豇豆（Vigna unguiculata L.）铝毒害及耐性机理
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摘要
铝是地壳中含量最丰富的金属元素,通常以难溶性硅酸盐或氧化铝的形式存在,对植物没有毒害,但在酸性条件下(pH<5),可溶性的铝(主要是Al3+)对大多数植物都会产生毒害。酸性上壤中的铝毒害抑制植物生长,降低作物品质和产量。因此对植物耐铝胁迫机理研究具有重要的理论价值和实际意义。本文以不同耐铝性豇豆品种‘T6’(耐铝型)和‘S3’(铝敏感型)为试验材料,对两份品种耐铝差异进行了比较研究。通过研究不同浓度铝胁迫下两份豇豆品种种子萌发、幼苗生长、营养元素吸收、光合荧光特性、抗氧化系统、细胞壁特性和有机酸分泌等方面的变化,来探讨豇豆铝毒害及耐铝机理。全文研究结果如下：
1.铝敏感品种‘S3’种子的发芽率、发芽势、淀粉酶活性和可溶性糖含量在50μmol·L-1以下的低浓度铝(10、50μmol·L-1)处理下与对照相比差异不显著；而在100μmol·L-1以上的高浓度铝(100、500、1000μmol·L-1)胁迫下显著下降。耐铝品种‘T6’种子的相关指标在低浓度铝处理下显著提高；在500、1000μmol·L-1铝胁迫下显著下降。
低浓度铝处理时,两份豇豆品种幼苗的株高、地上部干重与对照相比差异不显著,500μmol·L-1的铝胁迫显著降低了两份豇豆品种的株高和地上部干重。耐铝品种‘T6’在10μmol·L-1铝处理时,根长和根系干重与对照相比差异不显著,在高浓度铝胁迫时显著下降；铝敏感品种‘S3’随着铝处理浓度的提高,根长、根系干重一直呈下降趋势。
2.铝胁迫增加了两份豇豆品种植株体内的铝含量,且铝主要集中在根系,铝敏感品种‘S3’的增加量大于耐性品种‘T6’；两份品种的根系和地上部分的大量元素N、P、Mg、Ca、K的含量在高浓度的铝胁迫时有不同程度下降,除K含量外,对铝敏感品种'S3'N、P、Mg、Ca的含量影响大于耐性品种‘T6’。铝胁迫对豇豆微量元素的吸收也有不同程度的抑制作用,以Cu和Fe受到的抑制作用最大,Zn次之,Mn吸收受到的抑制作用最小。
铝胁迫后,铝主要集中在豇豆根尖的0～10mm处,铝敏感品种‘S3’铝积累量大于耐铝品种‘T6’；苏木精染色后发现铝敏感品种‘S3’根尖染色较深,而耐铝品种‘T6’根尖染色较浅；X-射线能谱分析发现,铝胁迫后,两份豇豆品种的铝主要分布在表皮和皮层组织,而中柱分布较少。
3.10μmol·L-1的铝处理提高了耐性品种‘T6’的叶绿素a、叶绿素a/b、净光合速率(Pn)和非光化学淬灭系数(NPQ),降低了胞间CO2浓度(Ci),增强了其光合能力；而高浓度的铝胁迫破坏了两品种的叶绿体、线粒体和细胞核结构,降低了叶绿素a、叶绿素a/b、Pn,气孔导度(Gs)、植物水分利用效率(WUE)、PSⅡ光合电子传递量子效率(ΦPSⅡ)、光化学淬灭系数(qP)和NPQ,提高了Ci,抑制了其光合作用,且对敏感品种‘S3’的影响大于耐铝品种‘T6’。
4.随着铝处理浓度的增加,豇豆根尖细胞壁果胶和纤维素中的糖醛酸含量和总糖含量显著增加,半纤维素中的总糖含量显著增加,且铝敏感品种‘S3’增加幅度大于耐铝品种‘T6’；豇豆根尖细胞壁果胶、半纤维素和纤维素中的铝含量在高浓度铝胁迫时显著增加,铝敏感品种‘S3’增加量显著高于耐铝品种‘T6’。高浓度铝胁迫下,两品种根尖细胞壁的果胶甲酯酶(PME)活性显著提高,‘S3’的升高幅度大于‘T6’,根尖细胞壁果胶甲酯化程度显著降低,‘S3’的降低幅度大于‘T6’。
随着铝处理浓度的升高,两份豇豆品种的根尖O2.-产生速率、H202含量、MDA含量及质膜透性都逐渐增加,且铝敏感品种‘S3’的增加幅度大于耐铝品种‘T6’。抗氧化系统的超氧化物歧化酶(SOD)、过氧化物酶(POD)、过氧化氢酶(CAT)、抗坏血酸过氧化物酶(APX)、谷胱甘肽还原酶(GR)、抗坏血酸(AsA)、还原型谷胱甘肽(GSH)含量都有所上升,其中铝敏感品种‘S3’的SOD活性上升幅度比耐性品种‘T6’高,而POD、CAT、APX、ASA的上升幅度相对较低,从而引起H2O2过量积累而导致氧化胁迫,使细胞的脂质过氧化程度加剧,最终严重影响根系的生长。
5.铝胁迫可诱导豇豆根系分泌柠檬酸和苹果酸,柠檬酸的分泌量在不同铝处理浓度和不同品种间没有显著变化。苹果酸的分泌量随着铝处理浓度的增加而升高,耐铝品种苹果酸的分泌量高于铝敏感品种,两份品种根系有机酸的分泌均存在一个6小时的滞后期,属于分泌模式Ⅱ型。缺P处理可以诱导豇豆根系分泌有机酸,但分泌量不随处理时间的增加而增加,缺P处理加铝24小时后苹果酸分泌量增加了约10倍,镧处理不能诱导豇豆根系分泌有机酸,铝诱导豇豆有机酸的分泌主要发生在根尖0～10mm。
随着铝处理浓度的增加,根系苹果酸分泌量显著增加,根尖苹果酸含量显著减少,但与其代谢相关酶的活性没有显著变化；阴离子通道抑制剂A-9-C和蛋白合成抑制剂CHM可以完全抑制铝诱导的豇豆根系有机酸的分泌,但根尖有机酸的含量没有显著变化。因此推测铝诱导的豇豆根系有机酸的分泌与其代谢无关,可能与新蛋白的合成和通道的开启有关。
外源水杨酸(SA)、脱落酸(ABA)对铝诱导的豇豆根系有机酸的分泌影响不显著,而外源生长素(IAA)可显著提高铝诱导的豇豆根系苹果酸的分泌,Ca2+通道抑制剂和螯合剂则显著抑制铝诱导的豇豆根系苹果酸的分泌,同时豇豆根尖有机酸的含量和其代谢相关酶的活性均没有显著变化。因此推测IAA和Ca2+可能参与调控铝诱导的豇豆根系苹果酸的分泌,但这个调控过程可能与其代谢无关。
Aluminum (Al) is the most abundant metal element in the earth's curst, usually in the form of insoluble silicate or alumina which was not toxic to plants. But in the acidic soil (pH<5), soluble A1(mainly Al3+) was of toxic to most of plant. In the acidic soil, Al toxicity inhibited the growth of plant, decrease the quality and output of crops. So the researches on Al resistant mechanism are of important theoretical and practical significance. In this research., the seed germinate, seedling growth, nutrient uptake, photosynthetic characteristics, reactive oxygen system, cell wall characteristic and the secretion of organic acid of two cowpea (Vigna unguiculata L.) cultivars 'T6'(Al-toleranct) and 'S3'(Al-sensitive) were studied, to reveal the Al resistance mechanism of cowpea. The results were reported as follows:
1. There were no significant change on the germination energy, germination percentage, amylase activity and soluble sugar content of 'S3' after exposure to10and50μmol·L-1Al, but significantly decreased after exposure to100、500、1000μmol·L-1Al. These indexes of 'T6' significantly increased after exposure to10and50μmol·L-1Al, and significantly decreased after exposure to500、1000μmol·L-1.
There was no significant change on the plant height and shoot dry weight of'T6'and 'S3' after exposure to10and50μmol·L-1Al, and significant decrease after exposure to500μmol·L-1Al. Compared with control, there was no significant change on the root length and root dry weight of 'T6' after exposure to10μmol·L-1Al, significantly decreased after exposure to50,100and500μmol·L-1Al. The root length and root dry weight of'S3' decreased significantly by Al increase from10to500μmol·L-1.
2. The Al content in 'T6' and 'S3' increased after Al stress, and Al mainly accumulated in roots, the increase of 'T6' was lower than 'S3'; The content of N, P, K, Ca, Mg in'T6' and'S3'decreased after exposure to500μmol·L-1Al. Besides K, the decrease of N, P, Ca, Mg in'T6'was lower than'S3'. The uptake of trace elements was inhibited in cowpea, the decrease of Cu and Fe was highest, Zn next, the decrease of Mn was lowest.
Al mainly accumulated in root apex (0～10mm) after A1stress, the content of A1in 'S3'was higher than 'T6'.The root apex of'S3'was stained more deep than 'T6' by hematoxylin after Al treatment. The results of X-ray analysis indicated that A1mainly distributed in epidermal cells and subepidermal cortical cells.
3. The chlorophylla a, chlorophyll a/b, net photosynthetic rate (Pn) and non-photochemical quenching coefficient (NPQ) of 'T6' increased and the intercellular CO2concentration (Ci) of 'T6' decreased after exposure to10umol·L-1Al, so the photosynthetic capacity was strengthened. The structure of chloroplast, mitochondria and cell nucleus of 'T6' and 'S3' was destroyed; the chlorophylla a, chlorophyll a/b, Pn, stomatal conductance (Gs), plant water use efficiency (WUE), PSⅡ photosynthetic electron transport quantum efficiency (ΦPSⅡ), photochemical quenching (qP) and NPQ of 'T6' and 'S3' decreased; the Ci of 'T6' and 'S3' increased after exposure to100and500μmol·L-1Al, so the photosynthetic capacity was inhibited, and the inhibition in'S3'was larger than'T6'.
4. The uronic acid and sugar of pectin and cellulose in cell wall of 'T6' and 'S3' root apex increased significantly with the increase of Al, and the sugar in hemicellulose (HC) increased significantly with the increase of Al, and increase of'S3'was significantly higher than 'T6'. The A1content of pectin, HC, and cellulose increase significantly after exposure to100and500μmol·L-1Al, and the increase of'S3'was significantly higher than'T6'.The cell wall PME activity of 'T6' and 'S3' root apex increased significantly after exposure to100,500μmol·L-1Al, the increase of 'S3' was significantly higher than 'T6'.The degree of cell wall pectin methyl of 'T6' and 'S3' root apex decrease significantly after exposure to100and500μmol·L-1Al, the decrease of'S3'was significantly higher than'T6'.
The superoxide anion radical, H2O2content, MDA content and plasma-membrane permeability of'T6' and 'S3' root tip increase significantly by A1increase from0to500μmol·L-1, and the increase range of 'S3' was higher than 'T6'.The SOD, POD, CAT, APX, GR, AsA, GSH of 'T6' and 'S3' root tip increased after exposure to0to500μmol·L-1Al. The increase range of SOD in'S3'root tip was higher than 'T6', but increase range of POD、CAT、APX in 'S3' root tip were significant lower than 'T6'.We presume that 'S3' root tip accumulated more H2O2and increased the oxidative stress then result in the inhibited of root growth.
5. After exposure to Al, both malate and citrate were secreted from 'T6' and 'S3' roots. There was no significant change of citrate between'T6'and'S3'. The amount of malate secreted from 'T6' and 'S3' increase significantly by Al increase, the secretion of malate from 'T6' root was higher than'S3'. The secretion type of malate and citrate from 'T6' and 'S3'roots was similar, and both of them have a6h lag-phase. Potassium (P) starvation also induced the secretion of malate and citrate, but the secretion amount did not increase by P starvation time increase, and Al stimulate10times secretion than P starvation. Exposure to La (Lanthanum)did not induce organic acid secretion from cowpea root. The secretion site was mainly located in the0～10mm segment from root tip.
The secretion of malate increase from cowpea root, and the malate content of cowpea tips decrease by Al increase, there was no significant change in enzyme activity relate to malate metabolism. The anion-channel inhibitor (A-9-C) and protein-synthesis inhibitor (CHM) completely inhibited the organic acid secretion of cowpea root under Al treatment, but the organic acid content in root tip did not changed. So we presume that there was no relative between the organic secretion and organic metabolism under A1treatment, and maybe related to the new protein synthesis.
Exogenous SA and ABA did not changed the organic acid secretion in cowpea root induced by A1treatment, exogenous IAA significantly increased the malate secretion, and exogenous Ca2+channel inhibitors and chelator significantly inhibited the malate secretion. Meanwhile the organic acid content and enzyme activity relate to organic acid metabolism did not changed. So we presume that IAA and Ca2+may involved in the regulation of malate secretion induced by Al, but the regulation may be not related to malate metabolism.
1.铝敏感品种‘S3’种子的发芽率、发芽势、淀粉酶活性和可溶性糖含量在50μmol·L-1以下的低浓度铝(10、50μmol·L-1)处理下与对照相比差异不显著；而在100μmol·L-1以上的高浓度铝(100、500、1000μmol·L-1)胁迫下显著下降。耐铝品种‘T6’种子的相关指标在低浓度铝处理下显著提高；在500、1000μmol·L-1铝胁迫下显著下降。
低浓度铝处理时,两份豇豆品种幼苗的株高、地上部干重与对照相比差异不显著,500μmol·L-1的铝胁迫显著降低了两份豇豆品种的株高和地上部干重。耐铝品种‘T6’在10μmol·L-1铝处理时,根长和根系干重与对照相比差异不显著,在高浓度铝胁迫时显著下降；铝敏感品种‘S3’随着铝处理浓度的提高,根长、根系干重一直呈下降趋势。
2.铝胁迫增加了两份豇豆品种植株体内的铝含量,且铝主要集中在根系,铝敏感品种‘S3’的增加量大于耐性品种‘T6’；两份品种的根系和地上部分的大量元素N、P、Mg、Ca、K的含量在高浓度的铝胁迫时有不同程度下降,除K含量外,对铝敏感品种'S3'N、P、Mg、Ca的含量影响大于耐性品种‘T6’。铝胁迫对豇豆微量元素的吸收也有不同程度的抑制作用,以Cu和Fe受到的抑制作用最大,Zn次之,Mn吸收受到的抑制作用最小。
铝胁迫后,铝主要集中在豇豆根尖的0～10mm处,铝敏感品种‘S3’铝积累量大于耐铝品种‘T6’；苏木精染色后发现铝敏感品种‘S3’根尖染色较深,而耐铝品种‘T6’根尖染色较浅；X-射线能谱分析发现,铝胁迫后,两份豇豆品种的铝主要分布在表皮和皮层组织,而中柱分布较少。
3.10μmol·L-1的铝处理提高了耐性品种‘T6’的叶绿素a、叶绿素a/b、净光合速率(Pn)和非光化学淬灭系数(NPQ),降低了胞间CO2浓度(Ci),增强了其光合能力；而高浓度的铝胁迫破坏了两品种的叶绿体、线粒体和细胞核结构,降低了叶绿素a、叶绿素a/b、Pn,气孔导度(Gs)、植物水分利用效率(WUE)、PSⅡ光合电子传递量子效率(ΦPSⅡ)、光化学淬灭系数(qP)和NPQ,提高了Ci,抑制了其光合作用,且对敏感品种‘S3’的影响大于耐铝品种‘T6’。
4.随着铝处理浓度的增加,豇豆根尖细胞壁果胶和纤维素中的糖醛酸含量和总糖含量显著增加,半纤维素中的总糖含量显著增加,且铝敏感品种‘S3’增加幅度大于耐铝品种‘T6’；豇豆根尖细胞壁果胶、半纤维素和纤维素中的铝含量在高浓度铝胁迫时显著增加,铝敏感品种‘S3’增加量显著高于耐铝品种‘T6’。高浓度铝胁迫下,两品种根尖细胞壁的果胶甲酯酶(PME)活性显著提高,‘S3’的升高幅度大于‘T6’,根尖细胞壁果胶甲酯化程度显著降低,‘S3’的降低幅度大于‘T6’。
随着铝处理浓度的升高,两份豇豆品种的根尖O2.-产生速率、H202含量、MDA含量及质膜透性都逐渐增加,且铝敏感品种‘S3’的增加幅度大于耐铝品种‘T6’。抗氧化系统的超氧化物歧化酶(SOD)、过氧化物酶(POD)、过氧化氢酶(CAT)、抗坏血酸过氧化物酶(APX)、谷胱甘肽还原酶(GR)、抗坏血酸(AsA)、还原型谷胱甘肽(GSH)含量都有所上升,其中铝敏感品种‘S3’的SOD活性上升幅度比耐性品种‘T6’高,而POD、CAT、APX、ASA的上升幅度相对较低,从而引起H2O2过量积累而导致氧化胁迫,使细胞的脂质过氧化程度加剧,最终严重影响根系的生长。
5.铝胁迫可诱导豇豆根系分泌柠檬酸和苹果酸,柠檬酸的分泌量在不同铝处理浓度和不同品种间没有显著变化。苹果酸的分泌量随着铝处理浓度的增加而升高,耐铝品种苹果酸的分泌量高于铝敏感品种,两份品种根系有机酸的分泌均存在一个6小时的滞后期,属于分泌模式Ⅱ型。缺P处理可以诱导豇豆根系分泌有机酸,但分泌量不随处理时间的增加而增加,缺P处理加铝24小时后苹果酸分泌量增加了约10倍,镧处理不能诱导豇豆根系分泌有机酸,铝诱导豇豆有机酸的分泌主要发生在根尖0～10mm。
随着铝处理浓度的增加,根系苹果酸分泌量显著增加,根尖苹果酸含量显著减少,但与其代谢相关酶的活性没有显著变化；阴离子通道抑制剂A-9-C和蛋白合成抑制剂CHM可以完全抑制铝诱导的豇豆根系有机酸的分泌,但根尖有机酸的含量没有显著变化。因此推测铝诱导的豇豆根系有机酸的分泌与其代谢无关,可能与新蛋白的合成和通道的开启有关。
外源水杨酸(SA)、脱落酸(ABA)对铝诱导的豇豆根系有机酸的分泌影响不显著,而外源生长素(IAA)可显著提高铝诱导的豇豆根系苹果酸的分泌,Ca2+通道抑制剂和螯合剂则显著抑制铝诱导的豇豆根系苹果酸的分泌,同时豇豆根尖有机酸的含量和其代谢相关酶的活性均没有显著变化。因此推测IAA和Ca2+可能参与调控铝诱导的豇豆根系苹果酸的分泌,但这个调控过程可能与其代谢无关。
Aluminum (Al) is the most abundant metal element in the earth's curst, usually in the form of insoluble silicate or alumina which was not toxic to plants. But in the acidic soil (pH<5), soluble A1(mainly Al3+) was of toxic to most of plant. In the acidic soil, Al toxicity inhibited the growth of plant, decrease the quality and output of crops. So the researches on Al resistant mechanism are of important theoretical and practical significance. In this research., the seed germinate, seedling growth, nutrient uptake, photosynthetic characteristics, reactive oxygen system, cell wall characteristic and the secretion of organic acid of two cowpea (Vigna unguiculata L.) cultivars 'T6'(Al-toleranct) and 'S3'(Al-sensitive) were studied, to reveal the Al resistance mechanism of cowpea. The results were reported as follows:
1. There were no significant change on the germination energy, germination percentage, amylase activity and soluble sugar content of 'S3' after exposure to10and50μmol·L-1Al, but significantly decreased after exposure to100、500、1000μmol·L-1Al. These indexes of 'T6' significantly increased after exposure to10and50μmol·L-1Al, and significantly decreased after exposure to500、1000μmol·L-1.
There was no significant change on the plant height and shoot dry weight of'T6'and 'S3' after exposure to10and50μmol·L-1Al, and significant decrease after exposure to500μmol·L-1Al. Compared with control, there was no significant change on the root length and root dry weight of 'T6' after exposure to10μmol·L-1Al, significantly decreased after exposure to50,100and500μmol·L-1Al. The root length and root dry weight of'S3' decreased significantly by Al increase from10to500μmol·L-1.
2. The Al content in 'T6' and 'S3' increased after Al stress, and Al mainly accumulated in roots, the increase of 'T6' was lower than 'S3'; The content of N, P, K, Ca, Mg in'T6' and'S3'decreased after exposure to500μmol·L-1Al. Besides K, the decrease of N, P, Ca, Mg in'T6'was lower than'S3'. The uptake of trace elements was inhibited in cowpea, the decrease of Cu and Fe was highest, Zn next, the decrease of Mn was lowest.
Al mainly accumulated in root apex (0～10mm) after A1stress, the content of A1in 'S3'was higher than 'T6'.The root apex of'S3'was stained more deep than 'T6' by hematoxylin after Al treatment. The results of X-ray analysis indicated that A1mainly distributed in epidermal cells and subepidermal cortical cells.
3. The chlorophylla a, chlorophyll a/b, net photosynthetic rate (Pn) and non-photochemical quenching coefficient (NPQ) of 'T6' increased and the intercellular CO2concentration (Ci) of 'T6' decreased after exposure to10umol·L-1Al, so the photosynthetic capacity was strengthened. The structure of chloroplast, mitochondria and cell nucleus of 'T6' and 'S3' was destroyed; the chlorophylla a, chlorophyll a/b, Pn, stomatal conductance (Gs), plant water use efficiency (WUE), PSⅡ photosynthetic electron transport quantum efficiency (ΦPSⅡ), photochemical quenching (qP) and NPQ of 'T6' and 'S3' decreased; the Ci of 'T6' and 'S3' increased after exposure to100and500μmol·L-1Al, so the photosynthetic capacity was inhibited, and the inhibition in'S3'was larger than'T6'.
4. The uronic acid and sugar of pectin and cellulose in cell wall of 'T6' and 'S3' root apex increased significantly with the increase of Al, and the sugar in hemicellulose (HC) increased significantly with the increase of Al, and increase of'S3'was significantly higher than 'T6'. The A1content of pectin, HC, and cellulose increase significantly after exposure to100and500μmol·L-1Al, and the increase of'S3'was significantly higher than'T6'.The cell wall PME activity of 'T6' and 'S3' root apex increased significantly after exposure to100,500μmol·L-1Al, the increase of 'S3' was significantly higher than 'T6'.The degree of cell wall pectin methyl of 'T6' and 'S3' root apex decrease significantly after exposure to100and500μmol·L-1Al, the decrease of'S3'was significantly higher than'T6'.
The superoxide anion radical, H2O2content, MDA content and plasma-membrane permeability of'T6' and 'S3' root tip increase significantly by A1increase from0to500μmol·L-1, and the increase range of 'S3' was higher than 'T6'.The SOD, POD, CAT, APX, GR, AsA, GSH of 'T6' and 'S3' root tip increased after exposure to0to500μmol·L-1Al. The increase range of SOD in'S3'root tip was higher than 'T6', but increase range of POD、CAT、APX in 'S3' root tip were significant lower than 'T6'.We presume that 'S3' root tip accumulated more H2O2and increased the oxidative stress then result in the inhibited of root growth.
5. After exposure to Al, both malate and citrate were secreted from 'T6' and 'S3' roots. There was no significant change of citrate between'T6'and'S3'. The amount of malate secreted from 'T6' and 'S3' increase significantly by Al increase, the secretion of malate from 'T6' root was higher than'S3'. The secretion type of malate and citrate from 'T6' and 'S3'roots was similar, and both of them have a6h lag-phase. Potassium (P) starvation also induced the secretion of malate and citrate, but the secretion amount did not increase by P starvation time increase, and Al stimulate10times secretion than P starvation. Exposure to La (Lanthanum)did not induce organic acid secretion from cowpea root. The secretion site was mainly located in the0～10mm segment from root tip.
The secretion of malate increase from cowpea root, and the malate content of cowpea tips decrease by Al increase, there was no significant change in enzyme activity relate to malate metabolism. The anion-channel inhibitor (A-9-C) and protein-synthesis inhibitor (CHM) completely inhibited the organic acid secretion of cowpea root under Al treatment, but the organic acid content in root tip did not changed. So we presume that there was no relative between the organic secretion and organic metabolism under A1treatment, and maybe related to the new protein synthesis.
Exogenous SA and ABA did not changed the organic acid secretion in cowpea root induced by A1treatment, exogenous IAA significantly increased the malate secretion, and exogenous Ca2+channel inhibitors and chelator significantly inhibited the malate secretion. Meanwhile the organic acid content and enzyme activity relate to organic acid metabolism did not changed. So we presume that IAA and Ca2+may involved in the regulation of malate secretion induced by Al, but the regulation may be not related to malate metabolism.
引文
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