Citrate Secretion Induced by Aluminum Stress May Not be a Key Mechanism Responsible for Differential Aluminum Tolerance of Some Soybean Genotypes

Abstract

Eighteen soybean genotypes differing in aluminum (Al) tolerance were used to investigate genotypic differences in Al-induced citrate exudation and its role in Al tolerance. Aluminum accumulation and localization in soybean roots were examined by analysis of total Al and hematoxylin staining. Soybean genotypes exhibited a wide range of Al tolerance. Based on relative root elongation, several Al-tolerant genotypes from Brazil such as B1, B10, and B15 were more tolerant than the Al-tolerant PI 416937 (PI) and Perry. All soybeans exuded citrate in response to Al stress, and some Al-sensitive genotypes secreted more citrate than tolerant ones, showing no correlation between the Al tolerance and Al-induced citrate exudation. Further study found that both copper (Cu) and cadmium (Cd) stimulated citrate and malate exudation in soybean, indicating that organic acid secretion is not specifically induced by Al. Aluminum concentrations were significantly higher in 2–3 and 3–4 cm of segments than that in 0–1 and 1–2 cm segments under 15 μM AlCl₃. Both the root mature zone and apex were heavily stained by hematoxylin after exposure to 10, 15, or 20 μM AlCl₃ (24 h), whereas root elongation zone was not stained. After exposure to 50 μM AlCl₃ for 20 min, the Al-tolerant PI was less stained by hematoxylin than the Al-sensitive Young, suggesting that Al accumulation in root apices seem to be an immediate response to Al stress, and related to differential Al sensitivity. Present results suggest that citrate secretion induced by Al stress may not be a key mechanism responsible for the differential Al tolerance of some soybean genotypes and other mechanism(s) conferring Al exclusion should exist and operate immediately after exposure to Al stress.     

铝胁迫下大豆根系脱落酸累积与柠檬酸分泌的关系研究

为探讨铝(Al)胁迫条件下脱落酸(ABA)调控植物根系有机酸分泌的机制,进行了ABA与Al诱导大豆根系柠檬酸分泌的关系试验。结果表明:1)外源ABA和ABA合成抑制剂fluridone分别提高和降低了Al诱导的大豆根尖ABA含量的增加,但对根系柠檬酸分泌量均无影响,ABA对根系内源柠檬酸含量和柠檬酸合成酶的活性也没有影响;2)分根试验表明,与Al直接接触的根部(Part A)内源ABA含量发生变化,且有柠檬酸的分泌,而不与Al直接接触的根部(Part B)内源ABA含量也发生变化,但没有柠檬酸分泌;3)Al胁迫下,大豆耐Al基因型柠檬酸分泌量远高于敏感基因型,但二者的内源ABA含量却没有差异;4)30μmol AlCl3处理,在0~12 h柠檬酸分泌速率和内源ABA含量随Al处理时间增加而增加,去除Al胁迫时(12~18 h),柠檬酸分泌速率继续增加,但内源ABA含量则迅速下降。综合以上结果,推测ABA不是通过提高Al诱导柠檬酸分泌来调控大豆耐Al性。     

铝和镉胁迫对两个大麦品种矿质营养和根系分泌物的影响

A hydroponic experiment was carried out to study the effect of aluminum （Al） and cadmium （Cd） on Al and mineral nutrient contents in plants and Al-induced organic acid exudation in two barley varieties with different Al tolerance. Al- sensitive cv. Shang 70-119 had significantly higher Al content and accumulation in plants than Al-tolerant cv. Gebeina, especially in roots, when subjected to low pH （4.0） and Al treatments （100 μmol L^-1 Al and 100 μmol L^-1 Al ＋1.0 μmol L^-1 Cd）. Cd addition increased Al content in plants exposed to Al stress. Both low pH and Al treatments caused marked reduction in Ca and Mg contents in all plant parts, P and K contents in the shoots and leaves, Fe, Zn and Mo contents in the leaves, Zn and B contents in the shoots, and Mn contents both in the roots and leaves. Moreover, changes in nutrient concentrations were greater in the plants exposed to both Al and Cd than in those exposed only to Al treatment. A dramatic enhancement of malate, citrate, and succinate was found in the plants exposed to 100 μmol L^-1 Al relative to the control, and the Al-tolerant cultivar had a considerable higher exudation of these organic acids than the Al-sensitive one, indicating that Al-induced enhancement of these organic acids is very likely to be associated with Al tolerance.     

Aluminum tolerance of two wheat cultivars (Brevor and Atlas66) in relation to their rhizosphere pH and organic acids exuded from roots

Phytotoxicity of aluminum (Al) has become a serious problem in inhibiting plant growth on acid soils. Under Al stress, the changes of rhizosphere pH, root elongation, absorption of Al by wheat roots, organic acids exuded from roots, and some main factors related to Al-tolerant mechanisms have been studied using hydroponics, fluorescence spectrophotometry, and high performance liquid chromatography (HPLC). Two wheat cultivars, Brevor and Atlas66, differing in Al tolerance are chosen in the study. Accordingly, the rhizosphere pH has a positive effect on Al tolerance. Atlas66 (Al-tolerant) has higher capability to maintain high rhizosphere pH than Brevor (Al-sensitive) does. High pH can reduce Al3+ activity and toxicity, and increase the efficiency of exuding organic acids from the roots. More inhibition of root elongation has been found in Brevor because of the exposure of roots to Al3+ solution at low pH. Brevor accumulate more Al in roots than Atlas66 even at higher pH. Al-induced exudation of malic and citric acids has been found in Atlas66 roots, while no Al-induced organic acids have been found in Brevor. These results indicate that the Al-induced secretion of organic acids from Atlas66 roots has a positive correlation with Al tolerance. Comprehensive treatment of Al3+ and H+ indicates that wheat is adversely influenced by excess Al3+, rather than low pH.     

Aluminum tolerance associated with enhancement of plasma membrane H+-ATPase in the root apex of soybean

Seventeen soybean cultivars were screened to discern differences in aluminum (Al) sensitivity. The Sowon (Al-tolerant) and Poongsan (Al-sensitive) cultivars were selected for further study by simple growth measurement. Aluminum-induced root growth inhibition was significantly higher in the Poongsan cultivar than in the Sowon cultivar, although the differences depended on the Al concentration (0, 25, 50, 75 or 100 μmol L^–1) and the amount of exposure (0, 3, 6, 12 or 24 h). Damage occurred preferentially in the root apex. High-sensitivity growth measurements using India ink implicated the central elongation zone located 2–3 mm from the root apex. The Al content was lower 0–5 mm from the root apices in the Sowon cultivar than in the apices of the Poongsan cultivar when exposed to 50 μmol L^–1 Al for 12 h. Furthermore, the citric acid exudation rate was more than twofold higher in the Sowon cultivar. Protein production of plasma membrane (PM) H⁺-ATPase from the root apices (0–5 mm) was upregulated in the presence of Al for 24 h in both cultivars. This activity, however, decreased in both cultivars treated with Al and the Poongsan cultivar was more severely affected. We propose that Al-induced growth inhibition is correlated with changes in PM H⁺-ATPase activity, which is linked to the exudation of citric acid in the root apex.     

Aluminum tolerance associated with enhancement of plasma membrane H+-ATPase in the root apex of soybean

Abstract

Seventeen soybean cultivars were screened to discern differences in aluminum (Al) sensitivity. The Sowon (Al-tolerant) and Poongsan (Al-sensitive) cultivars were selected for further study by simple growth measurement. Aluminum-induced root growth inhibition was significantly higher in the Poongsan cultivar than in the Sowon cultivar, although the differences depended on the Al concentration (0, 25, 50, 75 or 100?μmol?L^–1) and the amount of exposure (0, 3, 6, 12 or 24?h). Damage occurred preferentially in the root apex. High-sensitivity growth measurements using India ink implicated the central elongation zone located 2–3?mm from the root apex. The Al content was lower 0–5?mm from the root apices in the Sowon cultivar than in the apices of the Poongsan cultivar when exposed to 50?μmol?L^–1 Al for 12?h. Furthermore, the citric acid exudation rate was more than twofold higher in the Sowon cultivar. Protein production of plasma membrane (PM) H⁺-ATPase from the root apices (0–5?mm) was upregulated in the presence of Al for 24?h in both cultivars. This activity, however, decreased in both cultivars treated with Al and the Poongsan cultivar was more severely affected. We propose that Al-induced growth inhibition is correlated with changes in PM H⁺-ATPase activity, which is linked to the exudation of citric acid in the root apex.     

The plasma membrane intactness of root-tip cells is a primary factor for Al-tolerance in cultivars of five species

We investigated the role of the cell wall and plasma membrane (PM) of root-tip cells in Al tolerance in Al-tolerant and Al-sensitive cultivars of five plant species (rice, maize, pea, wheat, and sorghum). No correlation was found between the differences in Al tolerance and the cation exchange capacity of cell walls isolated from root-tips (0–1 em). Preliminary exposure to Al for 1 h was sufficient to inhibit subsequent root re-elongation in an Al-free solution, and the inhibitory effect was more pronounced in the Al-sensitive cultivars than in the Al-tolerant ones. Together with the inhibition of root re-elongation, the PM of the root-tip cells of all the Al-sensitive cultivars was more permeabilized than that of the Al-tolerant cultivars, based on the FDA-PI fluorescence staining technique. Exposure for 30 min to Al treatment at 100 µM significantly increased the PM permeability of protoplasts isolated from the root-tips for the Al-sensitive pea cultivar placed in a moderately hypotonic medium. Protoplasts from root-tip portions of all the Al-sensitive cultivars took up more Al than those of the Al-tolerant ones when treated with 100 pM Al under isotonic conditions for 30 min. The co-existence of DNP or hypotonic conditions led to a larger increase of Al uptake by the protoplasts from Al-sensitive maize cultivars. These results suggest that Al ions rapidly alter the PM of the root-tip portion in the Al-sensitive cultivars, irrespective of plant species, resulting in an increase of the PM permeability.     

小麦的铝毒及耐性

为探明Al的毒性和忍耐机理 ,比较了Scout 66和Atlas 66Al敏感和抗性的 2个小麦品种的根对Al的积累模式、根细胞壁对Al的吸附以及Al诱导的有机酸的分泌。结果表明 ,Al对Scout 6 6根伸长的抑制作用较Atlas 66明显。根系吸收的Al主要积累于 0至 5mm根尖处。Scout 6 6的根尖及Al处理后分离的根尖细胞壁对Al的积累量大于Atlas 6 6。但是 ,Al处理前分离根尖细胞壁 ,Al处理后细胞壁对Al的吸附量两品种间无显著差异。Al可诱导Atlas 6 6的根系分泌苹果酸 ,而Scout 6 6的分泌物中未发现Al诱导的有机酸。这些结果表明 ,Atlas 6 6的根尖及其细胞壁较Scout 66积累较少的Al,这种差异与Al诱导的有机酸分泌有关 ,而与根尖细胞壁固有的吸附Al的能力无关     

The significance of organic‐anion exudation for the aluminum resistance of primary triticale derived from wheat and rye parents differing in aluminum resistance

Eight primary octoploid triticale genotypes (xTriticosecale Wittmack) derived from four wheat cultivars (Triticum aestivum L.) and two rye inbred lines (Secale cereale L.) differing in aluminum (Al) resistance were investigated with respect to their response to Al supply. Aluminum‐induced inhibition of root elongation (48 h, 25 µM Al supply), callose formation, and the accumulation of Al in root tips (4 h, 25 µM Al supply) were used as parameters to assess Al resistance. Using these parameters, the existing information on Al resistance of the wheat and rye cultivars was generally confirmed. The triticale cultivars showed a wide range of Al resistance amongst the Al‐sensitive wheat and the Al‐resistant rye cultivars. The rye parents and the Al‐resistant wheat parent Carazinho were characterized by Al‐induced exudation particularly of citrate but also of malate from whole root systems of 14 d old seedlings (8 h, 50 µM Al supply). Regression analysis revealed that the degree of Al resistance of the triticale genotypes was closely related to the Al‐induced citrate exudation, which was mainly controlled by the Al resistance of the wheat parent.     

Abstracts of nippon Dozyohiryogaku Zasshi

Aluminum toxicity and boron deficiency are the major factors that limit plant growth and development in acid soils and in B-deficient soils. Root growth inhibition is an early symptom of AI toxicity and B deficiency. Effects of AI and B supply and their interaction on the growth of wheat (Triticum aestivum L.) seedlings were investigated using hydroponics. Fifteen wheat cultivars commonly grown in Bangladesh were used and found to differ considerably in their tolerance to AI toxicity and B deficiency. The relative root length of all the wheat cultivars at 50 µM AI (pH 4.5) ranged from 27 to 71% relative to the control (0 µM AI). Among the cultivars, Inia66 and Kalyansona were found to be the most Al-tolerant and sensitive cultivars, respectively, based on the data of relative root length, malate exudation and AI content of roots. Malate was detected in all the cultivars in the presence of 100 µM AI (pH 4.3). Inia66 exuded a 6-fold larger amount of malate and the AI content of roots was 4 times lower than that in Kalyansona. The vigorous seedling growth was observed at 40 µM B among the series of B treatments. Considerable cultivar differences in response to 40 µM B were observed among the 15 cultivars. Kalyansona was considered to be the most sensitive and Kheri the most tolerant to B deficiency. The interaction effects of B ( 40 and 200 µM) and AI (50 µM) on seedling growth were also examined in Inia66 and Kalyansona. Root growth was inhibited in the presence of Al but B supply especially at 200 µM B in the Kalyansona cultivar caused a slight improvement.     

Ammonium Enables Aluminum-Induced Stimulation of Nitrogen Assimilation in Roots of Al-Tolerant Maize Genotypes

One aluminum (Al)-sensitive (B-73) and two Al-tolerant (F-2 and L-2039) maize genotypes were subjected to Al stress (100 μM Al) under two nitrogen (N) treatments [13.2 mM nitrate (NO₃^?) and 8.3 mM NO₃^? + 4.9 mM ammonium (NH₄⁺)]. Growth parameters, chlorophyll, root N and NO₃^? contents, root nicotinamide adenine dinucleotide (NADH-) and nicotinamide adenine dinucleotide phosphate (NADPH)-nitrate reductase, glutamine synthetase, and glutamate dehydrogenase activities were determined. Aluminum significantly decreased growth and chlorophyll content in Al-sensitive genotype. Nitrate accumulation in roots was increased by Al in tolerant plants. In the sensitive genotype, Al suppressed all enzymes in NO₃^? plants, while in NO₃^?/NH₄⁺ plants the suppression was less severe, and NADPH-nitrate reductase was even stimulated. In tolerant NO₃^?plants, glutamate dehydrogenase was stimulated in F-2 and glutamine synthetase suppressed in L-2039 genotype. In tolerant NO₃^?/NH₄⁺- plants, all enzymes were stimulated by Al, which may be attributed to their participation in defense mechanisms.     

铝诱导大豆根系有机酸分泌的能量消耗的定量研究

铝诱导大豆根系有机酸分泌是其解铝毒的一种重要机制,该过程需要消耗能量,然而有关能量消耗的定量研究还未见报道。本文比较了铝胁迫条件下,两个大豆品种根尖有机酸分泌、 腺苷酸、 无机磷、 细胞质pH值等指标的变化。结果表明,铝处理（25 mol/L）明显诱导大豆根系苹果酸和柠檬酸的分泌。与对照相比,铝胁迫条件下中豆32和本地2号的根尖ATP含量分别降低40.1%和13.2%,根系细胞质子跨膜电化学势差分别增加1711.8和570.6 J/mol,然而,根尖无机磷浓度变化差异不大。运用Nernst-Gibbs方程定量计算自由能变化,发现中豆32和本地2号根尖细胞自由能分别消耗16.13 kJ/mol和14.59 kJ/mol, 中豆32分泌单位苹果酸和柠檬酸的能量消耗分别为0.96 kJ/mol和3.15 kJ/mol,本地2号则为2.01 kJ/mol和5.68 kJ/mol。这表明不同耐铝性大豆品种分泌有机酸消耗的能量存在差异,该结果为筛选耐铝作物品种提供了新思路。     

Phosphorus Alleviates Aluminum Toxicity in Al-Sensitive Wheat Seedlings

The role of phosphorus (P) in the amelioration of aluminum (Al) toxicity to plants is still unclear. The aim of this study was to examine the amelioration of Al toxicity by P supply. The study involved growing Al-sensitive wheat seedlings for 13 days in an acidic soil [pH 4.5 in calcium chloride (CaCl₂)] with increasing added rates of P (0, 20, 40, and 80 mg P kg^?1 soil) and Al [0, 50, and 150 mg aluminum chloride (AlCl₃) kg^?1 soil]. The results indicated that the effects of Al toxicity in this soil could be fully alleviated by the application of P at 50 mg AlCl₃ kg^?1. The 150 mg kg^?1 AlCl₃ treatment significantly reduced root growth, but this was partially overcome by the 80 mg kg^?1 P treatment. High P significantly reduced the concentration of Al in the apoplast, root, and shoot. It is possible that an insoluble Al-P complex forms in the soil and this decreases Al bound in apoplast as well as uptake into the roots. High P decreased the translocation of Al from root to shoot. This study also concluded that detoxification of Al³⁺ by P mainly occurs in soil but not within the plant tissue.     

Predicting Initial Tall Fescue Root Growth Response to Calcium/Aluminum Solution Concentrations

A study was conducted to quantify effects of soluble aluminum (Al) and gypsum (CaSO₄) on initial root growth of three varieties of tall fescue (Festuca arundinacea). Experiments were performed in a growth chamber using hydroponic solutions containing Al from 0 to74 µM in combination with CaSO₄ at 0 to10 mM. Seedlings were grown for 7 d, harvested, air dried, scanned, and weighed for treatment comparisons. Significant differences in root length existed between varieties in Al‐only solutions at low Al concentrations. All varieties showed reduced root growth at concentrations greater than 37 µM Al. Increased calcium (Ca²⁺) and sulfate (SO₄ ^2?) at given concentrations of Al resulted in greater root growth. Relative root growth increased approximately 30% to >80% at 37 µM Al as CaSO₄ increased from 2.5 to 10 mM. A simple logistic model adequately described the effects of Al and CaSO₄ on root growth (r² = 0.86, 0.95, and 0.96 for the three varieties).     

Impact of Boron on Biomass Production and Nutrition of Aluminum‐Stressed Apple Rootstocks

Abstract

The aim of the experiment was to examine the effect of boron (B) on biomass production and nutrition of aluminum (Al)‐stressed apple (Malus sp.) rootstocks. The study was carried out under greenhouse conditions on Polish rootstock (P22) and Malling 26 (M.26) planted singly into 1‐L plastic pots filled with perlite and supplied with Hoagland's medium at pH 4.5 without or with Al (100 µM as AlCl₃). Boron was added into the Al‐containing medium at 20, 40, or 60 µM whereas into the medium without Al only at 20 µM as boric acid. The results showed that the presence of Al in the medium reduced biomass production of P22 and M.26 rootstocks by 22% and 41%, respectively. Rates of uptake and translocation of phosphorus (P), magnesium (Mg), and calcium (Ca) to aerial plant parts were decreased for Al‐treated rootstocks. Aluminum‐stressed P22 rootstocks grown in nutrient solution at 40 and 60 µM B had higher dry weight of leaves and roots, and also higher ability to take up P, Mg, and Ca and lower Al than those grown in the presence of Al at 20 µM B in the medium. Rates of absorption and transport of B to aerial plant parts corresponded with B level in the medium. These results suggest that on acid soils with high Al availability, supra‐optimal B concentrations in soil solution (40–60 µM) can prevent/alleviate Al toxicity in apple trees grafted on P22 rootstocks.     

Soybean root growth in relation to ionic composition in magnesium-amended acid subsoils: Implications on root citrate ameliorating aluminum constraints

Abstract

Hydroponic studies with soybean (Glycine max [L.] Merr.) have shown that µmol L^?1 additions of Mg²⁺ were as effective in ameliorating Al rhizotoxicity as additions of Ca²⁺in the mmol L^?1 concentration range. The objectives of this study were to assess the ameliorative effects of Mg on soybean root growth in acidic subsoils and to relate the soil solution ionic compositions to soybean root growth. Roots of soybean cultivar Plant Introduction 416937 extending from a limed surface soil compartment grew for 28 days into a subsurface compartment containing acid subsoils from the Cecil (oxidic and kaolinitic), Creedmoor (montmorillonitic) and Norfolk (kaolinitic) series. The three Mg treatments consisted of native equilibrium soil solution concentrations in each soil (50 or 100 µmol L^?1) and MgCl₂ additions to achieve 150 and 300 µmol L^?1 Mg (Mg150 and Mg300, respectively) in the soil solutions. Root elongations into Mg-treated subsoils were compared with a CaCO₃ treatment limed to achieve a soil pH value of 6. Subsoil root growth responses to the Mg treatments were less than for the lime treatments. Root length relative to the limed treatments for all subsoils (RRL) was poorly related to the activity of the soil solution Al species (Al³⁺ and Al-hydroxyl species) and Mg²⁺. However, the RRL values were more closely related to the parameters associated with soil solution Ca activity, including (Ca²⁺), (Al³⁺)/(Ca²⁺) and (Al³⁺)/([Ca²⁺] + [Mg²⁺]), suggesting that Ca could be a primary factor ameliorating Al and H⁺ rhizotoxicity in these subsoils. Increased tolerance to Al rhizotoxicity of soybean by micromolar Mg additions to hydroponic solutions, inducing citrate secretion from roots to externally complex toxic Al, may be less important in acid subsoils with low native Ca levels.     

Effects of nitrogen source and aluminum on growth of tropical tree seedlings adapted to low pH soils

The effects of N-source and Al on the growth of seedlings of Melastoma malabathricum, Acacia mangium, and Melaleuca cajuputi, which are tropical woody plants and are very tolerant to Al, and barley (Hordeum vulgare), which is a typical Al-sensitive plant, were investigated. The Al and N treatments consisted of the application of either 0 or 0.5 mM Al, and 2 mM NH₄ ⁺ or N0₃ ^-, respectively. Growth of the tropical plants was enhanced by Al and NH₄ application. In all the plant species, the pH of the culture solution decreased and the concentrations of soluble Al and P increased with the + NH₄ treatment, which positively affected the growth of the tropical plant species. Excised roots of M. malabathricum dissolved insoluble Al with NH₄ application and absorbed Al mainly from root tips. Al did not affect the leaf N concentration except in the case of barley. Roots of M. cajuputi exuded a large amount of citrate, which slightly increased by the + Al treatment. In A. mangium, the reactivity of soluble Al to PCV (pyrocatecholviolet) decreased in the culture solution of the + Al + NH₄. treatment and Al concentration of roots in this treatment was very low. Roots of M. malabathricum released H+ along with Al uptake as well as NH₄ ⁺ uptake. It is concluded that Al and NH₄ ⁺ exert beneficial effects on the growth of tropical tree seedlings.     

Differential Responses of Anti-Oxidative Enzymes to Aluminum Stress in Tolerant and Sensitive Soybean Genotypes

Seedlings of two soybean genotypes, BX10 [aluminum (Al)-tolerant] and BD2 (Al-sensitive), were treated with Al to evaluate the relative root growth (RRG), callose content, Al-sensitive zone, lipid peroxidation, and the anti-oxidative enzyme activities by histochemical and biochemical assays. Under Al toxicity, the RRG reduction of BD2 was more significant than that of BX10, while callose content displayed a contrary trend. The 2–5 mm zone of root apex was the main Al-sensitive zone for soybeans. The activities of superoxide dismutase (SOD), peroxidase (POD), catalase (CAT), and ascorbate peroxidase (APX) were found to be higher in the two genotypes under Al toxicity than that of the controls. The SOD, POD, and CAT activities of BX10; however, were significantly lower than those of BD2. These results implied that producing low quantity of ROS may be one aspect of the Al-tolerant mechanism for soybeans, which in turn helps them adapt to Al stress.     

Organic Acid Metabolism in Roots of Various Grapevine (Vitis) Rootstocks Submitted to Iron Deficiency and Bicarbonate Nutrition

Abstract

The effects of Fe limitation and bicarbonate addition to the nutrient medium on the organic acid metabolism were investigated in the root tips of various grapevine genotypes. Cuttings of two limestone‐tolerant and two limestone‐susceptible Vitis genotypes were grown for four weeks in nutrient solutions containing 10 or 0.5 µM Fe. The effect of bicarbonate addition (5 mM) was studied for two of these genotypes. Compared to 10 µM, Fe limitation (0.5 µM) significantly increased citrate concentration in root tips after 2 weeks, and malate concentration after 4 weeks. When Fe limitation and bicarbonate addition were combined, citrate and malate concentrations were significantly increased after 2 weeks. Fe limitation or addition of 5 mM bicarbonate had a larger effect on citrate than on malate concentrations. Addition of 5 mM bicarbonate discriminated more clearly tolerant and susceptible genotypes than Fe limitation. High malate and citrate concentrations in the roots were associated to high PEPC activities. These results confirm that root organic acid metabolism is involved in grapevine response to Fe deficiency stress. If verified on a larger range of genotypes, a procedure using bicarbonate effect on root tip citrate concentration could be proposed to screen limestone‐tolerant Vitis rootstocks.     

Comparison of aluminum tolerance and phosphate absorption between rape (Brassica napus L.) and Tomato (Lycopersicum esculentum Mill.) in relation to organic acid exudation

Aluminum (Al) tolerance and phosphate absorption in rape and tomato were compared under water culture and field conditions. The relative growth rate in the Al treatment compared with -A1 treatment was similar in the two crops under water culture conditions, while under field conditions, the growth rate was 2- to 3-fold higher in rape than in tomato in spite of the higher Al concentration in the soil solution than in the culture solution. The relative amount of phosphate absorbed in the Al treatment compared with - Al was not appreciably different between rape and tomato under water culture conditions, while under field conditions, it was 3- to 6-fold larger in rape than in tomato. The exudation rate of citric acid by roots was much higher in rape than in tomato. The plant growth, root elongation, and amount of phosphate absorbed in rape were inhibited in the 150 µM Al in the culture solution. However, the inhibition was alleviated by the addition of 200 µM citric acid or 500 µM malic acid. The P concentration in the culture solution decreased by the presence of Al as aluminum phosphate. However, addition of citric and malic acids increased the amount of phosphate released from the precipitated aluminum phosphate. In conclusion, one of the mechanisms for the higher Al tolerance and larger phosphate absorption in rape than in tomato under field conditions was ascribed to the higher concentration of exuded citric acid by Al in the rhizosphere. It was suggested that the exudation of citric acid might contribute to the detoxification of Al and to the increase phosphate availability in the rhizosphere in rape.