Differential responses of alfalfa cultivars to aluminum stress

Abstract

Toxic levels of aluminum can cause severe yield reduction in alfalfa (Medicago sativa L.), especially in the presence of drought stress. Reactions to Al stress of alfalfa cultivars and germplasms, representing a broad genetic base and the entire range of dormancy types, were evaluated in a Monmouth soil study [26.2% Al saturation (pH 4.8) vs 2.8% Al saturation (pH 5.7)] and in two nutrient solution experiments (0 vs 111 μmol Al; pH 4.5). The soil study, Experiment 1, and Experiment 2 were harvested 28, 40, and 25 d after seeding, respectively.

In all studies, entries differed significantly in vigor and yields were reduced significantly by Al stress. In the soil study, only ‘Lahontan’ was not affected significantly by Al stress, although Lahontan, ‘Atlantic’, ‘B13‐A14’ (tolerant check), ‘Ladak 65’, and ‘Mesa‐Slrsa’ had comparable relative weights (dry weight stressed/dry weight unstressed). There were no statistically significant differential responses to Al stress in Experiment 1, however the relative weight of B13‐A13 (tolerant check) was considerably larger that those of the other entries. Many entries were not affected significantly by Al stress in Experiment 2; B13‐A14, ‘Moapa 69’, ‘Saranac’, and ‘Teton’ had the largest relative weights. Relative weights for Experiment 1 and Experiment 2 were significantly correlated (r=0.46?) as was mean dry matter production in the soil study and Experiment 2 (r=0.73??).     

VARIATION IN PHOSPHORUS EFFICIENCY AMONG BRASSICA CULTIVARS I: INTERNAL UTILIZATION AND PHOSPHORUS REMOBILIZATION

Plants have adapted a number of mechanisms to cope with widespread phosphorus (P) deficiency in arable lands. Crop species and even cultivars differ widely in one or more of these adaptive mechanisms hence, in P efficiency. Identification of these mechanisms is pre-requisite for long term breeding programs. Two independent experiments were conducted to study the possible mechanisms of P efficiency in Brassica cultivars. Eight Brassica cultivars (‘B.S.A.’, ‘Toria’, ‘Toria Selection’, ‘Brown Raya’, ‘Peela Raya’, ‘Dunkeld’, ‘Rainbow’, and ‘CON-1’) were selected on the basis of differences in growth under P deficiency from preliminary experiment. In the first experiment, cultivars were grown for 40 days in sand supplied either with sparingly soluble phosphate rock (PR) or soluble mono-ammonium phosphate (MAP). Cultivars differed significantly (P<0.05) for biomass production, P contents and P use efficiency. Low P availability in PR treatment resulted in significantly lower dry weights and P contents than those grown with MAP. The cultivars ‘Rainbow’, ‘Brown Raya’ and ‘Dunkeld’ accumulated more biomass (3.2 g/pot) and P contents (3.0 mg/pot) than other cultivars when grown with PR. Root dry weight was significantly correlated with shoot dry weight, shoot P content and total P content (r > 0.65) indicating significance of improved root growth for P acquisition. While in the second experiment cultivars were grown with adequate P for 30 days and then P was withdrawn from the nutrient solution by replacing fresh P free nutrient solution for 10 days. Induced P deficiency increased P contents in young leaves by two folds indicating remobilization of P from older leaves and shoot. Nonetheless cultivars varied for remobilization but differences in P remobilization could not explain the differences in P utilization efficiency among cultivars. Hence further experimentation to study root morphology, P uptake, and organic acid exudation by these cultivars in relation to P deficiency is recommended.     

Aluminum toxicity in tomato. Part 1. growth and mineral nutrition

Aluminum (Al) toxicity was studied in two tomato cultivars (Lycopersicon esculentum Mill. ‘Mountain Pride’ and Floramerica') grown in diluted nutrient solution (pH 4.0) at 0, 10, 25, and 50 μM Al levels. In the presence of 25 and 50 μM Al, significant reduction was found in leaf area, dry weight, stem length, and longest root length of both cultivars. Growth of ‘Floramerica’ was less sensitive to Al toxicity than growth of ‘Mountain Pride’. Elemental composition of the nutrient solutions were compared immediately after the first Al addition and four days later. The uptake of micronutrients copper (Cu), manganese (Mn), molybdenum (Mo), zinc (Zn), boron (B), and iron (Fe) from the nutrient solution was reduced in both cultivars with increasing Al levels. Nutrient solution Al gradually decreased in time for every treatment; less in cultures of ‘Floramerica’ than in ‘Mountain Pride’. Aluminum treatments decreased the calcium (Ca), potassium (K), magnesium (Mg), Mn, Fe, and Zn content in the roots, stems, and leaves. Aluminum treatment promoted the accumulation of P, Mo, and Cu in the roots, and inhibited the transport of these nutrients into stems and leaves. At 25 and 50 μM levels of Al, lower Al content was found in the roots of cv. “Floramerica’ than in the roots of cv. ‘Mountain Pride’.     

Aluminum tolerances of two snapbean cultivars related to organic acid content evaluated by high‐performance liquid chromatography

High‐performance liquid chromatography (HPLC) was used to determine aluminum (Al)‐induced changes in organic acid (OA) concentrations of Al‐tolerant ‘Dade’ and Al‐sensitive ‘Romano’ snapbean cultivars. Two week old ‘Dade’ and ‘Romano’ snapbean were grown in 1/5‐strength Steinberg nutrient solution for 10 days and then subjected to 0, 2, 4, 6, and 8 mg L^‐1 Al treatments at pH 4.5 for an additional 3–15 days. Current studies confirmed earlier findings that the Dade cultivar was significantly more tolerant to Al than the Romano variety. Organic acid analyses were performed on extracts of root and leaf, and on stem exudates. The organic acids were separated on an ion exclusion column using a mobile phase of 0.01 N H₃PO₄. Individual OA were quantified with a variable wavelength detector operating at 210 nm. Aluminum stress tended to reduce the concentrations of citric, malonlc, malic, glycolic, fumaric, and acetic acids in the roots and increased the OA concentrations in stem exudates. In the presence or absence of Al stress, the Al tolerant Dade cultivar contained higher OA concentrations than did the Al‐sensitlve Romano. Aluminum stress reduced total OA levels in root extracts from Al‐sensltive Romano plants to a greater extent than in those of the Al‐tolerant Dade. Malic and citric acid concentrations were decreased more than those of the other organic acids examined. Results indicate that the Al‐tolerant Dade snapbean cultivar has a higher potential for Al‐chelation and detoxification than does the Al‐sensitive Romano. Hence, an Al‐chelation mechansism may be involved in differential Al tolerance within this species.     

Differential tolerances of oat cultivars to aluminum in nutrient solutions and in acid soils of Poland

Aluminum tolerant oat cultivars are needed for use on acid soil sites where neutralization of soil acidity by liming is not economically feasible. Oat germplasm in Poland has not been examined for range of Al tolerance. Eleven Polish oat cultivars were screened for Al tolerance in nutrient solutions containing 0, 5 and 15 mg L^‐1 Al. Three of these cultivars showing high to moderate tolerance to Al in nutrient solutions were also grown in greenhouse pots of soil and in field plots of soil over a pH range of 3.8 to 5.5 as determined in 1 N KC1.

The eleven oat cultivars differed significantly in tolerance to Al in nutrient solutions. Based on relative root yield (15 mg L^‐1 Al/no A1%), the cultivars ‘Solidor’ and ‘Diadem’ were most tolerant and ‘Pegaz’ and ‘B‐20’ were least tolerant. For these three cultivars, the order of tolerance to acid soil agreed with the order of tolerance to Al in nutrient solution ‐ namely, Solidor > Diadem > Leanda. Hence, for these cultivars, the nutrient solution methods used appear adequate for selecting plants that are more tolerant to Al in strongly acid soils. Additional study is needed to assess the value of this method for screening a broad range of germplasm.

Superior tolerance of the Solidor cultivar to acid soil was associated with significantly higher concentrations of N in the grain. Hence, results suggest that selecting for acid soil or Al tolerance may increase N efficiency in oats.     

Differential responses of red clover germplasms to aluminum stress

Toxic levels of aluminum can cause severe yield reduction in red clover (Trifolium pratense L.), especially in the presence of drought stress. Aluminum tolerances of 17 red clover cultivars and germplasms representing a broad genetic base were evaluated in a Monmouth soil [26.2% Al saturation (pH 4.8) vs. 2.8% Al saturation (pH 5.7)] and in nutrient solutions (0 vs 111 μM Al; pH 4.5). The soil and nutrient culture studies were harvested 29 and 27 d after seeding, respectively.

Aluminum stress reduced shoot and root growth significantly in soil but not in nutrient culture. Entries differed significantly in shoot vigor in both media and in root vigor in nutrient culture; responses to the two media were positively correlated. Relative weights (dry weight stressed/dry weight unstressed) in soil and nutrient culture were not correlated.

In soil, Al stress significantly reduced shoot growth of all entries except ‘Tristan’, whereas root growth was not affected significantly in ‘Atlas’, ‘Lakeland’, ‘Persist’, ‘Reddy’, ‘Redman’, or Tristan. Reddy, ‘Redland II’, Redman, and Tristan had the highest relative shoot and root weights whereas ‘Kenstar’ had the lowest. In nutrient culture, only the shoot growth of Atlas, Lakeland, Redman and ‘YKYC’ and the root growth of Redman were significantly reduced under Al stress. Atlas, ‘Kenland’, and Redman had among the lowest relative shoot and root weights and Kenstar among the highest. Some entries exhibited a positive growth response to Al.     

Screening wheat and sorghum cultivars for aluminum sensitivity at low aluminum levels

Screening cultivars for aluminum (Al) tolerance is often conducted in acid soils or in complete nutrient solutions. The former method lacks precise measurements of Al, and the second requires high Al concentrations because of precipitation and chelation of the Al and is less representative of the actual environmental stresses to which plants must adapt. These experiments were designed to determine Al tolerance of wheat (Triticum aestivum L. em Thell) and sorghum (Sorghum bicolor L. Moench) using incomplete solutions with very low Al concentrations. Six wheat and five sorghum cultivars were screened for Al tolerance in solution culture with 0 to 10 μM Al and only Ca, K, Mg, NO₃, and Cl in the solutions. Plants were subjected to the solutions for 4 d, and the change in relative root length was measured. Solution Al levels and pH were measured after the termination of the experiments. ‘Atlas’ 66 and ‘Stacy’ were the most tolerant wheat cultivars ('Atlas 66’ = ‘Stacy’ ≥ ‘Monon’ ≥ ‘Scout 66’ ≥ ‘Arthur 71’ = ‘Oasis'). The wheat cultivars were effectively separated on a genetic response basis at 2 μM Al. Sorghum cultivars were uniform in their Al tolerance, but did show some separation at 1 μM Al (SC56 > Tx430 > ‘Funk GS22DR’ > SC283 = SC599). The pH and Al variations did not account for any of the differences observed, indicating that root length differences were caused by genetic control of response to high Al.     

Organic acids in root exudates from Picea abies seedlings influenced by mycorrhiza and aluminum

Organic acid anions exuded from roots of Picea abies (Norway spruce) seedlings grown on glass beads in the presence and absence of mycorrhiza (Laccaria bicolor) and aluminum (Al) at pH 3.9 were measured. We wanted to test if the roots exuded more organic acid anions when exposed to Al and if mycorrhization influenced the exudation. Oxalate was exuded in far higher amounts than any other organic acid anion, with a maximum rate of 1.7 nmol (mg root DW)^–1 d^–1. Mycorrhizal roots exuded significantly more oxalate than nonmycorrhizal roots. The presence of Al did not enhance oxalate exudation. We conclude that the oxalate exuded constitutively by Picea abies/Laccaria bicolor may lead to rhizosphere oxalate concentrations that are relevant for Al resistance.     

Responses of Kentucky bluegrass cultivars to excess aluminum in nutrient solutions

Kentucky bluegrass, Poa pratensis L., is generally regarded as an acid‐soil‐sensitive species. However, previous studies in our laboratory showed that cultivars within the species differed widely in tolerance to acid Tatum subsoil (pH 4.6) which is used routinely to screen plants for aluminum (Al) tolerance. In the early studies, specific differential Al tolerance was not demonstrated. The objective of the current study was to test the hypothesis of differential Al tolerance more precisely in nutrient solutions. In one experiment, acid‐soil‐tolerant Victa and Fylking and acid‐soil‐sensitive Windsor and Kenblue cultivars were grown for 35 days in nutrient solutions containing 0, 2, 4, 6, 12, and 24 mg Al L^‐1, at initial pH 4.5, with no subsequent adjustment. In a second experiment, Victa and Windsor were grown for 30 days in solutions containing 0, 4, and 6 mg Al L^‐1, at initial pH 4.5, with no further adjustment. For Victa and Windsor, tolerance to Al in nutrient solution corresponded with tolerance to acid Tatum subsoil, however, the cultivar difference in tolerance, based on relative root dry weight, was only about 2‐fold, compared with 20‐fold in acid Tatum subsoil. Fylking and Kenblue cultivars, which showed a wide difference in tolerance to acid Tatum subsoil, did not show distinct differences in tolerance to Al in nutrient solutions. Possible reasons for this discrepancy are discussed. Superior Al tolerance of Victa (compared with Windsor) was associated with a greater plant‐induced increase in the pH of its nutrient solutions and a corresponding decrease in concentrations of soluble Al in the filtered solutions at the end of the experiments. Greater Al sensitivity in Windsor (compared with Victa) was not related to reduced uptake of phosphorus (P) or excessive uptake of Al; neither cultivar accumulated appreciable Al concentrations in its shoots. The observed differential acid soil and Al tolerance among bluegrass cultivars appears worthy of further study. Improved understanding of Al tolerance mechanisms would contribute to fundamental knowledge of plant mineral nutrition and could aid plant breeders in tailoring plants for greater tolerance to acid subsoils.     

Screening Kentucky bluegrass for aluminum tolerance

Aluminum (Al) toxicity is a growth‐limiting factor in acid soils for many turfgrasses. The genetic diversity among turfgrass cultivars for Al tolerance is not well known. One hundred‐fifty Kentucky bluegrass (Poa pratensis L.) genotypes (cultivars, selections, and breeding lines) belonging to seven ecotypes were selected to screen for Al tolerance under greenhouse conditions using solution culture, sand culture, and an acid Tatum subsoil (Clayey, mixed, thermic, typic, Hapludult). This soil had 69% exchangeable Al and a pH of 4.4. An Al concentration of 320 μM and a pH of 4.0 in a modified 1/4 strength Hoagland nutrient solution was used in solution screening and sand screening. The grasses were seeded and grown four to five weeks before harvesting. Differences were identified among cultivars and the seven ecotypes by measuring relative growth. ‘Battan’, ‘Viva’, and ‘Nassau’ were the most Al‐tolerant cultivars based on the rank average of the three screening methods. Among the seven ecotypes, BVMG, which refers to cultivars such as ‘Baron’, ‘Victa’, ‘Merit’, and ‘Gnome’, were most Al tolerant while Midwest ecotypes, which are frequently referred to as common Kentucky bluegrasses, consistently exhibited the least Al tolerance. The results indicate that the Kentucky bluegrass cultivars vary genetically in Al tolerance and that there is potential to improve such tolerance with breeding and to refine cultivar‐specific management recommendations regarding soil pH.     

Tolerances of wheat germplasm to acid subsoil

Aluminum toxicity is a major growth limiting factor for plants in many acid soils of the world. Correcting the problem by conventional liming is not always economically feasible, particularly in subsoils. Aluminum tolerant plants provide an alternative and long‐term supplemental solution to the problem. The genetic approach requires the identification of Al tolerance sources that can be transferred to cultivars already having desirable traits. Thirty‐five cultivars and experimental lines of wheat (Triticum aestivum L. em. Thell) were screened for Al tolerance on acid Tatum soil (clayey, mixed thermic, typic Hapludult) receiving either 0 or 3500 mg CaCO₃/kg (pH 4.1 vs. pH 7.1). Entries showed a wide range of tolerance to the acid soil. On unlimed soil at pH 4.3, absolute shoot dry weights differed by 5‐fold, absolute root dry weights by 6.5‐fold, relative shoot weights (wt. at pH 4.3/wt. at pH 7.1 %) by 4.7‐fold and relative root dry weights by 7‐fold. Superior acid soil (Al) tolerance of ‘BH‐1146’ from Brazil and extreme sensitivities of cultivars ‘Redcoat’ (Indiana, USA) and ‘Sonora 63’ (Mexico) were confirmed. Seven experimental (CNT) lines from Brazil showed a range of acid soil tolerance but were generally more tolerant than germplasm from Mexico and the USA. One line, ‘CNT‐1’, was equal to BH‐1146 in tolerance and may be useful in transferring Al tolerance to existing or new cultivars. Five durum cultivars (Triticum, durum, Desf.) were extremely sensitive to the acid Tatum subsoil at pH 4.3 compared with pH 7.1.     

Magnetic resonance imaging of absorbed aluminum in alfalfa roots

An estimated 30% of the world's arable soils are acidic and aluminum (Al) toxicity is often the primary growth‐limiting factor. Excess Al is especially undesirable in sub‐soils because it reduces rooting depth and branching and predisposes plants to drought injury. Liming the plow layer does not generally neutralize subsoil phytotoxicity and Al‐tolerant cultivars offer an alternative or supplemental solution to the problem. Genetic diversity for acid soil tolerance in alfalfa (Medicago sativa L.) is limited and a better understanding of the basic tolerance mechanisms would facilitate the design of more efficacious breeding procedures. Evidence is accumulating that organic acids and proteins elicited by Al stress may complex and detoxify Al either within, or external to, the root. Because Al is a paramagnetic element that can reduce T2 relaxation times (inter‐proton interactions) markedly, the mechanism of Al tolerance in alfalfa was investigated through T2‐based Magnetic Resonance Imaging (MRI) of young lateral root sections of an Al‐sensitive and an Al‐tolerant alfalfa clone grown in nutrient solution (0 or 111 μmol Al; pH 4.5). Root sections that developed under phytotoxic levels of Al accumulated considerable Al in the epidermis and internal root tissue. Aluminum may have been complexed by low molecular weight proteins and organic acids in the tolerant clone whereas the sensitive clone appeared to have abundant free Al; however, variation among replications indicates that free Al may still have been present in tolerant roots and that other tolerance mechanisms may also be important. Root buds accumulated little Al compared to the remainder of the root, indicating that the pronounced effects of Al on lateral root development are indirect. Magnetic Resonance Imaging images evaluated in this study provided clues to the basic mechanisms of Al tolerance in alfalfa and, with further refinement, could be used as one criterion for selecting Al‐tolerant plants.     

Aluminum toxicity in tomato. Part 2. leaf gas exchange,chlorophyll content,and invertase activity

The effect of aluminum (Al) toxicity on leaf gas exchange, leaf chlorophyll content, and sucrose metabolizing enzyme activity of two tomato cultivars (Lycopersicon esculentum Mill. ‘Mountain Pride’ and ‘Floramerica') was studied to determine the mechanism of growth reduction observed in a related study (Simon et al., 1994, Part 1). Plants were grown in diluted nutrient solution (pH 4.0) with 0, 10, 25, or 50 μM. Al for 16 days. Leaf gas exchange was reduced 2–3 fold in both cultivars as Al concentration increased. Gas exchange of ‘Mountain Pride’ was more sensitive to Al toxicity than ‘Floramerica’, agreeing with growth responses observed. Reductions in carbon dioxide (CO₂) assimilation rate appeared to be due to nonstomatal factors in ‘Floramerica’, but stomatal and non‐stomatal limitations in ‘Mountain Pride’. Chlorophyll content of leaves was not affected by Al. Acid invertase (AI) and neutral invertase (NI) activity of roots responded consistently to Al concentration in both cultivars. Root AI and NI activity decreased to a greater extent for ‘Mountain Pride’ than for ‘Floramerica’.     

Role of water stress in differential aluminum tolerance of two barley cultivars grown in an acid soil

Two cultivars of barley (Hordeum vulgare L.), Al‐sensitive ‘Dayton’ and Al‐tolerant ‘Kearney’, were grown under controlled environmental conditions to determine the influence of Al stress and water stress imposed separately and in combination with one another. Plants were grown for 4 weeks in polyethylene‐lined, waxed cartons containing 1 kg of acid, Al‐toxic, Tatum subsoil (clayey, mixed, thermic, Typic Hapludult) at high (pH 4.7) or low (pH 6.6) Al stress. During the final 2 weeks they were also subjected to low (‐20 to ‐40 kPa) or high (‐60 to ‐80 kPa) water stress. Under low water stress, little difference in the growth or appearance of the two cultivars was found, even in the presence of low Al stress (pH 6.6). When high water stress treatment was superimposed on low Al stress treatment, however, significant differences between the two cultivars in biomass production, leaf enlargement, and tillering resulted. When high water stress was combined with high Al stress (pH 4.7), these differences in vegetative growth were further magnified. Thus, drought exacerbates the stress effects of Al toxicity in plants and may account for a significant portion of the reduction in yield commonly observed in acid soils under field conditions and formerly attributed to Al toxicity alone. By increasing soil moisture level, the growth suppressive effect of Al toxicity was significantly reduced.     

Role of water stress in differential aluminum tolerance of six sunflower cultivars grown in an acid soil

Six cultivars of sunflower (Helianthus annuus L.), were screened under controlled environmental conditions for tolerance to Al stress and water stress imposed separately and in combination with one another. Plants were grown for 4 weeks in waxed cartons containing 1 kg of acid, Al‐toxic Tatum, subsoil (clayey, mixed, thermic, Typic Hapludult) at high (pH 4.3) or low (pH 6.3) Al stress. During the final 2 weeks they were also subjected to low (‐20 to ‐40 kPa) or high (‐60 to ‐80 kPa) water stress. Plant growth responses and symptoms of Al toxicity suggested that a wide range of cultivar sensitivity existed. ‘Manchurian’, ‘S‐212’, ‘S‐254’, and ‘S‐265’ were relatively tolerant to Al toxicity while cultlvars ‘Romania HS‐52’ and ‘RM‐52’ were extremely sensitive. Under high Al stress and high water stress, chloroplasts in cells from the Al‐sensitive cultivar ‘Romania HS‐52’ were smaller and contained less starch than chloroplasts from the Al‐tolerant cultivar ‘Manchurian’. Furthermore, the smaller chloroplasts tended to have fewer grana stacks per unit area than did the chloroplasts from tolerant plants. These differences were not apparent when the Al‐sensitive cultivar was grown either in the absence of Al or water stress. In general, Al‐sensitive cultivars of sunflower were more tolerant to water stress than were Al‐tolerant cultivars. Increasing the soil moisture level reduced Al toxicity in Al‐sensitive cultivars. Similarly, decreasing Al stress partially overcame the detrimental effects of high water stress. Hence, Al stress and water stress are interrelated factors which must be considered in the characterization and breeding of plants for better adaptation to acid soils.     

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.     

Manganese Supply and pH Influence Growth,Carboxylate Exudation and Peroxidase Activity of Ryegrass and White Clover

ABSTRACT

The effects of differential manganese (Mn) supply (0 to 355 μ M) and pH (4.8 and 6.0) on dry weight (DW), tissue concentrations of Mn, exudation of carboxylates, and the peroxidase activity were studied in ryegrass (Lolium perenne L.) and white clover (Trifolium repens L.) grown in nutrient solution. In both plant species, the increase in Mn supply caused a significant reduction in DW due to severe Mn toxicity, especially at pH 4.8. The critical toxicity concentration of Mn in shoots was 421 mg kg^{? 1} for ryegrass and 283 mg kg^{? 1} for white clover. For both plant species, an increase in Mn supply levels stimulated the exudation of carboxylates and the activity of peroxidase, which was related to stress conditions. The highest amount of carboxylates was exuded at pH 4.8. There was no clear effect of carboxylates on the complexation of Mn^{2 +}.     

VARIATION IN PHOSPHORUS EFFICIENCY AMONG BRASSICA CULTIVARS II: CHANGES IN ROOT MORPHOLOGY AND CARBOXYLATE EXUDATION

Increased phosphorus (P) efficiency is needed to sustain agriculture productivity on soils with low available P. Significant differences were found among Brassica cultivars for growth, P utilization, and remobilization under P deficiency (see our companion paper, Aziz et al., 2011a). To identify the possible mechanisms of P acquisition from low soluble P compounds, four cultivars (‘Rainbow’, ‘CON-1’, ‘Dunkeld’, and ‘Peela Raya’) were selected to ascertain the relationship of their differential P acquisition and growth with their root length in soil and with organic acid release pattern in solution culture experiments. For this purpose their growth and P acquisition from phosphate rock (PR) was compared with calcium di-hydrogen phosphate (Ca-P) when adding uniform dose of 100 mg P kg^?1 soil separately from the two sources. Biomass accumulation, root length, root fineness, plant P uptake and ash alkalinity was significantly (P < 0.01) different in plants of all the four cultivars when supplied with PR or Ca-P in soil. Minimum biomass produced by ‘Peela Raya’ grown with either P source was followed by ‘CON-1’, ‘Dunkeld’, and ‘Rainbow’ in ascending order. Shoot dry matter production had a significant positive correlation with root dry matter production (r = 0.85, P < 0.01), root length (r = 0.59, P < 0.05) and root P uptake (r = 0.95, P < 0.01). Cultivars varied significantly for organic acid secretion in solution culture experiment. Higher quantities of secreted citric acid, malic acid and tartaric acid in solution culture experiment were measured for ‘Rainbow’ and ‘Dunkeld’ cultivars. Efficient performance of these two cultivars for growth and P uptake was associated with their longer roots and more secretion of organic acids especially citric acid.     

Acid soil tolerances of two wheat cultivars related to soil Ph,KCl‐extractable aluminum and degree of aluminum saturation

Aluminum toxicity, associated with soil acidity, is a major growth‐limiting factor for plants in many parts of the world. More precise criteria are needed for the identification of potential Al toxicity in acid soils. The objective of the current study was to relate the acid soil tolerances of two wheat cultivars to three characteristics of an acid Tatum subsoil (clayey, mixed, thermic, typic Hapludult): pH in a 1:1 soil to water suspension; KCl‐extractable Al; and degree of Al saturation. Aluminum‐tolerant ‘BH 1146’ (Brazil) and Al‐sensitive ‘Sonora 63’ (Mexico) wheat cultivars were grown in greenhouse pots of soil treated with CaCO₃ to establish final soil pH levels of 4.1, 4.6, 4.7, 4.9, 5.2 and 7.3. Soil Al, Ca and Mg were extracted with 1 N KCl, and Al saturation was calculated as KCl‐Al/KCl Al + Ca + Mg%.

Within the soil pH range of 4.1 to 4.9, BH 1146 tops and roots produced significantly more dry matter than did those of Sonora 63; however, at pH 5.2 and 7.3, the top and root yields of the two cultivars were not significantly different. Significant cultivar differences in yield occurred over a range of 36 to 82% saturation of the Tatum soil. Graphs of relative top or root yields against soil pH, KCl‐extractable Al and Al saturation indicated that the two cultivars could be separated for tolerance to Tatum soil under the following conditions: pH less than 5.2 (1:1 soil‐water); KCl‐Al levels greater than 2 c mole kg^‐1 and Al saturations greater than 20%. Results demonstrated that any soil test used to predict Al toxicity in acid soils must take into account the Al tolerances of the plant cultivars involved.