影响. 麦类作物学报, 2011, 31: 735-740 Guo W, Wang Q X, Yu L H. Effects of salinity alkalinity stress on cation absorption and distribution in wheat seedlings. J Triticeae Crops, 2011, 31: 735-740 (in Chinese with English abstract) [本文引用:1]
19	Hasegawa P M, Bressan R A, Zhu J K, Bohnert Paulm H J. Plant cellular and molecular responses to high salinity. Plant Physiol Plant Mol Biol, 2000, 51: 463-499 [本文引用:1] [JCR: 25.962]
20	辛承松, 董合忠, 唐薇, 温四民. 棉花盐害与耐盐性的生理和分子机理研究进展. 棉花学报, 2005, 17: 309-313 Xin C S, Dong H Z, Tang W, Wen S M. Physiological and molecular mechanisms of salt injury and salt tolerance in cotton. Cotton Sci, 2005, 17: 309-313 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.462]
21	李品芳, 白文波, 杨志成. NaCl胁迫对苇状羊茅离子吸收与运输及其生长的影响. 中国农业科学, 2005, 38: 1458-1565 Li P F, Bai W B, Yang Z C. Effects of NaCl stress on ions absorption and transportation and plant growth of tall fescue. Sci Agric Sin, 2005, 38: 1458-1565 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.889]
22	张海燕. 盐胁迫下盐地碱蓬体内无机离子含量分布特点的研究. 西北植物学报, 2002, 22: 129-135 Zhang H Y. A study on the characters of content of inorganic ions in salt-stressed Suaeda salsa. Acta Bot Boreali-Occident Sin, 2002, 22: 129-135 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.321]
23	赵学良, 张彦才. NaCl胁迫对棉花苗期营养元素吸收与含量的影响. 河北农业大学学报, 1992, 15(2): 41-44 Zhao X L, Zhang Y C. Influence of NaCl stress on the uptake and concentrations of nutriments in cotton Seedlings. J Agric Univ Hebei, 1992, 15(2): 41-44 (in Chinese with English abstract) [本文引用:1] [CJCR: 0.669]
24	孙小芳, 刘友良. NaCl胁迫下棉花体内Na^{+、K^{+分布与耐盐性}}. 西北植物学报, 2000, 20: 1027-1033 Sun X F, Liu Y L. Distribution of Na^{+ and K^{+ in cotton plant under NaCl stress and salt tolerance}}. Acta Bota Boreali-Occident Sin, 2000, 20: 1027-1033 (in Chinese with English abstract) [本文引用:1]
25	赵旭, 王林权, 周春菊, 尚浩博. 盐胁迫对四种基因型冬小麦幼苗Na^{+、K^{+吸收和累积的影响}}. 生态学报, 2007, 27: 205-213 Zhao X, Wang L Q, Zhou C J, Shang H B. Effects of salt stress on the absorption and accumulation of Na^{+ and K^{+ in seedlings of four winter wheat (Tritium aestivum) genotypes}}. Acta Ecol Sin, 2007, 27: 205-213 (in Chinese with English abstract) [本文引用:1]
26	Mittler R. Oxidative stress, antioxidants and stress tolerance. Trends Plant Sci, 2002, 7: 405-410 [本文引用:1] [JCR: 11.808]
27	杨帆, 丁菲, 杜天真. 盐胁迫下构树幼苗各器官中K^{+、Ca²+、Na^{+和Cl^{-含量分布及吸收特征}}}. 应用生态学报, 2009, 20: 767-772 Yang F, Ding F, Du T Z. Absorption and allocation characteristics of K^{+, Ca²+, Na^{+ and Cl^{- in different organs of Broussonetia papyrifera seedlings under NaCl stress}}}. Chin J Appl Ecol, 2009, 20: 767-772 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.742]
28	Parida A K, Das A B. Salt tolerance and salinity effects on plants: a review. Ecol Toxicol Environ Safety, 2005, 60: 324-349 [本文引用:1] [JCR: 2.203]
29	郁万文, 曹帮华, 吴丽云. 盐胁迫下刺槐无性系生长和矿质营养平衡研究. 西北植物学报, 2005, 25: 2097-2102 Yu W W, Cao B H, Wu L Y. Growths and mineral nutrient balance of black locust clones under salt stress. Acta Bot Boreali- Occident Sin, 2005, 25: 2097-2102 (in Chinese with English abstract) [本文引用:1]
30	Yang C W, Chong J N, Li C Y, Kim C M, Shi D C, Wang D L. Osmotic adjustment and ion balance traits of an alkali resistant halophyte Kochia sieversiana during adaptation to salt and alkali conditions. Plant Soil, 2007, 294: 263-276 [本文引用:1] [JCR: 2.638]
31	郑青松, 王仁雷, 刘友良. 钙对盐胁迫下棉花离子吸收分配的影响. 植物生理学报, 2001, 27: 325-330 Zheng Q S, Wang R L, Liu Y L. Effects of Ca²+ on absorption and distribution of ions in salt-treated cotton seedlings. Acta Phytol Sin, 2001, 27: 325-330 (in Chinese with English abstract) [本文引用:1]

2009

0.657

0.0

Ann. Bot.. 2009, 104(7):1263-80 DOI:10.1093/aob/mcp251

Arbuscular mycorrhizal fungi in alleviation of salt stress: a review.

Evelin H, Kapoor R, Giri B

Applied Mycology Laboratory, Department of Botany, University of Delhi, Delhi-110 007, India.

Salt stress has become a major threat to plant growth and productivity. Arbuscular mycorrhizal fungi colonize plant root systems and modulate plant growth in various ways.

PMID: 19815570

... 盐渍化土壤严重影响作物生长和产量^[1] ...

2004

1.925

0.0

Trees-struct Funct. 2004, 18(2):167 - 174 DOI:10.1007/s00468-003-0293-8

Effects of salt on growth, ion accumulation, photosynthesis and leaf anatomy of the mangrove,Bruguiera parviflora

AsishKumar Parida(1) A.B. Das(12) B. Mittra(3)

1.National Institute for Plant Biodiversity Conservation and Research Nayapalli 751015 Bhubaneswar Orissa India
2.Regional Plant Resource Centre 751015 Bhubaneswar Orissa India
3.Centre for Environmental Management of Degraded Ecosystems, School of Environmental Studies University of Delhi North Campus 110 007 Delhi India

The effects of a range of salinity (0, 100, 200 and 400 mM NaCl) on growth, ion accumulation, photosynthesis and anatomical changes of leaves were studied in the mangrove, Bruguiera parviflora of the family Rhizophoraceae under hydroponically cultured conditions. The growth rates measured in terms of plant height, fresh and dry weight and leaf area were maximal in culture treated with 100 mM NaCl and decreased at higher concentrations. A significant increase of Na⁺ content of leaves from 46.01 mmol m^-2in the absence of NaCl to 140.55 mmol m^-2in plants treated with 400 mM NaCl was recorded. The corresponding Cl^- contents were 26.92 mmol m^-2and 97.89 mmol m^-2. There was no significant alteration of the endogenous level of K⁺ and Fe²⁺ in leaves. A drop of Ca²⁺and Mg²⁺content of leaves upon salt accumulation suggests increasing membrane stability and decreased chlorophyll content respectively. Total chlorophyll content decreased from 83.44 mgr

g cm^-2 in untreated plants to 46.56 mgr

g cm^-2 in plants treated with 400 mM NaCl, suggesting that NaCl has a limiting effect on photochemistry that ultimately affects photosynthesis by inhibiting chlorophyll synthesis (ca. 50% loss in chlorophyll). Light-saturated rates of photosynthesis decreased by 22% in plants treated with 400 mM NaCl compared with untreated plants. Both mesophyll and stomatal conductance by CO₂diffusion decreased linearly in leaves with increasing salt concentration. Stomatal and mesophyll conductance decreased by 49% and 52% respectively after 45 days in 400 mM NaCl compared with conductance in the absence of NaCl. Scanning electron microscope study revealed a decreased stomatal pore area (63%) in plants treated with 400 mM NaCl compared with untreated plants, which might be responsible for decreased stomatal conductance. Epidermal and mesophyll thickness and intercellular spaces decreased significantly in leaves after treatment with 400 mM NaCl compared with untreated leaves. These changes in mesophyll anatomy might have accounted for the decreased mesophyll conductance. We conclude that high salinity reduces photosynthesis in leaves of B. parviflora, primarily by reducing diffusion of CO₂to the chloroplast, both by stomatal closure and by changes in mesophyll structure, which decreased the conductance to CO₂within the leaf, as well as by affecting the photochemistry of the leaves.

... 由于Na⁺和K⁺有相似的离子半径和水合能^[2], 高浓度Na⁺竞争K⁺, 使植物体内K⁺高亲和力AKT1等离子通道及HKT1等转运蛋白结合位的K⁺吸收及转运受到抑制^[3] ...

2003

3.633

0.0

Plant Cell Tiss Org. 2003, 73(2):101 - 115 DOI:10.1023/A:1022849200433

Developing salt tolerant plants in a new century: a molecular biology approach

Borsani

,Valpuesta

,Botella

M A

.Developing salt tolerant plants in a newcentury: a molecular biologyapproach. Plant Cell,Tissue Organ Cult,2003,73:101-115

1.Facultad de Agronomía
2.Universidad de Málaga Campus de Teatinos

Soil salinity is a major abiotic stress in plant agriculture strongly, influencing plant productivity world-wide. Classical breeding for salt tolerance in crop plants has been attempted to improve field performance without success. Therefore, an alternative strategy is to generate salt tolerant plants through genetic engineering. Several species and experimental approaches have been used in order to identify those genes that are important for salt tolerance. Due to high level of salt tolerance, halophytes are good candidates to identify salt tolerance genes. However, other species such as yeast and glycophytes have also been employed. Three approaches are commonly used to identify genes important for salt tolerance. The first approach is to identify genes involved in processes known to be critical for salt tolerance (osmolyte synthesis, ion homeostasis, etc.). The second approach is to identify genes whose expression is regulated by salt stress. This is relatively simply and applicable to any plant species. Genetic amenability of some species allows the third approach, which consists in the identification of salt tolerance determinants based on functionality. At the moment, there is a large number of reports in the literature claiming that plants with increased salt tolerance have been obtained. The main problem is that different plant species, stage of development, organs, promoters and salt conditions used it is difficult to compare the degree of salt tolerance conferred by different genes. In this review, we discuss progress made towards understanding the molecular elements involved in salt stress responses that have been used in transgenic approaches to improve salt tolerance.

1990

0.0

... 在盐碱胁迫下, 由于阳离子竞争效应, 植物对Mg²⁺的吸收和向地上部运输受Na⁺、K⁺、Ca²⁺抑制^[4,5], 而这4种离子的平衡吸收选择与分配能力与其耐盐碱性有关^[6] ...

... Mg²⁺是叶绿素分子的重要组成部分, 而盐碱胁迫下作物Na⁺增加Ca²⁺降低进一步限制了K⁺吸收, Ca²⁺吸收的降低和Na⁺的大量积累进一步限制Mg²⁺吸收, 由于K⁺与Mg²⁺间存在拮抗作用, 且主要发生在由根系向地上部的运输过程中^[4,5], 所以在盐碱胁迫下作物叶片中保持较低Na⁺/Mg²⁺值是衡量植物耐盐碱的重要指标之一^[27,28] ...

2007

0.0

0.3109

2010

0.0

0.605

2011

0.0

1.742

Chin J Appl Ecol. 2011, 22(11):2875 - 2882

Oat growth and cation absorption characteristics under salt and alkali stress.

盐碱胁迫下燕麦生长及阳离子吸收特征

FAN Yuan1, REN Chang-zhong2, LI Pin-fang1, REN Tu-sheng1

范远1,任长忠2,李品芳1,任图生1**

1College of Resources and Environmental Sciences, China Agricultural University, Beijing 100193, China;2Baicheng Academy of Agricultural Science, Baicheng 137000, Jilin, China

This paper monitored the oat growth and cation absorption characteristics on a saline-alkali soil in the Baicheng region of Jilin Province under low, medium, and high levels of salt stress. No significant differences were observed in the shoot growth and yield components under the three levels of salt stress, but the root biomass and root/shoot ratio decreased significantly with increasing salt stress level. At maturing stage, the root/shoot ratio under medium and high salt stresses was 77.2% and 64.5% of that under low salt stress, respectively. Under the three levels of salt stress, the K⁺/Na⁺ and Ca²⁺/Na⁺ ratios in oat plant had significant differences at trefoil stage, but no significant differences at heading stage. With the increase of salt stress level, the cation absorption selectivity coefficient of oat at filling stage decreased significantly, but the transportation selectivity coefficient had no significant difference under the three levels of stress. It was concluded that oat could adapt to the salt and alkali stress of soda-alkaline soil to some extent, and the adaptation capability decreased with the increasing level of stress. The decrease of oat root biomass and the stronger ion selective absorption capacity at heading stage under salt and alkali stress could benefit the shoot growth and yield components of oat.

在吉林白城地区盐碱土上监测了3个盐分胁迫水平下燕麦的生长动态及对土壤阳离子的吸收特征.结果表明：在低度、中度和高度3个胁迫水平下燕麦地上部生长和产量构成因素没有显著差异，但根部干质量和根冠比随盐碱胁迫程度的加重而显著降低，在成熟期，中度和高度胁迫下的根冠比分别是低度胁迫下的77.2%和64.5%.3个胁迫水平下燕麦的K⁺/Na⁺和Ca²⁺/Na⁺在三叶期差异最大，而在抽穗期无显著差异.随着盐分胁迫程度的加重，燕麦的离子吸收选择性系数在灌浆期显著降低，而运输选择性系数无显著变化.因此，燕麦能够在一定程度上适应苏打碱土的盐碱胁迫，适应能力随着胁迫水平的增加而降低；燕麦根部生物量的减少和抽穗期较强的离子选择吸收能力可能有利于地上部的生长和产量构成.

... 燕麦(Arena sativa)作为粮饲兼用的一年生草本植物, 具有抗寒、耐贫瘠和耐盐碱等特性, 被称为盐碱地改良的先锋作物^[7] ...

... 研究表明, 燕麦在盐碱胁迫下合成和积累有机溶质(脯氨酸和可溶性糖)以及吸收和积累无机离子(Na⁺)来进行渗透调节, 以适应盐碱胁迫, 且不同时期、不同燕麦品种间存在差异^[7,8,9] ...

2006

0.0

1.2453

2009

0.0

0.687

2002

2.699

0.0

... 盐碱胁迫下, 植物对Na⁺、K⁺、Ca²⁺、Mg²⁺吸收选择及分配能力不仅与胁迫时间和胁迫程度有关, 且与品种类型有关^{[10,11,12,13]} ...

2008

0.0

1.667

Acta Agron Sin. 2008, 34(2):333 - 336 DOI:10.3724/SP.J.1006.2008.00333

Relationship between Changes of Na⁺, K⁺, and Ca²⁺ Contents during Seed Germination and Salt Tolerance in Maize

玉米种子萌发过程中Na⁺、K⁺和Ca²⁺含量变化与耐盐性的关系

SHANG Xue-Fang¹²,DONG Shu-Ting^1*,ZHENG Shi-Ying²,WANG Li-Yan²

商学芳^1,2;董树亭^1*;郑世英²;王丽燕²

1 Key Laboratory of Crop Biology, Shandong Agricultural University, Tai’an 271018, Shandong; 2 Dezhou College, Dezhou 253023, Shandong, China

Maize (Zea mays L.) seeds of a salt tolerant cultivar Denghai 9 and a salt sensitive cultivar Jundan 18 were germinated and grown in 1/4 and 1/2 Hoagland solutions with 0, 50, 100, 150, and 200 mmol L^-1 NaCl, respectively. The contents of Na⁺, K⁺, and Ca²⁺ in testa, embryo, endosperm, roots, root crown, and leaf were measured by ICP Mass. The Na⁺ content increased accordantly with the NaCl concentration in culture solution. In young seedlings, Na⁺ content in root and root crown was higher than that in leaf; in germinating seeds, it was significantly higher in embryo than in testa and endosperm. The capacity of Na⁺ accumulation in roots was strong, while the capacity of Na⁺ rejection in embryo was weak. The testa could also accumulate some Na⁺. The K⁺and Ca²⁺ contents in germinating seeds and young seedlings of maize decreased obviously under NaCl stress, and the Ca²⁺ content showed changes in a large scale, which was 38.4%–55.9% in Denghai 9 and 65.6%–78.2% in Jundan 18. The capacities of Na⁺ accumulation and rejection as well as the Ca²⁺ selective absorption might be related to salt-tolerance of maize cultivars

玉米耐盐品种登海9号和盐敏感品种浚单18在含0、50、100、150和200 mmol L^-1 NaCl的营养液中萌发生长, 采用等离子质谱分别测定其萌动种子种皮、胚、胚乳和幼苗根、根颈、叶中Na⁺、K⁺、Ca²⁺的含量。结果表明, 随着培养液中NaCl浓度的增加, 玉米体内Na⁺含量逐渐升高, 在幼苗中表现地下部(根和根颈)显著高于地上部(叶); 在萌动种子中, 胚中Na⁺积累量显著高于种皮和胚乳。根系积累Na⁺能力较强, 胚拒Na⁺能力较弱, 种皮具有一定的Na⁺累积能力。随NaCl浓度的增加, K⁺和Ca²⁺含量逐渐降低, 尤其是Ca²⁺含量急剧减少, 达38.4%~55.9%(登海9号)和65.6%~78.2%(浚单18)。玉米根、根颈、种皮的Na⁺积累能力、叶的拒Na+能力和幼苗选择吸收Ca²⁺的能力可能与品种耐盐性有关。

... 盐碱胁迫下, 植物对Na⁺、K⁺、Ca²⁺、Mg²⁺吸收选择及分配能力不仅与胁迫时间和胁迫程度有关, 且与品种类型有关^{[10,11,12,13]} ...

... 本研究中, 碱胁迫降低了两品种的Ca²⁺含量, 但随着胁迫时间的延长, Ca²⁺总量增加, 这与范远等^[11]在白燕2号上试验结果不一致, 可能不同基因型品种间存在差异导致适应胁迫的机制有所不同 ...

2010

0.0

1.889

Sci Agric Sin. 2010, 43(7):1423 - 1432

Study on the Characteristics of Absorption, Distribution and Selective Transport of Na+ and K+ in Tomato Plants Under Salt Stress

NaCl胁迫下栽培型番茄Na+、K+吸收、分配和转运特性

WANG Xue-zheng, LI Qiu-hong, WU Feng-zhi

王学征,李秋红,吴凤芝

（东北农业大学园艺学院）

【Objective】 This experiment was carried out to elucidate the effects of salt stress on absorption, distribution and selective transport of Na+ and K+ in tomato plants. 【Method】 The absorption, distribution and selective transport of Na+ and K+in tomato plants under salt stress were analyzed by atomic absorption spectrometry. 【Result】 The results showed that with salt stress concentration and time increased, Na+ absorption of tomatoes increased, the fundamental order of accumulation in various organs was root＞stem＞leaf. The total accumulation of Na+ in salt-tolerant cultivar was lower than that in salt-sensitive one. Under salt stress, Na+ contents in tomato leaves, stems and roots were increased with the increase of NaCl concentration. Na+ contents in salt-tolerant cultivar were lower than that in salt-sensitive one. From the regional distribution of ions in the tomato plant, there was a higher Na+ proportion in rhizomes of salt-tolerant cultivar; the distribution of potassium was more focused in leaves. The salt-sensitive cultivar tended to allocate Na+ to the leaves. Under early salt stress, the salt concentration was 200 mmol•L-1 and more. The choice of K+ transport in tomato plants to shoot with the stress showed a downward trend. Below 200 mmol•L-1 showed a good alternative transportation, Sk/Na (transport) of salt-tolerant cultivar were higher than that of salt-sensitive cultivar. Root of salt-tolerant cultivar showed a greater K+ transport ability. 【Conclusion】 The lower Na+ content in leaves and better alternative transportation of K+ under salt stress are the most important characteristics of salt tolerant tomato varieties.

【目的】明确盐分胁迫下栽培型番茄离子吸收、分配和转运特性。【方法】以栽培番茄为试材,以NaCl溶液为盐分胁迫条件,通过苗期耐盐性鉴定,采用原子吸收光谱法测定不同耐盐性番茄品种体内离子含量,对盐分胁迫下番茄体内离子积累、分布和转运机制进行系统分析。【结果】番茄对Na+的吸收随盐分处理浓度和时间的增加而增加,在各器官的积累量顺序为根＞茎＞叶。对于较耐盐品种,Na+在体内的积累总量低于盐敏感品种。盐分胁迫后,番茄叶片、茎和根系中Na+/K+比均随NaCl浓度的升高而升高。耐盐品种的Na+/K+比低于盐敏感品种。离子在体内的区域化分布情况是,较耐盐品种的Na+在根茎中的分配比例较高,盐敏感品种趋向于向叶片分配。K+在较耐盐品种的分布集中于叶片。在盐胁迫初期,盐分处理浓度超过200 mmol•L-1时,番茄植株对K+向地上部的选择运输性随着胁迫时间的延长呈现出下降的趋势。低于200 mmol•L-1时,表现出很好的选择运输性,耐盐品种的Sk/Na(运输)高于盐敏感品种,根系表现出更强的向地上部运输K+的能力。【结论】盐分胁迫下,叶片中较低的Na+含量和更强的向地上部运输K+的能力是番茄耐盐性的重要特征。

... 盐碱胁迫下, 植物对Na⁺、K⁺、Ca²⁺、Mg²⁺吸收选择及分配能力不仅与胁迫时间和胁迫程度有关, 且与品种类型有关^{[10,11,12,13]} ...

2011

0.0

... 盐碱胁迫下, 植物对Na⁺、K⁺、Ca²⁺、Mg²⁺吸收选择及分配能力不仅与胁迫时间和胁迫程度有关, 且与品种类型有关^{[10,11,12,13]} ...

2011

0.0

... 3 讨论渗透调节在作物耐盐碱机制中有重要作用^[14,15,16] ...

2011

0.0

... 3 讨论渗透调节在作物耐盐碱机制中有重要作用^[14,15,16] ...

2001

0.0

1.667

Acta Agron Sin. 2001, 27(6):776 - 780

Effects of Salt Stress on Growth and Ionic Distribution of Salt-born Glycine soja

盐胁迫对盐生野大豆生长和离子分布的影响

Yu Bingjun;Luo Qingyun;Liu Youliang

於丙军;罗庆云; 刘友良

以一年生栽培大豆品种(Lee68,耐盐性较强)和野生大豆品系(N23232,耐盐性较弱)为参照,以水培结合盐土种植的方法,从生长速率、电解质渗漏率和离子分布等方面,研究了盐生野大豆品系(BB52)的耐盐机理.结果表明,盐胁迫下,该品系幼苗生长受抑程度最轻,电解质渗漏率升幅最小.根系吸收的Na+、Cl-向地上部(尤其是叶片)的运输较少,而对

... 3 讨论渗透调节在作物耐盐碱机制中有重要作用^[14,15,16] ...

2001

0.0

0.59

... 参与渗透调节的阳离子主要是Na⁺、K⁺和Ca²⁺, 其在植物器官中的分配比例因物种、品种和生育时期而异^[17,18,19] ...

2011

0.0

... 参与渗透调节的阳离子主要是Na⁺、K⁺和Ca²⁺, 其在植物器官中的分配比例因物种、品种和生育时期而异^[17,18,19] ...

2000

25.962

0.0

Annu. Rev. Plant Physiol. Plant Mol. Biol.. 2000, 51:463-499

PLANT CELLULAR AND MOLECULAR RESPONSES TO HIGH SALINITY.

Hasegawa PM, Bressan RA, Zhu JK, Bohnert HJ

Center for Plant Environmental Stress Physiology, 1165 Horticulture Building, Purdue University, West Lafayette, Indiana 47907-1165; e-mail: paul.m.hasegawa.1@purdue.edu, Departments of 1 Plant Sciences and 2Biochemistry, University of Arizona, Tucson, Arizona 85721; e-mail: bonerth@u.arizona.edu

Plant responses to salinity stress are reviewed with emphasis on molecular mechanisms of signal transduction and on the physiological consequences of altered gene expression that affect biochemical reactions downstream of stress sensing. We make extensive use of comparisons with model organisms, halophytic plants, and yeast, which provide a paradigm for many responses to salinity exhibited by stress-sensitive plants. Among biochemical responses, we emphasize osmolyte biosynthesis and function, water flux control, and membrane transport of ions for maintenance and re-establishment of homeostasis. The advances in understanding the effectiveness of stress responses, and distinctions between pathology and adaptive advantage, are increasingly based on transgenic plant and mutant analyses, in particular the analysis of Arabidopsis mutants defective in elements of stress signal transduction pathways. We summarize evidence for plant stress signaling systems, some of which have components analogous to those that regulate osmotic stress responses of yeast. There is evidence also of signaling cascades that are not known to exist in the unicellular eukaryote, some that presumably function in intercellular coordination or regulation of effector genes in a cell-/tissue-specific context required for tolerance of plants. A complex set of stress-responsive transcription factors is emerging. The imminent availability of genomic DNA sequences and global and cell-specific transcript expression data, combined with determinant identification based on gain- and loss-of-function molecular genetics, will provide the infrastructure for functional physiological dissection of salt tolerance determinants in an organismal context. Furthermore, protein interaction analysis and evaluation of allelism, additivity, and epistasis allow determination of ordered relationships between stress signaling components. Finally, genetic activation and suppression screens will lead inevitably to an understanding of the interrelationships of the multiple signaling systems that control stress-adaptive responses in plants.

PMID: 15012199

... 参与渗透调节的阳离子主要是Na⁺、K⁺和Ca²⁺, 其在植物器官中的分配比例因物种、品种和生育时期而异^[17,18,19] ...

2005

0.0

1.462

... 通过调节阳离子在不同器官、组织或细胞内的区域(隔)化的分布, 有助于减轻离子毒害^[20] ...

2005

0.0

1.889

Sci Agric Sin. 2005, 38(7):1458 - 1465

Effects of NaCl Stress on Ions Absorption and Transportation and Plant Growth of Tall Fescue

NaCl胁迫对苇状羊茅离子吸收与运输及其生长的影响

李品芳,白文波,杨志成

中国农业大学资环学院

The absorption and transportation of inorganic ions and growth of Tall Fescue were studied in a hydroponics experiment. The results showed that, (1) NaCl stress reduced the number of K+ contents, K+/ Na+ and Ca2+/ Na+ ratios in shoots and roots, and dry weights of Tall Fescue, but increased Na+ and Cl- contents, and enhanced the selectivity of Cl--Na+ and K+-Na+; (2) The increments of Na+ and Cl- contents in shoot were higher than those in roots, the increment of Cl- contents were more than that of Na+ contents, and the Cl-/ Na+ ratios were almost 1.6 in shoots and about 0.8 in roots; (3) NaCl stress time affected the K+ and Na+ contents significantly and reduced the transportation selectivity ratios of K+ and Na+, but NaCl concentration mainly influenced the absorption ratios of K+ and Na+; (4) NaCl stress inhibited the growth of shoots more severely than that of roots, and the stress effect of Cl- was more significant than that of Na+.

通过室内水培试验，探讨了NaCl胁迫对苇状羊茅离子吸收与运输及生长的影响及其随时间的变化。结果表明，NaCl胁迫浓度的增加使苇状羊茅茎叶和根系的K+含量、K+/ Na+、Ca2+/Na+、干物质重下降，而使Na+、Cl-含量、Cl-/Na+ 及对K+/Na+的吸收与运输选择性增高。茎叶Na+含量和Cl-含量增大的幅度大于根系，且Cl-大于Na+，茎叶中Cl-/Na+基本保持在1.6左右，而根系在0.8左右。胁迫时间对植株K+ 、Na+含量的影响最为显著。K+－Na+吸收选择性主要受环境盐分浓度的影响，而其运输选择性却主要受胁迫时间的影响。盐分对苇状羊茅地上部生长的抑制作用大于根系，且Cl-的影响程度大于Na+。

... 这一方面是因为Vao-9通过地上部积累大量离子, 增大地上部和根部的渗透势差, 促进水分从根部向地上部运输, 改善地上部的水分状况, 促进生长^[21,22] ...

2002

0.0

1.321

1992

0.0

0.669

... 另一方面是通过把吸收的大部分Na⁺分配在茎部, 从而减少叶片受伤害^[23,24] ...

2000

0.0

... 另一方面是通过把吸收的大部分Na⁺分配在茎部, 从而减少叶片受伤害^[23,24] ...

2007

0.0

... 这与赵旭等^[25]在小麦耐盐性研究中的结果一致 ...

2002

11.808

0.0

Trends Plant Sci.. 2002, 7(9):405-10

Oxidative stress, antioxidants and stress tolerance.

Mittler R

Dept of Botany, Plant Sciences Institute, 353 Bessey Hall, Iowa State University, Ames, IA 50011, USA. rmittler@iastate.edu

Traditionally, reactive oxygen intermediates (ROIs) were considered to be toxic by-products of aerobic metabolism, which were disposed of using antioxidants. However, in recent years, it has become apparent that plants actively produce ROIs as signaling molecules to control processes such as programmed cell death, abiotic stress responses, pathogen defense and systemic signaling. Recent advances including microarray studies and the development of mutants with altered ROI-scavenging mechanisms provide new insights into how the steady-state level of ROIs are controlled in cells. In addition, key steps of the signal transduction pathway that senses ROIs in plants have been identified. These raise several intriguing questions about the relationships between ROI signaling, ROI stress and the production and scavenging of ROIs in the different cellular compartments.

PMID: 12234732

... Ca²⁺不仅对于保持植物细胞膜的完整性和稳定性有重要作用, 而且与植物的耐盐碱能力有重要关联^[26], 由于胁迫下高活性Na⁺置换质膜上的Ca²⁺, 细胞中Ca²⁺浓度降低, 导致质膜上Ca²⁺失去平衡, 失去保护细胞的作用, 使细胞膜的稳定性和选择性下降而破坏作物体内离子平衡 ...

2009

0.0

1.742

Chin J Appl Ecol. 2009, 20(4):767 - 772

Absorption and allocation characteristics of K⁺, Ca²⁺, Na⁺and Cl^-in different organs ofBroussonetia papyrifera seedlings under NaCl stress.

盐胁迫下构树幼苗各器官中K⁺、Ca²⁺、Na⁺和Cl^-含量分布及吸收特征

YANG Fan;DING Fei;DU Tian-zhen

杨帆;丁菲;杜天真

College of Garden and Art, Jiangxi Agricultural University, Nanchang 330045, China

One-year-old Broussonetia papyrifera seedlings were subjected to 04, 1, 2, 3, and 4 g·kg-1 of soil NaCl stress, and their biomass accumulation, leaf plasma membrane permeability, and the absorption, allocation and translocation of K+, Ca2+, Na+, and Cl-, as well as the symptoms of salt injury, were studied and investigated. The leaf plasma membrane permeability increased with the increase of soil NaCl concentration and of the duration of soil NaCl stress, and the seedling’s root/shoot ratio also increased with increasing soil NaCl concentration. When the soil NaCl concentration exceeded 3 g·kg-1, leaf plasma membrane permeability and seedling’s biomass accumulation were affected significantly. The Na+ and Cl- concentrations in different organs of seedlings increased with increasing soil NaCl concentration while the K+ and Ca2+ concentrations were in adverse, and the ion contents in leaves were always much higher than those in other organs, illustrating that soil NaCl stress affected the K+ and Ca2+ absorbing capability of roots, and inhibited the selective translocation of K+ and Ca2+ to aboveground parts. As a result, the K+ and Ca2+ concentrations in leaves and stems decreased. The study showed that B. papyrifera could effectively resist the injury of osmotic stress from soil salt via absorbing and accumulating Na+ and Cl-, but excessive accumulation of Na+ and Cl- could induce salt toxicity. As a non-halophyte species with relatively strong salt resistance, the aboveground parts of B. papyrifera did not have significant salt-exclusion effect.

将当年生构树幼苗置于含有不同浓度（04、1、2、3、4 g·kg^-1）NaCl的土壤中，研究其生物量积累、叶片细胞质膜透性和K⁺、Ca²⁺、Na⁺、Cl^-等离子的吸收、分布及运输，并观察盐害症状.结果表明：构树幼苗的叶片质膜透性随着NaCl浓度的增加和胁迫时间的延长而升高，根冠比随NaCl浓度的升高而增加，大于3 g·kg^-1的土壤盐胁迫对构树叶片的质膜透性及植株的生物量积累影响显著.构树幼苗各器官中Na⁺和Cl^-含量随土壤NaCl浓度升高而显著增加，K⁺和Ca²⁺则随之降低，叶片各离子含量均明显高于根和茎.说明盐胁迫影响根系对K⁺和Ca²⁺的吸收，并抑制了它们向地上部分的选择性运输，使叶和茎的K⁺和Ca²⁺含量下降.构树通过吸收积累Na⁺和Cl^-抵御土壤盐分带来的渗透胁迫，但过量的Na⁺和Cl^-积累会造成单盐毒害.作为抗盐性较高的非盐生植物，构树地上部分的拒盐作用不显著.

2005

2.203

0.0

Ecotox Environ Safe. 2005, 60(3):324 - 349 DOI:10.1016/j.ecoenv.2004.06.010

Salt tolerance and salinity effects on plants: a review

Parida

A K

,Das

A B

.Salt tolerance and salinity effects on plants: a review.Ecol Toxicol Environ Safety,2005,60:324-349

Abstract

Plants exposed to salt stress undergo changes in their environment. The ability of plants to tolerate salt is determined by multiple biochemical pathways that facilitate retention and/or acquisition of water, protect chloroplast functions, and maintain ion homeostasis. Essential pathways include those that lead to synthesis of osmotically active metabolites, specific proteins, and certain free radical scavenging enzymes that control ion and water flux and support scavenging of oxygen radicals or chaperones. The ability of plants to detoxify radicals under conditions of salt stress is probably the most critical requirement. Many salt-tolerant species accumulate methylated metabolites, which play crucial dual roles as osmoprotectants and as radical scavengers. Their synthesis is correlated with stress-induced enhancement of photorespiration. In this paper, plant responses to salinity stress are reviewed with emphasis on physiological, biochemical, and molecular mechanisms of salt tolerance. This review may help in interdisciplinary studies to assess the ecological significance of salt stress.

2005

0.0

... 这证实了不同刺槐品种存在Mg²⁺吸收选择能力差异的结果^[29] ...

2007

2.638

0.0

Plant Soil. 2007, 294(1-2):263 - 276 DOI:10.1007/s11104-007-9251-3

Osmotic adjustment and ion balance traits of an alkali resistant halophyteKochia sieversiana during adaptation to salt and alkali conditions

Chunwu Yang(1) Jianna Chong(1) Changyou Li(1) Changmin Kim(2) Decheng Shi(13) Deli Wang(1)

1.Ministry of Education Key Laboratory of Vegetation Ecology, Northeast Normal University, 130024 Changchun, Jilin Province, China
2.Department of Horticulture, Sariwon Agricultural University, Sariwon, Democractic Peoples Republic of Korea
3.Life Science College, Northeast Normal University, 130024 Changchun, Jilin Province, China

Kochia sieversiana (Pall.) C. A. M., a naturally alkali-resistant halophyte, was chosen as the test organism for our research. The seedlings ofK. sieversiana were treated with varying (0–400 mM) salt stress (1:1 molar ratio of NaCl to Na₂SO₄) and alkali stress (1:1 molar ratio of NaHCO₃ to Na₂CO₃). The concentrations of various solutes in fresh shoots, including Na⁺, K⁺, Ca²⁺, Mg²⁺, Cl⁻, SO₄²⁻, NO₃⁻, H₂PO₃⁻, betaine, proline, soluble sugar (SS), and organic acid (OA), were determined. The water content (WC) of the shoots was calculated and the OA components were analyzed. Finally, the osmotic adjustment and ion balance traits in the shoots ofK. sieversiana were explored. The results showed that the WC ofK. sieversiana remained higher than 6 [g g⁻¹ Dry weight (DW)] even under the highest salt or alkali stress. At salinity levels >240 mM, proline concentrations increased dramatically, with rising salinity. We proposed that this was not a simple response to osmotic stress. The concentrations of Na⁺ and K⁺ all increased with increasing salinity, which implies that there was no competitive inhibition for absorption of either inK. sieversiana. Based on our results, the osmotic adjustment feature of salt stress was similar to that of alkali stress in the shoots ofK. sieversiana. The shared essential features were that the shoots maintained a state of high WC, OA, Na⁺, K⁺ and other inorganic ions, accumulated largely in the vacuoles, and betaine, accumulated in cytoplasm. On the other hand, the ionic balance mechanisms under both stresses were different. Under salt stress,K. sieversiana accumulated OA and inorganic ions to maintain the intracellular ionic equilibrium, with close to equal contributions of OA and inorganic ions to anion. However, under alkali stress, OA was the dominant factor in maintaining ionic equilibrium. The contribution of OA to anion was as high as 84.2%, and the contribution of inorganic anions to anion was only 15.8%. We found that the physiological responses ofK. sieversiana to salt and alkali stresses were unique, and that mechanisms existed in it that were different from other naturally alkali-resistant gramineous plants, such asAneurolepidium chinense,Puccinellia tenuiflora.

... 在棉花、碱地肤上的研究表明, 在盐、碱胁迫下, 作物对Na⁺的区域化分配, K⁺、Ca²⁺、Mg²⁺的吸收, 与叶片细胞质膜H⁺-ATPase、液泡膜H⁺-ATPase、H⁺-PPase和Na⁺/H⁺逆向运输蛋白等活性有关^[30,31] ...

2001

0.0