Natural suppression of take-all disease of wheat in Montana soils

This research was initiated to determine whether soils suppressive to take-all of wheat caused by Gaeumannomyces graminis var. tritici (Ggt) occur in Montana, and to identify the organisms most likely involved in this suppression. From an initial screening of eight soils collected from different wheat growing areas of Montana, two were highly suppressive to take-all. Microbial characterization of these soils indicated that different mechanisms were involved in the suppression. In Larslan soil, mycoparasitism appeared to be the main mechanism. Two different fungi with exceptional ability to reduce the severity of take-all were isolated from this soil. One of these fungi could parasitize the hyphae of Ggt. Field tests with these fungi in Ggt infested soil showed increases of over 100% in both harvestble tillers and grain yield as compared to treatments without these two fungi. In tests with 48 different bacteria and 10 actinomycetes from Larslan soil, none were able to consistently reduce severity of take-all alone, or in mixtures. In Toston soil, antibiosis by actinomycetes and perhaps the involvement of Pseudomonas spp. in production of antibiotics and/or siderophores appeared to be the most likely mechanisms involved in take-all suppression. Increases in shoot dry weight over that in the Ggt infested control using mixtures of pseudomonads and actinomycetes ranged from 25% to 87%. Actinomycetes added individually or in mixtures to soil infested with Ggt consistently reduced the severity of the disease to a greater extent than did mixtures of Pseudomonas spp.     

The effect of take-all disease on gas-exchange rates and biomass in two winter wheat lines with different drought response

There have been no studies of the effect of take-all on leaf gas-exchange rates, despite the fact that take-all severely restricts plant water and nutrient uptake, which results in significant biomass and grain yield reduction. Here we describe the effect of inoculation with Gaeumannomyces graminis (Sacc.) var. tritici (Ggt) on carbon assimilation rate (A) and biomass production of wheat plants grown under two water regimes. We show that the impact of Ggt inoculation on plant growth and leaf A may be through reduced photosynthetic capacity of the leaves and not water stress per se. The nature of this reduced photosynthetic capacity remains uncertain but may involve nutrient deficiency and different enzymes produced by the fungus. In each of the 3 years the experiment was conducted, Ggt significantly reduced A, i.e. at anthesis by 18% in 2000, 15% in 2001, and 12% in 2002. In agreement with other field studies, Ggt reduced tiller number and production of all plant components, mostly root dry mass and grain mass per plant. Highly significant negative correlations were found between disease rating and A in all years, showing that at disease ratings equal or higher than 3 (on a scale from 1 to 4) A could practically be zero. While A decreased, intercellular CO₂ concentration increased or did not change, and stomatal conductance was relatively high. In addition, A was more reduced under high than under low soil moisture content. These results support the idea that water stress per se did not contribute to the observed reduction of A. The mechanism of photosynthetic capacity reduction due to the Ggt root-rotting fungus is of interest as it may lead to the molecular mechanisms of plant resistance and ultimately to the development of take-all resistant plants.     

Plant genotype, micronutrient fertilization and take-all infection influence bacterial populations in the rhizosphere of wheat

The relationship between micronutrient efficiency of four wheat (Triticum aestivum L.) genotypes, tolerance to take-all disease (caused by Gaeumannomyces graminis (Sacc.) Arx and Olivier var. tritici Walker), and bacterial populations in the rhizosphere was tested in soil fertilized differentially with Zn and Mn. Plant growth was reduced by Mn or Zn deficiency and also by take-all. There was an inverse relationship between micronutrient efficiency of wheat genotypes when grown in deficient soils and the length of take-all lesions on roots (efficient genotypes had shorter lesions than inefficient ones). In comparison to the rhizosphere of control plants of genotypes Aroona and C8MM receiving sufficient Mn and Zn, the total numbers of bacterial cfu (colony forming units) were greater in the rhizosphere of Zn-efficient genotype Aroona under Zn deficiency and in Mn-efficient genotype C8MM under Mn deficiency. These effects were not observed in other genotypes. Take-all decreased the number of bacterial cfu in the rhizosphere of fully-fertilized plants but not of those subjected to either Mn or Zn deficiency. In contrast, the Zn deficiency treatment acted synergistically with take-all to increase the number of fluorescent pseudomonads in the rhizosphere. Although numbers of Mn-oxidising and Mn-reducing bacteria were generally low, take-all disease increased the number of Mn reducers in the rhizosphere of Mn-efficient genotypes Aroona and C8MM. Under Mn-deficiency conditions, the number of Mn reducers in the rhizosphere increased in Aroona but not in C8MM wheat. The results suggest that bacterial microflora may play a role in the expression of Mn and Zn efficiency and tolerance to take-all in some wheat genotypes.     

A role for managanese in the resistance of wheat plants to take-all

Summary The hypothesis that wheat plants deficient in managenese are predisposed to infection byGaeumannomyces graminis is outlined, and a test of the hypothesis in a soil system is reported. The results supported the hypothesis: wheat plants growing in managanese-deficient soil, although not showing foliar symptoms, were markedly more susceptible to infection; plant analysis confirmed the nutrient status of the plants. A review of the literature on take-all in wheat coupled with the results of our experiments suggests a reinterpretation of the etiology of this disease, since those edaphic factors which promote infection by this organism are those which also render managese unavailable to the host. Managenese nutrition is proposed as a common factor in many of the environmental conditions which influence the host-pathogen balance.     

Intermittent wetting of soils at high temperature reduces survival of the take-all fungus

Two pot experiments using naturally infested soil and a range of watering regimes were conducted to study the possible effect of level and frequency of wetting of hot soil (to simulate the period between growing seasons in Western Australia) on inoculum of the take-all fungus (Gaeumannomyces graminis var.tritici). In combination with the high soil temperatures, all watering regimes reduced infectivity and propagule number of the take-all fungus, this reduction being absent in dry soils.     

Control of Mn status and infection rate by genotype of both host and pathogen in the wheat take-all interaction

Take-all is a world-wide root-rotting disease of cereals. The causal organism of take-all of wheat is the soil-borne fungus Gaeumannomyces graminis var tritici (Ggt). No resistance to take-all, worthy of inclusion in a plant breeding programme, has been discovered in wheat but the severity of take-all is increased in host plants whose tissues are deficient for manganese (Mn). Take-all of wheat will be decreased by all techniques which lift Mn concentrations in shoots and roots of Mn-deficient hosts to adequate levels. Wheat seedlings were grown in a Mn-deficient calcareous sand in small pots and inoculated with four field isolates of Ggt. Infection by three virulent isolates was increased under conditions which were Mn deficient for the wheat host but infection by a weakly virulent isolate, already low, was further decreased. Only the three virulent isolates caused visible oxidation of Mn in vitro. The sensitivity of Ggt isolates to manganous ions in vitro did not explain the extent of infection they caused on wheat hosts. In a similar experiment four Australian wheat genotypes were grown in the same Mn-deficient calcareous sand and inoculated with one virulent isolate of Ggt. Two genotypes were inefficient at taking up manganese and were very susceptible to take-all, one was very efficient at taking up manganese and was resistant to take-all, and the fourth genotype was intermediate for both characters. All genotypes were equally resistant under Mn-adequate conditions.     

Effect of certain herbicide treatments on pasture composition and inoculum of the take-all fungus

The improvement of pastures by the use of a range of herbicides to eliminate grasses, and their effect on populations of the take-all fungus (Gaeumannomyces graminis vartritici=Ggt) were studied in the field (at Esperance Downs, on the south-coast of Western Australia) from 1982 to 1985. Field trials were conducted to evaluate three herbicide treatments (2,4-D amine+propyzamide; 2,4-D amine+paraquat; paraquat/ diquat) and an unsprayed control. A pot trial involving these treatments with two levels of nitrogen was undertaken to confirm treatment effects observed in the field trial. All herbicide treatments resulted in reduced grass composition of pastures, in both the year of spraying and in the second year of pasture, but reduced dry matter production in the year of spraying. In the year of spraying, however, inoculum ofGgt was reduced (P<0.1) only following the 2,4-D amine+propyzamide treatment and was greater (P<0.1) after 2,4-D amine+paraquat treatment than the unsprayed treatment. Despite reduced grass levels in the herbicide-treated plots in the second year of pasture,Ggt inoculum did not differ between treatments, nor did it after a wheat crop which followed a second year pasture. There was high correlation (P<0.001) between disease levels and dry weights of grasses in the pot trial. There was significantly less (P<0.001) grass in pots treated with herbicides compared to the unsprayed control but no difference (P>0.05) was evident between treatments. Inoculum levels were lower (P<0.05) in the treated pots than the unsprayed control with no evidence of differences among treatments (P>0.05). Nitrogen level had no effect on disease (P>0.05). All herbicide treatments tested reduced grass level and total dry matter, both in the field and in pots. Whereas in the pot trial reduced grass levels resulted in reducedGgt inoculum, in the field such a reduction occurred only with the 2,4-D amine+propyzamide treatment and only in the year of spraying. Herbicide treatments had no effect onGgt inoculum in second year of pasture or crop. Unknown soil and environmental factors in the field precluded a simple relationship between grass level in pasture and subsequent level ofGgt inoculum, and where such a relationship did occur (2,4-D amine+propyzamide treatment) it appeared to be shortlived.     

Manganese nutrition and accumulation of phenolics and lignin as related to differential resistance of wheat genotypes to the take-all fungus

Differential resistance of four Triticum aestivum L. genotypes to isolates of take-all fungus (Gaeuman-nomyces graminis var. ritici Walker) was tested in a complete factorial experiment set up in a growth chamber using Mn-deficient Wangary sand amended with four rates of Mn. Mn-efficient cultivars produced more dry matter at low supply of Mn. Fertilization with Mn significantly increased its accumulation in roots and shoots. The most sensitive measure of take-all infection was the total length of root stellar lesions; these lesions were reduced by Mn fertilization and were shorter in Mn-efficient genotypes. The resistance-enhancing effect of Mn was the most obvious in the Mn-inefficient genotype (Bayonet) and the least obvious in the Mn-efficient one (C8MM). Phenolics biosynthesis in roots was clicited by fungal infection, especially in the case of the highly virulent isolate. The weakly virulent isolate increased phenolics concentration in roots much more if no Mn was added, indicating that the resistance-enhancing effect of Mn may not be directly exerted through the effects on phenolics biosynthesis. Lignin concentration in roots decreased due to Mn fertilization, while no effect of take-all infection was noted. It appears that biosynthesis of phenolics and lignin in wheat roots has a low Mn requirement which can be satisfied at environmental Mn concentrations below those necessary for optimum plant growth. ei]Section editor: A C Borstlap ei]Section editor: H Lambers     

Soil conduciveness to take-all of wheat: Influence of the nitrogen fertilizers on the structure of populations of fluorescent pseudomonads

In a field cropped with wheat, a high and low level of soil conduciveness to take-all were induced by applying a nitrogen fertilizer with either calcium nitrate or ammonium sulphate. From these two soils, two representative populations of fluorescent pseudomonads were tested for their in situ behaviour. Take-all index and root dry weight were assessed on plants cropped in soils infested with Gaeumannomyces graminis var tritici (Ggt) and each bacterized with one of the isolates of fluorescent pseudomonads. The bacteria tested can be split into three groups: antagonists which reduce take-all, deleterious isolates which aggravate the disease and neutral without evident effect on the disease. The predominance of antagonistic fluorescent pseudomonads in the NH₄-treated soil and the predominance of deleterious ones in the NO₃-treated soil was confirmed after statistical analysis. The microbial impact on take-all must be more considered as the resulting effect of divergent activities of both rhizobacteria types than the only consequences of the presence of antagonistic pseudomonads. All the high cyanogenic pseudomonads were antagonists in situ and were more numerous in the NH₄-treated soil than in the NO₃-treated soil.     

Relationships between take-all,soil conduciveness to the disease,populations of fluorescent pseudomonads and nitrogen fertilizers

Take-all of wheat, caused by Gaeumannomyces graminis var tritici (Ggt), is reduced by ammoniacal fertilizers as compared to nitrate sources. This influence of nitrogen on the disease is only observed on nodal roots at flowering. But soil conduciveness to take-all, as measured in a soil bioassay, is modified earlier. Forty days after nitrogen application at early tillering, the NH₄-treated soil became less conducive than the NO₃-treated one. When nitrogen applications are done at sowing and at tillering, differences in disease propagation between the two soils are enhanced. Results from four years of experimentation show that when the level of natural soil inoculum is high, disease severity is reduced by ammonium, showing an effect on the parasitic phase of Ggt. At a low level of natural inoculum the effect of the source of nitrogen is mainly observed on the percent of infected plants, indicating that the saprophytic and preparasitic phases are affected. Rhizospheric bacterial populations increase from sowing to tillering, but differences on take-all conduciveness after tillering are not correlated with differences in the amounts of aerobic bacteria or fluorescent pseudomonads isolated from soils treated with different sources of nitrogen. Qualitative changes in fluorescent Pseudomonas spp. populations, like in vitro antagonism, are more likely to explain differences in soil conduciveness to take-all than are quantitative changes in this group. Nevertheless, the introduction of Ggt in a cropped soil leads to a greater increase in fluorescent pseudomonads populations than in total aerobic bacteria.The delay between reducing soil conduciveness and reducing disease in the field with ammonium nitrogen fertilization, the qualitative change of fluorescent pseudomonads populations and the role of necroses in rhizobacteria multiplication, provide information leading to our representation of a dynamic model based on the differentiation of the wheat root system into seminal and nodal roots.     

Inheritance of partial resistance to powdery mildew in spring wheat

Summary Four spring wheat (Triticum aestivum L.) cultivars exhibiting partial resistance to powdery mildew induced by Erysiphe graminis f.sp. tritici were crossed to a common susceptible cultivar to study the inheritance of resistance. The genetic parameters contributing to resistance were estimated by generation means analyses. Additive gene action was the most important genetic component of variation among generation means in all four crosses. Additive by additive effects were significant in one cross and both additive by additive and additive by dominance effects were significant in another. Dominance effects were not significant. The F2/F3 correlations in three crosses ranged from 0.27 to 0.43. Three additional crosses among resistant cultivars were employed to study the effectiveness of selection in improving resistance. By selecting the most resistant plants from the F2 and evaluating the progenies in the F4, increases in resistance ranging from 21% to 31% were obtained. In all crosses, there was transgressive segregation in both directions indicating that the genes conferring resistance to these cultivars differ and exhibit additive effects.     

Biological control of take-all in wheat by endophytic Bacillus subtilis E1R-j and potential mode of action

The bacterial strain E1R-j, isolated as an endophyte from wheat roots, exhibited high antifungal activity to Gaeumannomyces graminis var. tritici (Ggt). Strain E1R-j was identified as Bacillus subtilis based on morphological, physiological and biochemical methods as well as on 16S rDNA analysis. This strain inhibited mycelium growth in vitro of numerous plant pathogenic fungi, especially of Ggt, Coniothyrium diplodiella, Phomopsis sp. and Sclerotinia sclerotiorum. In greenhouse experiments, soil drenches with cell densities of 10⁶, 10⁹ and 10¹² CFU ml⁻¹ E1R-j reduced significantly take-all disease, caused by Ggt, in wheat seedling by 62.6%, 68.6% and 70.7%, respectively, compared to the inoculated control, 4 weeks after sowing. Growth parameters such as lengths and fresh weights of roots and shoots of Ggt-inoculated control plants were significantly lower compared to Ggt-inoculated and E1R-j treated plants. Field experiments in the season 2006/2007, heights of wheat plants in the Ggt inoculated plots were significantly reduced compared to the non inoculated treatments. Yield parameters such as kernels per head and thousand kernel weight (TKW) in inoculated control plants were lower compared to the other treatments. In the experimental year 2007/2008, independent treatments with the bacterial strain E1R-j and the fungicide Triadimefon reduced take-all disease in wheat roots by 55.3% and 61.9%, compared to the inoculated control plants. In this season plant height in inoculated control was significantly lower and also the yield parameters seeds per head and especially TKW were drastically reduced compared to the other treatments. E1R-j treatment alleviated the detrimental effects of take-all on grain yield parameters to a similar extent as Triadimefon application. SEM studies revealed that in the presence of E1R-j, hyphae of Ggt showed leakage, appeared ruptured, swollen and shriveled. Following root drench, strain E1R-j was able to colonize endophytically roots and leaves of wheat seedlings. While the population of the bacterial strain in wheat roots steadily increased from the second to the fourth leaf stage, in the leaf tissue the population of the strain rapidly declined. TEM studies also showed that cells of E1R-j were present in roots of wheat seedlings and effectively retarded infection and colonization of Ggt in root tissue; suppression of Ggt by E1R-j was accompanied by disintegration of hyphal cytoplasm. In addition, in the presence of E1R-j cells in Ggt-infected root tissue morphological defense reactions were triggered such as formation of wall appositions and papillae. The results presented indicate that the endophytic strain E1R-j of B. subtilis meets demands required for biocontrol of take-all.     

Cadmium uptake and bioaccumulation in selected cultivars of durum wheat and flax as affected by soil type

Accumulation of cadmium (Cd) in crop plants is of great concern due to the potential for food chain contamination through the soil-root interface. Although Cd uptake varies considerably with plant species, the processes which determine the accumulation of Cd in plant tissues are affected by soil factors. The influence of soil type on Cd uptake by durum wheat (Triticum turgidum var. durum L.) and flax (Linum usitatissimum L.) was studied in a pot experiment under environmentally controlled growth chamber conditions. Four cultivars/lines of durum wheat (Kyle, Sceptre, DT 627, and DT 637) and three cultivars/lines of flax (Flanders, AC Emerson, and YSED 2) were grown in two Saskatchewan soils: an Orthic Gray Luvisol (low background Cd concentration; total/ABDTPA extractable Cd: 0.12/0.03 mg kg^-1, respectively) and a Dark Brown Chernozem (relatively high background Cd concentration; total/ABDTPA Cd: 0.34/0.17 mg kg^-1 respectively). Plant roots, stems, newly developed heads, and grain/seeds were analyzed for Cd concentration at three stages of plant growth: two and seven weeks after germination, and at plant maturity. The results showed that Cd bioaccumulation and distribution within the plants were strongly affected by both soil type and plant cultivar/line. The Cd concentration in roots leaves and stems varied at different stages of plant growth. However, all cultivars of both plant species grown in the Chernozemic soil accumulated more Cd in grain/seeds than plants grown in the Orthic Gray Luvisol soil. The different Cd accumulation pattern also corresponded to the levels of ABDTPA extractable and metal-organic complex bound soil Cd found in both soils. Large differences were found in grain Cd among the durum wheat cultivars grown in the same soil type, suggesting the importance of rhizosphere processes in Cd bioaccumulation and/or Cd transport processes within the plant. Distribution of Cd in parts of mature plants showed that durum grain contained up to 21 and 36% of the total amount of Cd taken up by the plants for the Orthic Gray Luvisol and Chernozemic soils, respectively. These results indicate the importance of studying Cd speciation, bioaccumulation and cycling in the environment for the management of agricultural soils and crops.     

Selenium uptake by plants as a function of soil type,organic matter content and pH

In pots containing sandy soils at two levels (pH 5 and 7) to which 0.5 mg Se L^-1 soil had been added, an increase in the proportion of clay soil or peat soil led to a decrease in the uptake of Se by spring wheat grain (Triticum aestivum L., var. Drabant) and winter rape plants (Brassica napus L., var. Emil). The effect was most pronounced for the smallest additions of clay and peat soils. Differences in Se uptake between the two pH levels were greatest in treatments where the additions of clay and peat soils were small. At the high pH, an increase in clay content from 7% to 39% resulted in a decrease in Se uptake of 79% for wheat and 70% for rape. At the low pH, the uptake decreased by 72% and 77%, respectively. At the higher pH, an increase in the content of organic matter from 1.4% to 39% resulted in decreases in Se uptake of 88% for wheat grain and 69% for rape. At the low pH, Se uptake decreased by 63% and 48%, respectively. Adding peat soil to clay soil had little effect on Se uptake. Among the limed, unmixed clay, sand and peat soils to which Se had not been added, uptake was highest from the sandy soil, i.e. 8.3 ng Se/g wheat grain and 42 ng Se/g rape. The lowest uptake rates were obtained in the clay soil, i.e. 3.0 ng Se/g for wheat grain and 9.0 ng Se/g for rape.     

Evaluation of populations of fluorescent pseudomonads related to decline of take-all patch on turfgrass

Take-all on turfgrass caused by Gaeumannomyces graminis var. avenae (Gga) occurs as patches of yellowish plants. On some patches the central zone was recolonized by the same grass species, Festuca sp., previously damaged by the fungus despite the centrifugal extension of the disease. This disease remission was assimilated to decline. Rhizosphere bacterial counts showed that total population of bacteria was nearly the same in all zones across the patches. However, the ratio of fluorescent Pseudomonas spp./ total bacteria was 1/22, 1/15.4, 1/3.5 and 1/2.9 in the disease free area, the front margin of the patch, in the damaged part of the patch, and in the recolonized central part respectively. Furthermore, in this last mentioned zone, 44 to 82% of the fluorescent Pseudomonas spp. were antagonistic in vitro to Gga, whereas only 12 to 34% from the disease free area were antagonistic. So the development of take-all on turf induced quantitative and qualitative changes in populations of fluorescent pseudomonads. The remission of the disease in the center was correlated to higher amount of antagonistic fluorescent pseudomonads in this part of the patches. This typical patch with the well defined zones can provide a good model for the study of changes in bacterial populations related to the build up of take-all decline.     

Role of antibiotics and siderophores in biocontrol of take-all disease of wheat

Both antibiotics and siderophores have been implicated in the control of soilborne plant pathogens by fluorescent pseudomonads. In Pseudomonas fluorescens 2–79, which suppresses take-all of wheat, the importance of the antibiotic phenazine-1-carboxylic acid was established with mutants deficient or complemented for antiobiotic production and by isolation of the antibiotic from the roots of wheat colonized by the bacteria. Genetic and biochemical studies of phenazine synthesis have focused on two loci; the first is involved in production of both anthranilic acid and phenazine-1-carboxylic acid, and the second encodes genes involved directly in phenazine synthesis. Because the antibiotic does not account fully for the suppressiveness of strain 2-79, additional mutants were analyzed to evaluate the role of the fluorescent siderophore and of an antifungal factor (Aff, identified as anthranilic acid) that accumulates when iron is limiting. Whereas strains producing only the siderophore conferred little protection against take-all, Aff⁺ strains were suppressive, but much less so than phenazine-producing strains. Iron-regulated nonsiderophore antibiotics may be produced by fluorescent pseudomonads more frequently than previously recognized, and could be partly responsible for beneficial effects that were attributed in the past to fluorescent siderophores.     

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.     

Variability among strains of Trichoderma harzianum in their ability to reduce take-all and to produce pyrones

Antagonism tests on agar-plates and glasshouse screening indicated that three isolates of Trichoderma harzianum varied in their ability to antagonize the take-all fungus (Gaeumannomyces graminis var. tritici). Isolate 71 which was the most effective in suppressing take-all of wheat, produced two pyrones and other undetermined analogues. Isolates of T. koningii and T. hamatum shown to suppress take-all, produced a simple pyrone compound. Although T. harzianum isolates 70 and 73 did not produce any pyrones, they reduced the disease albeit to a much lesser extent than isolate 71; with isolate 73 showing distinct host growth promotion effects. It is proposed that the success of isolate 71 of T. harzianum was related to the pyrones it produces and that the ability of isolates 70 and 73 to reduce take-all may be related to mechanisms other than those involving antibiotics.     

Salicylic acid induced physiological and biochemical changes in wheat seedlings under water stress

Salicylic acid (SA) is an important signal molecule modulating plantresponses to stress. It is recently reported to induce multiple stresstolerancein plants including drought. An experiment was, therefore, conducted toascertain the effect of salicylic acid on the growth and metabolic profile ofwheat seedlings under water stress. Irrespective of the SA concentration(1–3 mM) and water stress, SA treated plants showed, ingeneral, a higher moisture content, dry mass, carboxylase activity of Rubisco,superoxide dismutase (SOD) activity and total chlorophyll compared to those ofuntreated seedlings. SA treatment, under water stress, protected nitratereductase (NR) activity and maintained, especially at 3 mM SAconcentration, the protein and nitrogen content of leaves compared to watersufficient seedlings. Results signify the role of SA in regulating the droughtresponse of plants and suggest that SA could be used as a potential growthregulator, for improving plant growth under water stress.     

Effects of variations in soil water potential,depth of N placement,and cultivar on postanthesis N uptake by wheat

This study was conducted to investigate the influence of soil water potential, depth of N placement, timing, and cultivar on uptake of a small dose of labeled N applied after anthesis by wheat (Triticum aestivum L.) Understanding postanthesis N accumulation should allow better control of grain protein concentration through proper manipulation of inputs. Two hard, red spring-wheat cultivars were planted in early and late fall each yr of a 2-yr field experiment. Less than 1 kg N ha^–1 as K ¹⁵NO₃ was injected into the soil at two depths: shallow (0.05 to 0.08 m) and deep (0.15 to 0.18 m). In both years an irrigation was applied at anthesis, and injections of labeled N were timed 4, 12, and 20 days after anthesis (DAA). Soil water potential was estimated at the time of injection. Mean recovery of ¹⁵N in grain and straw was 57% of the ¹⁵N applied. Recovery did not differ between the high-protein (Yecora Rojo) and the low-protein (Anza or Yolo) cultivars. Mean recovery from deep placement was 60% versus only 54% from shallow placement (p < 0.01). Delaying the time of injection decreased mean recovery significantly from 58% at 4 DAA to 54% at 20 DAA. This decrease was most pronounced in the shallow placement, where soil drying was most severe. Regressions of recovery on soil water potential of individual cultivar x yr x planting x depth treatments were significant only under the driest conditions. Stepwise regression of ¹⁵N recovery on soil water potential and yield parameters using data from all treatments of both years resulted in an equation including soil water potential and N yield, with a multiple correlation coefficient of 0.64. The translocation of ¹⁵N to grain was higher (0.89) than the nitrogen harvest index (0.69), and showed a highly significant increase with increase in DAA. This experiment indicates that the N uptake capacity of wheat remains reasonably constant between 4 and 20 DAA unless soil drying is severe.