Physiological basis of differential aluminum tolerance in rice genotypes

Abstract

Two rice genotypes, aluminum (Al)‐tolerant Co 37 and Al‐susceptible ADT 36, were evaluated for their physiological responses in the presence of Al stress in a hydroculture experiment. Two levels of Al (0 and 222 μmol/L) were supplied in the nutrient solution and the two rice genotypes were subjected to Al for two weeks. Root growth parameters, relative growth reduction in roots (RGRR), effects of calcium (Ca²⁺) nitrate metabolism, Al content in roots, and pH shift patterns were recorded. The Al treatments had a lesser effect on Co 37 in terms of reduction in root growth and root dry matter production as compared to ADT 36. While Co 37 did not show significant differences in response to various levels of Ca²⁺ added in the medium under Al stress, ADT 36 registered a dose‐dependent effect in circumventing the injurious effects of Al. Further, reduction in nitrate content and in vivo nitrate reductase (NR) activity in the leaf tissue in Co 37 under Al treatment was less than that in ADT 36. Moreover, Co 37 had a lower content of Al in its root tissue than did ADT 36. Co 37 was also able to shift the pH of the medium more efficiently than ADT 36, thereby minimizing the uptake of Al, and eventually reducing Al toxicity. The higher level of tolerance to Al of Co 37 seems to have emanated from an efficient nitrate metabolism and its capacity to shift the pH of the medium. It is also evident that Al toxicity in ADT 36 can be circumvented by a Ca treatment to a considerable extent. Our results offer a possible physiological basis for Al tolerance in crop plants.     

Genotypic Difference of Germination and Early Seedling Growth in Response to Cd Stress and its Relation to Cd Accumulation

A greenhouse experiment was conducted using twelve Japonica rice genotypes differing in grain Cd concentration, to study the genotypic difference in germination and early seedling growth in response to Cd stress and its relation to Cd accumulation. The results showed that under low Cd level (0.5 mM), germination was significantly enhanced for most rice genotypes, but higher Cd levels (2.5 and 10.0 mM) inhibited germination dramatically. When exposed to 10.0 mM Cd, a few genotypes could still germinate, but the seedlings of all genotypes would die in early stage. Shoot height and dry weight, and root length and dry weight of seedling were significantly inhibited at 0.5 and 2.5 mM Cd levels, irrespectively of genotypes, and the inhibition became more severe under higher Cd level. There was a significant genotypic difference in response of germination, early seedling growth and shoot Cd concentration to Cd stress. The difference was the largest for germination, followed by root growth, the least for shoot growth. A cluster analysis of tolerance indices, including germination rate, shoot height and weight, root length and weight of the plants exposed to 2.5 mM Cd level showed that 12 rice genotypes may be grouped into three clusters, i.e. high-, intermediate- and weak Cd-tolerance. No significant correlation was found between grain Cd concentration and Cd-tolerance in terms of germination and seedling growth, and shoot Cd concentrations, suggesting that it is possible to develop rice cultivars with high Cd-tolerance and low grain Cd accumulation.     

Effect of water stress on aluminum toxicity in pitch pine seedlings

A decrease in soil water content during droughts may increase aluminum (Al) to concentrations that are toxic to the growth of trees. The effects of water stress (WS) on the response of ectomycorrhizal pitch pine (Pinus rigida Mill.) seedlings to aluminum was determined by growing seedlings in sand irrigated with nutrient solution (pH 3.8) containing 0, 5, or 10 mg L^‐1 Al. Water stress was imposed for 41 days by withholding nutrient solution for five consecutive days each week. At harvest time, seedlings at high WS had 72% of mean gravimetric water contents of seedlings at low WS. Aluminum decreased growth of seedlings at high WS, but had no effect on growth of seedlings at low WS. Aluminum toxicity symptoms in roots (e.g., dark thickened tips) were observed at lower Al levels at high WS than at low WS. Stem dry weight was the only plant part decreased by water stress alone. Across Al levels, Al concentration in roots was higher at low WS than at high WS. Water stress alone reduced root [phosphorus (P), potassium (K), and calcium (Ca)] and foliar [P, K, and magnesium (Mg)] concentrations of mineral nutrients. Decreases of nutrients in roots with increasing Al was greater at low than at high WS. Calcium was the only foliar nutrient decreased by Al treatment.     

Amelioration of Aluminum Toxicity by Pretreatment with Phosphate in Aluminum‐Tolerant Rice Cultivar

Abstract

The phytotoxicity of aluminum (Al) in relation to preculture with phosphates was examined in the rice cultivar Arkansas fortuna. In plants precultured with phosphates, Al did not inhibit shoot growth, while Al retarded shoot growth in plants precultured without phosphates. In contrast, Al inhibited root elongation, irrespective of the presence of phosphates in the preculture solution. A large proportion of the Al in roots was in unknown, insoluble forms. In phosphate‐precultured plants, Al deposition was slightly increased, presumably due to the formation of aluminum phosphates in the roots, and phosphorus levels in shoots were markedly increased. Binding with phosphates may ameliorate the toxicity of Al when it enters the shoots and account for the uninhibited shoot growth in the presence of Al in plants precultured with phosphates.     

Identification of an aluminum tolerant tropical cowpea cultivar by growth and biomass accumulation parameters

Ten‐day‐old seedlings of four cowpea (Vigna unguiculata Walp) genotypes were subjected to six levels of aluminum (Al) (0, 74, 148, 222, 296, and 370 μM/L) to test their tolerance to Al toxicity in a nutrient solution at pH 4.0±0.1. Seedlings were grown in the presence of Al under controlled environmental conditions in a growth chamber. The nutrient solutions were replenished once a week. After 20 days, treatments were terminated and the differences in their growth patterns were compared. Standard growth parameters, such as plant growth, dry matter production, relative growth reduction in roots (RGRS) and shoots (RGRS), and root and shoot tolerance indices (RTI and STI) have been used as markers of Al toxicity. The cowpea genotypes studied exhibited a wide range of responses in their tolerance to Al. Though the genotypes were subjected to six levels of Al, a good degree of separation in their responses was observed only at the 222 μM Al/L treatment level. Therefore, this concentration was chosen to treat and compare the performances of the genotypes. The genotype Co 3 showed an increase in growth, while Paiyur 1 and other genotypes showed severe inhibitions in the presence of Al. Furthermore, for RTI and STI, Co 3 also registered its tolerance to Al by showing increased ratios in the presence of Al. Whereas, Paiyur 1 recorded severe reductions. The RGRR and RGRS data also substantiates this finding. Based on the growth parameters, the four cowpea genotypes were ranked based on their tolerance to Al: Co 3 > Co 4 > KM > Paiyur 1. Co 3 was the most Al‐tolerant genotype which performed extremely well in the presence of Al, while Paiyur 1was the most Al‐susceptible genotype. Therefore, the Al‐tolerant genotype can be used for future breeding programmes to produce Al‐tolerant genotypes, subsequently, can be recommended for acidic infertile soils in the tropics.     

Effect of aluminium and calcium on growth of wheat seedlings and germination of seeds

Two separate experiments were conducted to investigate the aluminium (Al) and calcium (Ca) effects on wheat seedling growth and on seed germination. Wheat (Tritcum aestivum L, cs Yangmai No. 5) seedlings were grown for a 15‐day period and treated with 0.5 mM Al with low Ca (1 mM Ca) or high Ca (5 mM Ca). The growth of seedlings was signficantly inhibited by Al. Supplement of Ca improved the growth of Al‐treated plants, increased dry matter weight of plant and leaf area, and decreased shoot/root ratio. This showed that Ca ameliorated Al toxicity in wheat. In experiments on seed germination, Al concentrations less than 2 mM in the germinating medium had little or no visible effect on length of shoot and root of germinating seed. The germinating rate of seed was not affected significantly by Al, when Al concentrations lower than 5 mM Al. The addition of 3 mM Ca did not increase the length of shoot and root and germination rate of seeds. Both pretreatments with 6 mM Ca and 1 μM GA had no significant effect on the length of shoot and root and amylolytic activity of Al‐treated germinating seeds. No significant differences were found in the total amylolytic activity in Al‐treated and control seeds two days and five days after germination. The results of Al and Ca effects on seedlings and seed germination showed that Al‐toxicity on germinating seeds was different from on seedling growth. The high concentrations of Al inhibit growth of roots and shoots of germinating seeds by other toxicity mechanism rather than interaction of Al with Ca and mobilization of carbohydrate reserves.     

不同水、氮条件对水稻苗生长及伤流液的影响

为探明不同水分供应和氮素形态对水稻根苗及伤流液的影响,设正常水分及50 g/L PEG模拟水分胁迫和3种不同质量比例的NH4+-N/NO3--N（9/1,5/5,1/9）氮素营养处理,测定了水稻幼苗生物量,根系形态指标,根系活力及根基伤流量。结果表明,正常水分条件下,NH4+-N促进水稻根系平均直径增大,有利于水稻地上部物质累积;NO3--N则使水稻根系总吸收面积增大,促进根系物质累积;NH4+-N/NO3--N为5/5处理的水稻活跃吸收面积最大,活跃吸收面积比亦最高。水分胁迫条件下,NH4+-N/NO3--N为5/5的处理更有利于水稻地上部分的生长,NO3--N有利于水稻鲜重和干重增加,促进根系平均直径增大,水稻的根系总吸收面积、活跃吸收面积均随NO3--N供应比例的增加呈上升趋势。正常水分条件下,水稻幼苗白天的耗水量随NH4+-N/ NO3--N比例降低呈下降趋势,水分胁迫条件降低了水稻对水分的吸收。水分胁迫显著降低各处理水稻伤流量,正常水分条件下,NH4+-N/NO3--N为5/5处理的水稻伤流量最大;水分胁迫后,9/1处理的水稻伤流量相对较多。     

Elemental toxicity effects on germination and growth of pearl millet seedlings 1

Variability in millet stands in West Africa is clearly visible as early as three weeks after planting. The objectives of this study were to determine the influence of pH and chemical toxicities on millet germination and seedling growth and to compare varietal tolerance of toxic conditions. A nutrient solution study was carried out with a series of Hoagland‐based nutrient solutions. Germination percentage was calculated, and root and shoot lengths were measured for one week. Critical values were determined for toxic elements. The only treatment which reduced germination percentage significantly was copper (Cu) concentrations >0.05M. Solution pH values between 5 and 7 resulted in the best root growth, though shoot growth was unaffected by pH. The roots were more sensitive than the shoots to several [aluminum (Al), boron (B), zinc (Zn)] of the elemental toxicities studied. Soil Al and manganese (Mn) levels may be high enough to have toxic effects on millet roots. However, natural soil iron (Fe), Cu, and Zn levels were much lower than the critical levels determined in the nutrient solution study. The improved varieties were more tolerant of Fe and Zn toxicity than the LOCAL variety, but the LOCAL variety was more tolerant of high B concentrations.     

Changes in nitrate reductase activity,plastid pigment content,and plant mineral composition of wheat,rye, and triticale grown in the presence of aluminum

The effects of aluminum (Al) on nitrate reductase activity, plastid pigment content, and mineral element composition in wheat, rye, and triticale seedlings were studied. Different responses of the plant species to Al content in the growth solution were observed. Under conditions of different Al concentrations (from 1 to 10 ppm), nitrate reductase (NR) activity increased in wheat and triticale, while in rye an interference with nitrate reductase by Al was observed. A definite tendency in plastid pigment content changes independent on Al levels was not found. The chlorophyll “a”;, chlorophyll “b”;, and carotenoid contents were influenced in a different way in wheat, rye, and triticale seedlings. A positive effect of 1, 5, and 10 ppm Al on the nitrogen (N) content in the shoots of wheat and rye was observed. The N content in the shoots of triticale was not affected by the presence of Al. The presence of Al in the nutrient solution led to a tendency toward reduction of potassium (K), calcium (Ca), and magnesium (Mg) contents in the shoots of rye seedlings, a reduction of K in the shoots and manganese (Mn) content in roots of wheat, and a reduction of K in both shoots and roots and an accumulation of Mn in triticale roots. In general, our investigation on the effect of Al in the early stages of wheat, rye, and triticale development showed that a large number of biochemical and physiological parameters are required to characterize the plant responses to Al stress.     

Mechanisms of Aluminum Tolerance in Phosphoenolpyruvate Carboxylase Transgenic Rice

Transgenic rice, in which C4-phosphoenolpyruvate carboxylase (PEPC) is overexpressed, was grown in hydroponic culture to evaluate its tolerance for high aluminum (Al) content in relation to organic acid synthesis and exudation into the rhizosphere. With regard to root growth in short-term experiments, PEPC-transgenic rice was more tolerant of Al than the wild type. At increased Al levels in the root medium, root tips of the transgenic rice accumulated less Al than those of the wild type. Aluminum-induced oxalate exudation from roots occurred at increased rates in the transgenic line. Overexpression of C4-PEPC drastically increased PEPC activity in leaves of the transgenic rice, causing higher organic acid concentrations in leaves and roots. The increased organic acid concentrations seemed to cause higher exudation rates of oxalate from roots, leading to a detoxification of Al in the root apoplast and rhizosphere.     

外源有机酸缓解水稻幼苗根系铝毒的生理机制

以4种杂交水稻品种为材料,进行溶液培养,测定了不同Al胁迫处理下发芽期幼苗的根长、根重、根电导率以及根系线粒体中重要的活性氧清除酶POD、CAT和APX的变化,研究了水稻幼苗对外源有机酸和Al毒响应的生理机制。结果表明:1.5mmol/L Al胁迫下水稻幼苗根系受到轻微伤害,同时POD和APX活性明显增加,CAT活性变化不大;而添加有机酸后,Al胁迫得到明显缓解,其中琥珀酸的缓解效果最为明显,柠檬酸、苹果酸、柠檬酸 琥珀酸的效果表现出品种的特异性;且加入外源有机酸后,POD和APX活性呈下降趋势,CAT的活性变化相对较小。表明Al胁迫引起水稻根细胞线粒体中活性氧代谢平衡发生变化,而有机酸的加入调节了Al胁迫下根系细胞线粒体中活性氧清除酶的活性,从而缓解了发芽期水稻幼苗的Al毒。     

不同硝响应型水稻品种苗期根系生长对增硝营养的响应

利用控制条件下的水培试验方法,研究了两种铵硝配比(NH4+/NO3-为100/0和75/25)营养条件对4种不同硝响应型水稻品种苗期根系生长的影响。结果表明,在增硝营养(NH4+/NO3-为75/25)条件下,不同水稻品种NO3-的反应差异明显。与全NH4+营养条件相比,增硝营养条件下对NO3-强响应的水稻品种南光的根系干重和氮积累量显著增加,增幅达50%和79%;同时南光的根系总根长、总不定根长和总侧根长增幅均达到显著水平;不定根数、新根数和侧根数亦显著增加;平均不定根长和平均侧根长差异不显著;对硝弱响应型的水稻品种上海97、辽粳和Elio在增硝营养培养下的根系不定根、新根和侧根的长度和数量差异均不显著。这表明增NO3-营养仅仅促进了对NO3-强响应型水稻南光根系的不定根和侧根的发生,进而促进根系对氮素的吸收,并没有促进不定根和侧根的伸长。从本试验的结果可推论,水稻根系对硝态氮的响应度强弱可能是水稻品种氮素效率差异性的因子之一。     

Calcium‐induced modification of aluminum toxicity in sorghum genotypes

Effects of calcium (1, 2 and 5 mM) and aluminum (0, 15 and 45 μM) on growth and internal nutrient concentrations were examined with 12 sorghum genotypes (Sorghum bicolor (L.) Moench) in a nutrient solution experiment with a factorial design. At 1 (or 2) mM Ca the severity of root damage induced by Al well reflected the genotypical variation in growth response to Al toxicity. Severity of Al‐induced root damage slightly decreased with increasing Ca level. Moreover, Ca at 5 mM amplified the Mg deficiency induced by Al, as seen from both heavier deficiency symptoms and lower internal Mg concentrations. Under conditions of Al stress at a high Ca supply, induced Mg deficiency apparently predominated the genotypical differentiation in growth response to Al toxicity. An antagonism between Al‐ and Ca ions for uptake was hardly found with the sorghum genotypes. However, the genotypes differed in Ca efficiency, a characteristic which may be relevant in assessing their sensitivity to Mg deficiency.     

Accumulation of amino acids,proline, and carbohydrates in response to aluminum and manganese stress in maize

Synthesis of amino acids, proline, and carbohydrates was studied in roots and shoots of 5 maize accessions, differing in aluminum (Al) and manganese (Mn) tolerance, in response to Al and Mn stress at the seedling stage in solution culture. The concentrations of these metabolites increased in roots and shoots of the seedlings in the nutrient solution with added Al (0.22 mM), and Mn (2.0 mM). Both Al and/or Mn tolerant and non‐tolerant accessions accumulated more metabolites under stress than control. Generally, the tolerant accessions accumulated more solutes than the non‐tolerant maize accessions examined.     

The effect of aluminum on the germination of wheat seeds 1

The effect of aluminum (Al) ions on the germination of wheat (Triticum aestivum L.) seeds has been investigated using two varieties, one sensitive to Al (Robin) and the other relatively Al‐tolerant (Carazinho). High concentrations of Al (i.e. greater than 1 mM) were required to inhibit the growth of the emerging roots and shoots of germinating seeds of both varieties. Moreover, Al at concentrations up to 10 mM did not affect the mobilization of carbohydrate reserves in the germinating seeds. It was concluded that germinating seeds of both wheat varieties were much less sensitive to the effects of Al than were established seedlings.     

Differential aluminum tolerance in some tropical rice cultivars: I. Growth performance

Ten‐day‐old seedlings of 22 rice (Oryza sativa.L) cultivars originated from various tropical countries were subjected to six levels of aluminum (Al) [0, 74, 148, 222, 296, and 370 μM] to test their tolerance to Al toxicity in nutrient solutions at pH 4.0±0.l. Seedlings were grown in the presence of Al under controlled environmental conditions in growth chambers. The nutrient solutions were replenished once a week. After 30 days, treatments were terminated and the differences in their growth patterns were compared. Standard growth parameters such as plant growth, dry matter production, relative growth reduction in roots (RGRS) and shoots (RGRS), root tolerance index (RTI) and shoot tolerance index (STI) have been used as markers of Al toxicity.

Rice cultivars studied exhibited wide range of responses in their tolerance to Al. Though, the rice cultivars were subjected to six levels of Al, a good degree of separation in their responses was observed only at 222 μM Al. Therefore, this concentration was chosen to analyze and compare the performances of the cultivars. Further, only six cultivars showed significant changes in their expression in the presence of Al compared to control, and so data have been presented only for those cultivars for clarity. The cultivars BW 196, Bhura Rata, Basmati 370 and Co 37 recorded increases in growth, while Damodar and ADT 36 showed severe inhibitions in the presence of Al. Furthermore, in RTI and STI also Co 37 and Basmati 370 registered their tolerance to Al by showing increased growth in the presence of Al. Whereas, Damodar and ADT 36 recorded severe reductions. The RGRR and RGRS data also substantiates this finding. Based on the growth parameters, the six rice cultivars were ranked based on their tolerance to Al: Co 37 > Basmati 370 > BW 196 > Bhura Rata > Damodar > ADT 36. Co 37 and Basmati 370 are the two most tolerant cultivars which performed extremely well in the presence of Al, and Damodar and ADT 36 are the most susceptible cultivars. Therefore, the Al‐tolerant cultivars can be used for future breeding programes to develop Al‐tolerant, cultivars that subsequendy can be recommended for planting in acidic, infertile soils of the tropics.     

Tea plant (Camellia sinensis L.) roots secrete oxalic acid and caffeine into medium containing aluminum

We examined the response of the tea plant (Camellia sinensis L.) to aluminum (Al) exposure under sterile conditions, focusing specifically on the secretion of low molecular mass organic compounds from roots. After germination in agar medium, tea seedlings together with medium were placed on agar containing 0.4?mM Al with 0.2% hematoxyline (hematoxylin-Al medium). The purple color of the hematoxylin-Al medium was observed to fade gradually, until none of the color remained 6 days later. The tea seedlings were then treated with simple calcium solution (0.2?mM, at pH 4.2) containing AlCl₃, which ranged in concentration from 0 to 0.8?mM, for 24?hrs. The amount of oxalate secreted into the medium increased as the external Al concentration increased, while the concentrations of malate and citrate in the medium remained unchanged. Oxalate secretion started within 30?min after Al exposure and increased linearly thereafter. The findings demonstrated that oxalate was a key compound in the Al-tolerance mechanism employed by the tea plant, which detoxifies Al³⁺ externally in the rhizosphere. In addition to oxalate, caffeine was also secreted by tea roots in response to Al exposure. It is possible that caffeine excretion from the roots of tea plants may stimulate root growth through the inhibition of callose deposition in root tips.     

Boron amelioration of aluminum toxicity in mungbean seedlings

Boron (B) amelioration of aluminum (Al) toxicity was studied for growth of mungbean (Phaseolus aureus Roxb.) seedlings and cuttings (without roots) in a growth chamber. Mungbean seedlings and cuttings were grown in the solution with combinations of three concentrations of B (0,5, and 50 μM) and three concentrations of Al (0, 2, and 5 mM) in randomized complete block design experiments for 16 days. Results showed that B significantly promoted elongation of epicotyls and hypocotyls, and increased height of seedlings grown under Al stress. Boron also increased fresh weight of seedlings in high Al solution. Treatment of plants grown with high B and Al stress had no apparent effect on fresh and dry weights of seedling roots. Seedling dry weight increased significantly by adding high B to solutions with 2 mM or 5 mM Al. No significant differences were observed between the high B treatment and the control (normal B, 5 μM) in lengths of epicotyls and fresh and dry weights of mungbean cuttings grown under Al stress. High concentrations of B decreased soluble protein and increased chlorophyll in seedlings treated with 2 mM Al. Boron had no amelioration effect on cuttings grown with Al, although Al increased soluble protein. Our results suggested that B alleviation of Al toxicity was related to root function and Al toxicity may possibly be due, in part, to B deficiency.     

Silicon effects on aluminum toxicity to mungbean seedling growth

The effects of silicon (Si) on the toxicity of aluminum (Al) to mungbean (Phaseolus aureus Roxb.) seedlings were studied in a growth chamber. Mungbean seedlings were grown in a nutrient solution with combinations of three concentrations of Si (0,1, and 10 mM) and three concentrations of Al (0, 2, and 5 mM) in randomized completely block design experiments for 16 days. Silicon at 1 mM in the solution decreased root length, fresh and dry weights, and chlorophyll content, and showed no significant effect on epicotyl length and seedling height, and protein contents of shoots or roots in mungbean seedling under no Al stress. But, Si at 10 mM showed marked toxic effects on mungbean seedling growth and increased protein contents of the shoots or roots. In contrast, under 2 mM Al stress, Si addition at 1 mM had significant increasing effect on root length, fresh and dry weights, and chlorophyll content. It also had decreasing effect on protein contents of the shoots or roots, and had no effect on epicotyl length and seedling height. Silicon addition at 10 mM showed no effect on morphological and physiological measurements of mungbean seedling. However, Si at 1 mM added to solution only increased seedling height, epicotyl length, fresh weight, and chlorophyll content, but decreased dry weight and protein content of the roots under 5 mM Al stress, significantly. Silicon addition at 10 mM showed similar toxic effects on mungbean seedling growth under 5 mM Al stress to that under no Al stress.