Influence of soil solution aluminum on root elongation of wheat seedlings

Wheat (Triticum aestivum L.) seedlings were grown for 4 days in an acid soil horizon treated with 10 levels each of Ca(OH)₂, CaSO₄ and CaCl₂. The treatments resulted in a wide range of Al levels and Al speciation in soil solution. Seedling root length in the Ca(OH)₂ treatments was significantly related (p<0.01) to calculated Al³⁺ activity in soil solution. The Al–SO₄ complex in soil solution had a negligible effect on the root growth of Hart wheat, thus confirming the previously reached conclusion concerning the nonphytotoxicity of Al–SO₄. The short-term seedling root growth technique used in this investigation allowed for separation of Al effects on root elongation from those on plant nutrition and should be useful for studying Al toxicity relationships in soil.     

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     

Effects on winter wheat seedling growth by toxin-producing rhizobacteria

Summary Root-colonizing pseudomonads capable of inhibiting seedling winter wheat (Triticum aestivum L.) root growth in an agar seedling bioassay also significantly inhibited wheat root growth in vermiculite; however, the inhibitory trait is quite labile in laboratory culturing. The extent of inhibition in both the agar and vermiculite medium depended on inoculum level. These pseudomonads were found to produce a toxin capable of inhibiting growth ofEscherichia coli C-la andBacillus subtilis. Field isolates that strongly inhibit growth of indicator bacteria also inhibited root growth. Toxin production by the bacteria appeared necessary for inhibition of root growth and indicator bacteria as toxin-negative (TOX^–) mutants no longer inhibited either. Antibiosis towardsE. coli as well as wheat seedling root inhibition in agar was reversed by L-methionine, providing further evidence that a toxin, produced by these organisms, is involved in growth retardation.Contribution in cooperation with the College of Agric. Res. Center, Washington State Univ., Pullman, WA 99164. Scientific Paper No. 6837.     

Growth and development of maxipak wheat as affected by soil salinity and moisture levels

The effect of soil salinity and soil moisture on the growth and yield of maxipak wheat (Triticum aestivum L.) was studied in a lath-house experiment in whih, chloride-sulphate salt mixtures were used to artificially salinize a sandy loam soil from Al-Jadyriah Baghdad. Five soil salinity levels of ECe's equal to 1.7 (Control) 4.2, 5.8, 8.1, 9.4 and 11.0dSm^–1 were prepared and used at 3 levels of available soil moisture depletion, namely, 25, 50, and 75% as determined by weight. Both growth (vegetative) and yield components were studied throughout the growing season.Results showed that increasing the soil salinity from 1.7 to 11.0 dSm^–1, and decreasing the available soil water from 75 to 25% resulted in independent and significant decreases in Mazipak wheat growth and yield components at different stages of plant development. Root growth showed more sensitivity to both available soil water and soil salinity level than other components. It has been concluded that at soil salinity levels of more than 8.0 dSm^–1, available soil water became a limiting factor on wheat growth and the maintenance of 75% of available soil water during the growth period is recommended to obtain satisfactory grain yield.     

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     

Prevention of aluminium toxicity with supplemental boron. I. Maintenance of root elongation and cellular structure

Aluminium toxicity is an important factor limiting plant growth mi acid soils. Symptoms of B deficiency and Al toxicity are very similar and generally associated with impaired membrane Function and root growth. Thus the objective of this study was to determine whether supplemental B prevents Al inhibition of root growth and development. Squash (Cucurbita pepo L. cv. Sunbar) was grown in hydroponic nutrient media with 44 mmol m^?3 free Al and B concentrations extending from 5 to 100 mmol m^?3. Our results establish that B protects against Al inhibition of root growth. Protection was apparent at all levels of organization examined: primary root and lateral root lengths; primary root cell elongation, cell production rate, tissue organization and cell structure; primary root morphology and maturation. Protection against Al inhibition was also apparent for shoot growth. These studies were undertaken in solution culture to limit the variables examined; however, the underlying motivation for this study is the problem of worldwide Al toxicity in soils. Therefore, the effect of adding additional B to a high-Al soil was also investigated and is the subject of the companion paper (Le Noble. Blevins & Miles 1996, Plant, Cell and Environment 19, 1143–1148).     

Preliminary results from a microscopic examination on the effects of aluminium on the root tips of wheat

Root tips from aluminium (Al) tolerant (Waalt) and Al sensitive (Warigal) wheat (Triticum aestivum (L). Thell.) cultivars exposed to low concentrations of Al (10 M) for 10, 24 and 72 hours were examined under the light and electron microscope. After fixing and embedding, longitudinal and transverse thin and ultrathin sections were cut. There was no evidence of Al damage to the root tips of the Al tolerant cultivar under both the light and electron microscope. For the Al sensitive cultivar, Al had no observable effect on the root tips 10 hours after Al addition when examined under the light microscope. When examined under an electron microscope, electron dense globular deposits were observed between the cell wall and cell membrane of the epidermal cells. There was not obvious damage to the cell cytoplasm. Two or 3 days after Al addition, light microscopy showed that the cells in the root tips had become swollen and extensively vacuolated. The tissues appeared disorganised and degenerate, particularly in the epidermis and outer cortical cells. The electron microscope also revealed a thickening of the cell wall. The cell wall was broken down, particularly in the epidermis in the region 4–6 mm from the root tip. The tissue in the meristematic area was largely intact.     

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.     

The contribution of different regions of the seminal roots of wheat to uptake of nitrate from soil

A technique was developed to determine the physiological activity of defined sections of seminal roots of wheat grown in sand. Wheat plants were grown for 2 weeks in narrow columns of N-deficient sand to which all other nutrients had been added. The columns were split longitudinally and ¹⁵N-labelled nitrate, in an agar medium, supplied to 2 cm sections of root. Shoots and roots were analysed after 24 h to determine the uptake of ¹⁵N. Three sections were examined on either the secondary or tertiary seminal root: 1 cm from the seed (basal segment), 35 cm from the seed (middle segment) and 4 cm from the root apex (apical segment). Total uptake was greatest from the basal and middle segments, declining by 50% from the apical segment. However, uptake per unit root length, including exposed sections of lateral roots, was not significantly different along the root.     

The cost of stress: Dry matter partitioning changes with seasonal supply of water and nitrogen to dryland wheat

Spring wheat cv. ‘Gutha’ was grown in continuous wheat (W/W) and narrow-leafed lupin (L. angustifolius L. cv. Yandee)-wheat (L/W) rotation on a yellow earth over mottled clay (Arenic Fragiudult) in a mediterranean climate for two years. The first year had a higher than average rainfall with adequate soil water until anthesis. The second year was very dry (only 232 mm total rainfall) and soil water contents were low throughout the growing season. Nitrogen fertilizer (+N) treatments were included in both years. In the first year an adjacent experiment compared the effects of loosening a pronounced traffic pan which existed on the site (LS)versus unloosened (US). In the first year roots contained more dry matter than tops in the early vegetative stage in all crops and then declined exponentially to a ratio of 0.1 in the US and LS treatments. In the second year however, the decline was both linear and much less, so that root:shoot ratios at harvest were still between 0.4 and 0.8. There was a consistent trend in root:shoot ratios from the most favourable (LS) to least favourable (W/W-N) treatments over the combined two years’ data, and this was also found in grain yield, with a higher yield in year one from the LS than US, and the lowest yield in year two from the W/W-N treatment. The proportion of total biomass recovered from below ground was substantially higher than is commonly reported from studies carried out in temperate, high fertility soils, but probably still under-estimates of the true amount of dry matter in roots because of inadequacies of sampling, washing and storage techniques. Root length densities were much greater in the drier year, especially in the surface 0.1-m, and based on theoretical considerations, much greater than required for extraction of available water. The effect of environmental conditions on the relative size of cereal crop carbon sinks are discussed in relation to these results.     

The effect of inoculation with Azospirillum brasilense on growth and nitrogen utilization by wheat plants

Azospirillium brasilense is a rhizosphere bacteria that has been reported to improve yield when inoculated on wheat plants. However, the mechanisms through which this effect is induced is still unclear. In the present work, we have studied the effects of inoculating a highly efficient A. brasilense strain on wheat plant grown in 5 kg pots with soil in a greenhouse, under three N regimes (0, 3 or 16 mM NO₃ ^–, 50 ml/pot once or twice-a -week), and in disinfected or non-disinfected soil. At the booting stage, the inoculated roots in both soils showed a similar colonization by Azospirillum sp. that was not affected by N addition. The plants grown in the disinfected soil showed a higher biomass, N content and N concentration than those in the non-disinfected soil, and in both soils the inoculation stimulated plant growth, N accumulation, and N and NO₃ ^– concentration in the tissues.At maturity, the inoculated plants showed a higher biomass, grain yield and N content than the uninoculated ones in both soils, and a higher grain protein concentration than the uninoculated. It is concluded that in the present experiments, A. brasilenseincreased plant growth by stimulating nitrogen uptake by the roots.     

Growth of peach as affected by decomposition of own root residues in soil

The effects of decomposing peach root residues in soil on peach growth were determined in two pot experiments. In the first, peach root residues, despite their high C:N ratio and lignin content, largely decomposed under experimental conditions, leading to an immobilization of inorganic N. Shoot growth of peach seedlings was depressed by the addition of peach root residues, an effect that depended on their size and concentration: fine-textured root fragments (0.45–1.00 mm) resulted in more severe effects than medium sized ones (2–8 mm), while growth depression occurred only at concentrations higher than 0.35%. Peach root growth was depressed by root residues regardless of their size and amount. In the second experiment, where nitrogen was added to all pots to minimize the effects of immobilization of N during decomposition of root fragments, the growth of peach roots in residue-supplemented soil almost stopped. Pre-planting phosphate enrichment was very effective in stimulating growth of peach in virgin soil but did not offset the depression caused by peach root residues. It thus appears that besides mineral deficiencies, there exist alternative explantations of poor growth of peach in replant soils, including growth-inhibiting substances from decomposing root residues.     

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.     

Consumption of wheat seed reserves during germination and early growth as affected by soil water potential

Seed size and weight are important criteria for determining seedling vigour and stand establishment. Evolution of seed dry weight of wheat (Triticum aestivum L.) during germination and early growth was examined because poor stands are often associated with the depletion and exhaustion of seed reserves. Two laboratory experiments were conducted on filter paper and in soil at three water potentials using wheat seeds. Seed, root, and shoot dry weights were recorded at approximately one-day intervals. Coleoptile and first leaf lengths were also measured at all sampling periods. Wheat seedlings grown on filter paper in the dark grew to a length of 90 to 100 mm with 50% of the initial seed weight remaining after eight days when the experiment was terminated. In soil, wheat seedlings grew 15 mm with 25% of the initial seed weight remaining. Seed reserves were depleted more quickly when the soil was wet because seedlings grew more quickly. There were significant and similar negative relationships between seed weight and coleoptile length of wheat seedlings grown on filter paper and in soil. There was no effect of soil water potential on the relationship between seed weight and shoot length. Therefore, it was concluded that poor wheat stands are not likely to occur due to depletion of seed reserves under field conditions without mechanical obstacles.     

The effect of parental divergence on F2 heterosis in winter wheat crosses

Summary In winter wheat (Triticum aestivum L.), the development of a methodology to estimate genetic divergence between parental lines, when combined with knowledge of parental performance, could be beneficial in the prediction of bulk progeny performance. The objective of this study was to relate F₂ heterosis for grain yield and its components in 116 crosses to two independent estimates of genetic divergence among 28 parental genotypes of diverse origins. Genetic divergence between parents was estimated from (a) pedigree relationships (coefficients of kinship) determined without experimentation, and (b) quantitative traits measured in two years of field experimentation in Kansas and North Carolina, USA. These distances, designated (1 -r) and G, respectively, provided ample differentiation among the parents. The 116 F₂ bulks were evaluated at four locations in Kansas and North Carolina in one year. Significant rank correlations of 0.46 (P = 0.01) and 0.44 (P = 0.01) were observed between G and grain yield and kernel number heterosis, respectively. Although (1 -r) was poorly associated with grain yield heterosis, G and midparent performance combined to account for 50% of the variation in F₂ yields among crosses when (1 -r) was above the median value, whereas they accounted for only 9% of the variation among crosses when (1-r) was below the median. Midparent and (1 -r) had equal effects on F₂ grain yield (R ²= 0.40) when G was greater than the median value. A breeding strategy is proposed whereby parents are first selected on the basis of performance per se and, subsequently, crosses are made between genetically divergent parents that have both large quantitative (G) and pedigree divergence (1 -r).Paper No. 12162 of the Journal Series of the North Carolina Agricultural Research Service, Raleigh, NC 27695-7643, and Contribution No. 89-396-J of the Kansas Agricultural Experiment Station, Manhattan, KS 66506     

Empirical models to approximate calcium and magnesium ameliorative effects and genetic differences in aluminium tolerance in wheat

The activities of inorganic, monomeric aluminium (Al) species in the root environment are important in the toxicity of Al to plant roots, which may be ameliorated by increased activities of basic cations. Additionally, it has been suggested that electro-chemical processes in walls of root cells play a role in Al tolerance. Empirical models were proposed to accomodate genetic and calcium (Ca) and magnesium (Mg) ameliorative effects on Al toxicity. The models were tested using data from a solution culture study (with ionic strength 1.6 to 8.6 mM) in which wheat (Triticum aestivum L.) cvv. Warigal (Al-sensitive) and Waalt (Al-tolerant) were grown for 28 d at 0, 10 and 20 M Al, in factorial combination with 200, 400, 800 and 1600 M Ca and 100, 200, 400 and 800 M Mg. There was a poor relationship between relative total dry mass (TDM) (calculated as a percentage of the average TDM of each cultivar in the absence of added Al) and the activity of Al³⁺ or the sum of the activities of the monomeric Al species in solution. A model based on the ratios of activities of cations in solution, taking valency into consideration, was more successful, accounting for ca 85% of the observed variation in relative TDM. There were no systematic variations between observed values and those estimated by the model.     

Interactive effects of Al, Ca and other cations on root elongation of rice cultivars under low pH

BACKGROUND AND AIMS: As with other crop species, Al tolerance in rice (Oryza sativa) is widely different among cultivars, and the mechanism for tolerance is unknown. The Ca2+-displacement hypothesis, that is, Al displaces Ca2+ from critical sites in the root apoplast, was predicted to be the essential mechanism for causing Al toxicity in rice cultivars. If displacement of Ca is an essential cause of Al toxicity in rice, Al toxicity may show the same trend as toxicities of elements such as Sr and Ba that are effective in displacing Ca. METHODS: The interactive effects of Al, Ca, Sr and Ba on root elongation of rice cultivars with different Al tolerances were evaluated in hydroponic culture. Al and Ca accumulation in root tips was also investigated. KEY RESULTS AND CONCLUSIONS: Not only Al but also Sr and Ba applications inhibited root growth of rice cultivars under low Ca conditions. As expected, rice cultivars more tolerant of Sr and Ba were also tolerant of Al (japonica > indica). Although Mg application did not affect Sr or Ba toxicity, Mg alleviated Al toxicity to the same level as Ca application. In addition, Ca application decreased the Al content in root tips without displacement. These results suggest that Ca does not have a specific, irreplaceable role in Al toxicity, unlike Sr and Ba toxicities. Alleviation of Al toxicity with increasing concentrations of Ca in rice cultivars is due to increased ionic strength, not due to decreased Al activity. The difference in Al tolerance between indica and japonica cultivars disappears under high ionic strength conditions, suggesting that different electrochemical characteristics of root-tip cells are related to the significant difference in Al tolerance under low ionic strength conditions.     

Effect of phosphorus supply to the surface roots of wheat on root extension and rhizosphere chemistry in an acidic subsoil

Seedlings of two cultivars of wheat (Triticum aestivum L.) differing in tolerance to aluminium (Al) were grown using a split-root sand/soil culture technique. Each culture tube was divided horizontally into a surface (0–150 mm) compartment and a subsurface (150–250 mm) compartment separated by a root-permeable paraffin wax barrier. Thus phosphorus (P) supplied to surface roots could not percolate or diffuse into the soil in the subsurface compartment. The soil in the subsurface compartment was divided into ‘rhizosphere’ and ‘non-rhizosphere’ zones using a porous (5 μm) membrane. Root growth of both cultivars into the subsurface zone was enhanced by increased P supply to surface roots, but did not conform to known relationships between root growth and soil pH, extractable-Al, or pH, Al or P concentrations in soil solution. Concentrations of Al in soil solution in the rhizosphere were greater than those in solution in the bulk soil. Concentrations of Al reactive with pyrocatechol violet (30s-RRAI) in the rhizosphere soil solution were generally greater than those in non-rhizosphere soil. With the Al-sensitive cultivar, root dry weight and length increased as concentrations of RRAl in the rhizosphere soil solution increased. Increased concentrations of Al in rhizosphere soil solutions were not related to the presence of organic ligands in solution. The effect of P in promoting root penetration into the acidic subsurface stratum was not related to differential attainment of maturity by the plant shoots, but appeared to be related to the effect of P in enhancing the rate of root growth. Thus, suboptimal supply of P to the surface roots of a plant, even at levels sufficient to preclude development of nutritional (P) stress symptoms, may seriously reduce tolerance to Al, and hence diminish the ability of roots to penetrate into acidic subsoils.     

Effect of soil salinity and water stresses on growth and content of nitrogen,chloride and phosphate of wheat and triticale

Summary Three wheats and one triticale were grown, up to flowering stage, in pots on calcareous soil adjusted to a range of salinities (S₁=3.5, S₂=6, S₃=8.5, and S₄=11 mmhos/cm, 20°C, soilpaste extract) by adding solution consisting of 3∶2∶1 of Na-, Ca- and Mg chlorides in chemical equivalent amounts. Moisture in the pots was kept at 100% (W₁), 40% (W₂) and 20% (W₃) of the available water. The vegetative growth, nitrogen and phosphate were affected by S and W treatments, chloride was affected only by S. The interaction S×W affected only dry weight. Varietal effect was observed between wheat as a group and triticale. Multiple quadratic regression equations of these properties on salinity and water revealed that the higher the available water the wider the range of tolerable salinity. Triticale was relatively more tolerant to water stress. Salinity increases Cl and decreases N, whereas water stress enhances N accumulation to a certain extent. However, in triticale at S₃ and S₄ the effect of water stress on N was overshadowed by the excessive salinity. This did not occur for the wheat (Florence). P trends were described. R² for P was low (0.7435–0.3603) which made interpretations rather difficult.     

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.