有机酸在植物解铝毒中的作用及生理机制

酸性土壤上铝毒是限制作物产量的一个重要障碍因子。具有螯合能力的有机酸在植物铝的外部排斥机制和内部耐受机制均具有重要作用。在铝的外部排斥解毒过程中,植物通过根系分泌有机酸进入根际,如柠檬酸、草酸、苹果酸等与铝形成稳定的复合体,阻止铝进入共质体,从而达到植物体外解除铝毒害效应的目的,且分泌的有机酸对铝的胁迫诱导表现出高度的专一性,分泌的关键点位于根尖。不同的物种间分泌的有机酸种类、分泌的模式及生理机理存在差异。在铝积累型植物的内部解毒过程中,有机酸与铝形成稳定的化合物,降低植物体内铝离子的生理活性,从而降低细胞内铝离子的毒害效应,如绣球花中铝与柠檬酸形成1:1的复合体,荞麦内铝与草酸形成1:3的复合体。本文就有机酸在植物忍耐和积累铝中的作用及生理机制作一简要综述。     

高等植物金属抗性中有机酸的作用及其机理

植物的金属抗性可通过避性和耐性途径获得。具有螯合能力的有机酸在植物的金属外部排斥(避性)机制和内部耐受(耐性)机制中均具有重要作用。在金属的外部排斥过程中,植物根系分泌有机酸,与金属离子形成稳定的复合体,降低土壤金属的移动性,达到体外解毒的目的。超积累型植物的内部耐受机制主要体现在,有机酸可与金属元素发生螯合作用,将离子态的金属转变成低毒或无毒的螯合态,从而降低细胞内金属离子的毒害效应。有机酸的种类受植物种类、金属类型等因素的影响。     

有机酸在植物对重金属耐性和解毒机制中的作用

植物对重金属的耐受和解毒机制可分为外部排斥和内部耐受两大类。该文综述了有机酸作为一类金属配位体, 在植物对重金属的这两大类机制中的重要作用。在重金属的外部排斥过程中, 植物根系分泌有机酸, 与金属离子形成稳定的金属配位体复合物, 改变重金属的移动性和生物可利用性, 阻止金属离子进入植物体内或避免其在根部敏感位点累积。此外, 有机酸还可与进入植物体内的金属离子螯合, 使其转化为无毒或毒性较小的结合形态, 缓解重金属的毒害效应, 实现植物对重金属的内部耐受。     

铝胁迫下植物根系的有机酸分泌及其解毒机理

酸性土壤中的铝毒害问题,已成为限制植物生长发育的主要因素之一.耐铝植物通过根系分泌有机酸来解除或减轻铝的毒害是外部解铝毒的重要机制.文章对铝胁迫下植物根系分泌有机酸的种类,有机酸解铝毒机理、解铝毒能力,有机酸分泌方式及调控其分泌的主要因素等相关研究进行综述.     

植物耐铝的生物化学与分子机理

某些耐铝植物在铝胁迫下分泌有机酸被认为是一个重要的抗性机制.从根系分泌出来的有机酸能与根际的Al3 结合,形成无毒性的螯合物,从而减轻了铝对根系的毒害.但是,铝诱导有机酸分泌的中间环节及调节机制至今仍不清楚.一些证据表明,铝能激活根尖细胞质膜内的阴离子通道,因而可以调节有机酸的分泌.近年来,人们开始注意一些信号分子如蛋白激酶、水杨酸等介导铝诱导有机酸的分泌,已经获得一些成果.同时,铝胁迫基因的分离和鉴定也为人们从分子水平上研究和认识铝胁迫下植物的抗性机制奠定了基础.     

大豆耐铝性品种差异及其与有机酸的关系

从 1 0个大豆品种中筛选出两个耐铝性差异显著的品种 ,研究了其耐铝性与有机酸的关系。经铝处理后 ,吴川品种的相对根长为 1 3 3 .5 % ,化州只有 68.9% ,表明吴川相对耐铝 ,化州对铝较敏感。将不同浓度的AlCl3 加入营养液中处理大豆 1 0d,化州较吴川根长受到较大影响 ,进一步证实吴川相对耐铝毒 ,而化州对酸铝敏感。机理研究发现大豆在铝胁迫下根系可分泌两种有机酸 (草酸、柠檬酸 ) ,其中吴川根系草酸分泌速率提高了 74% ,化州几乎没有提高 ,表明耐铝性大豆品种的根系草酸分泌速率明显提高 ,可增强其缓解酸铝毒性的能力。而二者分泌柠檬酸的速率虽然均有显著提高 ,但处理后感抗品种之间差异不大 ,表明柠檬酸在缓解铝毒性中的作用不大。铝处理下大豆根系虽然分泌两种有机酸 ,但草酸在大豆耐酸铝机制中的作用可能更为重要。     

排根的形成及其所分泌的有机酸的调节

排根是在磷、铁、氮等养分缺乏条件下形成的一种特殊根系结构,能够形成排根的植物种类有限.不同植物排根分泌的有机酸种类和数量可能不同,如在缺磷条件下白羽扇豆的排根所释放的柠檬酸可达植物总干重的23%,使根簇周围的柠檬酸浓度达50～90 μmol*g-1土壤.这是一种防止胞质过度酸化以及柠檬酸过度积累而超过液泡储存能力的解毒机制.其分泌可能受阴离子通道的调控.     

植物铝毒害及遗传育种研究进展

本文简单概述了目前植物铝毒害及遗传育种方面的研究进展。Al3+可以通过与细胞骨架的作用,影响根的正常生理功能和形态建成。 植物可以通过根尖分泌有机酸或磷酸等将离子态的铝变成螯合态的铝,通过吸收H+提高根尖周围的pH,将Al3+变成难溶性的 Al(OH)3或磷酸铝从而解除铝毒害, 也可以通过在细胞内与Al3+形成无毒害的复合结构从而解除铝毒害。国外通过基因工程和突变体筛选已经获得了一批耐铝的植物材料,国内一些研究者通过突变体筛选也获得了一些耐铝的植物材料。 对植物耐铝性的遗传研究表明, 植物的耐铝性既可以是受单基因控制的,也可以是受多基因控制的。     

植物铝毒害及遗传育种研究进展

本文简单概述了目前植物铝毒害及遗传育方面的研究进展,Al^3 可以通过与细胞骨架的作用,影响根的正常生理功能和形态建成,植物可以通过根尖分泌有机酸或磷酸等将离子态的为成螯合态的铝,通过吸收H^ 提高根尖周围的pH,将Al^3 变成难溶性的Al(OH)3或磷酸铝从而解 除铝毒害,也可以通过在细胞内与Ａl^3 形成无毒害的复合结构从而解除铝毒害,国外通过基因工程和突变体筛选已经获得了一批耐铝的植物材料,国内一些研究者通过变体筛选也获得了一些耐铝的植物材料,对植物耐铝性的遗传研究表明,植物的耐铝性既可以是受单基因控制的,也可以是受多基因控制的。     

铝胁迫下两种外生菌根真菌的有机酸分泌与吸收动力学特征

为探索菌丝体吸附及有机酸分泌在外生菌根真菌(ECMF)抗铝性中的作用,将抗铝能力不同的两种ECMF[彩色豆马勃(Pisolithus tinctorius 715,Pt 715)和松乳菇(Lactarius deliciosus 2,Ld 2)]在酸性含Al~(3+)溶液中培养,检测它们的有机酸分泌和吸收动力学特征。结果发现:(1)两种ECMF分泌的有机酸种类和多寡因菌种而异,且不因铝胁迫发生改变:Ld 2为甲酸草酸,Pt 715为酒石酸甲酸草酸丁二酸。铝胁迫下Ld 2分泌的两种有机酸均显著增加,而Pt 715分泌的酒石酸和甲酸显著增加,丁二酸显著降低,草酸未显著改变;(2)两种ECMF对Al~(3+)的吸收符合离子的吸收动力学模型,为主动吸收。Ld 2的CEC、C_(min)、I_(max)以及吸附性Al~(3+)都高于Pt 715,而K_m和吸收性Al~(3+)的值低于Pt 715。即抗铝性较强的Ld 2比抗铝性较弱的Pt 715有更多的阳离子吸附位点和更强的Al~(3+)吸附能力。因此,有机酸分泌和菌丝体对Al~(3+)的吸附都能提高ECMF的抗铝性;有机酸的分泌有利于养分的活化与利用,而菌丝体吸附能阻止Al~(3+)进入共质体而防止铝毒害。     

Role of organic acids in detoxification of aluminum in higher plants

Phytotoxicity of aluminum ion (Al3+) is a serious problem limiting crop production on acid soils. Organic acids with Al-chelating ability play an important role in the detoxification of Al both externally and internally. Al is detoxified externally by the secretion of organic acids such as citric, oxalic, and/or malic acids from the roots. The secretion of organic acids is highly specific to Al and the site of secretion is localized to the root apex. The kind of organic acids secreted as well as secretion pattern differ among plant species. There are two patterns of Al-induced secretion of organic acids: In pattern I, there is no discernible delay between the addition of Al and the onset of the release of organic acids. Activation of the anion channel seems to be involved in this pattern; In pattern II, there is a marked lag phase between the addition of Al and the onset of organic acid release. The action of genes related to the metabolism and secretion of organic acids seems to be involved in this pattern. Internal detoxification of Al in Al-accumulating plants is achieved by the formation of Al-organic acid complex. For instance, the complex of Al-citrate (1:1) in hydrangea and Al-oxalate (1:3) in buckwheat has been identified.     

Recent progress in the research of external Al detoxification in higher plants: a minireview

Aluminum (Al) is highly toxic to plant growth. The toxicity is characterized by rapid inhibition of root elongation. However, some plant species and cultivars have evolved some mechanisms for detoxifying Al both internally and externally. In this review, the recent progress made in the research of external detoxification of Al is described. Accumulating evidence has shown that organic acids play an important role in the detoxification of Al. Some plant species and cultivars respond to Al by secreting citrate, malate or oxalate from the roots. Recently, the anion channel of malate and citrate in the plasma membrane has been characterized and a gene encoding the malate channel has been cloned. The metabolism of organic acids seems to be poorly correlated with the Al-induced secretion of organic acid anions. A number of QTLs (quantitative trait loci) for Al resistance have been identified in rice, Arabidopsis, and other species. Transgenic plants with enhanced resistance to Al have also been reported, but introduction of multiple genes may be required to gain high Al resistance in future.     

AHA1转基因拟南芥的抗铝性生理解析

AHA1基因是植物体内编码质膜H⁺ATPase的一个重要基因,参与植物的生长发育与抗逆胁迫反应.本文以AHA1转基因型及其野生型拟南芥为材料,研究了铝胁迫对拟南芥养分吸收、抗氧化胁迫和有机酸分泌的影响.结果表明：Al降低了拟南芥根系对N、K、Ca和Mg的吸收,但增加了根系对P的吸收,且AHA1转基因型拟南芥比野生型积累更多的P和较少的Al.铝胁迫诱导拟南芥抗氧化酶SOD和POD活性增加,转基因型与野生型之间没有明显差异.Al对拟南芥有机酸分泌具有明显的诱导作用,且AHA1转基因型分泌较多的有机酸.质膜H⁺ATPase的活性抑制剂钒酸盐对拟南芥有机酸分泌具有明显的抑制作用;而Zn²⁺、Mg²⁺可促进Al对拟南芥有机酸分泌的诱导,并部分恢复钒酸盐的抑制效应.说明AHA1基因通过增加拟南芥根系对P的吸收以及有机酸分泌,提高了植物的抗铝性.     

The role of arbuscular mycorrhizas in decreasing aluminium phytotoxicity in acidic soils: a review

Soil acidity is an impediment to agricultural production on a significant portion of arable land worldwide. Low productivity of these soils is mainly due to nutrient limitation and the presence of high levels of aluminium (Al), which causes deleterious effects on plant physiology and growth. In response to acidic soil stress, plants have evolved various mechanisms to tolerate high concentrations of Al in the soil solution. These strategies for Al detoxification include mechanisms that reduce the activity of Al³⁺ and its toxicity, either externally through exudation of Al-chelating compounds such as organic acids into the rhizosphere or internally through the accumulation of Al–organic acid complexes sequestered within plant cells. Additionally, root colonization by symbiotic arbuscular mycorrhizal (AM) fungi increases plant resistance to acidity and phytotoxic levels of Al in the soil environment. In this review, the role of the AM symbiosis in increasing the Al resistance of plants in natural and agricultural ecosystems under phytotoxic conditions of Al is discussed. Mechanisms of Al resistance induced by AM fungi in host plants and variation in resistance among AM fungi that contribute to detoxifying Al in the rhizosphere environment are considered with respect to altering Al bioavailability.     

Aluminum resistance in two cultivars of Zea mays L.: Root exudation of organic acids and influence of phosphorus nutrition

Root exudation of organic acids as Al-chelating compounds and P nutrition have been suggested to play a major role in Al-resistance in higher plants. Effects of Al exposure on maize plant growth, and organic acid root content and root exudation under various levels of P nutrition were examined. Sikuani, a Colombian maize cultivar tolerant to acid soils with high Al saturation, and Corso, a Swiss cultivar, were grown in sterile hydroponic conditions for 21 days. Al-caused inhibition of root growth was lower in Sikuani than in Corso. Al effect on plant growth was decreased with increasing P content in roots. Al content in roots increased with increasing P content and was higher in Sikuani than in Corso. When exposed to Al, the contents in root apices as well as the root exudation of citric and malic acids in Corso and citric, malic and succinic acids in Sikuani increased, and were higher in Sikuani than in Corso. Increased PEP carboxylase (PEPC) activity in root apices after Al exposure partially explained the variations of organic acid content in the roots. These Al-induced changes in PEPC activity, organic acid content and exudation were reduced in plants supplied with higher P concentrations during the 21 days prior to treatment. Increased secretion of organic acids after exposure to Al appeared to be specific to Al and was not totally explained by increased root content in organic acids.