Aluminium/silicon interactions in higher plants

Aluminium and silicon are usually abundant in soil mineral matter,but their availability for plant uptake is limited by low solubilityand, in the case of Al, high soil pH causes precipitation ofthe element in insoluble forms. Al toxicity is a major problemin naturally occurring acid soils and in soils affected by acidicprecipitation. Al has no known role in higher plants, and isgenerally known as a toxic element, whereas Si is generallyregarded as a beneficial element. Recently, it has been suggestedthat Al toxicity can be ameliorated by Si in a variety of animalsystems. In this review the evidence that amelioration of Altoxicity by Si can also occur in plants is assessed. At presentsuch amelioration has been shown in sorghum, barley, teosinte,and soybean, but not in rice, wheat, cotton, and pea. Plantspecies vary considerably in the amounts of Al and Si that theytransport into their tissues, and it seems that very high Siaccumulation and very high Al accumulation are mutually exclusive.The mechanisms considered for amelioration are: solution effects;codeposition of Al and Si within the plant; effects in the cytoplasmand on enzyme activity; and indirect effects. Key words: Aluminium, silicon, biomineralization, codeposition, toxicity, tolerance     

Effects of silicon on aluminum toxicity in upland rice plants

Aims

This study aimed to determine the capacity of Si to mitigate Al toxicity in upland rice plants (Oryza sativa L.) by evaluating plant growth and the Si and Al uptake kinetics.

Methods

Plants were grown for 40 days, after which the Si and Al uptake kinetics (Cmin, Km and Imax) were analyzed. Then, the shoots and roots were separated, and the dry matter, root morphology and Si and Al concentration and accumulation in the plant were evaluated.

Results

Aluminum decreased plant growth and the Si uptake capacity by decreasing the root growth and Si transport system efficiency in the upland rice roots (> Km and > Cmin). Silicon mitigated Al toxicity in the upland rice plants by decreasing Al transport to the plant shoots, although it did not reduce the Al uptake rate (Imax). Si treatment increased the growth of upland rice plant shoots grown in the presence of Al without influencing the root growth. The alleviation of Al toxicity by Si is more evident in the susceptible upland rice cultivar Maravilha.

Conclusions

Silicon mitigated Al toxicity in the upland rice plants by decreasing Al transport to the plant shoots but did not reduce the Al uptake rate by roots.

     

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.     

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.     

Linear relation between leaf xylem water potential and transpiration in pearl millet during soil drying

Plant and Soil - Populus can tolerant high concentration Al stress. However, the mechanisms of Mg alleviation to Al toxicity in populus remain unknown. In the present study, adequate Mg was...     

Mechanisms of metal resistance in plants: aluminum and heavy metals

Plants have evolved sophisticated mechanisms to deal with toxic levels of metals in the soil. In this paper, an overview of recent progress with regards to understanding fundamental molecular and physiological mechanisms underlying plant resistance to both aluminum (Al) and heavy metals is presented. The discussion of plant Al resistance will focus on recent advances in our understanding of a mechanism based on Al exclusion from the root apex, which is facilitated by Al-activated exudation of organic acid anions. The consideration of heavy metal resistance will focus on research into a metal hyperaccumulating plant species, the Zn/Cd hyperaccumulator, Thlaspi caerulescens, as an example for plant heavy metal research. Based on the specific cases considered in this paper, it appears that quite different strategies are used for Al and heavy metal resistance. For Al, our current understanding of a resistance mechanism based on excluding soil-borne Al from the root apex is presented. For heavy metals, a totally different strategy based on extreme tolerance and metal hyperaccumulation is described for a hyperaccumulator plant species that has evolved on naturally metalliferous soils. The reason these two strategies are the focus of this paper is that, currently, they are the best understood mechanisms of metal resistance in terrestrial plants. However, it is likely that other mechanisms of Al and/or heavy metal resistance are also operating in certain plant species, and there may be common features shared for dealing with Al and heavy resistance. Future research may uncover a number of novel metal resistance mechanisms in plants. Certainly the complex genetics of Al resistance in some crop plant species, such as rice and maize, suggests that a number of presently unidentified mechanisms are part of an overall strategy of metal resistance in crop plants.     

Role of magnesium in alleviation of aluminium toxicity in plants

Magnesium is pivotal for activating a large number of enzymes; hence, magnesium plays an important role in numerous physiological and biochemical processes affecting plant growth and development. Magnesium can also ameliorate aluminium phytotoxicity, but literature reports on the dynamics of magnesium homeostasis upon exposure to aluminium are rare. Herein existing knowledge on the magnesium transport mechanisms and homeostasis maintenance in plant cells is critically reviewed. Even though overexpression of magnesium transporters can alleviate aluminium toxicity in plants, the mechanisms governing such alleviation remain obscure. Possible magnesium-dependent mechanisms include (i) better carbon partitioning from shoots to roots; (ii) increased synthesis and exudation of organic acid anions; (iii) enhanced acid phosphatase activity; (iv) maintenance of proton-ATPase activity and cytoplasmic pH regulation; (v) protection against an aluminium-induced cytosolic calcium increase; and (vi) protection against reactive oxygen species. Future research should concentrate on assessing aluminium toxicity and tolerance in plants with overexpressed or antisense magnesium transporters to increase understanding of the aluminium-magnesium interaction.     

Characteristics of boron accumulation by fly ash application in paddy soil

Fly ash has a high content of plant available silicate which is strongly needed for rice cultivation in Korea. One concern for plants grown on soils amended with fly ash is boron (B) toxicity because most of the fresh fly ash contains considerable B. This study was conducted in paddy soil to determine B uptake by rice and characteristics of B accumulation in soil after fly ash application (0, 40, 80, and 120 Mg fly ash ha⁻¹). In all fly ash treatments, B content in rice leaves and available B in soil at all growing stage were higher than those of control, but were not exceeded a toxicity levels. Boron occluded in amorphous Fe and Al oxides comprised ca. 20–39% of total B and was not affected by fly ash application. Most of the B was accumulated by fly ash application as a residual B which is plant-unavailable form, comprised >60% of the total B in soil. Thus, fly ash can be a good soil amendment for rice production without B toxicity.     

Interactions between aluminium,magnesium and calcium with different monocotyledonous and dicotyledonous plant species

Growth inhibition of plants suffering from Al toxicity is generally accompanied by impaired root development which can be quantitatively described by reduced specific root length (m g^-1 dry root). In addition, the uptake of nutrients such as Mg and Ca is inhibited. Increased supply of either Mg or Ca can significantly diminish the negative effect of Al on root development and improve the Mg or Ca nutrition of the plants. The positive effect of Ca is well established but the effect of Mg has been observed in only a few plan species. Therefore, the effects of increasing Mg and Ca supply on Al toxicity in plants of seven monocots and eight dicots have been now examined in nutrient solution experiments. In general, Mg appears to be more effective than Ca in alleviating Al toxicity with the monocots, whereas the reverse is true for the dicots. Increased concentrations of Mg and Ca in solution seem to protect the plants against Al toxicity by improving the Mg or Ca nutrition and by alleviating the toxic effect of Al on root development.     

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。     

How a microbial drug transporter became essential for crop cultivation on acid soils: aluminium tolerance conferred by the multidrug and toxic compound extrusion (MATE) family

Background

Aluminium (Al) toxicity is a major agricultural constraint for crop cultivation on acid soils, which comprise a large portion of the world''s arable land. One of the most widely accepted mechanisms of Al tolerance in plants is based on Al-activated organic acid release into the rhizosphere, with organic acids forming stable, non-toxic complexes with Al. This mechanism has recently been validated by the isolation of bona-fide Al-tolerance genes in crop species, which encode membrane transporters that mediate Al-activated organic acid release leading to Al exclusion from root apices. In crop species such as sorghum and barley, members in the multidrug and toxic compound extrusion (MATE) family underlie Al tolerance by a mechanism based on Al-activated citrate release.

Scope and Conclusions

The study of Al tolerance in plants as conferred by MATE family members is in its infancy. Therefore, much is yet to be discovered about the functional diversity and evolutionary dynamics that led MATE proteins to acquire transport properties conducive to Al tolerance in plants. In this paper we review the major characteristics of transporters in the MATE family and will relate this knowledge to Al tolerance in plants. The MATE family is clearly extremely flexible with respect to substrate specificity, which raises the possibility that Al tolerance as encoded by MATE proteins may not be restricted to Al-activated citrate release in plant species. There are also indications that regulatory loci may be of pivotal importance to fully explore the potential for Al-tolerance improvement based on MATE genes.     

Development of an electrostatic model predicting copper toxicity to plants

The focus of the present study was to investigate the mechanisms for the alleviation of Cu toxicity in plants by coexistent cations (e.g. Al(3+), Mn(2+), Ca(2+), Mg(2+), H(+), Na(+), and K(+)) and the development of an electrostatic model to predict 50% effect activities (EA50s) accurately. The alleviation of Cu(2+) toxicity was evaluated in several plants in terms of (i) the electrical potential at the outer surface of the plasma membrane (PM) (Ψ(0)(°)) and (ii) competition between cations for sites at the PM involved in the uptake or toxicity of Cu(2+), the latter of which is invoked by the Biotic Ligand Model (BLM) as the sole explanation for the alleviation of toxicity. The addition of coexistent cations into the bulk-phase medium reduces the negativity of Ψ(0)(°) and hence decreases the activity of Cu(2+) at the PM surface. Our analyses suggest that the alleviation of toxicity results primarily from electrostatic effects (i.e. changes in both the Cu(2+) activity at the PM surface and the electrical driving force across the PM), and that BLM-type competitive effects may be of lesser importance in plants. Although this does not exclude the possibility of competition, the data highlight the importance of electrostatic effects. An electrostatic model was developed to predict Cu(2+) toxicity thresholds (EA50s), and the quality of its predictive capacity suggests its potential utility in risk assessment of copper in natural waters and soils.     

Plant Root Responses to Three Abundant Soil Minerals: Silicon,Aluminum and Iron

Silicon (Si), aluminum (Al), and iron (Fe) are the three most abundant minerals in soil; however, their effects on plants differ because they are beneficial, toxic, and essential to plant growth, respectively. High accumulation of silicon in the shoots helps some plants to overcome a range of biotic and abiotic stresses. However, plants vary in their ability to take up Si from the soil and load it into the xylem and so the accumulation of silicon varies greatly between plant species. Aluminum toxicity is characterized by a rapid inhibition of root elongation but some species and even genotypes within species can tolerate Al toxicity better than others. While the mechanisms controlling this tolerance in most of the more resistant species are poorly understood, some plants are able to detoxify Al externally and/or internally by complexation with ligands or by pH changes in the rhizosphere. Iron is taken up from the soil by two efficient mechanisms called Strategy I and Strategy II, which operate in distinct phylogenic groups. Strategy I plants increase soil Fe solubility by releasing protons and reductants/chelators, such as organic acids and phenolics, into the rhizosphere, while Strategy II plants are characterized by the secretion of ferric chelating substances (phytosiderophores) coupled with a specific Fe³⁺: chelate uptake system. In this review, the molecular mechanisms underlying root response to Si, Al, and Fe are described.

     

OsFRDL1 Is a Citrate Transporter Required for Efficient Translocation of Iron in Rice

Multidrug and toxic compound extrusion (MATE) transporters represent a large family in plants, but their functions are poorly understood. Here, we report the function of a rice (Oryza sativa) MATE gene (Os03g0216700, OsFRDL1), the closest homolog of barley (Hordeum vulgare) HvAACT1 (aluminum [Al]-activated citrate transporter 1), in terms of metal stress (iron [Fe] deficiency and Al toxicity). This gene was mainly expressed in the roots and the expression level was not affected by either Fe deficiency or Al toxicity. Knockout of this gene resulted in leaf chlorosis, lower leaf Fe concentration, higher accumulation of zinc and manganese concentration in the leaves, and precipitation of Fe in the root's stele. The concentration of citrate and ferric iron in the xylem sap was lower in the knockout line compared to the wild-type rice. Heterologous expression of OsFRDL1 in Xenopus oocytes showed transport activity for citrate. Immunostaining showed that OsFRDL1 was localized at the pericycle cells of the roots. On the other hand, there was no difference in the Al-induced secretion of citrate from the roots between the knockout line and the wild-type rice. Taken together, our results indicate that OsFRDL1 is a citrate transporter localized at the pericycle cells, which is necessary for efficient translocation of Fe to the shoot as a Fe-citrate complex.     

Analysis of acid-soil stress in sorghum genotypes with emphasis on aluminium and magnesium interactions

Acid-soil stress in 12 sorghum (Sorghum bicolor (L.) Moench) genotypes was attributed mainly to aluminium (Al) toxicity. Root damage and magnesium (Mg) deficiency are two independent aspects of plant sensitivity to Al, either in acid soil or in nutrient solution. At moderate soil acidity, Mg deficiency dominantly limited growth whilst at high acidity root damage overruled the effect of Mg deficiency on the growth response. In nutrient solutions containing Al, increased Mg supply improved both root development and Mg nutrition of plants, whereas increased calcium (Ca) supply, or nutrition with ammonium (NH₄) instead of nitrate (NO₃), alleviated root damage but amplified Mg deficiency. At lowered pH the syndrome of Al toxicity was more profound. The implications of Mg-Al interactions, root damage, Mg supply and genotype selection are elucidated.     

Al-induced cell wall hydroxyproline-rich glycoprotein accumulation is involved in alleviating Al toxicity in rice

Cell wall components such as pectin and hemicelluloses have been proposed to be involved in aluminum resistance mechanisms in plants. However, whether hydroxyproline-rich glycoproteins (HRGPs), one of the most abundant proteins of the cell walls, are involved in Al resistance mechanisms remains elusive. In this study, two rice cultivars Xiushui 03 (Al resistant) and Xiushui 128 (Al sensitive) significantly differing in Al resistance were identified. In the absence of Al, no significant difference was observed in contents of glycoproteins and hydroxyproline in cell wall fractions of these two cultivars. At the early stage of Al toxicity, glycoproteins and hydroxyproline were significantly induced in these two cultivars, but levels of their accumulation in cell walls were much higher in cv. Xiushui 03 than in cv. Xiushui 128. At the late stage of Al toxicity, their accumulation in cell walls dramatically decreased in cv. Xiushui 128 and, however, still kept a high level in cv. Xiushui 03. The finding that Al-induced changes of glycoproteins and hydroxyproline were completely consistent indicates that Al-induced glycoproteins are HRGPs. Further observation utilizing transmission electron microscope showed that HRGPs were greatly accumulated in cell walls leading to thickening of cell walls in cv. Xiushui 03, however, HRGPs and cell walls greatly decreased in cv. Xiushui 128. These data suggest that Al-induced HRGP accumulation in cell walls is involved in alleviating Al toxicity in rice.