Seedling root response of some Kenyan bread wheat cultivars grown in acid nutrient culture solution containing aluminum

The objective of this study was to determine whether a series of Kenyan bread wheat cultivars differed in tolerance to aluminum toxicity. Fourteen Kenyan wheat cultivars representing current and former widely grown cultivars of diverse pedigree origin, and two control cultivars, Maringa (Al-tolerant) and Siete Cerros (Al-susceptible), were tested in solution cultures with 0 (control), 148, 593, and 2370 M Al at pH 4.6. Highly significant (p0.01) differences in seedling growth were observed among cultivars for root mass, root length and root tolerance index (RTI). Significant (p0.05) cultivar × treatment interactions were observed for root mass and RTI. All characters were negatively affected by increased Al concentration, with root length and root mass being affected the most. RTI is a commonly used index which measures the relative performance of individual cultivars with and without aluminum stress. High levels of tolerance to Al were identified in the Kenyan cultivars by evaluating RTI with this simple nutrient solution technique. Romany and Kenya Nyumbu had RTI values approaching those of the Al tolerant Brazilian cultivar Maringa, a spring wheat standard that has been used for high Al tolerance.     

The effect of phosphate rock dissolution on soil chemical properties and wheat seedling root elongation

Soils of the Appalachian region of the United States are acidic and deficient in P. North Carolina phosphate rock (PR), a highly substituted fluoroapatite, should be quite reactive in these soils, allowing it to serve both as a source of P and a potential ameliorant of soil acidity. An experiment was conducted to evaluate the influence of PR dissolution on soil chemical properties and wheat (Triticum aestivum cv. Hart) seedling root elongation. Ten treatments including nine rates of PR (0, 12.5, 25, 50, 100, 200, 400, 800, and 1600 mg P kg^-1) and a CaCO₃ (1000 mg kg^-1) control were mixed with two acidic soils, moistened to a level corresponding to 33 kPa moisture tension and incubated for 30 days. Pregerminated wheat seedlings were grown for three days in the PR treated soils and the CaCO₃ control. Root length was significantly (P<0.05) increased both by PR treatments and CaCO₃, indicating that PR dissolution was ameliorating soil acidity. The PR treatments increased soil pH, exchangeable Ca, and soil solution Ca while lowering exchangeable Al and 0.01 M CaCl₂ extractable soil Al. Root growth in PR treatments was best described by an exponential equation (P<0.01) containing 0.01 M CaCl₂ extractable Al. The PR dissolution did not reduce total soil solution Al, but did release Al complexing anions into soil solution, which along with increased pH, shifted Al speciation from toxic to nontoxic forms. These results suggest that North Carolina PR should contribute to amelioration of soil acidity in acidic, low CEC soils of the Appalachian region.     

Chlorophyll content and leaf elongation rate in wheat seedlings as a measure of manganese tolerance

After aluminum toxicity, manganese (Mn) toxicity is probably the second most important growth limiting factor in acid soils. The purpose of this study was to determine the feasibility of using chlorophyll content and leaf elongation rate (LER) for regrowth of Mn stressed seedlings as a rapid seedling based screening bioassay for Mn tolerance in segregating populations of wheat (Triticum aestivum L.). In one experiment, chlorophyll was determined for the cultivars Norquay (Mn-tolerant) and Columbus (Mn-sensitive) subjected to twelve Mn levels (2 to 2000 μM) in nutrient solutions. As Mn concentration increased, chlorophyll ‘a’ and ‘b’ contents of the Mn-tolerant cultivar decreased up to 9%, while in the Mn-sensitive cultivar it was reduced by as much as 43%. The chlorophyll ‘a/b’ ratio did not differ among Mn concentrations for either cultivar. In a second experiment, chlorophyll content and LER for regrowth of Mn stressed seedlings (1000 μM) was determined for Columbus and Katepwa (Mn-sensitive), Oslo (Mn-intermediate), and Norquay and Laura (Mn-tolerant). Manganese tolerance as assayed by chlorophyll ‘a’ and ‘b’ and LER was significantly correlated with Mn tolerance as assayed by the relative root weight methodology (RRW). Thus, chlorophyll content of Mn-stressed seedlings and LER of seedling regrowth appear to be suitable techniques for screening unreplicated selections of segregating populations for tolerance to Mn.     

The control of boron accumulation by two genotypes of wheat

The mechanism of boron (B) uptake in wheat was studied using two genotypes with known differences in their ability to accumulate B. Influx and efflux of B was measured in the roots of intact 21 d old plants.Roots grown in 15 M B, when transferred to solutions containing 1mM B showed a rapid increase in B content for up to 60 min, after which no further increase was evident up to 4 h. No genotypic difference in B influx was apparent over these time periods. Roots grown in 1mM B for 7 d and then rinsed in B-free solutions quickly lost most of B that they contained within 1 hour; little further efflux was observed over the following three hours. As with the influx, no genotypic difference in B flux was evident.It is suggested that the lack of genotypic difference in the short-term B fluxes could be due to a masking effect of extracellular B bound in the cell walls of the roots.Department of Botany, University of Adelaide     

Accelerated production of transgenic wheat (Triticum aestivum L.) plants

We have developed a method for the accelerated production of fertile transgenic wheat (Triticum aestivum L.) that yields rooted plants ready for transfer to soil in 8–9 weeks (56–66 days) after the initiation of cultures. This was made possible by improvements in the procedures used for culture, bombardment, and selection. Cultured immature embryos were given a 4–6 h pre-and 16 h post-bombardment osmotic treatment. The most consistent and satisfactory results were obtained with 30 g of gold particles/bombardment. No clear correlation was found between the frequencies of transient expression and stable transformation. The highest rates of regeneration and transformation were obtained when callus formation after bombardment was limited to two weeks in the dark, with or without selection, followed by selection during regeneration under light. Selection with bialaphos, and not phosphinothricin, yielded more vigorously growing transformed plantlets. The elongation of dark green plantlets in the presence of 4–5 mg/l bialaphos was found to be reliable for identifying transformed plants. Eighty independent transgenic wheat lines were produced in this study. Under optimum conditions, 32 transformed wheat plants were obtained from 2100 immature embryos in 56–66 days, making it possible to obtain R3 homozygous plants in less than a year.     

The fate of carbon and fertiliser nitrogen when dryland wheat is grown in monoliths of duplex soil

Triticum aestivum L. (cv. Gutha), a short-season wheat, was grown to maturity in large monoliths of duplex soil (sand over sandy-clay) in a daylight phytotron mimicking field conditions. Either ¹⁵N-labelled ammonium sulphate ((NH₄)₂SO₄) or urea was banded into the soil at a rate of 30 kg N ha^–1: even though roots were about 20% heavier when grown in the presence of (NH₄)₂SO₄ for 86 d (P<0.05), above-ground mass was not affected by the source of nitrogen. At four times through crop development up to grain-filling (50, 56, 70 and 86 d after sowing) shoots were labelled heavily with ¹⁴CO₂ with two purposes. First, to trace `instantaneous' assimilate movement over 24 h, revealing relative sink strengths throughout plants. This, in turn, allowed precise measurements of live root mass and the proportion of recent photoassimilates deposited in the rhizosphere. Although root systems were sparse, even in surface soil layers, they were strong sinks for photoassimilates early in development (0–50 d), supporting the conversion of inorganic applied nitrogen (N) to soil organic forms. In the presence of roots, up to 28% of ¹⁵N was immobilised, whereas only 12% of labelled ammonium sulphate was immobilised in unplanted plots in spite of a favourable moisture status in both treatments. The effect of plants on rates of ¹⁵N transformation is ascribed to recently imported photoassimilates sustaining rhizosphere metabolism. Not more than 15% of recently fixed carbon imported by roots was recovered from the rhizoplane, suggesting that a highly localised microbial biomass supported vigorous immobilisation of soil N. Thus, more than twice as much applied N was destined for soil organic fractions as for root material. By these processes, root- and soil-immobilised N become substantial stores of applied N and together with shoot N accounted for all the applied N under dryland conditions.     

Importance of seed Zn content for wheat growth on Zn-deficient soil

Seed nutrient reserves may be important for an early establishment of crops on low-fertility soils. This glasshouse pot study evaluated effects of seed Zn content on vegetative growth of two wheat (Triticum aestivum L.) genotypes differing in Zn efficiency. Low-Zn (around 250 ng Zn per seed) and high-Zn seed (around 700 ng Zn per seed on average) of Excalibur (Zn efficient) and Gatcher (Zn inefficient) wheats were sown in a Zn-deficient siliceous sand fertilised with 0, 0.05, 0.2, 0.8 or 3.2 mg Zn kg^-1 soil. After 3 weeks, plants derived from the high-Zn seed had better root and shoot growth; the cv. Excalibur accumulated more shoot dry matter than the cv. Gatcher. After 6 weeks, greater root and shoot growth of plants grown from the high-Zn seed compared to those from the low-Zn seed was obvious only at nil Zn fertilisation. A fertilisation rate of 0.2 mg Zn kg^-1 soil was required for achieving 90% of the maximum yield for plants grown from the high-Zn seed compared to 0.8 mg Zn kg^-1 soil for plants derived from the low Zn seed. The critical Zn level in youngest expanded leaves for 90% maximum yield was 16 mg Zn kg^-1 dry matter for both genotypes. Zn-efficient Excalibur had greater net Zn uptake rates compared to Zn-inefficient Gatcher after 3 weeks but they were not different at the 6-week harvest. Zinc-deficient plants had greater net uptake rates of Cu, Mn, B, P, and K but a reduced uptake rate of Fe. It is concluded that higher seed Zn content acted similar to a starter-fertiliser effect by improving vegetative growth and dissipating differences in Zn efficiency of wheat genotypes.     

Modelling potassium uptake by wheat (Triticum aestivum) crops

Spring wheat was grown in the field under deficient and sufficient levels of soil K and with high and low supplies of fertiliser nitrogen. Measurements were made of K uptake, soil nutrient supply parameters, root growth and, in solution culture, root influx parameters. Mechanistic models predicted uptake reasonably well under K-deficient conditions, but over-predicted uptake, by as much as 4 times, under K-sufficient conditions. The over-prediction was apparently due to poor characterisation of plant demand.     

Influence of minerals on cytoplasmic streaming in root hairs of intact wheat seedlings (Triticum aestivum L.)

The calcium dependency of the cytoplasmic streaming of wheat root hairs was demonstrated by adding the Ca-Ionophore A 23187. Within three minutes the streaming velocity was decreased dramactically. The influence of ammonium on the cytoplasmic streaming is highly pH-dependent. While at a pH of 9.0 an inhibitory effect was observed even at low ammonium concentrations (0.5 mM) no effect could be measured at a pH of 6.5. Nitrate, independently of medium pH had no effect on the cytoplasmic streaming. The same is true for aluminium. It is suggested that at pH 9 ammonium permiates the plasmalemma as NH₃. Due to higher cytoplasmic pH ( 7.5), NH₃ is protonated leading to an increase in cytoplasmic pH. Ammonium may displace sorbed calcium leading to an increase in the free cytoplasmic calcium responsible for the cessation of the streaming. Alternative explanations are discussed.Abbreviations HEPES N-2-Hydroxyethylpiperazine-N-2-ethanesulfonic acid     

Identification of new sources of aluminum resistance in wheat

Aluminum (Al) toxicity is a major constraint for wheat production in acidic soils. An Al resistance gene on chromosome 4DL that traces to Brazilian wheat has been extensively studied, and can provide partial protection from Al damage. To identify potentially new sources of Al resistance, 590 wheat accessions, including elite wheat breeding lines from the United States and other American and European countries, landraces and commercial cultivars from East Asia, and synthetic wheat lines from CIMMYT, Mexico, were screened for Al resistance by measuring relative root elongation in culture with a nutrient solution containing Al, and by staining Al-stressed root tips with hematoxylin. Eighty-eight wheat accessions demonstrated at least moderate resistance to Al toxicity. Those selected lines were subjected to analysis of microsatellite markers linked to an Al resistance gene on 4DL and a gene marker for the Al-activated malate transporter (ALMT1) locus. Many of the selected Al-resistant accessions from East Asia did not have the Al-resistant marker alleles of ALMT1, although they showed Al resistance similar to the US Al-resistant cultivar, Atlas 66. Most of the cultivars derived from Jagger and Atlas 66 have the Al-resistant marker alleles of ALMT1. Cluster analysis separated the selected Al-resistant germplasm into two major clusters, labeled as Asian and American–European clusters. Potentially new germplasm of Al resistance different from those derived from Brazil were identified. Further investigation of Al resistance in those new germplasms may reveal alternative Al-resistance mechanisms in wheat. Electronic supplementary material The online version of this article (doi:contains supplementary material, which is available to authorized users. Responsible Editor: Thomas B. Kinraide.     

Tolerance of wheat (<Emphasis Type="Italic">Triticum aestivum</Emphasis> L.) to high soil and solution selenium levels

The fertilisation of wheat crops with Se is a cost-effective method of enhancing the concentration of organic Se in grain, in order to increase the Se intake of animals and humans. It is important to avoid phytotoxicity due to over-application of Se. Studies of phytotoxicity of Se in wheat grown in Australia, where rainfall and grain yield are usually relatively low, have not been reported previously, and overseas studies have had varied results. This study used trials conducted in the field, glasshouse and laboratory to assess Se phytotoxicity in wheat. In field trials that used rates of up to 120 g ha^–1Se as selenate, and in pilot trials that used up to 500 g ha^–1 Se soil-applied or up to 330 g ha^–1 Se foliar-applied, with soils of low S concentrations (2–5 mg kg^–1), no Se toxicity symptoms were observed. In pot trials of four weeks duration, the critical tissue level for Se toxicity was around 325 mg kg^–1 DW, a level attained by addition to the growth medium of 2.6 mg kg^–1 Se as selenate. Solution concentrations above 10 mg L^–1 Se inhibited early root growth of wheat in laboratory studies, with greater inhibition by selenite than selenate. For selenite, Se concentrations around 70 mg L^–1 were required to inhibit germination, while for selenate germination % was unaffected by a solution concentration of 150 mg L^–1 Se. Leaf S concentration and content of wheat increased three-fold with the addition of 1 mg kg^–1 Se as selenate to the growth medium. This effect is probably due to the induction of the S deficiency response of the main sulphate transporter. This study found wheat to be more Se-tolerant than did earlier studies of tobacco, soybeans and rice. We conclude that Se phytotoxicity in wheat will not be observed at the range of Se application rates that would be used to increase grain Se for human consumption (4–200 g ha^–1 Se as selenate, which would result in soil and tissue levels well below those seen in the above studies), even when – as is common in Australia – soil S concentration and grain yield are low.     

WGA reduces the level of oxidative stress in wheat seedlings under salinity

Protective effect of exogenous wheat germ agglutinin (WGA) on wheat seedling (Triticum aestivum L.) during salinity stress was studied. In particular, we examined the state of pro- and antioxidant systems as well as the level of peroxide oxidation of lipids and electrolyte leakage under control conditions and when stressed with NaCl. Generation of superoxide anions and activity of both superoxide dismutase (SOD) and peroxidase increased during saline stress. Accumulation of O₂ ^·− resulted in peroxide oxidation of lipids and electrolyte leakage in response to stress. The injurious effect of salinity on root growth of seedlings was manifested by a decreased mitotic index (MI) in apical root meristem. This study show that WGA pretreatment decreased salt-induced superoxide anion generation, SOD and peroxidase activities, levels of lipid peroxidation and electrolytes leakage as well as correlating with a reduction in the inhibition of root apical meristem mitotic activity in salt-treated plants. This suggests that exogenous WGA reduced the detrimental effects of salinity-induced oxidative stress in wheat seedlings. Thus WGA effects on a balance of reactive oxygen species (ROS) and activities of antioxidant enzymes may provide an important contribution to a range of the defense reactions induced by this lectin in wheat plants.     

Nutrient accumulation and translocation in maturing wheat plants grown on waterlogged soil

Wheat plants (Triticum aestivum L., cv. Arina) growing in large pots (perforated at the bottom for controls, intact for flooding) were embedded in the field in spring. Waterlogging was initiated at anthesis and was maintained throughout the maturation period. Grain yield as well as potassium, phosphorus and magnesium contents in the shoot were decreased on flooded soil, while manganese and iron contents increased considerably. Total calcium and zinc contents per shoot remained comparable to those in controls. The reduction of potassium, phosphorus and magnesium contents by waterlogging was greatest in the grains, while manganese and iron accumulated mostly in the vegetative parts and the glumes. Zinc contents were also lowered in the grains during waterlogging due to an inhibited redistribution from the vegetative parts to the grains. Our results indicate that flooding caused not only an accumulation of manganese and iron in the shoot, but also affected the redistribution of macro- and micronutrients to the maturing gains.     

Identification of C-banded chromosomes in meiosis of common wheat,Triticum aestivum L.

Summary The C-banding pattern of nine meiotic chromosomes of common wheat (Triticum aestivum L.) as described. In F₁s of crosses between monosomics of Chinese Spring and two Spanish wheat cultivars, univalent chromosomes were used to aid the recognition and analysis of the C-banding pattern for the individual chromosomes. The identification of one chromosome involved in one translocation in Chinese Spring x Pané 247 has been made through heterochromatin bands observed in the chromosomes involved in multivalents.     

Genotype and fungicide effects on late-season root growth of winter wheat

The aim of this work was to investigate differences among genotypes in post-anthesis root growth and distribution of modern UK winter wheat cultivars, and the effects of fungicide applications. Post-anthesis root growth of up to six cultivars of winter wheat (Triticum aestivum L.), given either one or three applications of fungicide, was studied in field experiments during two seasons. Total root mass remained unchanged between GS63 (anthesis) and GS85, but root length increased significantly from 14.7 to 31.4 km m⁻² in one season. Overall, there was no evidence for a decline in either root mass or length during grain filling. Root mass as a proportion of total plant mass was about 0.05 at GS85. There were significant differences among cultivars in root length and mass especially below 30 cm. Malacca had the smallest root length and Savannah the largest, and Shamrock had a significantly larger root system below 40 cm in both seasons. Fungicide applied at ear emergence had no significant effect on root mass in either season but increased root length (P<0.01) in the more disease-prone season. By maintaining a green canopy for longer, fungicide applied at flag leaf emergence may have resulted in delayed senescence of the root system and contributed to the post-anthesis maintenance of root mass and length. Section Editor: R. W. Bell     

Interaction of salinity and enhanced ammonium and potassium nutrition in wheat

Wheat (Triticum aestivum L.) was grown to maturity in a pot experiment in a calcareous silty sand soil. N was applied at two levels as granulated N-P fertilizers, amended or not with nitrification inhibitors (1% and 5% DCD, 1% N-serve). Potassium as KCl was given at three levels of application. P was applied at a uniform rate. Two levels of salinity were obtained by using the soil as such (EC= 0.3 mmho/cm) and by adding NaCl to the same soil (EC=2.4 mmho/cm). 1% DCD and 1% N-serve treatments gave significantly higher wheat grain yields and N-uptake than the other ones. Nitrate content of leachates indicated a prevalent nitrate nutrition in the treatment without nitrification inhibitors. The 5% DCD treatment showed a yield depression. In the lower N level treatments, a significant yield increase, generated by 1% DCD and N-serve was found in the salinized soil as compared to the non-saline soil. Soil salinity reduced N-uptake when nitrification inhibitors were not present. In treatments having the inhibitors, N-uptake was equal or greater in the salinized than in the non saline soil. An enhanced ammonium nutrition increased the P uptake.     

Aluminium speciation in the presence of wheat root cell walls: a wet chemical study

Hydrolysis of Al³⁺ was performed in the presence of isolated root cell walls from a series of wheat cultivars (Triticum aestivum L.) known to have differential tolerance to Al contamination. Aluminium speciation was dependent on the cell wall concentration. At low cell wall concentrations, significant amounts of the very toxic Al₁₃ species were formed. At higher cell wall concentrations, formation of the tridecamer was hindered or completely inhibited. The sensitive wheat cultivars displayed a higher affinity for aluminium than the tolerant cultivars. A possible Al tolerance mechanism based on cell wall permeability is discussed.     

Cycling of amino-nitrogen between shoots and roots in wheat seedlings

Summary One part of a split root system of wheat seedlings received full nutrient solution with¹⁵N-nitrate, the other received an identical solution with unlabelled nitrate. Appearance of labelled amino compounds was measured in the xylem sap exuding from roots not supplied directly with¹⁵N-nitrate after removal of the¹⁵N-nitrate-fed roots. This material indicates cycling of nitrogen from the shoots and through the roots. About 60 per cent of the nitrogen in the xylem appears to be cycling in this way.