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Organic acids, such as malate, citrate and oxalate, have been proposed to be involved in many processes operating in the rhizosphere, including nutrient acquisition and metal detoxification, alleviation of anaerobic stress in roots, mineral weathering and pathogen attraction. A full assessment of their role in these processes, however, cannot be determined unless the exact mechanisms of plant organic acid release and the fate of these compounds in the soil are more fully understood. This review therefore includes information on organic acid levels in plants (concentrations, compartmentalisation, spatial aspects, synthesis), plant efflux (passive versus active transport, theoretical versus experimental considerations), soil reactions (soil solution concentrations, sorption) and microbial considerations (mineralization). In summary, the release of organic acids from roots can operate by multiple mechanisms in response to a number of well-defined environmental stresses (e.g., Al, P and Fe stress, anoxia): These responses, however, are highly stress-and plant-species specific. In addition, this review indicates that the sorption of organic acids to the mineral phase and mineralisation by the soil's microbial biomass are critical to determining the effectiveness of organic acids in most rhizosphere processes.

Phosphorus deficiency-induced metabolic changes related to exudation of carboxylic acids and protons were compared in roots of wheat (Triticum aestivum L. cv Haro), tomato (Lycopersicon esculentum L., cv. Moneymaker), chickpea (Cicer arietinum) and white lupin (Lupinus albus L. cv. Amiga), grown in a hydroponic culture system. P deficiency strongly increased the net release of protons from roots of tomato, chickpea and white lupin, but only small effects were observed in wheat. Release of protons coincided with increased exudation of carboxylic acids in roots of chickpea and white lupin, but not in those of tomato and wheat. P deficiency-induced exudation of carboxylic acids in chickpea and white lupin was associated with a larger increase of carboxylic acid concentrations in the roots and lower accumulation of carboxylates in the shoot tissue compared to that in wheat and tomato. -Citric acid was one of the major organic acids accumulated in the roots of all investigated species in response to P deficiency, and this was associated with increased activity and enzyme protein levels of PEP carboxylase, which is required for biosynthesis of citrate. Accumulation of citric acid was most pronounced in the roots of P-deficient white lupin, chickpea and tomato. Increased PEP carboxylase activity in the roots of these plants coincided with decreased activity of aconitase, which is involved in the breakdown of citric acid in the TCA cycle. In the roots of P-deficient wheat plants, however, the activities of both PEP carboxylase and aconitase were enhanced, which was associated with little accumulation of citric acid. The results suggest that P deficiency-induced exudation of carboxylic acids depends on the ability to accumulate carboxylic acids in the root tissue, which in turn is determined by biosynthesis, degradation and partitioning of carboxylic acids or related precursors between roots and shoot. In some plant species such as white lupin, there are indications for a specific transport mechanism (anion channel), involved in root exudation of extraordinary high amounts of citric acid.

Description of the subject. The present study focuses on the description of the allelopathic interactions between wild and crop species that may occur in a given ecosystem. Objectives. The objective is the evaluation of the allo- and autoinhibition activity of root exudates of barley (Hordeum vulgare L. subsp. vulgare) and great brome (Bromus diandrus Roth.) seedlings by water-soluble allelochemicals. Method. The allelopathic activities of five Tunisian barley genotypes (modern varieties and landraces), one Saudi Arabian barley landrace and great brome were assessed using a modified laboratory bioassay named “seedling-after-seedling agar method”. Results. The barley or the great brome reduced, to a greater extent, the root growth compared to the shoot growth of receiver species. The response of the root system architecture of the great brome towards barley root exudates was studied in detail. All the measured root traits were highly sensitive to the presence of barley. In our conditions, the allelopathic activity of barley root exudates had no apparent relationship with the size of the root and a prominent action of genetic determinants in the allelopathic potential between genotypes is proposed. The alloinhibitory activity of barley or great brome root exudates deferred between the receiver species but was always higher than the autoinhibition potential. The autoinhibition in barley proved to depend on whether the genotypes used as donor and receiver are identical or different, suggesting a specific interaction of allelochemicals with the receiver plant. These molecules seem to be the main actors in the allelopathic barley potential as external factors such variations of pH have no evident relevance in the inhibition process. Conclusions. Barley and great brome exude molecules in their surroundings. This affects the growth of the receiver plants, suggesting that these compounds might contribute to the plant community dynamics. Évaluation du potentiel allélopathique des composés hydrosolubles de l’orge (Hordeum vulgare L. subsp. vulgare) et du grand brome (Bromus diandrus Roth.) moyennant un bio-essai modifié Description du sujet. La présente étude se focalise sur la description des interactions allélopathiques entre des espèces sauvages et cultivées qui peuvent survenir dans un écosystème donné. Objectifs. L’objectif est l'évaluation de l’activité d’auto- et d’alloinhibition des exsudats racinaires de l’orge (Hordeum vulgare L. subsp. vulgare) et du grand brome (Bromus diandrus Roth.) via les allélochimiques hydrosolubles. Méthode. Les activités allélopathiques de cinq génotypes tunisiens d’orge (variétés modernes et orges locales), d’une orge locale d’Arabie saoudite et du grand brome ont été évaluées moyennant un bio-essai modifié nommé « seedling-after-seedling agar method ». Résultats. L’orge et le grand brome ont réduit dans une grande mesure la croissance des racines des espèces receveuses comparativement à celle des pousses. La réponse de l’architecture du système racinaire du grand brome à l’égard des exsudats racinaires de l’orge a été étudiée en détail. Il s’est avéré que tous les traits des racines analysés ont été très sensibles en présence de l’orge. Dans nos conditions, l’activité allélopathique des exsudats racinaires de l’orge n’avait aucune relation apparente avec la taille des racines et une action prépondérante des déterminants génétiques dans le potentiel allélopathique entre les génotypes est ainsi proposée. L’activité d’alloinhibition des exsudats racinaires de l’orge ou du grand brome était différente entre les espèces receveuses mais toujours plus élevée par rapport au potentiel d’autoinhibition. L’autoinhibition chez l'orge a été dépendante des génotypes utilisés comme donneurs et receveurs qui étaient identiques ou différents, suggérant ainsi une interaction spécifique des allélochimiques avec la plante receveuse. Ces molécules semblent être les principaux acteurs du potentiel allélopathique chez l’orge, étant donné que les facteurs externes comme les variations du pH n’ont aucune pertinence évidente dans le processus d’inhibition. Conclusions. L’orge et le grand brome relâchent dans leur environnement des molécules qui affectent la croissance des plantes receveuses, suggérant ainsi que ces composés pourraient contribuer à la dynamique des communautés végétales.

The development of clustered tertiary lateral roots (proteoid roots) and the expression of phosphoenolpyruvate carboxylase (PEPC, EC 4.1.1.31) in roots were studied in white lupin (Lupinus albus L.) grown with either 1 mM P (+P-treated) or without P (-P-treated). The +P-treated plants initiated fewer clustered tertiary meristems and the emergence of these meristems was delayed compared with -P-treated plants. Proteoid root zones could be identified 9 d after emergence in both P treatments. Amounts of PEPC mRNA, PEPC specific activity, and enzyme protein were greater in proteoid roots than in normal roots beginning at 10, 12, and 14 d after emergence, respectively. The increases in PEPC mRNA, PEPC enzyme, and PEPC specific activity suggest that this enzyme is in part under transcriptional regulation. Recovery of organic acids from root exudates coincided with the increases in PEPC specific activity. The -P-treated plants exuded 40-, 20-, and 5-fold more citrate, malate, and succinate, respectively, than did +P-treated plants. Data presented support the hypothesis that white lupin has concerted regulation of proteoid root development, transcriptional regulation of PEPC, and biosynthesis of organic acids for exudation in response to P deficiency

Incorporation of cover crops into cropping systems may contribute to a more efficient utilization of soil and fertilizer P by less P-efficient crops through exudation of P-mobilizing compounds by the roots of P-efficient plant species. The main objective of the present work was to test this hypothesis. First a method has been developed which allows the quantification of organic anion exudation from individual cluster roots formed by P-deficient white lupin (Lupinus albus L.). Lupin plants were grown in nutrient solution at 1 µM P and in a low P loess in small rhizotrons. Organic anions exuded from intact plants grown in nutrient solution were collected from individual cluster roots and root tips sealed in small compartments by an anion-exchange resin placed in nylon bags (resin-bags). Succinate was the dominant organic anion exuded followed by citrate and malate. The mean of citrate exudation-rate was 0.06 pmol mm⁻¹ s⁻¹ with exudation highly dependent on the citrate concentration and on the age of the cluster roots. Exudates from cluster roots and root tips grown at the soil surface (rhizotron-grown plants) were collected using overlayered resin-agar (resin mixed with agar). Citrate exudation from cluster roots was 10 times higher than that from root tips. Fractionation of P in the cluster root rhizosphere-soil indicates that white lupin can mobilize P not only from the available and acid-soluble P, but also from the stable residual soil P fractions. In pot experiments with an acid luvisol derived from loess low in available P, growth of wheat was significantly improved when mixed-cropped with white lupin due to improved P uptake. Both in mixed culture and in rotation wheat could benefit from the P mobilization capacity of white lupin, supporting the hypothesis above. Nine tropical leguminous cover crops and maize were grown in a pot experiment using a luvisol from Northern Nigeria low in available P. All plant species derived most of their P from the resin and bicarbonate-extractable inorganic P. Organic P (Po) accumulated particularly in the rhizosphere of all plant species. There was a significant negative correlation between the species-specific rhizosphere acid phosphatase activity and Po accumulation. Growth and P uptake of maize grown in rotation after legumes were enhanced indicating that improved P nutrition was a contributing factor.

Two different types of root response to Fe deficiency (strategies) have been identified in species of the Plant Kingdom. In Strategy I which occurs in all plant species except grasses, a plasma membrane-bound reductase is induced with enhanced net excretion of protons. Often the release of reductants/chelators is also higher. In Strategy II which is confined to grasses, there is an increase in the biosynthesis and secretion of phytosiderophores which form chelates with FeIII. Uptake of FeIII phytosiderophores is mediated by a specific transporter in the plasma membrane of root cells of grasses. From results based mainly on long-term studies under non-axenic conditions this classification into two strategies has been questioned, and the utilization of Fe from microbial siderophores has been considered as an alternative strategy particularly in grasses. Possible reasons for controversial results are discussed in some detail. The numerous effects of microorganisms in non-axenic cultures, and the as yet inadequate characterization of the so-called standard (basic) reductase present major limitations to understanding different mechanisms of Fe acquisition. In comparison with the progress made in identifying the cellular mechanisms of root responses in Strategy I and Strategy II plants, our understanding is poor concerning the processes taking place in the apoplasm of root rhizodermal cells and of the role of low-molecular-weight root exudates and siderophores in Fe acquisition of plants growing in soils of differing Fe availability.

The role of organic acids in the mobilization of plant nutrients from the rhizosphere was assessed in seven contrasting soil types. The results indicated that malate was poor at mobilizing micronutrients from all the test soils, whilst citrate was capable of mobilizing significant quantities. Citrate was also capable of mobilizing P from one soil which possessed a large Ca-P fraction. This mobilization of P was due to both the complexing action of the citrate anion and due to the dissolution properties of the protons released from citric acid upon equilibrium with the soil solution. The reaction of citrate with cations was found to be near instantaneous with significant absorption to the solid phase in some soils at low concentrations. Soil decomposition studies indicated that citrate was rapidly broken down in organic soils but was more resistant to degradation in subsoil horizons. It was concluded that organic acids can be expected to be of little consequence in nutrient mobilization from high pH soils, whilst in acid soils they may be involved both in a more general mechanism for micronutrient uptake or as a potential Al detoxification mechanism.

Roots of taro (Colocasia esculenta [L.] Schott cvs Bun-long and Lehua maoli) exuded increasing concentrations of oxalate with increasing Al stress. This exudation was a specific response to excess Al and not to P deficiency. Addition of oxalate to Al-containing solutions ameliorated the toxic effect of Al

Aluminium (Al) stimulates the efflux of malate from the apices of wheat (Triticum aestivum L.) roots (Delhaize et al. 1993, Plant Physiol. 103, 695-702). The response was five to tenfold higher in Al-tolerant seedlings than Al-sensitive seedlings and the capacity for Al-stimulated malate efflux was found to co-segregate with Al tolerance in a pair of near-isogenic wheat lines differing in Al-tolerance at a single dominant locus. We have investigated this response further using excised root apices. Half-maximal efflux of malate from apices of Al-tolerant seedlings was measured with 30 micromolar Al in 0.2 mM CaCl2, pH 4.2, while saturating rates of 2.0 nmol.apex(-1).h(-1) occurred with concentrations above 100 micrmolar Al. The stimulation of malate efflux by Al was accompanied by an increase in K+ efflux which appeared to account for electroneutrality. The greater stimulation of malate efflux from Al-tolerant apices compared to Al-sensitive apices could not be explained by differences in the activities of phosphoenolpyruvate carboxylase or NAD-malate dehydrogenase. Several other polyvalent cations, including gallium, indium and the tridecamer Al13, failed to elicit malate efflux. Aluminium-stimulated efflux of malate was correlated with the measured concentration of total monomeric Al present, and with the predicted concentrations of Al3 + and AlOH2 + ions in the solution. Several antagonists of anion channels inhibited Al-stimulated efflux of malate with the following order of effectiveness: niflumic acid approximately NPPB > IAA-94 approximately A-9-C > ethacrynic acid. Lanthanum, chlorate, perchlorate, zinc and alpha-cyano-4-hydroxycinnamic acid inhibited malate release by less than 30% at 100 micromolar while 4,4'-diisothiocyanatostilbene-2,2' disulphonate (DIDS) had no effect. These results suggest that the Al3+ cation stimulates malate efflux via anion channels in apical cells of Al-tolerant wheat roots.

Although it is well known that aluminum (Al) resistance in wheat (Triticum aestivum) is multigenic, physiological evidence for multiple mechanisms of Al resistance has not yet been documented. The role of root apical phosphate and malate exudation in Al resistance was investigated in two wheat cultivars (Al-resistant Atlas and Al-sensitive Scout) and two near-isogenic lines (Al-resistant ET3 and Al-sensitive ES3). In Atlas Al resistance is multigenic, whereas in ET3 resistance is conditioned by the single Alt1 locus. Based on root-growth experiments, Atlas was found to be 3-fold more resistant in 20 micromolar Al than ET3. Root-exudation experiments were conducted under sterile conditions; a large malate efflux localized to the root apex was observed only in Atlas and in ET3 and only in the presence of Al (5 and 20 micromolar). Furthermore, the more Al-resistant Atlas exhibited a constitutive phosphate release localized to the root apex. As predicted from the formation constants for the Al-malate and Al-phosphate complexes, the addition of either ligand to the root bathing solution alleviated Al inhibition of root growth in Al-sensitive Scout. These results provide physiological evidence that Al resistance in Atlas is conditioned by at least two genes. In addition to the alt locus that controls Al-induced malate release from the root apex, other genetic loci appear to control constitutive phosphate release from the apex. We suggest that both exudation processes act in concert to enhance Al exclusion and Al resistance in Atlas