G. K. McDonald

Additive effects of Na+ and Cl– ions on barley growth under salinity stress

Soil salinity affects large areas of the worlds cultivated land, causing significant reductions in crop yield. Despite the fact that most plants accumulate both sodium (Na+) and chloride (Cl–) ions in high concentrations in their shoot tissues when grown in saline soils, most research on salt tolerance in annual plants has focused on the toxic effects of Na+ accumulation. It has previously been suggested that Cl– toxicity may also be an important cause of growth reduction in barley plants. Here, the extent to which specific ion toxicities of Na+ and Cl– reduce the growth of barley grown in saline soils is shown under varying salinity treatments using four barley genotypes differing in their salt tolerance in solution and soil-based systems. High Na+, Cl–, and NaCl separately reduced the growth of barley, however, the reductions in growth and photosynthesis were greatest under NaCl stress and were mainly additive of the effects of Na+ and Cl– stress. The results demonstrated that Na+ and Cl– exclusion among barley genotypes are independent mechanisms and different genotypes expressed different combinations of the two mechanisms. High concentrations of Na+ reduced K+ and Ca2+ uptake and reduced photosynthesis mainly by reducing stomatal conductance. By comparison, high Cl– concentration reduced photosynthetic capacity due to non-stomatal effects: there was chlorophyll degradation, and a reduction in the actual quantum yield of PSII electron transport which was associated with both photochemical quenching and the efficiency of excitation energy capture. The results also showed that there are fundamental differences in salinity responses between soil and solution culture, and that the importance of the different mechanisms of salt damage varies according to the system under which the plants were grown.

Drought-adaptive attributes in the Seri/Babax hexaploid wheat population

Agronomic and physiological traits associated with drought adaptation were assessed within the Seri/Babax recombinant inbred line population, which was derived from parents similar in height and maturity but divergent in their sensitivity to drought. Field trials under different water regimes were conducted over 3 years in Mexico and under rainfed conditions in Australia. Under drought, canopy temperature (CT) was the single-most drought-adaptive trait contributing to a higher performance (r2 = 0.74, P < 0.0001), highly heritable (h2 = 0.65, P < 0.0001) and consistently associated with yield phenotypically (r = -0.75, P < 0.0001) and genetically [R (g) = -0.95, P < 0.0001]. CT epitomises a mechanism of dehydration avoidance expressed throughout the cycle and across latitudes, which can be utilised as a selection criteria to identify high-yielding wheat genotypes or as an important predictor of yield performance under drought. Early response under drought, suggested by a high association of CT with estimates of biomass at booting (r = -0.44, P < 0.0001), leaf chlorophyll (r = -0.22 P < 0.0001) and plant height (r = -0.64, P < 0.0001), contrast with the small relationships with anthesis and maturity (averaged, r = -0.10, P < 0.0001), and with osmotic potential (r = -0.20, P < 0.0001). Results suggest that the ability to extract water from the soil under increasing soil water deficit is a major attribute of drought adaptation. The genetic variation and transgressive segregation suggest further genomic and transcriptomic studies for unravelling the complex relationship between drought adaptation and performance under drought.

High concentrations of Na+ and Cl– ions in soil solution have simultaneous detrimental effects on growth of faba bean under salinity stress

Despite the fact that most plants accumulate both sodium (Na+) and chloride (Cl–) ions to high concentration in their shoot tissues when grown in saline soils, most research on salt tolerance in annual plants has focused on the toxic effects of Na+ accumulation. There have also been some recent concerns about the ability of hydroponic systems to predict the responses of plants to salinity in soil. To address these two issues, an experiment was conducted to compare the responses to Na+ and to Cl– separately in comparison with the response to NaCl in a soil-based system using two varieties of faba bean (Vicia faba), that differed in salinity tolerance. The variety Nura is a salt-sensitive variety that accumulates Na+ and Cl– to high concentrations while the line 1487/7 is salt tolerant which accumulates lower concentrations of Na+ and Cl–. Soils were prepared which were treated with Na+ or Cl– by using a combination of different Na+ salts and Cl– salts, respectively, or with NaCl. While this method produced Na+-dominant and Cl–-dominant soils, it unavoidably led to changes in the availability of other anions and cations, but tissue analysis of the plants did not indicate any nutritional deficiencies or toxicities other than those targeted by the salt treatments. The growth, water use, ionic composition, photosynthesis, and chlorophyll fluorescence were measured. Both high Na+ and high Cl– reduced growth of faba bean but plants were more sensitive to Cl– than to Na+. The reductions in growth and photosynthesis were greater under NaCl stress and the effect was mainly additive. An important difference to previous hydroponic studies was that increasing the concentrations of NaCl in the soil increased the concentration of Cl– more than the concentration of Na+. The data showed that salinity caused by high concentrations of NaCl can reduce growth by the accumulation of high concentrations of both Na+ and Cl– simultaneously, but the effects of the two ions may differ. High Cl– concentration reduces the photosynthetic capacity and quantum yield due to chlorophyll degradation which may result from a structural impact of high Cl– concentration on PSII. High Na+ interferes with K+ and Ca2+ nutrition and disturbs efficient stomatal regulation which results in a depression of photosynthesis and growth. These results suggest that the importance of Cl– toxicity as a cause of reductions in growth and yield under salinity stress may have been underestimated.

Quantitative trait loci on chromosome 4B for coleoptile length and early vigour in wheat (Triticum aestivum L.)

The Norin-10 dwarfing genes, Rht-B1b (Rht1) and Rht-D1b (Rht2), are commonly used to reduce plant height and increase grain yield in wheat breeding programs. These dwarfing genes lower sensitivity of vegetative tissue to endogenous gibberellin to reduce cell and subsequent stem elongation. This reduction in cell elongation capacity reportedly results in a concomitant reduction in coleoptile length and early vigour (leaf area) thereby affecting seedling establishment and growth. A detailed genetic map from a cross between tall Halberd (Rht-B1a) and semidwarf Cranbrook (Rht-B1b) wheat cultivars was used to assess genetic factors affecting seedling growth. Parental and 150 doubled haploid progeny lines were characterised for seedling and height-related traits in controlled and field environments. Genotypic variation was large and predominantly under additive genetic control with evidence for transgressive segregation for some traits. Narrow-sense heritabilities were moderate to high (h2 = 0.31–0.91) indicating a strong genetic basis for differences between progeny. Molecular marker analyses identified a number of significant (P < 0.05) quantitative trait loci (QTL) for each trait. A major QTL, mapping directly to the Rht-B1 locus on chromosome arm 4BS, accounted for up to 49% of the genotypic variance in peduncle length and plant height, and 27–45% of the genotypic variance in coleoptile length across different temperatures. Another QTL, located close to the RFLP marker XksuC2 on the long arm of chromosome 4B, accounted for 15–27% of the genotypic variance in coleoptile length. The influence of the XksuC2-linked QTL on coleoptile length was greatest at 19˚C and decreased with cooler temperatures. The same QTL affected reductions in leaf size, and both coleoptile tiller size and presence to affect overall seedling vigour. There was also some evidence for epistatic interactions influencing coleoptile tiller growth. Reductions in plant size at the Rht-B1b and XksuC2 loci were associated with presence of the Cranbrook 4B allele. The negative genetic effect of the Rht-B1b dwarfing gene on early growth of wheat confirms phenotypic evidence of a pleiotropic effect of Rht-B1b on establishment and early vigour. Genetic increases in coleoptile length and early leaf area development are likely to be limited in wheat populations containing the Rht-B1b dwarfing gene.

Sodium exclusion QTL associated with improved seedling growth in bread wheat under salinity stress

Worldwide, dryland salinity is a major limitation to crop production. Breeding for salinity tolerance could be an effective way of improving yield and yield stability on saline-sodic soils of dryland agriculture. However, this requires a good understanding of inheritance of this quantitative trait. In the present study, a doubled-haploid bread wheat population (Berkut/Krichauff) was grown in supported hydroponics to identify quantitative trait loci (QTL) associated with salinity tolerance traits commonly reported in the literature (leaf symptoms, tiller number, seedling biomass, chlorophyll content, and shoot Na+ and K+ concentrations), understand the relationships amongst these traits, and determine their genetic value for marker-assisted selection. There was considerable segregation within the population for all traits measured. With a genetic map of 527 SSR-, DArT- and gene-based markers, a total of 40 QTL were detected for all seven traits. For the first time in a cereal species, a QTL interval for Na+ exclusion (wPt-3114-wmc170) was associated with an increase (10%) in seedling biomass. Of the five QTL identified for Na+ exclusion, two were co-located with seedling biomass (2A and 6A). The 2A QTL appears to coincide with the previously reported Na+ exclusion locus in durum wheat that hosts one active HKT1;4 (Nax1) and one inactive HKT1;4 gene. Using these sequences as template for primer design enabled mapping of at least three HKT1;4 genes onto chromosome 2AL in bread wheat, suggesting that bread wheat carries more HKT1;4 gene family members than durum wheat. However, the combined effects of all Na+ exclusion loci only accounted for 18% of the variation in seedling biomass under salinity stress indicating that there were other mechanisms of salinity tolerance operative at the seedling stage in this population. Na+ and K+ accumulation appear under separate genetic control. The molecular markers wmc170 (2A) and cfd080 (6A) are expected to facilitate breeding for salinity tolerance in bread wheat, the latter being associated with seedling vigour.

The response of barley to salinity stress differs between hydroponic and soil systems.

Many studies on salinity stress assume that responses in hydroponics mimic those in soil. However, interactions between the soil solution and the soil matrix can affect responses to salinity stress. This study compared responses to salinity in hydroponics and soil, using two varieties of barley (Hordeum vulgare L.). The responses to salinity caused by high concentrations of Na+ and Cl– were compared to assess any consistent differences between hydroponics and soil associated with a cation and an anion that contribute to salinity stress. Concentrated nutrient solutions were also used to assess the effects of osmotic stress. The effects of salinity differed between the hydroponic and soil systems. Differences between barley cultivars in growth, tissue moisture content and ionic composition were not apparent in hydroponics, whereas significant differences occurred in soil. Growth reductions were greater under hydroponics than in soil at similar electrical conductivity values, and the uptake of Na+ and Cl– was also greater. The relative importance of ion exclusion and osmotic stress varied. In soil, ion exclusion tended to be more important at low to moderate levels of stress (EC at field capacity up to 10 dS m–1) but osmotic stress became more important at higher stress levels. High external concentrations of Cl– had similar adverse effects as high concentrations of Na+, suggesting that Cl– toxicity may reduce growth. Fundamental differences in salinity responses appeared between soil and solution culture, and the importance of the different mechanisms of damage varies according to the severity and duration of the salt stress.

A simple method to evaluate genetic variation in grain zinc concentration by correcting for differences in grain yield

Increasing the grain zinc (Zn) concentration of staple food crops will help alleviate chronic Zn deficiency in many areas of the world. Significant variation in grain Zn concentration is often reported among collections of cereals, but frequently there is a concomitant variation in grain yield. In such cases grain Zn concentration and grain yield are often inversely related. Without considering the influence of the variation in grain yield on Zn concentration, the differences in grain Zn concentration may simply represent a yield dilution effect. Data from a series of field and glasshouse experiments was used to illustrate this effect and to describe an approach that will overcome the yield dilution effect. In experiments with a wide range of bread wheat, synthetic hexaploids and accessions of durum wheat, variation in grain yield among the genotypes accounted for 30–57% of the variation in grain Zn concentration. Variation in kernel weight also occurred, but was poorly correlated with grain Zn concentration. To account for the influence of variation in grain yield on grain Zn concentration grain Zn yield was plotted against grain yield. By defining the 95% confidence belt for the regression genotypes that have inherently low or high grain Zn concentrations at a given yield level can be identified. This method is illustrated using two data sets, one consisting of bread wheat and one comprising a collection of synthetic hexaploids.

High temperature effects on photosynthesis and water relations of grain legumes

Effects of high temperature on photosynthesis, and its interaction with water relations in common bean (Phaseolus vulgaris), cowpea (Vigna unguiculata), faba bean (Vicia faba), and five cultivars of field pea (Pisum sativum) were investigated. Responses of all species were compared at 20/15, 30/15, or 30/25 °C day/night, and cowpea and pea were compared at 20/15 and 30/25 °C under well-watered and limited-water conditions. Response of pea to 20/15 and 30/25 °C during flowering was ascertained, and sensitivity of the photosystem of pea and faba bean to 40 °C was determined.High temperature decreased chlorophyll variable fluorescence (Fv), a measure of injury to photosynthesis, in all species except cowpea, which was highly tolerant. Leaf chlorophyll and most measures of growth were favored by high day temperature but not by high night temperature, and photosynthetic rates were enhanced by high temperatures that increased leaf chlorophyll and nitrogen (N) contents. High temperature diminished growth less than water deficiency and increased water use of all three species but only lowered the water potential in faba bean. Water deficiency generally decreased growth, water use, and water potential more at 30/25 °C than at 20/15 °C. Stress from high temperature during flowering of pea decreased all components of yield at maturity, particularly at nodes that flowered latest. Whole-chain photosynthetic activity in thylakoids of pea, faba bean, and wheat (Triticum aestivum) were equally sensitive to high temperature, suggesting that Photosystem Il was the most labile component. The results show that high temperature affects photosynthesis, growth, and water relations of grain legumes, and sensitivity to the stress differs among species and genotypes.

Subsoil constraints to crop production on neutral and alkaline soils in south-eastern Australia: a review of current knowledge and management strategies

Crop yield variability and productivity below potential yield on neutral and alkaline soils in the semiarid Mediterranean-type environments of south-eastern Australia have been attributed, in part, to variable rooting depth and incomplete soil water extraction caused by physical and chemical characteristics of soil horizons below the surface. In this review these characteristics are referred to as subsoil constraints. This document reviews current information concerning subsoil constraints typical of neutral and alkaline soils in south-eastern Australia, principally salinity, sodicity, dense soils with high penetration resistance, waterlogging, nutrient deficiencies and ion toxicities. The review focuses on information from Australia (published and unpublished), using overseas data only where no suitable Australian data is available. An assessment of the effectiveness of current management options to address subsoil constraints is provided. These options are broadly grouped into three categories: (i) amelioration strategies, such as deep ripping, gypsum application or the use of polyacrylamides to reduce sodicity and/or bulk density, deep placement of nutrients or organic matter to overcome subsoil nutrient deficiencies or the growing of ‘primer’ crops to naturally ameliorate the soil; (ii) breeding initiatives for increased crop tolerance to toxicities such as salt and boron; and (iii) avoidance through appropriate agronomic or agro-engineering solutions. The review highlights difficulties associated with identifying the impact of any single subsoil constraint to crop production on neutral and alkaline soils in south-eastern Australia, given that multiple constraints may be present. Difficulty in clearly ranking the relative effect of particular subsoil constraints on crop production (either between constraints or in relation to other edaphic and biological factors) limits current ability to develop targeted solutions designed to overcome these constraints. Furthermore, it is recognised that the task is complicated by spatial and temporal variability of soil physicochemical properties and nutrient availability, as well as other factors such as disease and drought stress. Nevertheless, knowledge of the relative importance of particular subsoil constraints to crop production, and an assessment of impact on crop productivity, are deemed critical to the development of potential management solutions for these neutral to alkaline soils.

Chickpea genotypes differ in their sensitivity to Zn deficiency

Zinc (Zn) deficiency is common in most of the chickpea growing areas of the world and growing Zn-efficient genotypes on Zn-deficient soil is a benign approach of universal interest. Response of 13 chickpea genotypes (10 desi and 3 kabuli) to Zn nutrition was studied in a pot experiment under glasshouse conditions. Plants were grown in a Zn-deficient siliceous sand for 6 weeks and fertilized with 0 (Zn−) and 2.5 mg Zn per kg soil (Zn+). When grown with no added Zn, Zn deficiency symptoms (chlorosis of younger leaves and stipules followed by necrosis of leaf margins) appeared 3–4 weeks after planting and were more apparent in cultivars Tyson, Amethyst and Dooen than Kaniva and T-1587. Zn deficiency reduced shoot growth, but it was less affected in breeding lines T-1587 and CTS 11308 than cultivars Tyson, Dooen, Amethyst and Barwon. Among all genotypes, Tyson produced the lowest root dry weight in Zn– treatment. Zinc efficiency based on shoot dry weight showed marked differences among genotypes; breeding lines CTS-60543, CTS-11308 and T-1587 were 2-fold more Zn-efficient than cultivars Tyson and Dooen. A higher Zn accumulation per plant and higher Zn uptake per g. of root dry weight were recorded in T-1587 and CTS-11308 when compared with Tyson. Root:shoot ratio was increased and proportionally more Zn was transported to the shoot when the soil was deficient. Cultivars that were very sensitive to Zn deficiency tended to have their root:shoot ratio increased by Zn deficiency more than less sensitive cultivars. The insensitive lines T-1587 and CTS-11308 transported more than 70% of the total absorbed Zn to the shoot. It is concluded that chickpea genotypes vary in their sensitivity to Zn deficiency. Advanced breeding lines T-1587 and CTS-11308 are relatively more Zn-efficient compared with Australian chickpea cultivar Tyson. Zn efficiency in chickpea genotypes is probably related to an efficient Zn absorption coupled with a better root to shoot transport.