Tolerance of durum wheat lines to an acid,aluminum‐toxic subsoil

Durum wheat, Triticum durum Desf., is reportedly more sensitive to aluminum (Al) toxicity in acid soils than hexaploid wheat, Triticum aestivum L. em. Thell. Aluminum‐tolerant genotypes would permit more widespread use of this species where it is desired, but not grown, because of acid soil constraints. Durum wheat germplasm has not been adequately screened for acid soil (Al) tolerance. Fifteen lines of durum wheat were grown for 28 days in greenhouse pots of acid, Al‐toxic Tatum subsoil at pH 4.5, and non‐toxic soil at pH 6.0. Aluminum‐tolerant Atlas 66 and sensitive Scout 66 hexaploid wheats were also included as standards. Based on relative shoot and root dry weight (wt. at pH 4.5/wt. at pH 6.0 X 100), durum entries differed significantly in tolerance to the acid soil. Relative shoot dry weight alone was an acceptable indicator of acid soil tolerance. Relative dry weights ranged from 55.1 to 15.5% for shoots and from 107 to 15.8% for roots. Durum lines PI 195726 (Ethiopia) and PI 193922 (Brazil) were significantly more tolerant than all other entries, even the Al‐tolerant, hexaploid Atlas 66 standard. Hence, these two lines have potential for direct use on acid soils or as breeding materials for use in developing greater Al tolerance in durum wheat. Unexpectedly, the range of acid soil tolerance available in durum wheat appears comparable to that in the hexaploid species. Hence, additional screening of durum wheat germplasm for acid soil (Al) tolerance appears warranted. Durum lines showing least tolerance to the acid soil included PI 322716 (Mexico), PI 264991 (Greece), PI 478306 (Washington State, USA), and PI 345040 (Yugoslavia). The Al‐sensitive Scout 66 standard was as sensitive as the most sensitive durum lines. Concentrations of Al and phosphorus were significantly higher in shoots of acid soil sensitive than in those of tolerant lines, and these values exceeded those reported to cause Al and phosphorus (P) toxicities in wheat and barley.     

Aluminum,manganese, and iron tolerance improves performance of wheat genotypes in waterlogged acidic soils

Waterlogging results in high shoot concentrations of iron (Fe), aluminum (Al), and manganese (Mn) in wheat grown in acidic soil. The verification of this observation in several acidic soils, development of screening techniques, and identification of genotypes differing in tolerance made it possible to test whether tolerance of ion toxicities improves performance of wheat in waterlogged acid soils. Six wheat varieties selected for tolerance/intolerance of Al, Mn, and Fe were grown in three acidic soils (pH_CaCl2 4.1–4.3) with or without waterlogging for 40 d. In terms of relative shoot dry weight, Al‐, Mn‐, and Fe‐tolerant genotypes tolerated waterlogging better, outperforming intolerant genotypes by 35%, 53%, and 32%, respectively, across the soils. The Al‐tolerant genotype had up to 1.8‐fold better root growth than the intolerant genotype under waterlogging. Waterlogging increased DTPA‐extractable soil Mn (71%) and Fe (89%), and increased shoot Fe (up to 7.6‐fold) and Al (up to 5.9‐fold) for different genotypes and soils. The Al‐tolerant genotype maintained lower tissue concentrations of Al as compared to intolerant genotypes during waterlogging. Waterlogging delayed crop development but distinctly less so in the tolerant than in the intolerant genotypes, thus jeopardizing the capacity of intolerant genotypes to produce yield in Mediterranean climates with dry finish of the season. Pyramiding multiple ion tolerances into current wheat varieties with desirable agronomic and quality characteristics to enhance their performance under waterlogged acid soils should be considered.     

Acid soil tolerance of soybean (Glycine Max L. Merr.) germplasm from the USSR

Nineteen soybean genotypes (ten from the former USSR, two from Brazil and seven from USA) were tested for aluminum (Al) tolerance by growing them for 21 days in greenhouse pots of acid, Al‐toxic, unlimed Tatum (Typic Hapludult) subsoil at pH 4.0 and in limed subsoil at pH 5.1. Aluminum tolerance ranking depended upon the plant traits used in the screening process. Based on absolute dry shoot weights at pH 4.0, Giessener, Brunatna, and St.‐59 (USSR), and Biloxi (USA) were most tolerant; least tolerant entries included Yantarnaya and Smena (USSR), and Davis (USA). Based on relative shoot dry weights (pH 4.0/pH 5.1 %), Giessener, Brunatna, and St.‐59 (USSR) were among the most tolerant, Bossier, Biloxi, Essex, and Perry were intermediate, and Salute 216 (USSR), Chief (USA), and Santa Rosa and IAC‐9 (Brazil) were more sensitive to the acid soil. Based on absolute root dry weights, Giessener, and St.‐59 (USSR), and Biloxi (USA) were among the most tolerant and Smena, Yantarnaya and Salute 216 (USSR), and Chief (USA) were most sensitive. Based on relative root dry weights (pH 4.0/ pH 5.1 %), Giessener was most tolerant and Smena and Salute 216 least tolerant.

Preliminary evidence indicated that soybean entries screened for Al tolerance on acid Tatum soil also differed in tolerance to naturally occurring levels of ambient ozone in greenhouses at Beltsville. The Russian entries VNIIS‐2, Giessener, and Brunatna appeared more sensitive than USA entries Perry, Biloxi, Davis, and Bossier (USA), and Santa Rosa (Brazil). Aluminum tolerance and ozone tolerance appeared to coincide in the Perry genotype. Studies on Al‐ozone‐soybean genotype relationships are being continued at Beltsville.     

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.     

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.     

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.     

Aluminum tolerance of segregating wheat populations in acidic soil and nutrient solutions

Abstract

Increased demand for wheat (Triticum aestivum L.) cultivars tolerant to acid‐soil stress has accelerated genetic research on aluminum (Al) tolerance in soil and solution media. Our objective was to characterize the genetic segregation of tolerant and susceptible plants from two populations in an Al‐toxic Porters soil (coarse‐loamy, mixed, mesic Umbric Dystrochrepts), and in nutrient solutions with 0.09, 0.18, 0.36, 0.72, and 0.90 mM Al. Rapid bioassays were applied to determine seedling responses of two Al‐tolerant (Cardinal and Becker) and two susceptible cultivars (GK Zombor and GK Kincso) and their F₂ progenies. In the Al‐toxic soil, Becker/Kincso F₂ and Cardinal/Zombor F₂ exhibited contrasting segregation patterns but with similar heritability values (0.60 and 0.57, respectively). Higher values of root length in soil were dominant in Cardinal/Zombor F₂ (degree of dominance, d = 0.98), but dominance was absent (d = 0.07) for Becker/Kincso F₂. The results of the soil and nutrient‐solution experiments were not entirely consistent; gene expression appeared to be influenced by the concentration of Al in the nutrient solution. The frequency of susceptible F₂ plants increased proportionately to the increase in Al concentration for both populations. This unexpected pattern provides further evidence that segregation in wheat populations cannot always be explained by single‐gene inheritance.     

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.     

Acid soil tolerance of leucaena species in greenhouse trials

Abstract

Lines of Leucaena leucocephala (Lam.) de Wit were grown in greenhouse pots of an acid, Al‐toxic Tatum subsoil (clayey, mixed, thermic typic Hapludult) treated with 0 or 3000 ppm CaCO₃ to give final soil pH values of 4.1 and 5.3, respectively. Lines of L. leucocephala, plus those of other Leucaena species, were also tested on an acid, Monmouth soil (clayey, mixed, mesic, typic Hapludult) treated with 0 or 1500 ppm CaCO₃ to give final soil pH values of 4.8 and 6.6, respectively. The major index of acid soil tolerance used was relative root yield (unlimed/limed %).

Relative root yields of 117 L. leucocephala lines on Tatum soil ranged from 34 to 246%. Hence, liming the soil from pH 4.1 to 5.3 was highly beneficial to some lines and highly detrimental to others. Because Tatum subsoil is 89% Al saturated at pH 4.1, line tolerance to unlimed soil indicates tolerance to Al. Causes of yield depression at pH 5.3 were not determined.

On Monmouth soil, in a test involving 148 lines of 6 Leucaena species, relative root yields (unlimed/limed %) ranged from 23 to 386%. The line showing highest tolerance to the acid soil (P.I. 279578) and that showing lowest tolerance (P.I? 281636) are both L,. leucocephala. The majority of lines used on Monmouth soil (124 of a total of 148) were from this species. Average performances of the 6 species indicated that L. diversifolia Benth. (5 lines) was most tolerant to the acid Monmouth soil and liming the soil from pH 4.8 to 6.6 actually decreased root yields. The species L.. leucocephala (124 entries) and L. pulverulenta Benth. (4 lines) were intermediate, and L. lanceolata S. Wats. (3 lines) and I., retusa Benth. (1 line) appeared more sensitive to acid Monmouth soil. The Al saturation of Monmouth soil at pH 4.8 was only 23% (compared with 89% for Tatum at pH 4.1). The major growth limiting factor in acid Monmouth soil is believed to be Al toxicity, but this soil has not been as throughly characterized as has Tatum, and other factors may well be involved in explaining differential tolerances of Leucaena lines on the unlimed versus limed soil.

Results of these studies indicate that Leucaena species and lines within species differ significantly in tolerance to acid soils having high levels of exchangeable Al. Acid soil tolerant lines of Leucaena may be useful in expanding the acreage of this crop on oxisols and ultisols of the tropics and subtropics.     

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.     

Tolerance of barley cultivars to an acid,aluminum‐toxic subsoil related to mineral element concentrations in their shoots

Abstract

Barley, Hordeum vulgare L., is extremely sensitive to excess soluble or exchangeable aluminum (Al) in acid soils having pH values below about 5.5. Aluminum tolerant cultivars are needed for use in rotations with potatoes which require a soil pH below 5.5 for control of scab disease. They are also potentially useful in the currently popular “low input, sustainable agriculture (LISA)”; in which liming even the plow layer of soil is not always possible or cost effective, or in situations where surface soils are limed but subsoils are acidic and Al toxic to roots. Ten barley cultivars were screened for Al tolerance by growing them for 25 days in greenhouse pots of acid, Al‐toxic Tatum subsoil (clayey, mixed, thermic, typic Hapludult) treated with either 750 or 4000 μg?g^‐1 CaCO₃ to produce final soil pH values of 4.4 and 5.7, respectively. Based on relative shoot dry weight (weight at pH 4.4/weight at pH 5.7 X 100), Tennessee Winter 52, Volla (England), Dayton and Herta (Denmark) were significantly more tolerant to the acid soil than Herta (Hungary), Kearney, Nebar, Dicktoo, Kenbar and Dundy cultivars. Relative shoot dry weights averaged 28.6% for tolerant and 14.1% for sensitive cultivar groups. Comparable relative root dry weights were 41.7% and 13.7% for tolerant and sensitive cultivars, respectively. At pH 4.4, Al concentrations were nearly three times as high in shoots of sensitive cultivars as in those of the tolerant group (646 vs. 175 μg?g^‐1), but these differences were reduced or absent at pH 5.7. At pH 4.4, acid soil sensitive cultivars also accumulated phosphorus concentrations that were twice as high as those in tolerant cultivars (1.2% vs. 0.64%). At pH 5.7, these P differences were equalized at about 0.7% for both tolerant and sensitive groups. At pH 4.4, shoots of the Al‐sensitive cultivar Nebar contained 1067 μg?g^‐1 Al and 1.5% P. Concentrations of Al and P in the shoots of acid soil sensitive cultivars grown at pH 4.4 exceeded levels reported to produce toxicity in barley. The observed accumulation of such concentrations of Al and P in the shoots of plants grown under Al stress is unusual and deserves further study.     

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.     

Differential tolerances of two old world bluestem genotypes to excess aluminum in acid soil and nutrient solution

Two genotypes of Old world bluestems from the species Bothriochloa intermedia (R. Br.), A. Camus, shown earlier to differ in tolerance to acid, Al‐toxic Tatum subsoil at pH 4.1, were characterized further with respect to growth in pots of Tatum soil over a wider pH range and tolerance to Al in nutrient solutions. The two genotypes studied were acid‐soil tolerant P. I. 300860 (860) and acid soil sensitive P. I. 300822 (822).

The soil experiment confirmed earlier rankings of acid soil tolerance in these two genotypes. For example, with 0, 375 or 750 ug CaCO₃ g^‐1 soil (final pH 4.0, 4.3 and 4.6), the 860 genotype produced significantly more dry top weight than 822, but these differences were precluded with 1500 or 3000 ug g^‐1 CaCO₃ added (pH 4.7 and 5.4). At pH 4.3 and 4.6, the root dry weights of the two genotypes were also significantly different and weights were equalized at pH 4.7 and 5.4. The 860 genotype made fairly good top growth (67% of maximum) at pH 4.3 and a soil Al saturation of 63%; this situation was lethal for 822. When grown in greenhouse pots, the acid‐soil tolerant 860 genotype required only about one fourth as much CaCO₃ as 822 to produce good growth of forage on acid Tatum subsoil. If confirmed under field conditions, such a difference could be economically significant in reclaiming acidic marginal land and in producing forage at low cost.

Differential Al tolerance in the two genotypes was confirmed in nutrient solutions. For example, with 8 mg Al L^‐1 added, both top and root dry weights of 860 were significantly higher than those of 822, but with no Al added, these growth differences disappeared.

Mineral analyses of plants did not shed much light on mechanisms of differential acid soil or Al tolerance. For example, Al concentrations in plant tops associated with toxicity varied from 33–43 ug g^‐1 in nutrient solutions containing Al to 119–283 ug g^‐1 in acid soil It appears that elucidation of Al‐adaptive mechanisms will require physiological and biochemical studies at the cellular level.     

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.     

Tolerance of Australian wheat varieties to aluminium toxicity

Abstract

This paper reports the reaction of 24 Australian wheats and 16 overseas cultivars to high aluminium (Al) in solution culture. These results are compared with those from a rapid haematoxylin stain test. The relationship between the haematoxylin stain test results and performance in the field was also determined.

The dry matter yields in solution culture confirmed tolerances previously reported for the non‐Australian cultivars, with only two exceptions. The Australian varieties vary in tolerance but none were as tolerant as those from Brazil. The tolerances of the Australian varieties were not related to the breeding origins of the varieties. Exposure to Al in solution differentially reduced the concentration of calcium (Ca), magnesium (Mg), and phosphorus (P) in both shoots and roots. The more Al‐tolerant varieties were less affected.

The results obtained in solution culture and in the haematoxylin stain test generally agreed, but more differences between varieties were noted in solution culture results. The haematoxylin stain test was then used to classify cultivars and advanced lines in the breeding programme, and the results were compared with yield performance on acid (8 sites) and non‐acid soils (20 sites). The lines in haematoxylin class 4 had a 20% yield advantage over the acid sites.

We concluded that tolerance was useful in the field, that the haematoxylin stain test is useful as a rapid preliminary assessment of Al tolerance, and that the prospect of breeding cultivars with improved tolerance was rewarding.     

Variation of tolerance to manganese toxicity in Australian hexaploid wheat

High concentrations of manganese (Mn), iron (Fe), and aluminium (Al) induced in waterlogged acid soils are a potential constraint for growing sensitive wheat cultivars in waterlogged‐prone areas of Western Australian wheat‐belt. Tackling induced ion toxicities by a genetic approach requires a good understanding of the existing variability in ion toxicity tolerance of the current wheat germplasm. A bioassay for tolerance to high concentration of Mn in wheat was developed using Norquay (Mn‐tolerant), Columbus (Mn‐intolerant), and Cascades (moderately tolerant) as control genotypes and a range of MnCl₂ concentrations (2, 250, 500, 750, 1000, 2000, and 3000 μM Mn) at pH 4.8 in a nutrient solution. Increasing solution Mn concentration decreased shoot and root dry weight and intensified the development of toxicity symptoms more in the Mn‐intolerant cv. Columbus than in Norquay and Cascades. The genotypic discrimination based on relative shoot (54% to 79%) and root dry weight (17% to 76%), the development of toxicity symptoms (scores 2 to 4) and the shoot Mn concentration (1428 to 2960 mg kg^–1) was most pronounced at 750 μM Mn. Using this concentration to screen 60 Australian and 6 wheat genotypes from other sources, a wide variation in relative root dry weight (11% to 95%), relative shoot dry weight (31% to 91%), toxicity symptoms (1.5 to 4.5), and shoot Mn concentration (901 to 2695 mg kg^–1) were observed. Evidence suggests that Mn tolerance has been introduced into Australian wheat through CIMMYT germplasm having “LERMO‐ROJO” within their parentage, preserved either through a co‐tolerance to Mn deficiency or a process of passive selection for Mn tolerance. Cultivars Westonia and Krichauff expressed a high level of tolerance to both Mn toxicity and deficiency, whereas Trident and Janz (reputed to be tolerant to Mn deficiency) were intolerant to Mn toxicity, suggesting that tolerance to excess and shortage of Mn are different, but not mutually exclusive traits. The co‐tolerance for Mn and Al in ET8 (an Al‐tolerant near‐isogenic line) and the absence of Mn tolerance in BH1146 (an Al‐tolerant genotype from Brazil) limits the effectiveness of these indicator genotypes to environments where only one constraint is induced. Wide variation of Mn tolerance in Australian wheat cultivars will enable breeding genotypes for the genetic solution to the Mn toxicity problem.     

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.     

Aluminum toxicity and high bulk density: Role in limiting shoot and root growth of selected aluminum indicator plants and eastern gamagrass in an acid soil

Abstract

Shallow rooting and susceptibility to drought are believed to be caused, at least in part, by strongly acidic (pH <5.5, 1:1 soil‐water), aluminum (Al)‐toxic subsoils. However, this hypothesis has not been clearly confirmed under field conditions. The Al toxicity hypothesis was tested on a map unit of Matawan‐Hammonton loam (0–2% slope) on unlimed and limed field plots (pH range 5.1 to 5.8) at Beltsville, MD, during 1994 to 1998. Aluminum‐tolerant and sensitive pairs of barley (Hordeum vulgare L.), wheat [Triticum aestivum (L.)], snap bean (Phaseolus vulgaris L.), and soybean [Glycine max (L.) Merr.] cultivars were used as indicator plants. Eastern gamagrass [Tripsacum dactyloides (L.) L.], cultivar ‘Pete’, reported to tolerate both chemical and physical stress factors in soils, was grown for comparison. Shoots of Al‐sensitive ‘Romano’ snap beans showed a significant response to liming of the 0–15 cm surface layer, but those of Al‐tolerant ‘Dade’ did not, indicating that Al toxicity was a growth limiting factor in this acid soil at pH 5.1. Lime response of the Al‐tolerant and sensitive cultivars of barley, wheat, and soybean were in the same direction but not significant at the 5% level. Aluminum‐tolerant and sensitive cultivars did not differ in abilities to root in the 15–30 cm soil depth. Only 9 to 25% of total roots were in this layer, and 75 to 91% were in the 0–15 cm zone. No roots were found in the 30–45 cm zone which had a pH of 4.9. Soil bulk density values of 1.44 and 1.50 g cm^?3 in the 15–30 and 30–45 cm zones, respectively, indicated that mechanical impedance was a primary root barrier. Results indicated that restricted shoot growth and shallow rooting of the Al‐indicator plants studied in this acid soil were due to a combination of Al toxicity and high soil bulk density. Confounding of the two factors may have masked the expected response of indicator plants to Al. These two growth restricting factors likely occur in many, if not most acid, problem subsoils. Studies are needed to separate these factors and to develop plant genotypes that have tolerance to multiple abiotic stresses. Unlike the Al indicator cultivars, eastern gamagrass showed high tolerance to acid, compact soils in the field and did not respond to lime applications (pH 5.1–5.8).     

Short‐term effects of pH and aluminium on mineral nutrition in maize varieties differing in proton and aluminium tolerance

In short‐term (24 h) nutrient solution experiments, the influence of different proton (pH 6.0 and pH 4.3) and aluminium (Al) (0, 20, and 50 μM) concentrations on root and coleoptile elongation, dry weight, and the uptake of selected mineral nutrients was studied in maize (Zea mays L.) varieties that differ in acid soil tolerance under field conditions. The acid‐soil‐tolerant maize varieties, Adour 250 and C525M, proved to be hydrogen (H⁺) ion sensitive, but Al tolerant, while the acid soil tolerant variety BR201F was H⁺ tolerant but Al sensitive. The acid soil sensitive variety HS 7777 was affected by both H⁺ and Al toxicity. The proton‐induced inhibition of root elongation was closely related to the proton‐induced decrease of the specific absorption rates (SAR) of boron (B), iron (Fe), magnesium (Mg), calcium (Ca), and phosphorus (P). In contrast, only the specific absorption rate of B (SARB) was significantly correlated to the Al‐induced inhibition of root elongation. It is concluded, that alterations of nutrient uptake may play an important role in H⁺ toxicity, while at least after short‐term exposure to Al, alterations of Ca, Fe, Mg, or P uptake do not seem to be responsible for Al‐induced inhibition of root elongation. Further attention deserves the Al‐B interaction, moreover taking into account that a highly significant correlation between Al‐induced increase of callose concentration in root tips and Al‐induced decrease of SAR_B could be established.     

Differential response of cowpea lines to aluminum and phosphorus application

Two pot experiments were conducted, one to evaluate the levels of tolerance of fifteen cowpea [Vigna unguiculata (L.) Walp] lines to aluminum (Al) application, and the second to determine the effect of phosphorus (P) addition on the performance of Al‐tolerant lines (IT 91K‐93–10, IT 93K‐2046–1, and IT 90K‐2 77–2) and Al‐sensitive lines (IT 86D‐719, IT 90K‐284–2, and IT 89KD‐349) in an Alfisol with Al amendment. Fourteen of the fifteen lines tested showed decreased root biomass (between 19 to 81% reduction) with Al addition, but this effect was significant for eight of them. Fewer lines showed decreased shoot biomass and grain yield with Al application. Despite little change in nodule number following Al application, there was a significant decrease in nodule weight (between 24 and 53% reduction) for nearly all lines. Phosphorus fertilization increased shoot and root biomass, grain yield, nodule number, and weight, and nitrogen (N) and P content of nearly all lines. Al‐tolerant lines showed higher response in shoot and root biomass and nodulation to P fertilization than Al‐sensitive lines, with the highest response from IT 90K‐277–2. Increase in shoot dry weight as a result of P fertilization was from 64 to 107% for Al‐tolerant lines and from 44 to 48% for the Al‐sensitive lines, and increase in root dry weight was from 46 to 86% for the Al‐tolerant lines and from 7 to 42% for the Al‐sensitive lines. Results of these trials indicated that lines IT 91K‐93–10, IT 93K‐2046–1, and IT 90K‐277–2 have potential for good performance in soil with Al toxicity problems, and that cowpea lines with inherent genetic tolerance to Al will give higher response to P fertilization when grown in soil with Al toxicity problems.