外源水杨酸对Cu胁迫下水培烟草生长及营养元素吸收利用的影响

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徐根娣, 葛淑芳, 章艺, 吴玉环, 梅笑漫, 刘鹏. . 外源水杨酸对Cu胁迫下水培烟草生长及营养元素吸收利用的影响. 作物学报, 2015, 41(6): 956-962
XU Gen-Di, GE Shu-Fang, ZAHNG Yi, WU Yu-Huang, MEI Xiao-Man, LIU Peng. . Effect of Salicylic Acid on Growth and Nutrient Uptake in Hydroponic Tobacco under Cu Stress. ACTA AGRONOMICA SINICA, 2015, 41(6): 956-962 复制到剪切板

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外源水杨酸对Cu胁迫下水培烟草生长及营养元素吸收利用的影响

徐根娣^1,², 葛淑芳^1,², 章艺³, 吴玉环⁴, 梅笑漫⁵, 刘鹏^1,^2,^*

¹ 浙江师范大学生态学研究所, 浙江金华 321004

²浙江师范大学植物学实验室, 浙江金华 321004

³浙江旅游职业学院, 浙江杭州 3112313

⁴ 杭州师范大学生命与环境科学学院, 浙江杭州 330036

⁵ 杭州师范大学初等教育学院, 浙江杭州 330036

^* 通讯作者(Corresponding author): 刘鹏: E-mail:pliu99@vip.sina.com

第一作者联系方式: E-mail: sky113@zjnu.cn

收稿日期:2014-10-27 接受日期:2015-04-02 网络出版日期:2015-04-14

基金:本研究由国家自然科学基金项目(30970188, 30540056)和浙江省公益性技术应用研究计划项目(2011C22053)

摘要

以耐Cu性较强的烟草品种W38 ( Nicotiana tabacumcv. W38)和耐Cu性较弱的本氏烟( Nicotiana benthamiana)为试验材料, 设置水培处理组即CK (Cu²⁺ 0 mg L^-1, SA 0 μmol L^-1, 即对照组)、T1(Cu²⁺ 4 mg L^-1, SA 0 μmol L^-1)、T2 (Cu²⁺ 4 mg L^-1, SA 100 μmol L^-1)、T3 (Cu²⁺ 4 mg L^-1, SA 300 μmol L^-1)、T4 (Cu²⁺ 4 mg L^-1, SA 500 μmol L^-1), 探讨SA对烟草生长特性及根、茎、叶不同器官中元素吸收规律的影响。结果表明, 经过4 mg L^-1Cu胁迫15 d后, 2个品种的烟草生长均受到一定程度的抑制, 不同器官中Cu含量均显著升高, K、Ca、Mg、Fe、Zn、B、Mn的吸收受到抑制, 而在营养液中添加适宜浓度的SA能够有效缓解Cu胁迫对2个品种烟草根长、株高、鲜重的抑制作用, 并降低烟草体内Cu含量。与T1处理组相比, 本氏烟的根、茎、叶中Cu含量最大分别下降25.05%、39.78%和22.91%, W38的根、茎、叶中的Cu含量最大分别下降23.27%、37.30%和28.88%, 并促进了营养元素K、Ca、Mg、Fe、Mn的吸收, 但是对Zn、B元素的吸收影响并不明显。由此可知, 高浓度的Cu胁迫会抑制烟草的生长及营养元素的吸收和运输, 施加适宜浓度的外源水杨酸有利于营养元素的吸收和促进烟草的生长发育, 在本试验浓度范围内施加300 μmol L^-1的水杨酸可显著缓解Cu胁迫对烟草的抑制作用。

关键词: SA; Cu胁迫; 烟草; 微量元素

Effect of Salicylic Acid on Growth and Nutrient Uptake in Hydroponic Tobacco under Cu Stress

XU Gen-Di^1,², GE Shu-Fang^1,², ZAHNG Yi³, WU Yu-Huang⁴, MEI Xiao-Man⁵, LIU Peng^1,^2,^*

¹ Research Institute of Ecology, Zhejiang Normal University, Jinhua 321004, China

² Key Laboratory of Botany / Zhejiang Normal University, Jinhua 321004, China

³ Tourism College of Zhejiang, Hanzhou 311231, China

⁴ College of Life and Environmental Sciences, Hangzhou Normal University, Hangzhou 310036, China

⁵ College of Teacher Education, Hanzhou Normal University, Hangzhou 330036, China

Abstract

A hydroponic experiment was conducted to study the effects from treatmeats including CK ( Cu²⁺ 0 mg L^-1, SA 0 μmol L^-1 as control), T1 (Cu²⁺ 4 mg L^-1, SA 0 μmol L^-1), T2 (Cu²⁺ 4 mg L^-1, SA 100 μmol L^-1), T3 (Cu²⁺ 4 mg L^-1, SA 300 μmol L^-1), T4 (Cu²⁺ 4 mg L^-1, SA 500 μmol L^-1) on growth characteristics, elements absorption and accumulation in two tobacco cultivars, N. tabacum cv .W38 (Cu resistant genotype) and N. benthamiana (Cu sensitive genotype) under copper stress. The result showed that the growth of two tobacco cultivars under 4 mg L^-1Cu stress for 15 days was inhibited to some extent, meanwhile, the content of Cu in their organs was increased while the absorptions of K, Ca, Mg, Fe, Zn, B, and Mn were inhibited. The addition of exogenous SA could alleviate the inhibition of plant height, root length and fresh weight, and reduce Cu content of tobacco, but increase the absorption of nutritional elements K, Ca, Mg, Fe, and Mn. Compared with treatment T1, contents of Cu in root, stem and leaf were increased by 25.05%, 39.78%, 22.91% in N. benthamiana, and by 23.27%, 37.30%, 28.88% in N. tabacum cv. W38, respectively. In conclusion, appropriate concentration of SA promotes the growth and elements absorption in tobacco under copper stress. In this experiment, 300 μmol L^-1 SA could significantly alleviate copper inhibition to tobacco.

Keyword: Salicylic acid; Cu stress; Tobacco; Trace elements

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引言

近年来随着现代化工农产业的飞速发展, 土壤的重金属污染日趋严重。重金属在环境中具有难降解性和相对稳定性, 且会在植物的根、茎、叶中大量积累, 这不仅严重阻碍了植物的正常生长和发育, 而且会通过食物链进而危害动物和人类的健康^[1]。水杨酸(salicylic acid, SA), 即邻羟基苯甲酸, 植物体内广泛存在, 目前已证明能够调节植物体内的许多抗逆生理过程, 如抗盐性、抗寒性、抗热性、抗旱性以及抗病性^{[2, 3]}, 同时作为一种植物激素, SA也是用于增加作物收成的一种重要措施, 具有广阔的应用前景^{[4, 5, 6, 7]}。

重金属胁迫对植物造成毒害的机制主要是重金属离子取代植物必需营养元素与酶活性中心或蛋白中心的巯基结合, 或是干扰Ca²⁺等元素的运输并启动氧化还原反应产生自由基, 造成植物的氧化损伤, 进而造成植株代谢紊乱^[8]。铅、镉等重金属胁迫能使黄瓜、大麦等幼苗的内源SA水平增加^{[9, 10]}, Freeman等^[11]等发现拟南芥中自由态SA含量的升高, 能够激发丝氨酸乙酰转移酶活性, 进而使植株体内谷胱甘肽含量升高, 增强抗逆性。外源SA的施加还会影响植物对重金属和营养元素的吸收、运输。Drazic和Mihailovic^[12]研究SA对Cd胁迫下大豆幼苗对营养元素吸收规律的影响, 发现SA能减低K含量却促进了Fe的吸收, 同时也促进了Mg和Ca的吸收与运输, 使植物体内各元素间达到动态平衡, 提高机体抗胁迫的能力。总之, 目前有关SA诱导植物重金属抗性的机理研究仍然处于起步阶段, 已有的研究一般证实了SA能在一定程度上缓解重金属对植物的毒害, 但不同植物或相同植物的不同组织对SA抗重金属的机制可能有所不同, 甚至互相矛盾, 其中有许多问题亟待解决, 且有关SA对烟草Cu胁迫下营养元素吸收运输的影响国内外至今尚未见报道, 因此本文以烟草为试验材料, 旨在探讨外源SA对烟草营养元素吸收运输的调控机制及其可能的调控技术途径, 以期为进一步提高烟叶品质提供一定的理论依据, 且为抗性生理增添素材, 为重金属污染的防治提供科学参考。

1 材料与方法

1.1 供试材料与试验设计

以烟草根相对伸长率为指标, 从云烟85、云烟87、MS云烟85、W38、MSK326、NC89、双-70、89112、本氏烟等9个烟草品种中筛选出耐Cu性较强的W38和耐Cu性较弱的本氏烟, 设置施铜和不施铜2个处理。烟草种子购买于云南玉溪烟草种子公司。SA购买于金华医药有限公司。

先将烟草种子经10%的H₂O₂表面消毒15 min, 然后在25℃下黑暗浸种24 h, 之后将其播种到铺有滤纸的培养皿中, 等种子发芽后转移到沙子中继续培养, 待幼苗的第4片真叶展开后, 选取长势较为一致的幼苗移入Hoagland 1/2完全营养液, 预培养3 d, 再转入完全营养液预培养6 d, 然后按以下处理继续培养: CK (Cu²⁺ 0 mg L^-1, SA 0 μ mol L^-1, 即对照组)、T1 (Cu²⁺ 4 mg L^-1, SA 0 μ mol L^-1)、T2 (Cu²⁺ 4 mg L^-1, SA 100 μ mol L^-1)、T3 (Cu²⁺ 4 mg L^-1, SA 300 μ mol L^-1)、T4 (Cu²⁺ 4 mg L^-1, SA 500 μ mol L^-1)。其中, Cu²⁺以CuSO₄· 5H₂O的形式提供, SA采用直接添加到营养液中的方式, 每3 d更换1次营养液, 处理15 d后取样测定相应生理指标, 每个处理3个重复。试验材料和试验所用浓度由前期大量预试验所得。

1.2 测定指标和方法

根系长度、株高用直尺直接测量; 植株鲜重用电子天平称量。收集烟草的根、茎、叶, 用去离子水反复冲洗, 茎、叶部分先在105℃下杀青15 min, 后与根一起于70℃的烘箱中烘干至恒重, 再研磨成粉末。精确称取每个样品0.3 g加入HNO₃-HClO₄消化, 待溶液变得澄清透明时, 高温烘干, 此时有白色晶体析出, 再用去离子水将其定容至50 mL, 即得待测溶液。采用7500a型电感耦合等离子体质谱仪(ICP-MS) (美国Agilent公司), 测定Cu、K、Ca、Mg、Fe、Zn、B和Mn等8种元素。

1.3 数据处理

所有测定均设3次重复, 计算平均值和标准误差, 采用SPSS 18.0软件中的Duncan’ s法进行显著性差异分析, 用Microsoft Excel 2007软件制图。

2 结果与分析

2.1 烟草生长特性

外源SA对Cu胁迫下烟草生长特性的影响如图1所示, T1处理下, W38和本氏烟的根长显著降低, 与对照组相比分别降低8.56%和12.23%, 由此可知, Cu胁迫对本氏烟的根长抑制率大于W38, 表现出其对Cu毒的敏感性。施加一定浓度的外源SA可以缓解根长的抑制作用, 当SA浓度为300 μ mol L^-1时能有效缓解Cu毒, 与T1相比, W38和本氏烟的根长分别上升5.95%和13.50%。由图2和图3可知, 2个品种烟草的株高和鲜重在不同处理的变化趋势大致是一致的, T1处理条件下其株高和鲜重均显著下降, 在一定范围内随着施加外源SA浓度的升高而不断增加, 但是当外源SA浓度大于500 μ mol L^-1时, 却对烟草的生长产生抑制作用, 其中本氏烟表现更为突出, 这可能是因为高浓度的外源SA对烟草产生了一种新的胁迫作用。

图1

SA对铜胁迫下烟草根长的影响

Fig. 1

Effects of SA on the root length (mean± SE) of tobacco under copper stress

Bars superscripted by different letters are significantly different at the 5% probability level. CK: Cu²⁺ 0 mg L^-1, SA 0 μ mol L^-1, T1: Cu²⁺ 4 mg L^-1, SA 0 μ mol L^-1; T2: Cu²⁺ 4 mg L^-1, SA 100 μ mol L^-1, T3: Cu²⁺ 4 mg L^-1, SA 300 μ mol L^-1, T4: Cu²⁺ 4 mg L^-1, SA 500 μ mol L^-1.

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	图2 SA对铜胁迫下烟草株高的影响Fig. 2 Effects of SA on plant height (mean± SE) of tobacco under copper stressBars superscripted by different letters are significantly different at the 5% probability level. Treatment codes are the same as those given in Fig. 1.

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	图3 SA对铜胁迫下烟草鲜重的影响Fig. 3 Effects of SA on fresh weight (mean± SE) of tobacco under copper stressBars superscripted by different letters are significantly different at the 5% probability level. Treatment codes are the same as those given in Fig. 1.

2.2 烟草Cu含量变化

如表1所示, 与对照组相比, 在4 mg L^-1Cu胁迫下2个品种烟草地上部和地下部Cu含量均呈显著增加趋势。施加外源SA后, 2个品种烟草Cu含量均有明显的降低趋势。外源SA浓度为100 μ mol L^-1时对本氏烟的缓解效果较好, 与T1胁迫下相比, 其根、茎、叶中Cu含量分别下降25.05%、39.78%和22.91%。对于W38品种来说, 当外源SA浓度为300 μ mol L^-1时对其缓解效果较好, 其根、茎、叶中的Cu含量分别下降23.27%、37.30%和28.88%。

表1 SA对铜胁迫下烟草Cu含量的影响 Table 1 Effects of SA on the content of Cu (mean± SE) in tobacco under copper stress (μ g g^-1DW)

2.3 烟草K含量变化

由表2可以看出, 与对照相比, T1处理条件下2个品种烟草根、茎、叶中K含量均显著下降, W38分别下降46.01%、25.13%和35.87%, 本氏烟分别下降49.62%、32.56%和35.30%, 这可能是Cu毒抑制了根系对K的吸收, 进而降低了烟草地上部K含量。添加外源SA后, 2个品种烟草根、茎、叶中K含量均有所增加, 但是当SA浓度为500 μ mol L^-1时, 本氏烟各部分的K含量却呈现下降趋势, 说明高浓度的SA对烟草的生长发育有一定的抑制作用。

2.4 烟草Ca、Mg含量变化

从表3可以看出, T1处理与对照组相比, 烟草根、茎、叶Ca含量迅速下降, 这说明Cu对Ca的吸收有抑制作用。在烟草植株各部位的Ca含量分布不同, 叶> 茎> 根。外源SA浓度为300 μ mol L^-1时, W38各器官Ca含量均达最高, 与T1相比, 其根、茎、叶的Ca含量分别上升23.90%、17.22%和8.70%, 可见施加外源SA后对根部Ca的吸收促进作用最为明显。本氏烟的Ca含量随着外源SA浓度的增加而呈现递减趋势, 外源SA浓度100 μ mol L^-1时, 缓解效果较好, 与T1处理相比, 本氏烟根、茎、叶Ca含量分别上升16.94%、11.21%和5.52%。因此, 在Cu胁迫下, 施加一定浓度的SA可以促进烟草尤其是根部Ca元素的吸收。

由表3可知, 耐Cu性不同的W38和本氏烟相同部位中Mg含量的差异不大, 且在不同处理下对Mg的吸收变化趋势也是相似的。在Cu胁迫下2个烟草品种中Mg含量均有所下降, 但是在添加外源SA后并没有显著提高烟草各部位的Mg含量, 由此可知, 外源SA对Cu胁迫下Mg元素的吸收和运输影响不大。此外, Mg元素在2个烟草品种根中的含量均大于在茎和叶中。

表2 SA对铜胁迫下烟草K含量的影响 Table 2 Effects of SA on the content of K (mean± SE) in tobacco under copper stress (μ g g^-1 DW)

表3 SA对铜胁迫下烟草Ca、Mg含量的影响 Table 3 Effects of SA on contents of Ca and Mg (mean± SE) in tobacco under copper stress (μ g g^-1 DW)

2.5 烟草Fe、Zn、B、Mn含量变化

由表4可知, T1处理对烟草微量元素吸收有不同程度的抑制作用, 其中Fe和Mn元素受到的抑制作用最大, B, Zn受到的抑制作用最小。施加外源SA后Fe、Mn元素的含量有所回升但是仍然低于对照组, 这说明外源SA缓解Cu毒的能力是有限的。而施加外源SA后, Zn、B元素含量并没有显著的变化, 可见SA对Cu胁迫下Zn、B吸收和运输的影响不大。

表4 SA对铜胁迫下烟草Fe、Zn、B、Mn含量的影响 Table 4 Effects of SA on contents of Fe, Zn, B, and Mn (mean± SE) in tobacco under copper stress (μ g g^-1 DW)

微量元素 Trace element	处理 Treatment	W38 N. tabacumcv. W38			本氏烟 N. benthamiana
微量元素 Trace element	处理 Treatment	根Root	茎Stem	叶Leaf	根Root	茎Stem	叶Leaf
Fe	CK	524.59± 1.42 a	120.51± 0.44 a	132.85± 0.72 a	510.51± 0.43 a	102.62± 0.40 a	95.52± 0.50 a
	T1	410.84± 1.01 e	62.48± 0.46 d	59.58± 0.55 d	333.13± 0.16 e	82.38± 0.67 d	67.31± 0.29 c
	T2	494.36± 0.72 c	97.21± 0.20 b	117.22± 0.08 b	474.61± 0.52 c	91.44± 0.45 b	86.26± 0.29 b
	T3	499.44± 0.50 b	97.73± 0.63 b	132.14± 1.01 a	492.51± 0.44 b	87.44± 0.50 c	59.90± 0.28 d
	T4	444.12± 0.19 d	79.83± 0.18 c	100.41± 0.53 c	379.18± 0.07 d	81.76± 0.19 d	34.62± 0.40 e
Zn	CK	20.28± 0.20 b	13.10± 0.17 a	18.45± 0.06 a	22.83± 0.56 ab	17.26± 0.17 a	20.10± 0.43 ab
	T1	19.46± 0.06 c	12.69± 0.50 b	18.07± 0.20 b	21.82± 0.42 b	16.64± 0.40 b	19.19± 0.47 b
	T2	19.60± 0.06 c	12.82± 0.17 ab	18.20± 0.10 ab	22.95± 0.80 a	16.72± 0.23 b	19.76± 0.33 ab
	T3	21.29± 0.27 a	11.54± 0.23 d	18.20± 0.13 ab	22.19± 0.59 ab	16.42± 0.10 b	20.80± 0.86 a
	T4	19.51± 0.16 c	12.06± 0.24 c	17.33± 0.31 c	17.68± 0.30 c	15.10± 0.10 c	17.27± 0.83 c
B	CK	32.47± 0.10 a	17.35± 0.19 a	25.30± 0.05 a	30.13± 0.45 a	19.43± 0.18 a	21.49± 0.20 a
	T1	29.73± 0.44 d	15.22± 0.10 c	23.46± 0.22 c	28.66± 0.69 b	17.43± 0.12 c	20.80± 0.39 b
	T2	30.82± 0.34 c	16.40± 0.10 b	23.85± 0.23 b	28.75± 0.16 b	17.24± 0.06 c	20.57± 0.21 b
	T3	31.69± 0.39 b	16.40± 0.32 b	25.20± 0.16 a	29.76± 0.20 a	18.65± 0.18 b	21.68± 0.34 a
	T4	32.04± 0.08 ab	17.28± 0.14 a	25.39± 0.32 a	28.59± 0.08 b	16.38± 0.15 d	19.26± 0.27 c
Mn	CK	624.52± 0.43 a	181.87± 0.57 a	165.35± 0.10 a	611.48± 0.48 a	165.00± 0.61 a	115.24± 0.07 a
	T1	354.25± 0.08 d	74.20± 0.22 e	69.73± 0.45 d	401.51± 0.43 d	69.39± 0.78 e	74.16± 0.05 d
	T2	580.32± 1.77 b	169.67± 0.34 c	149.90± 0.29 b	584.92± 0.34 c	149.49± 0.45 b	112.27± 0.06 b
	T3	581.53± 0.43 b	172.36± 0.16 b	117.78± 0.67 c	587.33± 0.09 b	117.78± 0.67 c	108.32± 0.30 c
	T4	427.62± 0.53 c	95.51± 0.15 d	117.31± 0.10 c	401.25± 1.03 d	73.29± 1.01 d	60.74± 0.46 e

Values followed by different letters within a row are significantly different at the 0.05 probability level (P< 0.05). Treatment codes are the same as those given in Fig. 1.

同一元素、同列数据后的不同字母表示在0.05水平上差异显著。处理代码同图1。

表4 SA对铜胁迫下烟草Fe、Zn、B、Mn含量的影响 Table 4 Effects of SA on contents of Fe, Zn, B, and Mn (mean± SE) in tobacco under copper stress (μ g g^-1 DW)

3 讨论

Cu是植物正常生命活动所必需的微量元素之一, 对维持植物正常的新陈代谢及生长发育均具有极其重要的意义, 然而, 适合植物生长的铜浓度范围却很窄, 土壤中稍微过量的Cu便会干扰细胞的正常代谢及离子平衡继而会对植物造成严重的毒害作用^[13]。本研究表明, 烟草植株生长在Cu浓度为4 mg L^-1时, 与对照相比, 其生长受到抑制, 表现为根长、株高以及单株鲜重值显著下降。已有的研究表明, SA对植物抗性的诱导具有浓度效应, 表现为低促高抑^[14]。本研究表明, 100~300 μ mol L^-1的SA可以不同程度地缓解铜对烟草幼苗根长、株高及鲜重的抑制作用, 特别是SA质量浓度为300 μ mol L^-1时, 缓解效果最明显。说明SA可以诱导植物产生抗铜性。但当SA质量浓度达到500 μ mol L^-1时, 不仅不会缓解铜对烟草幼苗生长的抑制作用, 反而产生更严重的毒害作用, 这可能是高质量浓度的SA与Cu共同作用产生双重毒害的结果。

营养元素的吸收平衡被破坏是重金属植物毒理学效应的一个重要方面^[15]。植物Cu毒害的一个重要机制就是严重阻碍植物对养分的吸收及运输, 最终导致植物营养的亏缺或失衡。Cu、K、Ca、Mg、Fe、Zn、B和Mn都是植物正常生长和代谢所必需的营养元素, 其中Cu是许多氧化酶的成分及某些酶的活化剂, 在植物的光合作用、呼吸作用等方面的氧化还原过程中有着非常重要的作用^[16], K提高植物的抗逆性, 能增强碳水化合物的合成和转运, 能激活多种酶, 能调节渗透浓度, 另外, K能促进蛋白质和核酸的形成。

Ca是细胞代谢的总调和者, 被称为“ 第二信使” , 在多种细胞信号转导途径中起着重要的作用, 是参与细胞构建及代谢必需的营养元素, Mg是植物叶绿素的重要组成成分之一, 在光合作用和呼吸作用过程中可以活化各种磷酸变位酶和磷酸激酶, 且Mg也可以活化DNA和RNA的合成过程^[17], Fe作为植物必需的微量营养元素, 是许多重要酶的辅基, 如细胞色素、过氧化物酶等, Zn是植物多种代谢途径中关键酶的组成因子, 同样也是叶绿素合成的必需元素, B的主要生理功能是参与糖的转运与代谢, 可以调节植物细胞壁的形成、N代谢等生理过程, Mn是多种酶的活化剂, 可以直接参与植物的光合作用过程, 同时对氧化还原反应中的电子传递有着非常重要的影响。此外, Cu胁迫下植物的根原生质膜上的Ca²⁺通道会受阻, 从而导致根尖细胞Ca²⁺净吸收量下降, 原生质膜上Ca²⁺缺乏, 这种平衡被破坏, 进而对细胞的结构和功能造成严重的影响, 同时导致Mg在根系阳离子交换量的饱和度降低, 其吸收量的下降, 继而会导致叶绿素合成的受阻。K、Fe在植物体内参与多种酶的合成, 同时在植物体的吸收、代谢、蛋白质合成等过程中不可或缺^[18]。Cu胁迫阻碍了植物的正常生长发育, 同时也影响了营养元素的吸收和运输。本研究发现, 过量的Cu改变了烟草细胞阳离子的交换能力, 致使烟草根系和地上部的Cu含量增加, 抑制了植株的生长, 减少了烟草对K、Ca、Mg、Fe、Zn、B和Mn养分的吸收, 降低了烟草植株各部位K、Ca、Mg、Fe、Zn、B、Mn元素的含量。可能机制之一是Cu²⁺竞争性地抑制了其他离子的吸收, 如Cu²⁺通过与膜上特殊通道受体结合位点结合, 使根系原生质膜Ca²⁺通道受阻, 阻碍Ca²⁺在膜上的整合, 进而影响了Ca²⁺跨膜运输的GTP蛋白的活性, 降低了原生质体Ca²⁺通道的流量, Ca²⁺平衡遭到破坏, 进而影响细胞结构和正常的代谢功能, 导致Ca²⁺在根系阳离子交换量的饱和度下降。可能机制之二是Cu²⁺胁迫影响根系活力, Cu²⁺浓度超过一定值时会明显降低根系活力^{[19, 20]}, 对烟草发育产生毒害, 本试验中K、Ca、Mg、Fe、Zn、B和Mn的吸收在Cu²⁺胁迫时明显下降, 主要原因可能是根系活力减弱导致养分吸收受阻。由此可见, Cu胁迫造成的根系元素吸收受抑及养分分配平衡失衡都加剧了植株Cu毒害, 并且降低了植株的产量和品质。此外我们还发现, 外施SA有效的抑制了Cu胁迫下Cu含量的增加, 缓解了Cu这种重金属的过量吸收对烟草造成的不良影响, 同时, 外施SA可以有效缓解烟草对大量元素K、Ca、Mg和微量元素Fe、Mn吸收量的减少, 这可能是因为添加外源SA对溶液中Cu²⁺形成了络合物或螯合物, 从而降低了Cu的胁迫性。在植物体内蛋白质、核酸等重要物质的合成和代谢都离不开K、Mg的参与, 它们在植物体内含量的降低必将导致植物的代谢紊乱^[21], 这与SA减轻Cu对细胞膜结构的损伤有关。此外, 外源SA对Cu胁迫下W38和本氏烟营养元素吸收均有很好的促进效果, 且对铜敏感型品种本氏烟的营养元素吸收的促进作用更为显著, 这与周媛^[22]的结果较为一致。由此可知, 外源SA的添加能够有效缓解Cu胁迫下烟草对营养元素吸收的不足, 促进元素的吸收和由地下向地上部分的运输。

4 结论

烟草对营养元素的吸收是有平衡要求的, 营养元素在烟草的不同器官部位分配不均衡将严重危害其正常的代谢及生长发育, 而Cu胁迫下施加一定浓度的外源SA可以在一定程度上恢复烟草对营养吸收的动态平衡和缓解烟草营养缺乏和离子毒害症状。在本实验浓度范围内施加300 μ mol L^-1的SA可显著缓解Cu胁迫对烟草生长的抑制作用及促进烟草对营养元素的吸收利用, 进而提高烟叶的品质。

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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2003

0.0

... 重金属在环境中具有难降解性和相对稳定性, 且会在植物的根、茎、叶中大量积累, 这不仅严重阻碍了植物的正常生长和发育, 而且会通过食物链进而危害动物和人类的健康^[1] ...

2012

1.265

0.0

高培培, 章艺, 吴玉环, 徐根娣, 刘鹏. 外源水杨酸对铝胁迫下栝楼光合特性及耐铝性的影响. 水土保持学报, 2012, 26(6): 268-273

Gao P

, Zhang

, Wu Y

, Xu G

, Liu

Effect of exogenous SA on photosynthesis and Al tolerance of Trichosanthes kirilowii Maxim under aluminum stress.

J Soil Water Conserv, 2012, 26(6): 268-273 (in Chinese with English abstract)

A solution culture experiment was conducted to study the effects of different SA concentration on growth characteristics,photosynthesis,oxidation resistance,UV-B absorbing compounds and MDA content of two Trichosanthes kirilowii cultivars,Anguo(Al-tolerant genotype) and Pujiang(Al-sensitive genotype),under 800 μmol/L aluminum stress.The results showed that 800 μmol/L Al treated for 15 days,the relative root length,plant height and fresh shoot weight,chlorophyll content,the maximum energy conversion efficiency(Fv/Fm),maximum fluorescence yield(Fm) and electron transport rate(ETR) were significantly lower of the two T.kirilowii cultivars.SOD,CAT activity changed in the opposite trend,indicating the differences of the two T.kirilowii cultivars in aluminum tolerance,POD activity and UV-B absorbing compounds decreased,MDA content and initial fluorescence(F0) increased.Adding 10,30,50 μmol/L exogenous SA could reduce aluminum accumulation in the apical cell wall,MDA content and initial fluorescence(F0),also increased the relative root length,plant height,fresh shoot weight,chlorophyll content,chlorophyll fluorescence of the relevant indicators and antioxidant enzyme activity to a certain extent.By increasing antioxidant enzyme activities and photosynthesis-related indicators,SA enhanced the ability of T.kirilowii to resistance to aluminum toxicity.Comprehensive analyzing evaluation by the method of subjection function,the results showed that the external source of SA could alleviate aluminum toxicity to some extent,but it did not eliminate the inhibition of T.kirilowii growth caused by aluminum.

以耐铝品种安国栝楼和铝敏感品种浦江栝楼为试验材料,采用溶液培养法研究800μmol/L铝胁迫下,不同浓度水杨酸对2个栝楼品种的生长特性、光合特性及抗氧化酶活性、MDA和紫外吸收物含量的影响。结果表明,经过800μmol/L铝处理15d后,安国栝楼和浦江栝楼的相对根长、株高及地上部鲜重、叶绿素含量、最大光能转化效率（Fv/Fm）、最大荧光产量（Fm）及电子传递速率（ETR）明显下降,SOD、CAT活性却出现2种相反的变化趋势,显示出2个栝楼品种的耐铝性差异,POD活性和紫外吸收物含量降低,MDA含量及初始荧光（F0）升高。10,30,50μmol/L外源SA可不同程度地降低铝在根尖细胞壁中的积累、MDA含量及初始荧光（F0）,提高栝楼的相对根长、株高、地上部鲜重、叶绿素含量及叶绿素荧光的相关指标和抗氧化酶活性。由此可知,外源SA通过提高栝楼抗氧化酶活性及光合作用能力增强栝楼抗铝毒能力。由隶属函数分析得出,SA具有缓解铝胁迫的效果,但并不能消除铝胁迫对栝楼生长的抑制作用。

... 水杨酸(salicylic acid, SA), 即邻羟基苯甲酸, 植物体内广泛存在, 目前已证明能够调节植物体内的许多抗逆生理过程, 如抗盐性、抗寒性、抗热性、抗旱性以及抗病性^[2,3], 同时作为一种植物激素, SA也是用于增加作物收成的一种重要措施, 具有广阔的应用前景^[4,5,6,7] ...

2012

0.0

张营, 李法云, 严霞. 外源K+和水杨酸在缓解融雪剂对油松幼苗生长抑制中的效应与机理. 生态学报, 2012, 32: 4300-4308

Zhang

, Li F

, Yan

Alleviation effect and mechanism of exogenous potassium nitrate and salicylic acid on the growth inhibition of Pinus tabulaeformis seedlings induced by deicing salts.

Acta Ecol Sin, 2012, 32: 4300-4308 (in Chinese with English abstract)

Large quantities of deicing salts are applied in urban areas for clearing the pavement and ensuring the normal traffic flow during the winter seasons in northeastern China. The salt from deicing operation has been observed to accumulate in roadside soil and inhibit the growth of vegetation. It was reported that the application of exogenous K + and salicylic acid (SA) could protect plants against abiotic stress. In this study, pot experiments were performed in soil culture to test the efficacy of two soil additives, potassium nitrate (KNO 3 ) and SA, in alleviating the stress of deicing salts on growth and photosynthetic parameters of Pinus tabulaeformis seedlings. Three soil additive levels were used i.e. 10, 20, 40 mmol/L for KNO 3 and 2, 4, 8 mmol/L for SA, respectively, under the stress of 0.2% deicing salts per dry weight of soil. After the treatment for 20 days (d), relative water content, leakage rates of electrolyte, malondialdehyde (MDA), peroxidase (POD) and chlorophyll content in leaves were determined. Dry weight of roots, stems and leaves were measured respectively after 60 d of treatments. Three important parameters of photosynthesis, photosynthetic rates ( Pn ), intracellular CO 2 concentrations ( Ci ) and stomatal conductance ( Gs ) were measured every 7 d until 35 d.The results showed that the growth of seedlings was inhibited significantly by the stress of 0.2% deicing salts. The dry weight and relative water content of roots, stems and leaves under 0.2% deicing salt stress were decreased significantly in comparison with the control seedlings. The leaves of Pinus tabulaeformis were more sensitive to the stress of deicing salts compared with roots and stems. However, the treatment of 20 mmol/L KNO 3 and 2mmol/L SA to 0.2% deicing salts had high dry weight and water content. Meanwhile, the MDA content and leakage rates of electrolyte increased once exposure to 0.2% deicing salts, and salts exposure also had an inhibitory effect on the activities of the POD. However, with the addition of 20 mmol/L KNO 3 and 2 mmol/L SA, the high stability of membrane was improved by promoting the POD activities, with decreasing plasmalemma permeability and MDA content in leaves. On the other hand, the contents of Pn, Ci, Gs and the chlorophyll all decreased in the Pinus tabulaeformis seedlings by the treatment of 0.2% deicing salts. The addition of 20 mmol/L KNO 3 significantly increased photosynthetic rates from 5.77% to 20.6% with the prolonging of the exposure time. However, other treatments were found no obvious alleviation effect. It was also found that the contents of Ci and Gs remained unchanged in leaves of Pinus tabulaeformis seedlings in all alleviation treatments, but the enhancing of chlorophyll contents played a key role in maintaining the leaf photosynthetic rate under the treatment of KNO 3 and SA. The results indicated that the salt tolerance of Pinus tabulaeformis seedlings was improved by using soil additives of 20 mmol/L KNO 3 and 2 mmol/L SA. Exogenous K + and SA showed strong benefits against deicing salts-induced negative symptoms.

随着融雪剂在国内外寒冷地区的广泛应用及其在城市使用量的逐年增加,融雪剂对城市生态环境的危害引起了广泛的重视。其中,融雪剂在城市道路土壤中的积累对植物生长的影响已日益凸现。以油松幼苗为材料,通过分析0.2% 浓度融雪剂胁迫下外源钾(K + )和水杨酸(salicylic acid,SA)对油松幼苗各生长生理指标的影响,探讨外源K + 和SA在缓解融雪剂对油松幼苗生长抑制中的机理与剂量效应关系。结果表明,0.2% 浓度的融雪剂处理对油松生长有明显的抑制作用,而20 mmol/L KNO 3 和2 mmol/L SA能明显诱导过氧化物酶(peroxidase,POD)活性的增强,缓解膜脂过氧化作用,降低丙二醛(malondialdehyde,MDA)在叶片中的积累,维持细胞膜的稳定性。虽然外源K + 和SA对油松幼苗叶片胞间CO 2 浓度(intracellular CO 2 concentrations, Ci )和气孔导度(stomatal conductance, Gs )的缓解作用并不显著,但其可通过提高叶绿素含量促进光合作用的进行,缓解融雪剂胁迫对油松幼苗生长的抑制,分别增加生物量24.9%和63.6%。可见,20 mmol/L KNO 3 和2 mmol/L SA处理能有效缓解融雪剂对油松幼苗的伤害,为城市化学融雪剂的污染防治提供科学依据。

2010

2.578

0.0

2012

0.0

1.742

郝敬虹, 易旸, 尚庆茂, 董春娟, 张志刚. 干旱胁迫下外源水杨酸对黄瓜幼苗膜脂过氧化和光合特性的影响. 应用生态学报, 2012, 23: 717-723

Hao J

, Yi

, Shang Q

, Dong C

, Zhang Z

Effects of exogenous salicylic acid on membrane lipid peroxidation and photosynthetic characteristics of Cucumis sativus seedlings under drought stress.

Chin J Appl Ecol, 2012, 23: 717-723 (in Chinese with English abstract)

To approach the related mechanisms of exogenous salicylic acid (SA) in improving plant drought-resistance, this paper studied the effects of applying exogenous SA to the rhizosphere on the plant growth, membrane lipid peroxidation, proline accumulation, water use efficiency, net photosynthetic rate (Pn), and chlorophyll fluorescence parameters of cucumber (Cucumis sativus) seedlings under drought stresses (60% and 50% of saturated water capacity). Applying SA relieved the inhibitory effects of drought stress on plant growth, Pn, and water use efficiency, decreased membrane lipid peroxidation, and promoted proline accumulation. Meanwhile, the SA decreased the decrements of the maximum photochemical efficiency of PSⅡ, actual photochemical efficiency of PSⅡ, potential activity of PSⅡ, effective photochemical efficiency of PSⅡ, and photochemical quenching coefficient under drought stress significantly, and limited the increase of non-photochemical quenching coefficient. All the results suggested that applying exogenous SA could alleviate the oxidation damage of cell membrane resulted from the drought-caused membrane lipid peroxidation, improve the Pn by increasing PSⅡ activity to benefit water utilization, enhance the regulation capability of osmosis to decrease water loss and increase water use efficiency, and thereby, improve the plant drought-resistance.

为了探讨外源水杨酸(SA)提高植物抗旱性的相关机制，研究了干旱胁迫下(基质含水量为饱和持水量的60%和50%)，根际施用外源SA对黄瓜幼苗生长、膜脂过氧化、脯氨酸积累、水分利用效率、净光合速率(Pn)和叶绿素荧光参数的影响.结果表明： SA处理能够缓解干旱胁迫对黄瓜幼苗生长、Pn和水分利用效率的抑制，减小膜脂过氧化程度，促进脯氨酸的积累；添加外源SA显著减小了干旱胁迫下黄瓜幼苗的PSⅡ最大光化学效率、PSⅡ实际光化学效率、PSⅡ潜在活性、PSⅡ有效光化学效率和光化学猝灭系数的下降幅度，抑制了非光化学猝灭系数的升高.添加外源SA可以缓解干旱胁迫造成的膜脂过氧化对膜系统的氧化损伤，并通过增强PSⅡ反应中心活性提高了Pn，有助于水分的利用，同时增大渗透调节能力，减少水分的散失，提高水分利用效率，从而增强植株对干旱的适应能力.

2011

0.0

易小林, 杨丙贤, 宗学凤, 李名扬. 信号分子水杨酸减缓干旱胁迫对紫御谷光合和膜脂过氧化的副效应. 生态学报, 2011, 31: 67-74

Yi X

, Yang B

, Zong X

, Li M

Signal chemical salicylic acid mitigates the negative effects of drought on photosynthesis and membrane lipid peroxidation of purple majesty.

Acta Ecol Sin, 2011, 31: 67-74 (in Chinese with English abstract)

为研究信号分子水杨酸(SA)对干旱胁迫下紫御谷光合和膜脂过氧化的影响,为SA应用于紫御谷抗旱育苗提供理论依据,测定分析了SA处理对干旱胁迫下紫御谷幼苗叶片光合和膜脂过氧化相关指标的变化.结果表明:(1)干旱胁迫破坏了紫御谷叶绿体的膜结构,使基粒数量明显减少,垛叠不明显,排列比较松散,而SA处理能在一定程度上保护叶绿体的膜结构.(2)干旱胁迫降低了紫御谷幼苗叶片的叶绿素b和总叶绿素含量,而SA处理能提高干旱胁迫下紫御谷幼苗叶片的叶绿素a、叶绿素b、总叶绿素和胡萝卜素含量.(3)干旱胁迫降低了叶片的净光合速率、气孔导度、蒸腾速率、光饱和点和暗呼吸速率,增加了叶片的光补偿点、CO2饱和点、CO2补偿点和光呼吸速率,而SA处理则增加了紫御谷幼苗叶片的净光合速率、气孔导度、光饱和点和暗呼吸速率,降低了紫御谷幼苗叶片的光补偿点、CO2饱和点、CO2补偿点和光呼吸速率.(4)干旱胁迫降低了紫御谷幼苗叶片的Fv/Fm和φPSⅡ,显著增加了叶片的相对电导率和丙二醛含量,而SA处理则增加了幼苗叶片的Fv/Fm和φPSⅡ,降低了叶片的相对电导率和丙二醛含量.表明:信号分子水杨酸能够有效减缓干旱胁迫对紫御谷光合和膜脂过氧化的影响.

2013

3.473

0.0

1999

0.599

0.0

... 重金属胁迫对植物造成毒害的机制主要是重金属离子取代植物必需营养元素与酶活性中心或蛋白中心的巯基结合, 或是干扰Ca²⁺等元素的运输并启动氧化还原反应产生自由基, 造成植物的氧化损伤, 进而造成植株代谢紊乱^[8] ...

2003

6.555

0.0

... 铅、镉等重金属胁迫能使黄瓜、大麦等幼苗的内源SA水平增加^[9,10], Freeman等^[11]等发现拟南芥中自由态SA含量的升高, 能够激发丝氨酸乙酰转移酶活性, 进而使植株体内谷胱甘肽含量升高, 增强抗逆性 ...

2006

0.0

0.451

刘素纯, 萧浪涛, 廖柏寒, 匡逢春, 黄运湘. 铅、镉胁迫对黄瓜幼苗内源水杨酸的影响. 湖南农业科学, 2006, (3): 25-28

Liu S

, Xiao L

, Liao B

, Kuang F

, Huang Y

Effects on endogenous salicylic acid of cucumber seedling sunder lead and cadmium stress.

Hunan Agric Sci, 2006, (3): 25-28 (in Chinese with English abstract)

以黄瓜幼苗为材料,采用高效液相色谱(HPLC)法测定黄瓜幼苗内源游离水杨酸含量,研究铅、镉胁迫下黄瓜幼苗内源游离水杨酸的动态水平.结果表明:HPLC法采用290 nm波长检测,有较高的灵敏度,适宜微量样品中水杨酸的检测.铅、镉胁迫下随着质量浓度增加黄瓜幼苗内源水杨酸含量逐渐增加,其POD活性逐渐下降.

2005

6.555

0.0

2005

2.922

0.0

... Drazic和Mihailovic^[12]研究SA对Cd胁迫下大豆幼苗对营养元素吸收规律的影响, 发现SA能减低K含量却促进了Fe的吸收, 同时也促进了Mg和Ca的吸收与运输, 使植物体内各元素间达到动态平衡, 提高机体抗胁迫的能力 ...

2009

0.0

张艳英, 周楠, 刘鹏, 徐根娣, 陈文荣, 蔡妙珍. 铜胁迫对烟草幼苗氮代谢的影响. 生态学报, 2009, 29: 6779-6784

Zhang Y

, Zhou

, Liu

, Xu G

, Chen W

, Cai M

Effect of Cu stress on nitrogen metabolism of Nicotiana tabacum L. seedling.

Acta Ecol Sin, 2009, 29: 6779-6784 (in Chinese with English abstract)

Under solution culture, used Cu-tolerant cultivar 89112 and sensitive cultivar Shuang-70 of Nicotiana labacum L. as plant material, five Cu levels (0, 0.5, 1, 5, and 10 mg?L- 1) were applied to study the effects of copper on nitrogen metabolism. The results showed that with the increase of Cu concentration, the contents of NO-3 -N in two Nicotiana tabacum cultivars were increased firstly and then decreased, and the NO-3 -N contents in 89112 were higher than in Shuang-70. The negative effects of Cu on the assimilation of NO-3 -N were increased with the treated length., The activities of GS and GOGAT were increased firstly and then were inhibited by the increased Cu. Then, as the treated length prolonged, the inhibitory effects were increased. In constrast to Shuang-70, the enzyme activities were higher in 89112, implicating that the 89112 is more tolerant to high Cu stress. At low Cu lever, activity of glutamate-pyruvate transaminase (GPT) was increased, and content of the free amino acids was reduced. The decline of the free amino acid contents in Shuang-70 was more rapidly. Compared with the control, protein content was increased by Cu stress. In short, it is indicated that Cu stress may enhance nitrogen transformation metabolism in Nicotiana tabacum L. seedling.

以耐铜的‘89112’与铜敏感的‘双-70’的两个烟草品种作为试验材料，在水培条件下研究了不同质量浓度的铜（0，0.5，1，5，10 mg?L-1）对烟草幼苗氮代谢的几个关键酶及相关物质含量的影响。结果表明：两个品种的烟草硝态氮(NO-3 -N)含量均随着铜质量浓度的增加呈先增加后降低的趋势，品种89112硝态氮的含量明显比双-70的高，且铜处理时间越长对烟草硝态氮的同化抑制越大；GS、GOGAT的活性均随着铜质量浓度的增加，呈先上升后降低的趋势，且处理时间越长活性越低，品种89112的酶活性的要高于品种双-70的，表明89112对铜胁迫的耐性较强些；低质量浓度铜条件下谷丙转氨酶(GPT)活性与对照相比有所提高，但却降低了游离氨基酸含量，且在品种双-70的降低幅度更明显；同时铜处理也提高了烟草幼苗蛋白质含量。

... 3 讨论Cu是植物正常生命活动所必需的微量元素之一, 对维持植物正常的新陈代谢及生长发育均具有极其重要的意义, 然而, 适合植物生长的铜浓度范围却很窄, 土壤中稍微过量的Cu便会干扰细胞的正常代谢及离子平衡继而会对植物造成严重的毒害作用^[13] ...

2012

0.0

张小梅, 王智鑫, 李广录. 水杨酸对铬胁迫下绿豆幼苗生长抑制的缓解效应. 河南农业科学, 2012, 41(11): 52-54

Zhang X

, Wang Z

, Li G

Mitigating effects of SA on inhibition of Cr²+ stress to mung bean seedling growth.

Henan Agric Sci, 2012, 41(11): 52-54 (in Chinese with English abstract)

为探讨水杨酸(SA)对铬胁迫下绿豆幼苗生长抑制的缓解效应,采用水培法研究了不同质量浓度SA对10 mmol/L铬胁迫下绿豆幼苗生长的影响.结果表明:0.25～0.50 mg/L SA均能提高铬胁迫下绿豆幼苗的株高、鲜质量、根长、根系活力指数、叶绿素含量,降低过氧化产物MDA含量,明显缓解铬胁迫对绿豆幼苗生长的抑制.其中,以0.50 mg/L的SA缓解效果最好.因此,利用SA可以缓解铬胁迫对绿豆幼苗的伤害.

... 已有的研究表明, SA对植物抗性的诱导具有浓度效应, 表现为低促高抑^[14] ...

2007

0.0

计汪栋, 施国新, 张慧, 徐勤松, 许晔, 杜开和. 菹草对Hg²+胁迫的生理和结构应答反应. 生态学报, 2007, 27: 2856-2863

Ji W

, Shi G

, Zhang

, Xu Q

, Xu

, Du K

Physiolgical and ultrastructural responses of Potamogeton crispus to Hg²+ stress.

Acta Ecol Sin, 2007, 27: 2856-2863 (in Chinese with English abstract)

Potamogeton crispus plants exposed to 1, 3, 5, 7mg/L Hg 2+ for 7d were analyzed with reference to its toxic impact on mineral nutrition adsorption, total chlorophyll content, soluble protein, lipid peroxidation (MDA), proline, GSH and ultrastructure under lab conditions. The results revealed that, with the rise of Hg 2+ concentration in culture medium, (1)the content of chlorophyll and soluble protein decreased gradually, while that of MDA and free proline increased and GSH content enhanced progressively up to 5 mg/L Hg 2+ and followed by a decline at higher dose. (2)The addition of Hg 2+ also affected the absorption of mineral nutrients; it mainly increased the absorption of Ca 2+ , Fe 3+ , Mg 2+ and Zn 2+ , but reduced that of P and K + . (3) The amount and intensity of protein/peptide band decreased gradually with augment of Hg 2+ concentration was observed in SDS-PAGE, respectively, and two polypeptides with apparent molecular weights 83.6kDa and 51.3kDa disappeared significantly in fronds treated with 5 and 7 mg/L Hg 2+ . (4) Transmission electron microscope observation indicated that Hg 2+ imposed injury action on ultrastructure of leaf cells, such as swelling of chloroplast, breakage and disappearance of chloroplast envelope and disaggregation in the end. The conclusion could be reached that the death of plant was resulted from destruction of structure foundation of physiological function, unbalance of ion equilibrium and disorder of physiological metabolism. The lethal concentration scope of Hg 2+ to Potamogeton crispus was 1～3 mg/L based on the experiment results.

研究了不同处理浓度Hg 2+ 对菹草叶片矿质营养吸收、叶绿素含量、可溶性蛋白、丙二醛含量、脯氨酸含量、谷胱甘肽含量以及细胞超微结构的毒害影响。结果表明，随着Hg 2+ 浓度的增加，(1)菹草叶片叶绿素含量和可溶性蛋白含量逐渐下降，丙二醛含量和游离脯氨酸含量逐渐上升，谷胱甘肽含量则先升后降；(2)Hg 2+ 对菹草的矿质营养吸收也产生了影响，主要是促进对Ca2+、Fe3+、Mg2+、Zn2+的吸收，降低对大量元素P、K+的吸收；(3)SDS-PAGE蛋白电泳图的条带随Hg 2+ 浓度增加而逐渐减少，亮度也随之减弱，并且在5mg/L和7mg/L Hg 2+ 时导致了分子量为83.6kDa和51.3kDa多肽明显丢失；(4)电镜观察发现：随Hg 2+ 浓度的增加，对细胞超微结构的损伤程度也加剧，表现为被膜断裂、消失和叶绿体膨大、解体。Hg 2+ 破坏了菹草正常生理活动的结构基础和离子平衡，并造成功能紊乱。这些都是Hg 2+ 对菹草产生毒害的重要原因。Hg 2+ 对菹草的致死浓度范围应在1~3mg/L。

... 营养元素的吸收平衡被破坏是重金属植物毒理学效应的一个重要方面^[15] ...

1995

6.555

0.0

... Cu、K、Ca、Mg、Fe、Zn、B和Mn都是植物正常生长和代谢所必需的营养元素, 其中Cu是许多氧化酶的成分及某些酶的活化剂, 在植物的光合作用、呼吸作用等方面的氧化还原过程中有着非常重要的作用^[16], K提高植物的抗逆性, 能增强碳水化合物的合成和转运, 能激活多种酶, 能调节渗透浓度, 另外, K能促进蛋白质和核酸的形成 ...

2010

1.265

0.0

郑国红, 胡婵娜, 刘鹏. 外源钾对铁胁迫下水稻元素吸收运输规律的影响. 水土保持学报, 2010, 24(5): 141-145

Zheng G

, Hu C

, Liu

Effects of exogenous kalium on elements absorption and transportation of rice under iron toxicity.

J Soil Water Conserv, 2010, 24(5): 141-145 (in Chinese with English abstract)

以耐铁毒型水稻品种协优9308和铁毒敏感型品种Ⅱ优838为实验材料.采用水培法研究250 mg/L Fe2+胁迫下,不同浓度的K+对水稻的生长特性以及不同器官和亚细胞的元素吸收运输规律的影响.结果表明,在250 mg/L Fe2+胁迫下,两品种水稻生长均受到严重抑制,Fe含量显著上升,K、Na、Ca、Mg、Zn的吸收与运输受到抑制.施加外源钾可以不同程度的缓解铁毒,有效解除铁胁迫对水稻生长的抑制作用,降低水稻体内Fe含量,并促进营养元素K、Na、Ca、Zn的吸收,但对Mg元素的影响并不明显.在亚细胞分布中,Fe、K主要分布于细胞壁组分F1和以液泡为主的可溶性组分F5中.由此可知,高Fe2+胁迫抑制水稻生长,营养元素的吸收以及向地上部的运输,施加适宜浓度的外源钾有利于营养元素的吸收运输从而促进水稻生长,在一定程度上缓解铁毒.

... , 在多种细胞信号转导途径中起着重要的作用, 是参与细胞构建及代谢必需的营养元素, Mg是植物叶绿素的重要组成成分之一, 在光合作用和呼吸作用过程中可以活化各种磷酸变位酶和磷酸激酶, 且Mg也可以活化DNA和RNA的合成过程^[17], Fe作为植物必需的微量营养元素, 是许多重要酶的辅基, 如细胞色素、过氧化物酶等, Zn是植物多种代谢途径中关键酶的组成因子, 同样也是叶绿素合成的必需元素, B的主要生理功能是参与糖的转运与代谢, 可以调节植物细胞壁的形成、N代谢等生理过程, Mn是多种酶的活化剂, 可以直接参与植物的光合作用过程, 同时对氧化还原反应中的电子传递有着非常重要的影响 ...

1993

6.555

0.0

... K、Fe在植物体内参与多种酶的合成, 同时在植物体的吸收、代谢、蛋白质合成等过程中不可或缺^[18] ...

1997

0.0

... 可能机制之二是Cu²⁺胁迫影响根系活力, Cu²⁺浓度超过一定值时会明显降低根系活力^[19,20], 对烟草发育产生毒害, 本试验中K、Ca、Mg、Fe、Zn、B和Mn的吸收在Cu²⁺胁迫时明显下降, 主要原因可能是根系活力减弱导致养分吸收受阻 ...

2004

2.922

0.0

2009

3.137

0.0

... 在植物体内蛋白质、核酸等重要物质的合成和代谢都离不开K、Mg的参与, 它们在植物体内含量的降低必将导致植物的代谢紊乱^[21], 这与SA减轻Cu对细胞膜结构的损伤有关 ...

2012

0.0

... 此外, 外源SA对Cu胁迫下W38和本氏烟营养元素吸收均有很好的促进效果, 且对铜敏感型品种本氏烟的营养元素吸收的促进作用更为显著, 这与周媛^[22]的结果较为一致 ...