光学无损分析技术在植物铝毒性研究中的应用

光学技术以它所特有的无损、快速和在位分析的优势,广泛应用于植物逆境生理监测和逆境胁迫机制研究.作为一类最主要的逆境胁迫,铝毒性是酸性土壤限制作物生长的主要原因.本文从光学分子成像和在位实时监测两个功能层面上概述了光学分析技术在铝毒生物学机制研究和铝毒性生理探测中的应用,并对其做了展望,同时,也讨论了铝毒性生物胁迫机制尚未涉及的问题,以期为铝胁迫机制研究提供有用的技术和理论参考.     

酸性土壤上植物应对铝胁迫的过程与机制

铝胁迫是酸性土壤上影响作物产量最重要的因素之一.目前,全球土壤酸化程度进一步加剧了铝胁迫.植物可通过将铝离子与有机酸螯合储藏于液泡和从根系中排出铝毒.排出铝毒主要通过苹果酸转运蛋白ALMT和柠檬酸转运蛋白MATE的跨膜运输来实现.编码ABC转运蛋白和锌指转录因子的基因与植物抗铝胁迫有关.这些抗铝毒基因的鉴别使得通过转基因和分子标记辅助育种等生物技术来提高农作物的抗铝毒能力成为可能.最后提出了植物抗铝胁迫研究中需要解决的关键问题及今后的研究方向.     

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.     

植物适应酸铝胁迫机理的研究进展

酸铝胁迫是限制植物正常生长发育的重要非生物胁迫因子,严重制约了我国酸性土壤地区的农业生产水平。植物抵御酸铝胁迫的形式复杂多样,如分泌有机酸、提高根际pH、分泌黏液、细胞壁对Al³⁺的固定、有机酸对细胞溶质中Al³⁺的螯合与液泡区隔化等。现有研究多集中于常规生理特征分析,缺乏深入的分子生物学解析。基于此,本文对国内外植物适应酸铝胁迫机理的相关研究进行了归纳和总结,从酸铝胁迫对植物生长与生理代谢的影响、植物适应酸铝胁迫最主要的两种生理机制(Al排除机制、Al耐受机制)以及分子水平上调控相关耐铝基因进行了综述。最后针对现有研究的不足提出了展望,以期为深入揭示植物适应酸铝胁迫的机理以及挖掘适于酸土生长的优质作物资源提供理论依据。     

植物铝胁迫响应基因的研究进展

铝毒是酸性土壤中植物生长和作物生产的主要限制因子.近年来的很多研究应用差异显示PCR、抑制差减cDNA文库和DNA微正列等技术,在一些铝耐受型和敏感型植物中鉴定了很多铝胁迫响应基因.本研究通过参阅国内外有关报道和结合本实验室的研究成果,从铝诱导的通道蛋白、代谢相关、胁迫和细胞死亡以及信号转导相关基因4个方面的研究进展进行了综述.     

微生物铝毒害和耐铝毒机制研究进展

在酸性土壤中,铝毒是限制农作物生产的关键问题之一,铝同样对微生物产生毒害作用。研究微生物的铝毒害和耐铝毒机制可以为植物耐铝毒机制的研究提供一种新视角。目前的研究表明,铝作用于微生物细胞的细胞壁、细胞膜、细胞核和细胞器,影响微生物的物质和能量代谢,抑制微生物的生长和发育。针对这些毒害作用,铝毒耐受微生物通过多途径全方位的机制适应外界的铝毒环境。该文结合作者的研究工作,综述了微生物的铝毒害和耐铝毒机制。     

植物地上部对铝毒的生理响应及其耐性

全世界50％以上潜在的可耕地属于酸性土壤,铝毒害是酸性土壤上植物生长最有害因素之一。近年来,为了阐明植物铝毒害及其耐性,前人已进行了大量的研究,并有一些综述性文章发表。然而,大多数文章主要综述铝对植物根系的影响及其耐性,因为根生长受抑是最早的铝毒害症状之一和溶液培养时最容易辨认的铝毒害症状。为此,本文综述了铝对植物地上部光合作用、光保护系统、水分利用效率、含水量、碳水化合物含量、矿质营养、有机酸和氮代谢的影响,并对富铝植物的解铝毒机制（铝与小分子有机酸螯合和把铝隔离在对铝不敏感的表皮细胞和液泡内）进行了综述。本文还对植物耐铝遗传学和分子生物学及今后需要研究的问题进行了讨论。     

细胞壁在植物抗营养逆境中的作用及其分子生理机制

营养逆境是指土壤中必需营养元素供应不足或元素含量过剩的现象,这种现象在世界范围内非常普遍,可导致作物减产和品质下降,因此,深入解析植物应对这种逆境的机制对提高作物产量、改善品质至关重要,也对保证农业生产的健康可持续发展意义重大.铝毒是限制酸性土壤上作物生产的主要因素,而镉又是重要的重金属污染元素.鉴于细胞壁是接触和响应环境因子的第一道屏障,也是结合金属离子的主要部位.结合本实验室近年来取得的一些研究成果,从细胞壁中的果胶、半纤维素组成及其修饰以及可能参与其中的相关基因等方面综述了细胞壁在植物抗铝和镉胁迫中的作用及其分子生理机制,以期引起对该领域更多的关注.     

丹波黑大豆GmbHLH30转录因子耐铝功能初步研究

铝毒是酸性土壤作物生长的主要限制因素。前期研究发现,铝胁迫下,耐铝型丹波黑大豆SSH(suppression subtractive hybridization,SSH)cDNA文库中bHLH30转录因子基因上调表达,推测该基因与丹波黑大豆耐铝性相关。克隆GmbHLH30基因,构建GmbHLH30植物表达载体pK2-35S-GmbHLH30,并在烟草中过量表达获得转GmbHLH30的转基因烟草植株。在铝胁迫下,转GmbHLH30的转基因烟草相对根伸长率比野生型烟草大,可溶性糖和脯氨酸含量高,H2O2水平低。表明GmbHLH30基因的过量表达可以增强植物的耐铝能力,暗示GmbHLH30转录因子参与调控植物的耐铝特性。     

铝诱导植物程序性细胞死亡信号转导的研究进展

铝是制约酸性土壤上作物生产的主要因素。铝诱导氧化胁迫产生大量活性氧/一氧化氮,引起胞质钙超载,通过线粒体信号转导途径激发相关凋亡基因,从而引起细胞主动死亡,以减轻铝对植物的进一步毒害。本文综述了铝诱导程序性细胞死亡的信号分子、相关基因以及信号转导途径,对未来的研究方向提出了展望,为深入研究植物铝毒害机理和耐铝机制提供参考。     

The Physiology, Genetics and Molecular Biology of Plant Aluminum Resistance and Toxicity

Aluminum (Al) toxicity is the primary factor limiting crop production on acidic soils (pH values of 5 or below), and because 50% of the world’s potentially arable lands are acidic, Al toxicity is a very important limitation to worldwide crop production. This review examines our current understanding of mechanisms of Al toxicity, as well as the physiological, genetic and molecular basis for Al resistance. Al resistance can be achieved by mechanisms that facilitate Al exclusion from the root apex (Al exclusion) and/or by mechanisms that confer the ability of plants to tolerate Al in the plant symplasm (Al tolerance). Compelling evidence has been presented in the literature for a resistance mechanism based on exclusion of Al due to Al-activated carboxylate release from the growing root tip. More recently, researchers have provided support for an additional Al-resistance mechanism involving internal detoxification of Al with carboxylate ligands (deprotonated organic acids) and the sequestration of the Al-carboxylate complexes in the vacuole. This is a field that is entering a phase of new discovery, as researchers are on the verge of identifying some of the genes that contribute to Al resistance in plants. The identification and characterization of Al resistance genes will not only greatly advance our understanding of Al-resistance mechanisms, but more importantly, will be the source of new molecular resources that researchers will use to develop improved crops better suited for cultivation on acid soils.     

Conserved functions of MATE transporters and their potential to enhance plant tolerance to aluminium toxicity

Almost half of the world’s arable land has acidic pH. Aluminum salts present in acid soils dissociate to release Al³⁺ ions in the soil solution that inhibit root growth causing severe loss in crop yields. Aluminium toxicity accounts for the second highest loss in plant productivity after drought. Aluminium in high doses causes damage to the plant cell wall, cytoskeleton and DNA. One of the ways by which plants alleviate aluminium toxicity is by the exudation of citrate from the roots that chelates the free Al³⁺ and prevents its entry into the plant. In several crop plants Multidrug and Toxic Compound Extrusion (MATE) transporters regulate citrate exudation from the roots. The MATE proteins are ubiquitously present in bacteria, archaea, fungi, animals and plants. The origin and evolution of these membrane transporters in plants is not well known. Here, using protein sequence information we identify MATE transporters in major groups of land plants and their algal ancestors. Our study indicates that the MATE family members expanded in number and functionally diverse among the land plants. We also identify motifs present across the streptophyte clade and a conserved aspartate residue that might regulate citrate exudation. This study can provide leads to engineer MATE transporters to confer enhanced tolerance in acid soils.

     

Toxicity and tolerance of aluminum in plants: tailoring plants to suit to acid soils

Aluminum (Al) stress is one of the serious limiting factors in plant productivity in acidic soils, which constitute about 50 % of the world’s potentially arable lands and causes anywhere between 25 and 80 % of yield losses depending upon the species. The mechanism of Al toxicity and tolerance has been examined in plants, which is vital for crop improvement and enhanced food production in the future. Two mechanisms that facilitate Al tolerance in plants are Al exclusion from the roots and the ability to tolerate Al in the symplast or both. Although efforts have been made to unravel Al-resistant factors, many aspects remain unclear. Certain gene families such as MATE, ALMT, ASR, and ABC transporters have been implicated in some plants for resistance to Al which would enhance the opportunities for creating crop plants suitable to grow in acidic soils. Though QTLs have been identified related to Al-tolerance, no crop plant that is tolerant to Al has been evolved so far using breeding or molecular approaches. The remarkable changes that plants experience at the physiological, biochemical and molecular level under Al stress, the vast array of genes involved in Al toxicity-tolerance, the underlying signaling events and the holistic image of the molecular regulation, and the possibility of creating transgenics for Al tolerance are discussed in this review.     

Aluminium speciation and phytotoxicity in alkaline soils

Aim

Highly alkaline soils (pH?>?9.0) may adversely affect agricultural crop productivity. Problems encountered include poor structure and nutrient deficiency. Research based on solution cultures suggests that aluminium (Al) phytotoxicity may occur in soils with pH?>?9.0, but little research has been undertaken on actual soils under controlled conditions. The nature of the Al species responsible and the pH regime of the soils when this occurs are unknown.

Methods

The charge and species of Al responsible for this toxicity was investigated using Zeta Potential measurement, Nuclear Magnetic Resonance (NMR) spectroscopy, Al precipitation characteristics and electrical conductivity as a function of pH. An anion exchange resin was used to evaluate Al availability to plants at alkaline pH. To verify Al phytotoxicity, a pot experiment was performed with plants grown at near neutral and high pH, with and without Al.

Results

The anionic aluminate species of aluminium was ubiquitous at highly alkaline pH, and was the dominant charged species at pH 9.2. Aluminium was phytotoxic at high pH, significantly reducing the stem and root development of field pea test plants over and above that caused by alkalinity alone. The effects of both alkalinity in general and aluminium in particular became noticeable at pH 9.0 and debilitating at pH?>?9.2.

Conclusion

As this corresponds to the pH where aluminate becomes dominant, it is probably responsible for the phytotoxicity.     

Studies on the manganese uptake of lettuce on steam-sterilised glasshouse soils

Summary During steam sterilisation of glasshouse soils appreciable amounts of easily reducible manganese are converted into exchangeable manganese. The reverse process takes place much more gradually. As a result, manganese toxicity occurs in several crops on newly steamed soils. In the Netherlands, lettuce has been found to be particularly prone to manganese toxicity. An investigation was carried out to obtain more information about the manganese status of steamed glasshouse soils in which lettuce was used as the test crop. The following results were noted.The uptake of exchangeable manganese is easier in the lighter soils than in heavy soils. Application of iron to the soil inhibits manganese uptake by the plant, but the iron must be applied in the form of chelate. The pH has a profound effect on manganese uptake on steamed as well as on unsteamed soils. However, the relationship between the pH and the manganese content of the crop on steamed soils is different from that found on unsteamed soils.The slow rate of oxidation of manganese in steamed glasshouse soil may be explained by the fact that the oxidising bacteria are killed during the steam sterilisation process. The fixation of manganese can be accelerated appreciably by inoculating the steamed soil with manganese-oxidising bacteria.The effect on manganese uptake of five soil desinfection chemicals used in the investigation proved to be very small.The lettuce varieties used in the Netherlands show a wide variation in susceptibility to manganese toxicity. This cannot be explained by different rates of manganese uptake. It is more likely that the varietal differences are based on different levels of resistance to manganese present in the plants.     

Si amelioration of Al toxicity in barley (Hordeum vulgare L.) growing in two Andosols

The influence of silicon on aluminium toxicity in barley (Hordeum vulgare L. cv. Shunrai) was studied in two Andosols. Silicon sources were a solution of sodium metasilicate with pH adjusted to 5.0, silica gel, and an industrial waste, porous hydrated calcium silicate. The waste is produced in large amounts in the manufacturing processes of autoclaved light concrete, and has been used as a silicon source for rice plants. The addition of the waste increased the concentration of Si in the soil solution, soil pH and amelioration of aluminium toxicity was observed. The addition of silica gel and sodium metasilicate solution to both soils increased significantly (p<0.05) the Si concentration of the soil solutions, but no amelioration of aluminium toxicity was observed. An amelioration of aluminium toxicity by the waste porous hydrated calcium silicate was probably due to the increase in soil pH rather than to the increase of silicon concentration in the soil solution.     

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.     

Boron Increases Synthesis of Glutathione in Sunflower Plants Subjected to Aluminum Stress

Ionic stress caused by high aluminum (Al) concentrations is one of the most widespread phytotoxicity problems globally in agricultural regions, greatly limiting crop yield in affected areas. The objective of this work was to examine a possible involvement of boron (B) in the detoxification of Al by stimulating glutathione (GSH) metabolism, a mechanism essential for the resistance of plants under stress conditions. Our results clearly demonstrate that increased application of B in the presence of high Al concentrations in the growth medium stimulates GSH biosynthesis, suggesting it could be an effective strategy to combat stress associated with the formation of active-oxygen species (AOS). In the specific case of Al toxicity, B reduces phytotoxicity by stimulating leaf biosynthesis of GSH and an increase in its concentration in the roots. Therefore, in this work, we also identify GSH metabolism as one of the key processes in Al detoxification. Finally, our results imply that greater B application leads to a greater resistance to Al toxicity, a fact that might be significant for higher productivity of agricultural plants grown in acid soils.