酸胁迫对夏枯草叶绿素荧光特性和根系抗氧化酶活性的影响

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张利霞, 常青山, 薛娴, 刘伟, 张巧明, 陈苏丹, 郑轶琦, 李景林, 陈婉东, 李大钊. 酸胁迫对夏枯草叶绿素荧光特性和根系抗氧化酶活性的影响. 草业学报, 2020,29(8): 134-142
ZHANG Li-xia, CHANG Qing-shan, XUE Xian, LIU Wei, ZHANG Qiao-ming, CHEN Su-dan, ZHENG Yi-qi, LI Jing-lin, CHEN Wan-dong, LI Da-zhao. Effects of acid stress on chlorophyll fluorescence characteristics and root antioxidant activity of Prunella vulgaris . Acta Prataculturae Sinica,2020,29(8): 134-142 复制到剪切板

Doi:10.11686/cyxb2019466
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酸胁迫对夏枯草叶绿素荧光特性和根系抗氧化酶活性的影响

张利霞¹, 常青山^2,^*, 薛娴¹, 刘伟¹, 张巧明², 陈苏丹², 郑轶琦², 李景林², 陈婉东², 李大钊³

1.河南科技大学农学院,河南洛阳 471000

2.河南科技大学林学院,河南洛阳 471000

3.河南科技大学食品与生物工程学院,河南洛阳 471000

*通信作者Corresponding author. E-mail: hkdcqs@126.com

作者简介:张利霞(1982-),女,河南荥阳人,讲师,博士。E-mail: hkdzlx@126.com

收稿日期: 2019-10-30

基金项目:国家自然科学基金项目(31870093,31800096),河南省自然科学基金项目(182300410082,182300410050,182300410017),河南省科技攻关计划项目(182102110401),河南科技大学培育基金(4024-13350073)和河南科技大学大学生研究训练计划(SRTP)项目(2018162,2019422)资助

摘要

为了阐明夏枯草幼苗对酸性环境的耐受性,采用不同pH值(pH=4.5、4.0、3.5、3.0)的Hoagland营养液模拟酸胁迫环境,研究酸胁迫对夏枯草幼苗叶绿素荧光特性及根系抗氧化酶活性的影响。结果表明,随着pH值下降,代表供体与受体侧参数的K点相对可变荧光( W_K)、J点相对可变荧光( V_J)和电子受体QA-被还原的最大速率(d V/d t_o)逐渐升高,光系统Ⅱ(PSⅡ)电子传递的量子效率( φE_o)降低;表示单位反应中心吸收( ABS/ RC)、捕获( TR_o/ RC)、热耗散( DI_o/ RC)的能量逐渐增加;用于电子传递的能量( ET_o/ RC)逐渐下降;表示PSⅡ的最大光化学效率( F_v/ F_m)、吸收光能为基础的光合性能指数( PI_abs)、表征光系统Ⅰ(PSⅠ)的最大氧化还原活性( ΔI/ I_o)及PSⅡ与PSⅠ协调性( Φ_PSⅠ/PSⅡ)随酸胁迫程度增加逐渐下降。夏枯草根系超氧化物歧化酶(SOD)与过氧化物酶(POD)活性呈下降趋势,而根系抗坏血酸过氧化物酶(APX)活性先升高后降低;夏枯草叶片可溶性糖、脯氨酸含量逐渐上升。酸胁迫下,夏枯草PSⅡ反应中心受损,电子传递受阻,光系统性能及PSⅡ与PSⅠ的协调性降低,根系SOD与POD活性受到抑制,在pH 3.0处理下表现较为明显。夏枯草通过提高单位反应中心的热耗散能力来减少过剩光能对叶片光合机构的破坏,通过提高根系APX抗氧化酶及叶片渗透调节物质含量来减少酸胁迫带来的伤害。

关键词: 夏枯草; 光系统; 酸胁迫; 抗氧化酶; 叶绿素荧光

Effects of acid stress on chlorophyll fluorescence characteristics and root antioxidant activity of Prunella vulgaris

ZHANG Li-xia¹, CHANG Qing-shan^2,^*, XUE Xian¹, LIU Wei¹, ZHANG Qiao-ming², CHEN Su-dan², ZHENG Yi-qi², LI Jing-lin¹, CHEN Wan-dong¹, LI Da-zhao³

1.College of Agriculture, Henan University of Science and Technology, Luoyang 471000, China

2.College of Forestry, Henan University of Science and Technology, Luoyang 471000, China

3.College of Food and Bioengineering, Henan University of Science and Technology, Luoyang 471000, China

Abstract

This research aimed to elucidate the tolerance of Prunella vulgaris seedlings to acid environments. Hoagland nutrient solutions with a range of pH values (pH=4.5, 4.0, 3.5, 3.0) were used to simulate environmental acid stress, and the effects of acid stress on chlorophyll fluorescence characteristics and root antioxidant activity were studied. It was found that at lower pH the quantum efficiency of PSⅡ electron transfer ( φE_o) in leaves of P. vulgaris decreased, while a number of other fluorescence parameters increased, including: the variable fluorescence F_k to the amplitude F_j- F_o ( W_K), relative variable fluorescence at J step ( V_J),QA- maximum rate of reduction (d V/d t_o). The absorption flux per reaction center ( ABS/ RC), trapped energy flux per RC ( TR_o/ RC) and dissipated energy flux per RC ( DI_o/ RC) increased, but electron transport flux per RC ( ET_o/ RC) decreased gradually. The maximum quantum yield of PSⅡ ( F_v/ F_m), the photosynthetic index based on absorption basis ( PI_abs), the maximum PSⅠ redox activity ( ΔI/ I_o), and the coordination of photosystem between Ⅰ and Ⅱ ( Φ_PSⅠ/PSⅡ) all decreased under acid stress. The activity of superoxide dismutase (SOD) and peroxidase (POD) in roots decreased, but the activity of ascorbic acid peroxidase (APX) in roots increased on initial exposure to acid stress and then decreased. The soluble sugar and proline concentrations in leaves of P. vulgaris increased with the decrease in pH value. In summary, the PSⅡ reaction center of P. vulgaris was damaged, the electron transfer was blocked, the photosystem performance and the coordination of photosystem between Ⅰ and Ⅱ were reduced under acid stress, and the activities of SOD and POD in roots were inhibited, the latter being most obvious in the pH 3.0 treatment. P. vulgaris was able to reduce the damage to the photosynthetic mechanism by increasing the heat dissipation capacity of unit reaction center, and reduce the damage caused by acid stress by increasing the APX activity in roots and the concentration of osmolytes in leaves.

Keyword: Prunella vulgaris; photosystem; acid stress; antioxidant enzymes; chlorophyll fluorescence

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近年来, 随着世界经济发展、工业化进程加快, 酸雨污染问题日益严重, 其对生态系统及人类生活环境已造成严重影响, 成为全球范围内亟待解决的重要环境问题之一^[1]。中国近年来酸雨发生面积不断扩大, 几乎成燎原之势, 继欧洲和北美之后, 跻身世界第三大酸雨区^[2]。我国强酸雨区的面积较大, 长江以南地区是我国主要的酸雨控制区^[3]。植物是陆地生态系统中酸雨污染最直接的受体, 酸雨对植物生长发育的作用机理及植物对酸雨的抗性机制一直以来都是研究的热点^[4]。研究表明, 酸雨直接通过积累活性氧、破坏细胞质膜与叶绿体超微结构、使植物酶失活、降解叶绿素、损坏光合系统, 从而导致植物生长发育不良或畸形^[5]。同时, 酸雨沉降至地面, 会导致土壤酸化, 抑制土壤呼吸和植物根系的生长, 加重土壤养分流失, 从而使植物根毛对水分和养分的吸收能力显著下降^[6], 造成植物的生长发育受到抑制, 并导致农作物的产量与品质下降。尤其是在幼苗期, 酸雨对植物生长发育的抑制作用尤为显著^[7]。

夏枯草(Prunella vulgaris)是唇形科夏枯草属的多年生草本植物, 因其果穗具有重要的功效与药理作用, 受到了人们青睐, 在临床上被广泛用于治疗高血压、糖尿病、癌症、肺结核和肝炎等重大疾病^{[8, 9, 10]}。夏枯草资源主要分布于长江中下游地区, 主产区集中分布于江苏、安徽、河南、江西和浙江等地区, 这些地区多是酸雨发生较多的地区^{[2, 11]}。近年来, 夏枯草野生资源急剧减少, 其人工栽培种植是保障夏枯草资源可持续发展的重要举措。目前, 关于环境因子对夏枯草生理特性、产量及品质影响的研究主要集中于水分、养分、光照等因子的调控作用^{[12, 13]}, 而对于酸胁迫如何影响夏枯草的内部光系统以及根系抗氧化酶的机制尚不清楚。因此, 本研究通过设置不同pH值营养液, 模拟土壤酸性环境, 研究苗期夏枯草叶绿素荧光特性及根系保护酶活性对酸胁迫环境的响应特征, 以期为阐明夏枯草对酸性环境的耐受机制提供基础数据。

1 材料与方法

1.1 试验材料与培养

2018年10月进行夏枯草育苗:将夏枯草种子(采于河南驻马店)消毒后, 播种于装有等量营养土基质的周转箱(60 cm× 42 cm× 15 cm)中进行育苗, 正常肥水管理。次年, 挑选长至4叶期且长势均衡的幼苗, 于自然光照, 温度为昼夜(30± 3) ℃/(20± 3) ℃的条件下, 放入装有1/2 Hoagland营养液的周转箱中进行泡沫板悬浮培养, 每隔34 d换一次营养液。

1.2 试验设计

待夏枯草幼苗长到68叶期时, 选择大小相当的幼苗进行酸胁迫试验, 每个处理30株。根据前期预实验结果对夏枯草幼苗进行相应的酸胁迫处理。用硫酸和硝酸的混合液(按摩尔浓度5∶ 1)调制到相应pH值配制模拟酸胁迫母液, 与1/2Hoagland营养液混合配制成pH分别为4.5, 4.0, 3.5, 3.0的模拟酸溶液, pH 6.5为不含酸液的1/2Hoagland营养液。每个处理的营养液均每3 d更换一次。连续胁迫处理7 d后进行夏枯草荧光与生理指标的测定。

1.3 测定指标及方法

快速荧光诱导动力学参数的测定:夏枯草叶片经过充分暗适应后, 利用多功能植物效率分析仪(M-PEA, Hansatech, 诺福克, 英国)经3000 μ mol· m^-2· s^-1的脉冲光诱导, 测定夏枯草叶片快速叶绿素荧光诱导动力学曲线, 根据文献[14]方法计算JIP-test参数:光系统Ⅱ (photosystem Ⅱ , PSⅡ )最大光化学效率(F_v/F_m)、以吸收光能为基础的光合性能指数(performance index on absorption basis, PI_abs)。PSⅡ 电子传递的量子效率(quantum efficiency of PSⅡ electron transfer, φ E_o)、可变荧光F_k占F_j-F_o振幅的比例(ratio of the variable fluorescence F_k to the amplitude F_j-F_o, W_K)、J点的相对可变荧光(relative variable fluorescence at J step, V_J)、电子受体 ${Q_{A}}^{-}$ 被还原的最大速率( ${Q_{A}}^{-}$ maximum rate of reduction, dV/dt_o)。单位反应中心吸收的能量(absorption flux per reaction center, ABS/RC)、单位反应中心捕获的能量(trapped energy flux per RC, TR_o/RC)、单位反应中心传递的能量(electron transport flux per RC, ET_o/RC)、单位反应中心耗散的能量(dissipated energy flux per RC, DI_o/RC)、光系统Ⅰ (photosystem Ⅰ , PSⅠ )的最大氧化还原活性Δ I/I_o=(I_o-I_m)/I_m; PSⅡ 与PSⅠ 的协调性Φ _{PSⅠ /PSⅡ}=(Δ I/I_o)/Ψ _o。

采用氮蓝四唑(nitro-blue tetrazolium, NBT)还原法测定幼苗根系超氧化物歧化酶(superoxide dismutase, SOD)活性, 采用愈创木酚法测定根系过氧化物酶(peroxidase, POD)活性, 利用分光光度计法测定根系抗坏血酸过氧化物酶(ascorbic acid peroxidase, APX)活性, 采用蒽酮硫酸法测定叶片可溶性糖含量, 采用磺基水杨酸法测定叶片脯氨酸含量^[15]。每个处理重复3次, 取平均值。

1.4 数据分析

采用Excel 2007录入数据并计算平均值和标准差, 利用SPSS 16.0进行多重比较(Duncan)分析, P< 0.05表示差异显著。

2 结果与分析

2.1 酸胁迫对夏枯草叶片PSⅡ 供体侧和受体侧的影响

随着pH值的降低, 反应中心吸收的光能用于电子传递的量子效率(φ E_o)呈逐渐下降的趋势, 在pH 4.5和4.0时低于对照, 但差异不显著(图1)。在pH 3.5和3.0时显著低于对照(P< 0.05), 说明酸胁迫处理不同程度地降低了光系统Ⅱ (PSⅡ )的相对电子传递能力。在pH 4.53.5时, 可变荧光F_k占F_j-F_o振幅的比例(W_k)高于对照, 但无显著差异; 只有在pH为3.0时, W_k显著高于其他处理(P< 0.05), 这表明放氧复合体(oxygen evolving complex, OEC)可能在pH 3.0处理下受到显著伤害。

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图1 酸胁迫下夏枯草叶片PSⅡ 电子传递的量子效率、K点的相对可变荧光、J点的相对可变荧光和Q_A^-被还原的最大速率的变化
不同小写字母表示各处理在P< 0.05水平上差异显著, 下同。Fig.1 Changes of chlorophyll fluorescence parameters of quantum efficiency of PSⅡ electron transfer (φ E_o), ratio of the variable fluorescence F_k to the amplitude F_j-F_o (W_k), relative variable fluorescence at J step (V_J) and Q_A^- maximum rate of reduction (dV/dt_o) in P. vulgaris leaves under acid stress Different lowercase letters indicate significant differences in the level of P< 0.05 for each treatment, the same below.

V_J反映了J点的相对可变荧光强度, 可以表征Q_A^-的积累量。随着pH值的降低, V_J值呈逐渐增加趋势, 在pH 3.0处理下显著高于对照(P< 0.05)。dV/dt_o反映Q_A^-被还原的最大速率, dV/dt_o在pH 4.53.5处理下与对照差异不显著, 在pH 3.0处理下显著高于对照(P< 0.05)。pH 4.53.5处理下V_J与dV/dt_o的上升幅度较小, 表明电子传递受到影响较小, 而pH 3.0处理下两者上升幅度较大, 说明该处理下电子传递能力受到显著抑制。

2.2 不同营养液pH值对夏枯草幼苗叶片PSⅡ 比活性参数的影响

酸胁迫增加了夏枯草叶片单位反应中心吸收能量(ABS/RC), pH 4.03.0处理与对照差异显著(P< 0.05), pH 3.0处理上升幅度最大(图2)。单位反应中心捕获能量(TR_o/RC)在pH 4.53.5处理下逐渐增加, 但与对照差异不显著, 在pH 3.0处理下上升幅度较大, 且显著高于对照(P< 0.05)。夏枯草叶片单位反应中心传递的能量(ET_o/RC)总体呈降低趋势, 各处理与对照相比均未达显著差异。随着pH值的降低, 夏枯草叶片单位反应中心的热耗散(DI_o/RC)呈逐渐升高的趋势, 其中pH 4.5处理与对照差异不显著, 其他处理均显著高于对照(P< 0.05)。

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	图2 酸胁迫下夏枯草叶片单位反应中心吸收、捕获、用于电子传递和热耗散的能量变化Fig.2 Changes of absorption flux per reaction center (ABS/RC), trapped energy flux per RC (TR_o/RC), electron transport flux per RC (ET_o/RC) and dissipated energy flux per RC (DI_o/RC) in P. vulgaris leaves under acid stress

2.3 酸胁迫对夏枯草叶片PSⅡ 活性的影响

F_v/F_m, 是指光合机构把吸收的光能用于光化学反应的最大效率。随着pH值的降低, 夏枯草F_v/F_m呈下降的趋势, pH 4.5处理与对照差异不显著, 而pH 4.03.0处理均显著低于对照(图3)。基于光吸收性能的指数(PI_abs)也呈逐渐降低趋势, 除pH 4.5处理与对照差异不显著外, 其他处理均显著低于对照(P< 0.05)。

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	图3 酸胁迫下夏枯草叶片最大光化学效率和光合性能指数的变化Fig.3 Changes of the maximum quantum yield of PSⅡ (F_v/F_m) and performance index on absorption basis (PI_abs) in P. vulgaris leaves under acid stress

2.4 酸胁迫对PSⅠ 性能及光系统协调性的影响

随着pH值的降低, 夏枯草叶片光系统Ⅰ (PSⅠ )的最大氧化还原活性(Δ I/I_o)呈降低趋势, pH 4.53.5处理与对照差异不显著, 而pH 3.0处理显著低于对照(P< 0.05), 但与pH 3.5差异不显著(图4)。酸胁迫降低了夏枯草叶片PSⅡ 与PSⅠ 之间的协调性(Φ _{PSⅠ /PSⅡ}), 其中pH 4.53.5处理与对照差异不显著, 而pH 3.0处理显著低于对照及其他处理(P< 0.05)。说明酸胁迫不但降低了夏枯草叶的PSⅠ 的活性, 也破坏两个光系统之间的协调性(Φ _{PSⅠ /PSⅡ})。

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	图4 酸胁迫下夏枯草叶片PSⅠ 的最大氧化还原活性及叶片PSⅡ 与PSⅠ 的协调性的变化Fig.4 Changes of the maximum PSⅠ redox acitity (Δ I/I_o) and the coordination of photosystem between Ⅰ and Ⅱ (Φ _{PSⅠ /PSⅡ}) in leaves of P. vulgaris under acid stress

2.5 不同pH值对夏枯草幼苗根系抗氧化酶活性的影响

随着pH值的降低, 夏枯草幼苗根系超氧化物歧化酶(SOD)活性呈逐渐下降的趋势(表1)。与对照相比, 不同酸胁迫处理下根系SOD活性均显著降低, 且不同酸处理之间差异显著(P< 0.05)。根系过氧化物酶(POD)活性的变化趋势和SOD活性相类似, 随着pH值下降逐渐降低, 各酸胁迫处理均显著低于对照(P< 0.05)。随着pH值的降低, 根系抗坏血酸过氧化物酶(APX)活性呈先上升后下降的趋势, 但均显著高于对照, 在pH值为3.5时达到最大值, 显著高于其他处理(P< 0.05)。

表1 不同pH值对夏枯草幼苗根系超氧化物歧化酶、过氧化物酶和抗坏血酸过氧化物酶活性的影响 Table 1 Effects of different pH values in nutrient solutions on activities of superoxide dismutase, peroxidase and ascorbic acid peroxidase in roots of P. vulgaris seedlings (U· g^-1· min^-1)

2.6 不同pH值对夏枯草幼苗叶片可溶性糖和脯氨酸含量的影响

夏枯草叶片中可溶性糖含量随着营养液pH值的降低, 呈逐渐上升的趋势, pH值为4.53.5时, 可溶性糖的含量高于对照, 但差异不显著(图5), 在pH值为3.0时达到最大值, 且显著高于对照(P< 0.05)。叶片中脯氨酸的含量变化趋势与可溶性糖含量势相似, 随着营养液pH值的降低呈逐渐上升趋势, pH为4.5时, 与对照无显著差异, pH为4.03.0时, 均显著高于对照。

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	图5 不同pH值对酸胁迫下夏枯草可溶性糖与脯氨酸含量的影响Fig.5 Effects of different pH values in nutrient solutions on the contents of soluble sugar and proline in leaves of P. vulgaris under acid stress

3 讨论

叶绿素荧光诱导动力学参数能够用于描述植物光合作用的机理及光合生理状况, 可作为检测植物遭受胁迫伤害程度的指标^[16]。有研究表明, PSⅡ 在逆境条件下因受胁迫活性下降, 线性电子传递链在传递电子受阻后会导致活性氧的产生与积累^[17]。本研究中, φ E_o不断下降, 说明PSⅡ 的相对电子传递能力下降; V_J与dV/dt_o均逐渐升高, 表明Q_A到Q_B之间的电子传递受阻^[18], 在pH 3.0酸胁迫下的φ E_o显著低于对照, 而V_J和dV/dt_o显著高于对照, 表明该胁迫下PSⅡ 的电子传递能力严重受阻, 在Q_A到Q_B电子传递受到的抑制较重, 并导致Q_A^-的明显积累。W_k, 即K点相对可变荧光, 被人们广泛用于分析逆境胁迫对光合机构PSⅡ 供体侧, 即放氧复合体(OEC)伤害的程度^[19]。在本试验中, W_k随着酸胁迫程度加重而逐渐增加, 表明夏枯草叶片PSⅡ 放氧复合体(OEC)受到了伤害。pH 3.0处理下的W_k的上升幅度高于V_J, 表明该处理下PSⅡ 供体侧伤害程度重于受体侧^[20]。酸胁迫下夏枯草叶片的PSⅡ 性能降低, 必然导致其向PSⅠ 传递电子的能力下降。

ABS/RC用来衡量PSⅡ 反应中心捕光天线色素的大小^[21], 该指标升高说明叶片捕光天线色素增加, 意味着部分PSⅡ 反应中心的失活。ABC/RC和TR/RC增加, 说明叶片通过增加捕光天线色素的大小以及对光的捕获能力, 从而增强单一反应中心对光的吸收与捕获能力。夏枯草叶片单位面积的热耗散(DI_o/RC)在酸胁迫下升高, 用于电子传递的能量(ET_o/RC)减少, 说明叶片减少用于电子传递的量子份额, 通过增加热耗散来将过多的光能耗散掉, 来减少光抑制对叶片产生的伤害。F_v/F_m和PI_abs两个指标反映了植物叶片PSⅡ 反应中心活性。在本研究中, φ E_o下降, V_J、dV/dt_o、W_k与ABS/RC升高, 说明酸胁迫导致PSⅡ 反应中心失活, PSⅡ 电子传递受阻, 进一步造成了F_v/F_m的降低, 夏枯草叶片发生了光抑制, 导致其叶片光能利用率降低^[22]。PI_abs, 光化学性能指数, 能从光能吸收、捕获和电子传递3个方面内容来综合反映PSⅡ 的活性^[23]。本试验中, 由于PSⅡ 的结构和功能受到破坏, 因此导致夏枯草叶片PI_abs的下降, 且叶片PI_abs的下降幅度明显大于F_v/F_m的下降幅度, 说明PI_abs对酸胁迫的反应比F_v/F_m更为灵敏, 本研究与前人研究一致^[16]。

Δ I/I_o表征的是PSⅠ 反应中心的最大氧化还原能力, 可以用来评价植物叶片PSⅠ 性能, 而Φ _{PSⅠ /PSⅡ}表征的是两个光系统之间的协调性^[24]。本试验研究表明, pH 3.0处理下夏枯草的Δ I/I_o显著下降, 说明该胁迫条件下PSⅠ 性能受到显著伤害, 这将抑制电子流从PSⅡ 流向PSⅠ , 引起Q_A^-大量积累, 阻碍Q_A^-之后的电子传递。因此, PSⅡ 与PSⅠ 之间电子传递受阻, 导致pH 3.0处理下的PSⅡ 与PSⅠ 之间的协同性(Φ _{PSⅠ /PSⅡ})明显失衡, 这可能是引起Q_A后续电子传递链效率下降以及整个电子传递链功能降低的主要原因^[25]。

酸胁迫会诱导作物体内产生大量活性氧(ROS), 过量活性氧会破坏细胞膜的结构与功能, 导致细胞生理功能紊乱^{[26, 27]}。植物在遭受酸胁迫时, 会通过维持几种抗氧化酶之间的协调和平衡来提高植株抗活性氧的能力^[28]。本研究中酸胁迫下夏枯草根系的SOD与POD活性不断下降, 可能是由于根系在酸胁迫下产生了大量的活性氧并超出了根系SOD与POD的清除能力, 致使根系SOD、POD活性下降; 这与平邑甜茶在酸化处理一定天数后根系SOD与POD活性下降^[27]的结果相似, 类似的结果也出现在萝卜(Raphanus sativus)与莴笋(Lactuca sativa)的相关研究中^[29]。APX参与抗坏血酸清除过氧化氢的过程^[16], 夏枯草根系APX活性在酸胁迫处理下均表现出高于对照的活性, 这说明夏枯草在酸胁迫条件下可以通过增强根系APX活性, 增强植株根系清除过氧化氢的能力, 缓解ROS对植物根系的伤害, 提高植株抵抗酸胁迫的能力。

前人研究表明, 当OEC活性受到抑制时会造成放氧过程中水的不完全裂解, 导致H₂O₂的产生; PSⅡ 受体侧电子传递受阻会使过剩电子泄露, 细胞中游离的O₂因受泄露的电子攻击而生成超氧阴离子, 这些活性氧与超氧阴离子会造成细胞膜质过氧化增加和电解质渗漏等问题^{[30, 31]}。本研究表明, 酸胁迫处理导致夏枯草叶片的OEC受到伤害, 电子传递受阻, 因此产生的活性氧造成夏枯草叶片细胞膜受到不同程度的伤害。在逆境胁迫条件下, 植物能够通过增加可溶性糖、脯氨酸等渗透调节物质的含量来适应逆境过程^{[32, 33]}。本试验发现夏枯草叶片通过提高酸胁迫条件下的脯氨酸与可溶性糖含量, 来调节细胞液渗透压, 保护细胞膜的稳定性。特别地, 脯氨酸对羟基自由基清除和保护酶也具有一定的保护和稳定作用^[34], 脯氨酸增幅大于可溶性糖增幅, 说明脯氨酸可能已成为主要的渗透调节物质^[35]。由此说明, 夏枯草在酸胁迫条件下, 通过增加可溶性糖、脯氨酸的积累来提高细胞液的浓度, 维持细胞的水势, 与根系的抗氧化酶协同作用, 清除活性氧, 保护光系统和增加细胞膜的稳定性, 以减少酸雨带来的伤害。

综上所述, 酸胁迫造成放氧复合体(OEC)受到伤害, PSⅡ 的光合电子传递受阻, 抑制了PSⅡ 与PSⅠ 的活性, 降低了PSⅡ 与PSⅠ 的协调性, 导致夏枯草SOD与POD抗氧化酶活性下降。夏枯草可以通过提高根系APX活性, 增加叶片可溶性糖与脯氨酸的含量来清除除活性氧, 保护光系统和细胞膜的稳定性, 通过热耗散机制降低过剩光能对光合机构的破坏, 从而减少酸胁迫所造成的伤害。

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2009

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. 2009, 29(9):4956-4966

Effects of UV-B irradiation and acid rain on photosynthetic rate and antioxidant enzyme activities of maize seedlings

酸雨和UV-B对玉米幼苗光合速率和抗氧化酶活性的影响

Long Y , Liu Y , Zhong Z C

龙云, 刘芸, 钟章成

In order to reveal effects of acid rain (AR) and UV-B radiation (UV-B) singly and in combination on plant photosynthesis and anti\|oxidant enzyme activities, the net photosynthetic rate, relative membrane permeability, malondialdehyde (MDA) content, and anti\|oxidative enzyme activities in maize ( Zea mays L. ) leaves were studied under simulated acid rain ( pH=6.5(A0), 3.5(A1), 4.5(A2) ) and enhanced UV-B radiation ( 0(B0), 2.88(B1), 5.76(B2) kJ?m-2?d-1), both singly and in combination treatment in the green\|house. The results indicated that the physiological traits in the maize breeds Yu Nuo 7 and Su YuNuo experienced similar changes under single AR and UV-B treatment. The treatments resulted in reduction of net photosynthetic rate and catalase (CAT) activity, increase of the relative membrane permeability, MDA content and peroxidase (POD) activity, while superoxide dismutase (SOD) activity increased first but decreased afterwards during the treatment. Under the combined stresses, compared with the control treatment (A0B0), the changes of physiological traits in both breeds of maize were similar to those under single factor treatment. For some traits, combined stress displayed a synergistic effect, with significantly stronger effects than those under single factor stress, whereas for most traits it displayed antagonistic effects. The results showed that both enhanced UV-B radiation and decreased pH of simulated acid rain accelerated ROS metabolism rates, which damaged the anti\|oxidative enzyme system and reduction of photosynthesis. Taken together, the dynamic changes of physiological and biochemical indices under the dual stresses (AR+UV-B) were not identical, and none of them showed a simple linear variation, but displayed additive, synergistic or antagonistic effects depending on the different treatment periods and/or degree of stresses, indicating that plants had different mechanisms of resistance and adaptability under different stresses. The two different breeds of maize had different sensitivity to AR and UV-B treatment at different stages. Yu Nuo 7 was in general more tolerant to stresses than Su YuNuo.

以玉米( Zea mays L.)的两个品种渝糯7号和苏玉糯为研究材料, 研究酸雨与UV-B辐射单一因子和复合处理对植物光合作用及抗氧化酶活性的影响机理, 模拟酸雨\[pH6.5(A0), 4.5(A1)和3.5(A2)\]和增强UV-B辐射\[0(B0), 2.88(B1)和576(B2)kJ?m-2?d-1\]及其复合作用下, 玉米叶片光合速率, 膜脂过氧化程度, 抗氧化酶活性的变化。结果表明, 与对照相比, 酸雨或增强UV-B辐射单因素处理下两个品种的玉米幼苗所测生理指标有相似的变化, 基本表现为: 净光合速率和过氧化氢酶（CAT）活性降低, 质膜透性与丙二醛（MDA）含量增加, 过氧化物酶（POD）活性亦增加, 超氧化物歧化酶（SOD）活性随处理时间延长先升高后降低。复合处理下, 两个品种的玉米幼苗所测生理指标与对照相比也与单因素处理表现相似, 且随处理组合的不同, 复合胁迫下各指标的效果可能表现为协同效应, 但更多地表现为拮抗作用。以上结果说明, 模拟UV-B辐射升高和酸雨pH值降低促进了玉米叶片活性氧的代谢速率, 导致了抗氧化酶系统的紊乱, 抑制了光合作用。在复合处理下, 两个因子对不同指标在不同处理时间和复合水平下, 表现出一定的协同或拮抗效应可能与作物对环境变化产生一定的抗逆性和适应性有关。在不同时期两玉米品种对紫外辐射与酸雨敏感性不同。与苏玉糯相比, 渝糯7号对外界环境变化的抗性更大。

... 近年来,随着世界经济发展、工业化进程加快,酸雨污染问题日益严重,其对生态系统及人类生活环境已造成严重影响,成为全球范围内亟待解决的重要环境问题之一^[1] ...

2010

0.0

... 中国近年来酸雨发生面积不断扩大,几乎成燎原之势,继欧洲和北美之后,跻身世界第三大酸雨区^[2] ...

... 夏枯草资源主要分布于长江中下游地区,主产区集中分布于江苏、安徽、河南、江西和浙江等地区,这些地区多是酸雨发生较多的地区^[2,11] ...

2013

0.0

. 2013, 33(24):7615-7622 DOI:10.5846/stxb201312172973

The problems in red soil ecosystem in southern of China and its countermeasures

中国南方红壤生态系统面临的问题及对策

Zhao Q G , Huang G Q , Ma Y Q

赵其国, 黄国勤, 马艳芹

Southern red soil plays an important role for sustainable development of agriculture, even the whole economy, and the rapid increase in people's living standard. But red soil has some weakness, such as acid, thin and sticky, the uneven temporal distribution of precipitation in the region, and the irrational exploitation which leading to soil erosion, degradation, contamination and deterioration of ecological environment in red soil areas, the potential production capacity of red soil resources are not properly exert, and the entire agriculture and sustainable economic development are severely affected. Currently, China Southern red soil ecosystem faces many problems, such as soil erosion, acidification, fertility degradation, seasonal drought, soil pollution, biological degradation, desertification, land desertification, reduction of arable land and system functions decay and so on. In response to these problems, we need to raise awareness of comprehensive measures, to adjust the agricultural structure, optimize the farming system, develop three-dimensional agriculture, control soil and water erosion and pollution, improve soil quality and soil fertility, reconstruct vegetate ecosystem of red soil, establish and improve red soil early-warning system, so as to ensure steady and healthy development of the southern red soil ecosystem.

南方红壤对于我国农业乃至整个国民经济的持续发展和人民生活水平的迅速提高发挥了重要作用，但由于红壤性质上的酸、瘦、粘等弱点，分布区域降水时空不均匀，以及不合理开发利用造成的水土流失、土壤退化、土壤污染等，导致红壤地区的生态环境恶化，红壤资源潜在的生产能力得不到应有的发挥，使整个地区农业及经济持续发展受到严重影响。目前，中国南方红壤生态系统面临水土流失、土壤酸化、肥力退化、季节性干旱、土壤污染、生物退化、石漠化及土地沙化、耕地面积缩减和系统功能衰减等问题。针对上述问题，在综合治理措施上必须提高认识，调整农业结构，优化耕作制度，大力发展立体农业，防治水土流失和土壤污染，改良土壤，培肥地力，重建红壤区植被生态系统，建立和完善红壤预警，从而保证南方红壤生态系统的持续稳定健康发展。

... 我国强酸雨区的面积较大,长江以南地区是我国主要的酸雨控制区^[3] ...

2000

0.0

... 植物是陆地生态系统中酸雨污染最直接的受体,酸雨对植物生长发育的作用机理及植物对酸雨的抗性机制一直以来都是研究的热点^[4] ...

2011

0.0

... 研究表明,酸雨直接通过积累活性氧、破坏细胞质膜与叶绿体超微结构、使植物酶失活、降解叶绿素、损坏光合系统,从而导致植物生长发育不良或畸形^[5] ...

2019

0.0

... 同时,酸雨沉降至地面,会导致土壤酸化,抑制土壤呼吸和植物根系的生长,加重土壤养分流失,从而使植物根毛对水分和养分的吸收能力显著下降^[6],造成植物的生长发育受到抑制,并导致农作物的产量与品质下降 ...

2017

0.0

. 2017, 37(16):5418-5427 DOI:10.5846/stxb201605230987

Effects of ectomycorrhizal fungi on seedlings of Pinus massoniana under simulated acid rain

野外模拟酸雨胁迫下接种外生菌根真菌对马尾松幼苗的缓解作用

Yu H , Chen Z , Shang H

于浩, 陈展, 尚鹤

Many tree species in forest ecosystems have symbiotic relationships with ectomycorrhizal fungi (ECM), which protects their hosts from environmental stress. Effects of ECM on growth, elemental nutrients, and rhizosphere soil of Pinus massoniana seedlings under simulated acid rain were analyzed in this study to provide a scientific basis for restoration of P. massoniana in an area with severe acid rain in China. Two-year-old seedlings of P. massoniana were grown in forest soil from one of six treatments, including pH 5.6 (CK) inoculated with ECM, CK without ECM, pH 4.5 inoculated with ECM, pH 4.5 without ECM, pH 3.5 inoculated with ECM, and pH 3.5 without ECM. The results showed that:(1) For the P. massoniana seedlings grown in soil not inoculated with ECM, the acid rain treatments significantly reduced leaf area, total biomass, and biomass of roots, stems, and needles, whereas plant height did not differ from that of plants grown in CK. Inoculation with ECM removed the negative effects of acid rain on growth. (2) For the P. massoniana seedlings grown in soil without ECM, the N, P, and Ca content was higher in needles and decreased in roots in the acid rain treatments than those of plants grown in CK. The content of Mg decreased in needles in acid rain treatments, whereas it increased at first and then decreased in roots with a decrease in pH with respect to that of plants grown in CK. Inoculation with ECM significantly increased N, P, Ca, and Mg content of roots; however, there was no effect on N, P, Ca, and Mg content of the needles in the pH 3.5 treatment. (3) For plants grown in soil without ECM, pH 3.5 treatment decreased the content of soil organic matter, available P, available K, soluble carbon, soluble nitrogen, nitrate nitrogen, and ammonium nitrogen, but inoculation with ECM improved soil quality. Acid rain treatments did not significantly affect the cation exchange capacity. These results indicated that inoculation with ECM could stimulate growth of P. massoniana seedlings and effectively alleviate adverse effects on nutrients and rhizosphere soil of P. massoniana seedlings caused by acid rain. Therefore, inoculation with ECM could be an effective way to decrease the hazard of acid rain to Pinus massoniana .

外生菌根真菌能够提高宿主植物对外界环境胁迫的抵抗力。主要探讨野外条件下外生菌根真菌对酸雨胁迫下马尾松（ Pinus massoniana ）幼苗生长、养分元素以及表层土壤的影响，以期为酸雨严重区马尾松林恢复提供科学依据。以2年生马尾松幼苗为材料，采用原位试验，共设置6个处理：pH5.6（对照）处理未接种、对照处理接种、pH4.5酸雨处理未接种、pH4.5酸雨处理接种、pH3.5酸雨处理未接种、pH3.5酸雨处理接种。研究表明：（1）酸雨处理与对照处理相比显著降低了非菌根苗总生物量及各部位生物量（根、茎、叶），对株高无显著影响，接种外生菌根真菌可以缓解酸雨对马尾松幼苗生长的不利影响；（2）与对照处理相比，酸雨处理的非菌根苗的针叶中N、P、Ca含量升高，Mg含量降低，根系中N、P、Ca含量降低，Mg含量随pH的降低先升高后降低。接种外生菌根真菌显著提高了pH3.5酸雨处理的马尾松幼苗根系中N、P、Ca、Mg含量，而对针叶中N、P、Ca、Mg含量无显著影响。（3）在非菌根土壤中，pH3.5酸雨处理与对照处理相比显著降低了土壤中有机质、速效磷、速效钾、可溶性碳、可溶性氮、铵态氮、硝态氮含量，而接种外生菌根真菌显著提高了上述指标。酸雨对土壤阳离子交换量无显著影响。总而言之，接种外生菌根真菌促进了酸雨处理的马尾松幼苗生长、缓解了酸雨对马尾松幼苗养分元素和表层土壤的不利影响，由此可见接种外生菌根真菌是减轻酸雨对马尾松危害的一个重要途径。

... 尤其是在幼苗期,酸雨对植物生长发育的抑制作用尤为显著^[7] ...

2012

0.0

... 夏枯草(Prunella vulgaris)是唇形科夏枯草属的多年生草本植物,因其果穗具有重要的功效与药理作用,受到了人们青睐,在临床上被广泛用于治疗高血压、糖尿病、癌症、肺结核和肝炎等重大疾病^[8,9,10] ...

2011

0.0

2018

0.0

2011

0.0

... 夏枯草资源主要分布于长江中下游地区,主产区集中分布于江苏、安徽、河南、江西和浙江等地区,这些地区多是酸雨发生较多的地区^[2,11] ...

2011

0.0

... 目前,关于环境因子对夏枯草生理特性、产量及品质影响的研究主要集中于水分、养分、光照等因子的调控作用^[12,13],而对于酸胁迫如何影响夏枯草的内部光系统以及根系抗氧化酶的机制尚不清楚 ...

2013

0.0

1995

0.0

... s^-1的脉冲光诱导,测定夏枯草叶片快速叶绿素荧光诱导动力学曲线,根据文献[14]方法计算JIP-test参数:光系统#cod#x02161 ...

2012

0.0

... 采用氮蓝四唑(nitro-blue tetrazolium, NBT)还原法测定幼苗根系超氧化物歧化酶(superoxide dismutase, SOD)活性,采用愈创木酚法测定根系过氧化物酶(peroxidase, POD)活性,利用分光光度计法测定根系抗坏血酸过氧化物酶(ascorbic acid peroxidase,APX)活性,采用蒽酮硫酸法测定叶片可溶性糖含量,采用磺基水杨酸法测定叶片脯氨酸含量^[15] ...

2018

0.0

... 3 讨论叶绿素荧光诱导动力学参数能够用于描述植物光合作用的机理及光合生理状况,可作为检测植物遭受胁迫伤害程度的指标^[16] ...

... 的结构和功能受到破坏,因此导致夏枯草叶片PI_abs的下降,且叶片PI_abs的下降幅度明显大于F_v/F_m的下降幅度,说明PI_abs对酸胁迫的反应比F_v/F_m更为灵敏,本研究与前人研究一致^[16] ...

... APX参与抗坏血酸清除过氧化氢的过程^[16],夏枯草根系APX活性在酸胁迫处理下均表现出高于对照的活性,这说明夏枯草在酸胁迫条件下可以通过增强根系APX活性,增强植株根系清除过氧化氢的能力,缓解ROS对植物根系的伤害,提高植株抵抗酸胁迫的能力 ...

2013

0.0

. 2013, 46(6):1191-1200 DOI:10.3864/j.issn.0578-1752.2013.06.012

Effects of different temperature and light treatments on photosynthetic system #cod#x02161; in Vitis vinifera L. cv. Cabernet Sauvignon

不同温度/光照组合对#cod#x02018;赤霞珠#cod#x02019;葡萄叶片光系统#cod#x02161;功能的影响

Sun Y J , Fu Y D , Du Y P

孙永江, 付艳东, 杜远鹏

College of Horticultural Science and Engineering, Shandong Agricultural University/State Key Laboratory of Crop Biology, Tai’an 271018, Shandong

【Objective】The combined effects of different temperature and light treatments on the operation of photosynthetic apparatus were investigated to reveal the mechanism of inhibition of photosynthesis.【Method】One-year-old Cabernet Sauvignon grapevines were treated with different temperatures（28, 34, 37 and 40℃）combined with two kinds of light intensities (800, 1600 μmol•m-2•s-1). Gas exchange and chlorophyll a fluorescence parameters were simultaneously analyzed to investigate the effects of different treatments on the PSII function【Result】Despite the treated light intensities, as temperature increased, net photosynthetic rate (Pn), stomatal conductance (Gs), actual PSⅡ efficiency (ΦPSII), electron transport rate (ETR), performance index (PIABS), density of RCs (RC/CS) and the probability that a trapped exciton moves an electron into the electron transport chain beyond QA (ΨEo) decreased; PSⅡ excitation pressure (1-qP), non-photochemical quenching (NPQ) and approximated initial slope of the fluorescence transient (Mo) increased. Under the moderate light intensities, the decrease of Pn was due mainly to stomatal limitation during the temperature from 28℃ to 37℃, and the xanthophyll cycle played an increased role to protect photosystem II. Under strong light stress, 28℃ treatment decreased Pn to the level of the 40℃ treatment of moderate light intensities, meanwhile PIABS reached a lower level.【Conclusion】Under moderate light intensities, the PSⅡacceptor sides were not injured until the temperature reached 37℃; strong light stress, however, began to injure the PSⅡ acceptor sides from 28℃, when the temperature reached or exceeded 37℃, the density of RCs and the acceptor sides were damaged severely which showed PSⅡcould be more susceptible to strong light than moderate heat temperature.

【目的】探讨温度与光照交叉处理对‘赤霞珠’葡萄叶片光系统功能的伤害机制。【方法】以一年生‘赤霞珠’为试材，设置不同温度（28、34、37、40℃）与光强（800、1600 μmol?m-2?s-1）组合，测定叶片气体交换和叶绿素荧光参数，分析不同温度和光强处理对葡萄叶片PSⅡ活性的影响。【结果】无论高光强还是适宜光强下，随着处理温度升高，叶片净光合速率（Pn）、气孔导度（Gs）、PSⅡ的实际光化学效率（ΦPSⅡ）、电子传递速率（ETR）、光合性能指数（PIABS）、捕获光能用于QA以后的电子传递的概率（ΨEo）、单位面积有活性反应中心的数量（RC/CS）均呈下降趋势；PSⅡ激发压（1-qP）、非光化学淬灭系数（NPQ）、相对荧光曲线的初始斜率（Mo）呈现上升趋势；在适宜光强条件下，28—34℃内，叶片PSⅡ的RC/CS以及受体侧未发生显著变化，Pn的下降主要是由于气孔限制引起的，温度达到和超过37℃时，PSⅡ反应中心和受体侧开始出现伤害，叶黄素循环在植物光破坏防御过程中作用增加；在高光照条件下，高光强/28℃时的Pn已降至适光/34℃时的水平， PIABS已低于适光/40℃时的水平，适宜温度范围内（28—34℃），PSⅡ电子受体侧已受到伤害，温度超过37℃时，反应中心和电子受体侧出现更加严重的伤害，叶黄素循环受到抑制，最大光化学效率明显降低。【结论】适宜光强时，温度达到或超过37℃时，PSⅡ受到伤害，发生较严重的光抑制；强光胁迫在28℃时对PSⅡ功能造成伤害，随着温度的升高，伤害显著加剧，说明强光胁迫比高温胁迫更易造成PSⅡ功能的伤害。

... 在逆境条件下因受胁迫活性下降,线性电子传递链在传递电子受阻后会导致活性氧的产生与积累^[17] ...

2016

0.0

... V_J与dV/dt_o均逐渐升高,表明Q_A到Q_B之间的电子传递受阻^[18],在pH 3 ...

2012

0.0

. 2012, 45(23):4794-4800 DOI:10.3864/j.issn.0578-1752.2012.23.005

Relationship between photochemistry activity and decrease in chlorophyll content during senescence in leaves of stay green and quick-leaf-senescence inbred line of maize

保绿玉米与早衰玉米叶片衰老过程中叶绿素降解与光合作用光化学活性的关系

Zhang Z S , Yang C , Gao H Y

张子山, 杨程, 高辉远

1.College of Life Science, Shandong Agricultural University/State Key Laboratory of Crop Biology, Taian 271018, Shandong 2.College of Agronomy, Shandong Agricultural University, Taian 271018, Shandong

【Objective】 The purpose of this study is to explore the relationship between the photosynthetic performance and the decrease in chlorophyll content, and to provide a theoretical basis to breed long duration of active photosynthesis hybrids.【Method】 By simultaneously analyzing chlorophyll a fluorescence transient and light absorbance at 820 nm as well as analyzing net photosynthetic rate and chlorophyll content, the photosynthetic behavior including light absorption, energy transformation and the photo-activities of PSⅡ and PSⅠ during senescence in the stay green inbred line of maize (Zea mays L.) Q319 and the quick-leaf-senescence inbred line of maize HZ4 was studied. 【Result】 Results showed that the net photosynthetic rate, light absorption, energy transformation, the activities of PSⅡ acceptor side and donor side and the PSⅠ activities decreased much slower in stay green inbred line Q319 than in quick-leaf-senescence inbred line HZ4 during the ethephon-induced senescence. Although the chlorophyll content decreased faster in HZ4 with decrease of chlorophyll content induced by ethephon, the photosynthetic performance of Q319 deteriorated much more severely than that of HZ4. 【Conclusion】 The above results indicate that the retention of light absorption, energy transformation and activity of electron transfer contribute to the extended duration of active photosynthesis in the stay green inbred line. It was concluded that attention should be paid to two favorable characteristics in breeding long duration of active photosynthesis hybrids: 1) maintaining more chlorophyll content during senescence and 2) maintaining higher photosynthetic activity during the loss of chlorophyll.

【目的】阐明玉米叶片衰老过程中叶片叶绿素降解与光合作用光反应活性的关系，为选育长效光合持续期的作物品种提供理论基础。【方法】本研究使用保绿型玉米品种“齐319”（Q319）和早衰型玉米品种“黄早4”（HZ4）为材料，在严格控制环境因素的条件下，用乙烯利诱导离体叶片衰老，通过快速叶绿素荧光诱导动力学曲线分析（OJIP test）和820 nm光吸收曲线技术，研究衰老过程中叶片叶绿素含量、光合速率、光系统1（PSI）和光系统2（PSII）光化学活性的变化，并分析光化学活性下降与叶绿素降解的关系。【结果】在衰老过程中，与保绿品种相比，早衰品种HZ4叶片的光能吸收能力、能量的转化效率以及PSI和PSII电子传递的活性均衰退较快。虽然早衰品种在衰老过程中光化学活性的衰退快于保绿品种，但是当叶片的叶绿素含量下降相同程度时，保绿品种Q319叶片的光能的捕获、能量的转化以及电子传递活性都比HZ4叶片下降得更明显。【结论】光化学活性的快速衰退是导致早衰品种HZ4叶片在生育后期光合功能快速衰退的一个重要原因，但它在叶绿素降解时能维持较高的光化学活性。因此在长效光合持续期品种的选育中，应当同时重视两个典型性状：①衰老过程中维持较高的叶绿素含量；②在叶绿素降解时维持较高的单位叶绿素光能吸收、能量转换以及电子传递活性。

... 供体侧,即放氧复合体(OEC)伤害的程度^[19] ...

2015

0.0

... 供体侧伤害程度重于受体侧^[20] ...

1997

0.0

... 反应中心捕光天线色素的大小^[21],该指标升高说明叶片捕光天线色素增加,意味着部分PS#cod#x02161 ...

2013

0.0

. 2013, 27(7):1054-1059

Effects of acid rain stress on antioxidant enzyme activity and chlorophyll fluorescence in leaves of Gazania Hybrids

酸雨对勋章菊保护酶活性及叶绿素荧光参数的影响

Hou W , Pan Y Z

侯维, 潘远智

1. College of Forestry, Sichuan Agricultural University, Ya'an, Sichuan 625014; 2. College of Landscape Architecture, Sichuan Agricultural University, Chengdu, Sichuan 611130

In this paper, pot experiments were conducted to study the effect of simulated acid rain that included four different pH values (5.6, 4.5, 3.5, 2.5) on the membrane permeability, malondialdehyde (MDA) content and antioxidant enzyme activity, chlorophyll (Chl) content, and chlorophyll fluorescence in of leaves Gazania Hybrids . The membrane permeability and MDA content increased significantly after acid rain stress and with an extremely significant correlation between them. The dismutase (SOD), catalase (CAT) and peroxidase (POD) activities were showed a single-peak curve which was firstly increased and then decreased, and the maximum activity of SOD, CAT and POD were observed at pH4.5. Chl a, Chl b, total chlorophyll content, Chl a/b, the PS II photochemical efficiency (Fv/Fm) and the PS II potential activity ( F v/ F o), actual PS II photochemical quantum yield (Φ PSⅡ ) and q P decreased accordingly with the decreasing of pH value, while q N increased. The study showed that G. Hybrids has strong resistance to acid rain stress and the visible damage threshold was less than pH 3.5.Therefore, G. Hybrids can be considered as one of the landscaping and vegetation constructing plants in the acid rain-hit areas.

采用盆栽方法,以不同pH值(5.6、4.5、3.5、2.5)的模拟酸雨胁迫试验,探讨其对勋章菊( Gazania Hybrids )叶片质膜透性、MDA含量、保护酶活性、叶绿素含量及叶绿素荧光参数的影响。结果表明,随pH值的降低,勋章菊叶片质膜透性和MDA含量呈逐渐升高的趋势,且二者呈显著正相关;SOD、CAT和POD活性呈先升高后下降的单峰曲线变化,其中SOD、CAT和POD活性最大值均出现在pH4.5处理;叶绿素a、叶绿素b、叶绿素a+b的含量、叶绿素a/b、PSⅡ最大光化学效率( F v/ F m)、PSⅡ的潜在活性( F v/ F o)、PSⅡ实际光化学量子产量(Φ PSⅡ )、光化学淬灭系数( q P)均随pH值的降低而下降,非光化学淬灭系数( q N)随pH值的降低而升高。模拟酸雨对勋章菊叶片隐形伤害的阈值为pH≤3.5,表明勋章菊对酸雨具有较强的耐受性,可作为酸雨危害较严重地区的园林绿化地被植物之一。

... 电子传递受阻,进一步造成了F_v/F_m的降低,夏枯草叶片发生了光抑制,导致其叶片光能利用率降低^[22] ...

2016

0.0

... 的活性^[23] ...

2019

0.0

... 表征的是两个光系统之间的协调性^[24] ...

2009

0.0

. 2009, 35(10):1916-1922 DOI:10.3724/SP.J.1006.2009.01916

Effects of drought stress on activity of photosystems in leaves of maize at grain filling stage

灌浆期干旱胁迫对玉米叶片光系统活性的影响

Li G , Gao H Y , Zhao B

李耕, 高辉远, 赵斌

1 College of Agronomy, Shandong Agricultural University; 2 College of Life Science, Shandong Agricultural University; 3 State Key Laboratory of Crop Biology, Tai’an 271018, China

At grain filling stage, the effects of drought stress on photosynthetic acivities of photosystem I (PS I) and photosystem II (PS II) in leaves of maize (Zhengdan 21, a cultivar with high starch content) were studied by simultaneously analyzing chlorophyll a fluorescence transient and light absorbance at 820 nm. The results, obtained from two years experiments, demonstrated that the drought stress significantly reduced photosynthesis ( P P A in PS II. The acceptor side of PSII was damaged more severely than the donor side of PSII. Furthermore, the maximal oxidation-reduction activity of PS I ( Δ I/I o ) was also significantly decreased by the drought stress, which inhibited the photosynthetic electron tranport from the PS II to PS I, destructing the coordination between PS I and PS II. We suggest that the inhibition of PS I and PS II and the destruction of the coordination between PS I and PS II by the drought stress is one of the main reasons to cause the decrease in photosyntheis and grain yield of maize.

以高淀粉玉米品种郑单 21 为材料，借助叶绿素荧光快速诱导动力学曲线和 820 nm 光吸收曲线，研究了灌浆期土壤干旱胁迫对玉米籽粒产量和对叶片光系统 I (PSI) 及光系统 II (PS II) 活性的影响。两年的研究结果均表明，干旱胁迫显著抑制叶片光合速率 ( P 和籽粒产量 ( P 。 JIP-test 分析发现干旱胁迫导致叶绿素荧光快速诱导动力学曲线中的 K 点和 J 点上升，表明 PS II 放氧复合体 (OEC) 和 Q A 之后的电子传递链受到抑制，且 PS II 受体侧受抑制的程度大于供体侧。此外，干旱胁迫也显著地抑制 PS I 的最大氧化还原活性 ( Δ I/I o ) ，阻碍光合电子从 PS II 向 PS I 的传递，破坏了 PS I 和 PS II 的协调性。我们认为干旱胁迫抑制 PS I 和 PS II 活性并破坏二者的协调性，是导致导致 P n 和籽粒产量下降的重要原因之一。

... )明显失衡,这可能是引起Q_A后续电子传递链效率下降以及整个电子传递链功能降低的主要原因^[25] ...

2012

0.0

. 2012, 32(24):7866-7873 DOI:10.5846/stxb201203080315

Effects of simulated acid rain stress on the PS#cod#x02161; reaction center and free radical metabolism in leaves of longan

模拟酸雨对龙眼叶片PS#cod#x02161;反应中心和自由基代谢的影响

Li Y Y , Pan T F , Yu D

李永裕, 潘腾飞, 余东

Injury mechanism of acid rain on the photosynthetic apparatus of plants has been one of the hot spots of the ecological research. To elucidate the relationship between the damage of reaction center and the accumulation of free radicals in photosynthetic system due to foliar acidification, one-year old longan ( Dimocarpus longana Lour. cv. Wulongling) seedlings grown in pots were used to study the effects of simulated acid rain stress on chlorophyll fluorescence parameters, free radical metabolism and their recovery. The results showed that, under acid rain stress, the curve of chlorophyll fluorescence was quickly changed and PSⅡ reaction center was damaged. Maximum photochemistry efficiency of PSⅡ( F v /F m ), the activity of PSⅡ reaction center (1/ F O -1/F M ) and the density of active reaction center (RC/CS O ) decreased significantly after 5d stress with acid rain of pH2.5. The closure degree of active reaction center ( V J ) and the percentage of inactive reaction center (Non-Q A and Non-Q B ) increased, whereas the Q A was reduced rapidly. The oxygen releasing complexes were destructed, and the number of electron transducer of PSⅡ receptor ( S m ), the electronic conversion efficiency ( ψ O ), the transduction rate of electron ( φ Eo ) all decreased significantly. These results displayed that foliar acidification inhibited the linear electronic transduction of PSⅡ reaction center. The results of free radical metabolism of longan indicated the contents of O 2 ·－ , H 2 O 2 and MDA increased significantly after 5d stress with acid rain of pH2.5. AsA and GSH converted to oxidized form, the contents of reduced form decreased, and the activities of SOD, APX, MDAR, DHAR and GR all reduced significantly. The inability of scavenging the free radicals in chloroplast in time resulted in the attack of photosynthetic organs by the redundant free radicals and thus the damage of PSⅡof longan leaves. Taken together, we concluded that acid rain stress damaged the oxidation and reduction sides of the reaction centers of PSⅡof the leaves in longan, and further decreased the linear electronic transduction to result in the shortage of assimilatory power which weakened the scavenging of free radicals, the accumulation of free radicals in chloroplasts destructed photosynthetic apparatus more.

酸雨对植物光合机构的伤害机理一直是生态学研究的热点之一,为了探讨叶面酸化导致的PSⅡ反应中心损伤和光合机构自由基累积之间的内在联系,以1年生龙眼( Dimocarpus longana Lour.)实生小苗为研究对象,采用盆栽试验,研究了模拟酸雨胁迫对龙眼叶片叶绿素荧光参数和自由基代谢的影响。结果表明:酸雨胁迫改变了龙眼叶片的快速叶绿素荧光诱导动力学曲线形状,伤害PSⅡ反应中心;pH2.5酸雨胁迫5d后最大光化学效率( F v /F m )、PSⅡ反应中心活性(1/ F O -1/F M )、反应中心含量(RC/CS O )急剧下降;有活性反应中心的关闭程度( V J )、失活反应中心的比例(Non-Q A 和Non-Q B )显著增加,Q A 迅速还原;放氧复合体(OEC)被破坏;PSⅡ受体侧电子传递体数( S m )、电子转化效率( ψ O )和电子传递速率( φ Eo )明显降低,叶面酸化导致光系统线性电子传递受损。pH2.5酸雨胁迫5d后叶片超氧阴离子自由基(O 2 ·－ )、过氧化氢(H 2 O 2 )和丙二醛(MDA)含量显著增加;抗坏血酸(AsA)和谷胱甘肽(GSH)转化为氧化型,还原型减少;超氧化物歧化酶(SOD)、抗坏血酸过氧化物酶(APX)、单脱氢抗坏血酸还原酶(MDAR)、脱氢抗坏血酸还原酶(DHAR)和谷胱甘肽还原酶(GR)活性下降,叶绿体内自由基不能被及时清除,过多的自由基损伤光合器官,导致龙眼叶片PSⅡ受伤害。模拟酸雨胁迫伤害龙眼叶片PSⅡ反应中心供体侧和受体侧的电子传递体,造成同化力不足,清除自由基能力下降,导致叶绿体自由基累积,光合机构受到伤害。

... 酸胁迫会诱导作物体内产生大量活性氧(ROS),过量活性氧会破坏细胞膜的结构与功能,导致细胞生理功能紊乱^[26,27] ...

2012

0.0

. 2012, 48(8):87-93 DOI:10.11707/j.1001-7488.20120814

Effect of calcium on the functions of antioxidant systems and mitochondria in Malus hupehensis var. pingyiensis roots under acid treatment

钙对酸化处理平邑甜茶根系抗氧化酶活性及线粒体功能的影响

Zhang Z , Liang Y C , Wang L

张召, 梁元存, 王利

1. Key Laboratory of Agricultural Ecology and Environment, Shandong Agricultural University Tai'an 271018;2. College of Plant Protection, Shandong Agricultural University Tai'an 271018;3. Post-Doctoral Mobile Station of Agricultural Resource Utilization, Shandong Agricultural University Tai'an 271018

The purpose of this paper is to explore the effects of acid treatment and exogenous application of calcium to nutrient solution on the functions of antioxidant systems and mitochondria in Malus hupehensis var. pingyiensis roots. The M. hupehensis var. pingyiensis rootstock seedlings were cultured in 1/2 Hoagland nutrient solutions of different pH (3, 4, 5 and 6) and different Ca 2+ concentrations (10, 20 and 30 mmol·L -1 ) with pH 3, respectively. The antioxidant enzyme activities and the characteristics of mitochondria were measured in day 0, 2, 3 and 4. Compared with the control (pH6), the maximum activity of SOD, CAT in the treatments occurred in day 2, and the change of POD activity was different, the maximum activity of POD occurred in day 2 in pH 3 and pH 4, and in day 3 pH 5. Meanwhile, the mitochondrial membrane permeability, potential (Δ ψ m ) and cytochrome c/a decreased with the increasing of the acidity and the treatment time and the content of H 2 O 2 was decreased at first and then increased. The exogenous application of calcium (10 and 20 mmol·L -1 ) in the nutrient solution under acid treatment (pH 3) increased the activities of SOD, POD and CAT, relieved the damage of the mitochondrial functions by the acidity and inhibited the accumulation of H 2 O 2 (the decline, with 10 and 20 mmol·L -1 , was 6.78% and 13.43% of the accumulation caused by the single acid treatment, respectively). The 20 mmol·L -1 Ca 2+ had better effect whereas the 30 mmol·L -1 Ca 2+ increased the damage and increased the content of H 2 O 2 by 8.96%. The above results suggested that the acid treatments would damage the roots of M. hupehensis var. pingyiensis and the exogenous application of dilute calcium in the nutrient solution could relieve the damage of acid treatment to the roots.

为探讨酸化处理及钙对酸化处理下平邑甜茶根系抗氧化酶活性及线粒体功能的影响,将平邑甜茶幼苗分2组分别置于pH 3,4,5,6(对照)的1/2 Hoagland营养液中及3个钙离子浓度梯度(10,20,30 mmol·L -1 )的pH值为3的1/2 Hoagland营养液中,于第0,2,3,4天分别测定抗氧化酶活性及线粒体指标。结果表明:与对照相比,各处理的SOD,CAT酶活性在处理的第2天达到最大值,POD酶活性变化不一,pH 3与pH 4处理在第2天达到最大值,pH5处理在第3天达到最大值; 根系线粒体膜通透性、膜电位及细胞色素c/a随着酸度增加和时间的延长逐渐降低; H 2 O 2 含量先降低后升高。低浓度外源钙(10,20 mmol·L -1 )处理能提高SOD,CAT,POD酶活性并维持一定时间,缓解线粒体各项功能的伤害,抑制H 2 O 2 的积累,主要表现在与单独酸处理相比H 2 O 2 含量降低了6.78%, 13.43%,且20 mmol·L -1 钙离子缓解效果最好; 30 mmol·L -1 钙离子处理却加剧了酸化伤害,H 2 O 2 含量上升了8.96%。酸化处理会对平邑甜茶根系造成伤害而营养液适量加钙可以缓解其伤害。

... 酸胁迫会诱导作物体内产生大量活性氧(ROS),过量活性氧会破坏细胞膜的结构与功能,导致细胞生理功能紊乱^[26,27] ...

... 这与平邑甜茶在酸化处理一定天数后根系SOD与POD活性下降^[27]的结果相似,类似的结果也出现在萝卜(Raphanus sativus)与莴笋(Lactuca sativa)的相关研究中^[29] ...

2018

0.0

... 植物在遭受酸胁迫时,会通过维持几种抗氧化酶之间的协调和平衡来提高植株抗活性氧的能力^[28] ...

2014

0.0

2016

0.0

... 受体侧电子传递受阻会使过剩电子泄露,细胞中游离的O₂因受泄露的电子攻击而生成超氧阴离子,这些活性氧与超氧阴离子会造成细胞膜质过氧化增加和电解质渗漏等问题^[30,31] ...

2004

0.0

2016

0.0

... 在逆境胁迫条件下,植物能够通过增加可溶性糖、脯氨酸等渗透调节物质的含量来适应逆境过程^[32,33] ...

2017

0.0

... 在逆境胁迫条件下,植物能够通过增加可溶性糖、脯氨酸等渗透调节物质的含量来适应逆境过程^[32,33] ...

2010

0.0

... 特别地,脯氨酸对羟基自由基清除和保护酶也具有一定的保护和稳定作用^[34],脯氨酸增幅大于可溶性糖增幅,说明脯氨酸可能已成为主要的渗透调节物质^[35] ...

2010

0.0

. 2010, 45(1):73-78 DOI:10.3969/j.issn.1674-3466.2010.01.010

Effects of various concentration of seawater on the growth and physiological characteristics of Lactuca indica seedlings

海水胁迫对苦荬菜幼苗生长及生理特性的影响

Li H Y , Zheng Q S , Liu Z P

李洪燕, 郑青松, 刘兆普

Key Laboratory of Marine Biology in Jiangsu Province, College of Resources and Environmental Sciences, Nanjing Agricultural University, Nanjing 210095, China

Planting of salt-tolerant plants is vital for the development of available coastal land resources. We studied the effects of different concentrations of seawater (diluted in 1/2 Hoagland’s solution) on the biomass and chlorophyll, ion, soluble sugar and soluble protein contents of Lactuca indica seedlings by sand culture in the greenhouse. The dry weight of L. indica shoots was more susceptible to seawater stress than that of roots. Compared with dry weight of controls, that of roots of L. indica seedlings was unchanged with seawater concentrations from 10% to 30%. With increasing seawater stress, the ratio of roots to shoots and Na + , Cl – and soluble sugar contents were increased significantly. With 10% and 20% seawater treatments, chlorophyll content did not differ from that of controls but was significantly decreased with further increased seawater stress. At a seawater concentration of 10%, K + content in the shoots and roots of L. indica seedlings did not differ from that of controls, but with increased seawater stress, the K + content was markedly dose-dependently decreased. However, as gradually increasing soluble sugar content significantly, soluble protein content first increased then decreased with increasing seawater concentration. Adaptation to seawater in L. indica was mainly caused by strong K + selective absorption and transport. Furthermore, accumulating soluble sugar and protein were important osmotica in L. indica seedlings and could be an indicator of salt tolerance.

抗盐耐海水植物的种植是有效利用和开发滩涂资源的措施之一。采用温室砂培方式, 研究了不同稀释配比的海水处理8天对苦荬菜( Lactuca indica )幼苗生物量、根冠比、叶绿素含量、离子含量、可溶性蛋白和可溶性糖含量的影响。结果表明: 苦荬菜幼苗地上部受海水胁迫较为显著, 而根在海水浓度小于30%时与对照相比没有显著差异; 根冠比随着海水浓度的增加而不断提高; 在10%和20%海水浓度处理下, 叶绿素含量与对照相比差异不显著, 但随着海水浓度的进一步增加,叶绿素含量显著下降; 在10%海水浓度处理下, 苦荬菜地上部分及根部的K + 含量与对照相比差异不显著, 而海水浓度高于10%时, 随着海水浓度的增加地上部和根部的K + 含量均逐渐降低; 海水处理下, 苦荬菜体内Na + 和Cl – 含量逐渐增加; 地上部可溶性糖含量逐渐增加, 而可溶性蛋白含量先升后降。海水胁迫下, 苦荬菜幼苗维持一定的K + 选择性吸收是其一定程度上盐适应的重要原因。同时, 积累的可溶性糖和可溶性蛋白是苦荬菜幼苗在盐胁迫下的重要渗透调节物质, 可作为其抗盐性的生理参数。