Aluminum Toxicity In Plants Research Articles

Aluminum Toxicity in Plants: Present and Future

Aluminum Toxicity in Plants: Present and Future

Phytohormone signalling and cross-talk to alleviate aluminium toxicity in plants.

Aluminium (Al) is one of the most abundant metals in earth crust, which becomes toxic to the plants growing in acidic soil. Phytohormones like ethylene, auxin, cytokinin, abscisic acid, jasmonic acid and gibberellic acid are known to play important role in regulating Al toxicity tolerance in plants. Exogenous applications of auxin, cytokinin and abscisic acid have shown significant effect on Al-induced root growth inhibition. Moreover, ethylene and cytokinin act synergistically with auxin in responding against Al toxicity. A number of studies showed that phytohormones play vital roles in controlling root responses to Al toxicity by modulating reactive oxygen species (ROS) signalling, cell wall modifications, organic acid exudation from roots and expression of Al responsive genes and transcription factors. This review provides a summary of recent studies related to involvement of phytohormone signalling and cross-talk with other pathways in regulating response against Al toxicity in plants.

Alleviating aluminum toxicity in plants: Implications of reactive oxygen species signaling and crosstalk with other signaling pathways.

Aluminum (Al) toxicity is a major limiting factor for plant growth and productivity in acidic soil. At pH lower than 5.0 (pH < 5.0), the soluble and toxic form of Al (Al3+ ions) enters root cells and inhibits root growth and uptake of water and nutrients. The organic acids malate, citrate, and oxalate are secreted by the roots and chelate Al3+ to form a non-toxic Al-OA complex, which decreases the entry of Al3+ into the root cells. When Al3+ enters, it leads to the production of reactive oxygen species (ROS) in cells, which are toxic and cause damage to biomolecules like lipids, carbohydrates, proteins, and nucleic acids. When ROS levels rise beyond the threshold, plants activate an antioxidant defense system that comprises of superoxide dismutase (SOD), catalase (CAT), ascorbate peroxidase (APX), glutathione S-transferase (GST), ascorbic acid (ASA), phenolics and alkaloids etc., which protect plant cells from oxidative damage by scavenging and neutralizing ROS. Besides, ROS also play an important role in signal transduction and influence many molecular and cellular process like hormone signaling, gene expression, cell wall modification, cell cycle, programed cell death (PCD), and development. In the present review, the mechanisms of Al-induced ROS generation, ROS signaling, and crosstalk with other signaling pathways helping to combat Al toxicity have been summarized, which will help researchers to understand the intricacies of Al-induced plant response at cellular level and plan research for developing Al-toxicity tolerant crops for sustainable agriculture in acid soil-affected regions of the world.

Extraction of metals from mildly acidic tropical soils: Interactions between chelating ligand, pH and soil type

Naturally occurring and synthetic chelating ligands can act as suppressants for fungal pathogens, nematodes and weeds, based on their ability to alter micronutrient bioavailability in soil, particularly iron. Chelators are also used as detergents, for remediation of heavy metal contamination and for supplying metals as fertiliser. The aim of this work was to test the ability of chelators to solubilise metals, in particular iron, in tropical soils over an environmentally relevant pH range. Six topsoils from farms in North Queensland, Australia were adjusted to pH 5, 6 and 7 and then extracted with CaCl2, EDTA, DTPA, EDDHA and mimosine. The extracts were analysed for concentrations of aluminium, copper, iron, magnesium, manganese, potassium, strontium and zinc. EDDHA solubilised iron effectively under all of the conditions tested, indicating its likely suitability for pest suppression. The concentration of aluminium in EDDHA extracts was positively correlated with pH, and at pH 7 the concentration of aluminium was far greater than that of iron. An increase in the mobility of aluminium from EDDHA application to soil may lead to aluminium toxicity in plants, which should be considered further in any practical application of EDDHA. Mimosine, which is also a strong chelator, was a poor extractor of all metals, possibly due to adsorption to the soil.

Importance of Mineral Nutrition for Mitigating Aluminum Toxicity in Plants on Acidic Soils: Current Status and Opportunities.

Aluminum (Al) toxicity is one of the major limitations that inhibit plant growth and development in acidic soils. In acidic soils (pH < 5.0), phototoxic-aluminum (Al3+) rapidly inhibits root growth, and subsequently affects water and nutrient uptake in plants. This review updates the existing knowledge concerning the role of mineral nutrition for alleviating Al toxicity in plants to acid soils. Here, we explored phosphorus (P) is more beneficial in plants under P-deficient, and Al toxic conditions. Exogenous P addition increased root respiration, plant growth, chlorophyll content, and dry matter yield. Calcium (Ca) amendment (liming) is effective for correcting soil acidity, and for alleviating Al toxicity. Magnesium (Mg) is able to prevent Al migration through the cytosolic plasma membrane in root tips. Sulfur (S) is recognized as a versatile element that alleviates several metals toxicity including Al. Moreover, silicon (Si), and other components such as industrial byproducts, hormones, organic acids, polyamines, biofertilizers, and biochars played promising roles for mitigating Al toxicity in plants. Furthermore, this review provides a comprehensive understanding of several new methods and low-cost effective strategies relevant to the exogenous application of mineral nutrition on Al toxicity mitigation. This information would be effective for further improvement of crop plants in acid soils.

Tropical soils with high aluminum concentrations cause oxidative stress in two tomato genotypes.

Tropical and subtropical soils are usually acidic and have high concentrations of aluminum (Al). Aluminum toxicity in plants is caused by the high affinity of the Al cation for cell walls, membranes, and metabolites. In this study, the response of the antioxidant-enzymatic system to Al was examined in two tomato genotypes: Solanum lycopersicum var. esculentum (Calabash Rouge) and Solanum lycopersicum var. cerasiforme (CNPH 0082) grown in tropical soils with varying levels of Al. Plant growth; activities of superoxide dismutase (SOD), catalase (CAT), ascorbate peroxidase (APX), guaiacol peroxidase (GPOX), and glutathione reductase (GR) enzymes; stress-indicating compounds (malondialdehyde (MDA) and hydrogen peroxide); and morphology (root length and surface area) were analyzed. Increased levels of Al in soils were correlated with reduced shoot and root biomass and with reduced root length and surface area. Calabash Rouge exhibited low Al concentrations and increased growth in soils with the highest levels of Al. Plants grown in soils with high availability of Al exhibited higher levels of stress indicators (MDA and hydrogen peroxide) and higher enzyme activity (CAT, APX, GPOX, and GR). Calabash Rouge absorbed less Al from soils than CNPH 0082, which suggests that the genotype may possess mechanisms for Al tolerance.

Molecular Basis of Aluminium Toxicity in Plants: A Review

Aluminium toxicity in acid soils having pH below 5.5, affects the production of staple food crops, vegetables and cash crops worldwide. About 50% of the world’s potentially arable lands are acidic. It is trivalent cationic form i.e. Al3+ that limits the plant’s growth. Absorbed Aluminium inhibits root elongation and adversely affects plant growth. Recently researches have been conducted to understand the mechanism of Aluminium toxicity and resistance which is important for stable food production in future. Aluminium resistance depends on the ability of the plant to tolerate Aluminium in symplast or to exclude it to soil. Physiological and molecular basis of Aluminium toxicity and resistance mechanism are important to understand for developing genetically engineered plants for Al toxicity resistance. This paper provides an overview of the state of art in this field.

Aluminum stress and its role in the phospholipid signaling pathway in plants and possible biotechnological applications

An early response of plants to environmental signals or abiotic stress suggests that the phospholipid signaling pathway plays a pivotal role in these mechanisms. The phospholipid signaling cascade is one of the main systems of cellular transduction and is related to other signal transduction mechanisms. These other mechanisms include the generation of second messengers and their interactions with various proteins, such as ion channels. This phospholipid signaling cascade is activated by changes in the environment, such as phosphate starvation, water, metals, saline stres, and plant-pathogen interactions. One important factor that impacts agricultural crops is metal-induced stress. Because aluminum has been considered to be a major toxic factor for agriculture conducted in acidic soils, many researchers have focused on understanding the mechanisms of aluminum toxicity in plants. We have contributed the last fifteen years in this field by studying the effects of aluminum on phospholipid signaling in coffee, one of the Mexico's primary crops. We have focused our research on aluminum toxicity mechanisms in Coffea arabica suspension cells as a model for developing future contributions to the biotechnological transformation of coffee crops such that they can be made resistant to aluminum toxicity. We conclude that aluminum is able to not only generate a signal cascade in plants but also modulate other signal cascades generated by other types of stress in plants. The aim of this review is to discuss possible involvement of the phospholipid signaling pathway in the aluminum toxicity response of plant cells.

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.

Aluminum triggers broad changes in microRNA expression in rice roots

MicroRNAs are small 21-nucleotide RNA molecules with regulatory roles in development and in response to stress. Expression of some plant miRNAs has been specifically associated with responses to abiotic stresses caused by cold, light, iron, and copper ions. In acid soils, aluminum solubility increases, thereby causing severe damage to plants. Although physiological aspects of aluminum toxicity in plants have been well characterized, the molecular mediators are not fully elucidated. There have been no reports about miRNA responses to aluminum stress. Modulation of miRNA expression may constitute a key element to explain the mechanisms implicated in aluminum toxicity and tolerance. We examined the expression of at least one miRNA member from each miRNA family in rice roots of Oryza sativa spp indica cv. Embrapa Taim and Oryza sativa spp japonica cv. Nipponbare under high concentrations of aluminum. Forty-six miRNA families were effectively detected by quantitative PCR. Among these, 13 were down-regulated and six were up-regulated in roots of the Nipponbare cultivar after 8 h of aluminum treatment. In roots of the Embrapa Taim cultivar, five miRNAs were down-regulated and three were up-regulated. Analyses of their putative targets suggest that these rice miRNAs are involved in the regulation of various metabolic pathways in response to high concentrations of aluminum.

Molecular Physiology of Aluminum Toxicity and Tolerance in Plants

ABSTRACT Aluminum being the third most abundant metal in the earth's crust poses a serious threat to crop productivity in acid soils, which comprise almost half of the arable land. This review travels across time and updates research done on aluminum stress in plants. In its phytotoxic forms, aluminum affects root growth by acting in the root apical zone, resulting in growth inhibition in a very short time at micromolar concentrations. The mechanisms of aluminum toxicity in plants may proceed by growth inhibition, callose accumulation, cytoskeletal distortion, disturbance of plasma membrane surface charge, and H+-ATPase activity, lipid peroxidation of membranes, production of reactive oxygen species in cytosol and mitochondria, respiratory dysfunction, opening of mitochondrial permeability transition pores, collapsing of inner mitochondrial membrane potential, activation of mitochondrial protease, and induction of nuclear apoptosis, resulting ultimately in programmed cell death. In contrast, the mechanism...

Aluminium toxicity in plants: internalization of aluminium into cells of the transition zone in Arabidopsis root apices related to changes in plasma membrane potential, endosomal behaviour, and nitric oxide production

The extent of aluminium internalization during the recovery from aluminium stress in living roots of Arabidopsis thaliana was studied by non-invasive in vivo microscopy in real time. Aluminium exposure caused rapid depolarization of the plasma membrane. The extent of depolarization depends on the developmental state of the root cells; it was much more extensive in cells of the distal than in the proximal portion of the transition zone. Also full recovery of the membrane potential after removal of external aluminium was slower in cells of the distal transition zone than of its proximal part. Using morin, a vital marker dye for aluminium, and FM4-64, a marker for endosomal/vacuolar membranes, an extensive aluminium internalization was recorded during the recovery phase into endosomal/vacuolar compartments in the most aluminium-sensitive cells. Interestingly, aluminium interfered with FM4-64 internalization and inhibited the formation of brefeldin A-induced compartments in these cells. By contrast, there was no detectable uptake of aluminium into cells of the proximal part of the transition zone and the whole elongation region. Moreover, cells of the distal portion of the transition zone emitted large amounts of nitric oxide (NO) and this was blocked by aluminium treatment. These data suggest that aluminium internalization is related to the most sensitive status of the distal portion of the transition zone towards aluminium. Aluminium in these root cells has impact on endosomes and NO production.

Aluminium toxicity in plants: a review

L'article fait le point sur la toxicite de l'aluminium, la tolerance des plantes a cet element et leurs performances ecologiques. La tolerance au stress provoque par l'aluminium varie d'un genotype de plante a l'autre et cette approche est prometteuse pour ameliorer notre comprehension de la tolerance a l'aluminium. L'induction de la tolerance a l'aluminium et sa caracterisation sont egalement passees en revue. Les effets cytogenetiques de l'aluminium sur les plantes font l'objet d'une discussion approfondie. On compare la sensibilite relative a l'aluminium des diverses especes de plantes, de la macroflore a la microflore. On traite de l'absorption et du transport de l'aluminium, ainsi que de sa phytotoxicite et de ses interactions avec les nutriments. La connaissance actuelle concernant la physiologie et la biochimie de l'aluminium en relation avec la phytotoxicite est discutee et offre certaines possibilites pour accroitre la tolerance a l'aluminium. Cette mise au point montre la complexite des mecanismes lies a la toxicite des elements traces.

Aluminum-induced 1-->3-beta-D-glucan inhibits cell-to-cell trafficking of molecules through plasmodesmata. A new mechanism of aluminum toxicity in plants.

Symplastic intercellular transport in plants is achieved by plasmodesmata (PD). These cytoplasmic channels are well known to interconnect plant cells to facilitate intercellular movement of water, nutrients, and signaling molecules including hormones. However, it is not known whether Al may affect this cell-to-cell transport process, which is a critical feature for roots as organs of nutrient/water uptake. We have microinjected the dye lucifer yellow carbohydrazide into peripheral root cells of an Al-sensitive wheat (Triticum aestivum cv Scout 66) either before or after Al treatment and followed the cell-to-cell dye-coupling through PD. Here we show that the Al-induced root growth inhibition is closely associated with the Al-induced blockage of cell-to-cell dye coupling. Immunofluorescence combined with immuno-electron microscopic techniques using monoclonal antibodies against 1-->3-beta-D-glucan (callose) revealed circumstantial evidence that Al-induced callose deposition at PD may responsible for this blockage of symplastic transport. Use of 2-deoxy-D-glucose, a callose synthesis inhibitor, allowed us to demonstrate that a reduction in callose particles correlated well with the improved dye-coupling and reduced root growth inhibition. While assessing the tissue specificity of this Al effect, comparable responses were obtained from the dye-coupling pattern in tobacco (Nicotiana tabacum) mesophyll cells. Analyses of the Al-induced expression of PD-associated proteins, such as calreticulin and unconventional myosin VIII, showed enhanced fluorescence and co-localizations with callose deposits. These results suggest that Al-signal mediated localized alterations to calcium homeostasis may drive callose formation and PD closure. Our data demonstrate that extracellular Al-induced callose deposition at PD could effectively block symplastic transport and communication in higher plants.

Windrow Composting of a Pulp and Paper Mill Sludge: Process Performance and Assessment of Product Quality

Pulp and paper mill sludges (PMS) are a significant by-product of the paper making industry worldwide, and composting with mineral nutrients in Tasmania is viewed as the most environmentally acceptable technology to convert this material into a useful horticultural or agricultural product, thereby eliminating the need for landfilling. The objective of this study was to determine the feasibility of composting PMS in large-scale periodically turned windrows based on optimal conditions previously determined in laboratory-scale reactors. Performance of the composting process and quality of the composted PMS is reported. PMS was composted with mineral nutrients at an initial C:N:P:K ratio of 35:1:0.6:0.1 in three windrows of 25 m in length, 3 m in width and 2 m in height with weekly turning for 21 weeks, following which the trial was terminated. Temperature at the pile center remained above 50°C for the duration of the trial, with pile pH peaking at 7.46 after 11 weeks. At termination, phytotoxicity was absent and pile volume had decreased by 45 percent, partly due to a 31.6 percent increase in bulk density and a decrease in gravimetric water content from 71.4 percent before to 63.7 percent after composting. Due to the moderate level of aluminum in the PMS compost, substitution of the aluminum sulfate flocculant used in primary clarification and sludge dewatering with a nonaluminum based flocculant may be required to reduce the potential for aluminum toxicity in plants. The C:N ratio was reduced from 218:1 prior to mineral nutrient addition to 23:1 after composting with a concomitant rise in electrical conductivity from 0.59 dSm−1 to 2.78 dSm−1, possibly making the material unsuitable for direct contact with plants sensitive to moderately saline conditions. The electrical conductivity could be reduced to less than 2 dSm−1 by substituting some of the nitrogen and potassium amendments with nonsalinizing nutrient forms, thereby substantially improving the quality of the composted PMS.

Some aspects of aluminum toxicity in plants

Aluminum toxicity is a major factor in limiting growth in plants in most strongly acid soils. Toxic effects on plant growth have been attributed to several physiological and biochemical pathways, although the precise mechanism is still not fully understood. In general, root elongation is hampered through reduced mitotic activity induced by Al, with subsequent increase in susceptibility to drought. The initial site of uptake is usually the root cap and the mucilaginous secretion covering the epidermal cells. Al ions bind very specifically to the mucilage by exchange adsorption on the polyuronic acid, complexing with the pectic substances and by the formation of polyhydroxy forms, increasing the number of Al atoms per positive charge. Toxicity has been suggested to be initiated at the sites of mucopolysaccharide synthesis. Al is absorbed on all Ca-binding sites on the cell surface. In the intact tissues, most of the Al is bound to the pectic substances of the cell wall and a part to the nucleic acids and cell membrane. Al is also reported to enter the plant by moving into meristematic cells via the cortex, bypassing the endodermal barrier. Being a polyvalent cation, it follows principally the apoplasmic pathway of transport through cortical cells, but may also enter the stele through the plasmalemma. Ultrastructural studies have shown the maximum accumulation to be in the epidermal and cortical cells. The interaction of Al with different systems follows different pathways. The plasma membrane at the outer boundary of the root cell is a potential target and its physical properties can be altered by Al through interaction with membrane-bound ATPase, lipids, carbohydrates and proteins. The Golgi apparatus has been suggested as the primary site of action, followed by damage to the plasmalemma. Aluminum interferes with the uptake, transport and use of several essential elements, including Cu, Zn, Ca, Mg, Mn, K, P and Fe. Excess of Al reduces the uptake of certain elements and increases that of others, the patterns being dependent on the element, the plant part and species involved. A major factor is the pH concentration. At an acid pH, below 5.5, the antagonism between Ca and Al is probably the most important factor affecting Ca uptake by plants. The molecular mechanism of tolerance of Al is as yet not clear. Tolerant plants reduce the absorption by the root or detoxify Al after absorption. Al tolerant plants may be grouped into those with higher Al concentrations in tops and those with less. In the latter, more Al is entrapped in roots. Uptake of Al may be reduced by binding to cell wall or to membrane lipid. Tolerance may be different in different species and seems to be controlled by one or more genes. Absorption of Al in non-metabolic conditions is affected only slightly by temperature. Anaerobic conditions, like the presence of nitrogen and metabolic inhibitors, damage the endodermal membrane barrier, increasing the uptake and enhancing injurious effects. Aluminum also causes morphological damage to plant parts. It affects photosynthesis by lowering chlorophyll content and reducing electron flow. Reduced respiratory activity might be due to reduced metabolic energy requirement. Protein synthesis is decreased probably due to effect on ribosome distribution at endoplasmic reticulum. Aluminum is known to bind to DNA and nuclei. However, its penetrance to DNA of mitotically active centers is slow. On accumulating in roots, it initially inhibits mitotic activity, possibly through affecting the integrated control function of the root meristem. Aluminum toxicity in acid soil is of special importance due to the destruction of components of forest ecosystems under specific conditions. It reduces biomass yield and tree growth and represses litter-degrading microflora. Further information is required on the factors affecting membrane permeability, distribution and accumulation of Al in different plant parts and different species. Al tolerance may be studied with relation to the presence of different ligands, nitrogen metabolism (nitrate reductase and protein accumulation), nitrogen tolerance in relation to pH change and metal ion activities, the role of Ca and P and interference with water relations and litter degradation.

Silvicultural Application of Alum Sludge

Various properties of two types of alum sludge were investigated to determine whether silvicultural application was an appropriate method of disposal. These particular sludges did not produce any undesirable leachates, values of which were within limits set out in the Connecticut Public Health Code, nor did they produce aluminum toxicity in plants—even those grown in pure sludge. Alum sludge binds phosphorus, making it unavailable or slowly available to plants and thus possibly limiting plant growth and even physiologically stressing plants. The results of analyses indicated that limited applications of this material on forested land would most likely not measurably affect tree growth. Land application of alum sludge appears to hold promise as a low‐cost disposal alternative.

Calmodulin Levels Are Not Responsible for Aluminium Tolerance in Wheat

Circumstantial evidence has suggested that the protein calmodulin may be the primary target of aluminium toxicity in plants. Experiments were carried out to test the hypothesis that genetic differences in aluminium tolerance in wheat (Triticum aestivum L.) are the consequence of different levels of calmodulin. The tolerant wheat, selection Waalt, differed from cv. Warigal by a single dominant gene for aluminium tolerance as indicated by analysis of the F2. The tolerance of cv. Carazinho also is reported to be genetically simple. Crosses between the two tolerant wheats, Carazinho and Waalt, showed that they have linked or allelic genes for aluminium tolerance. Calmodulin was extracted from the whole roots and from root tips alone, either after 3 days exposure to aluminium, or without aluminium treatment. Calmodulin levels were measured using a radioimmunoassay. No significant differences were observed between Carazinho (tolerant) and Sunstar (sensitive). Additionally the four genotypes (Waalt, Carazinho, Warigal and Sunstar) showed no consistent differences in sensitivity, as measured by seedling root growth, to three calmodulin antagonists, trifluoperazine, chlorpromazine and calmidizolium. Thus it appears that the different aluminium tolerances in these wheats are not due to different calmodulin levels.

Calmodulin dependent formation of membrane potential in barley root plasma membrane vesicles: A biochemical model of aluminum toxicity in plants

Micromolar concentrations of aluminum ions interfere with calmodulin‐stimulated, membrane bound ATPase activity which plays a role in the maintenance of the transmembrane potential of plasma membrane enriched vesicles isolated from barley roots. Calmodulin appears to be the major target for aluminum interaction resulting in pronounced changes in the exposure of a large, hydrophobic surface on this protein as determined with a fluorescent, hydrophobic surface probe. At a molar ratio of 3:1 [aluminum]/[calmodulin], the calmodulin stimulated enzymatic activity, probably associated with a Ca2++ Mg2+ATPase, is about 95% inhibited. Aluminum induced changes in calmodulin structure are reflected in reduced formation of the membrane potential when assayed with a fluorescent potential probe, oxonol VI. We hypothesize that the aluminum caimodulin complex represents a primary lesion in toxic responses of plants to this metal.

Aluminum toxicity in plants grown in solution culture

Abstract Earlirose rice (Oryza sativa L. ) and Hawkeye soybeans (Glycine max L.) were grown in solution culture with A12(SO4)3 in concentrations of 0, 10‐6, 10‐5, 10‐4, 10‐3 M. Only at 10‐4 (slightly) and at 10‐3 M were there yield depressions due to Al. The threshold concentration of Al for toxicity was about 20 μg/g in rice shoots and about 30 μg/g in soybean leaves. The solution level necessary for these concentrations was 8 μg Al/ml. Plant concentrations which caused severe toxicity were 70 μg Al/g plant with 81 μg Al/ml solution. Most Al remained in roots, but leaves contained more than did stems of soybeans. The high Al decreased Fe, Cu, and Mn concentrations in shoots of rice and decreased Fe, Cu, and Zn in roots of rice. The high Al resulted in decreased Fe and Zn in leaves of soybeans. No Fe deficiency symptoms were present due to the high Al.