Gary M. Paulsen

Interaction of drought and high temperature on photosynthesis and grain-filling of wheat

Drought and high temperature often occur simultaneously, but their effects on crops are usually investigated individually. Our objective was to compare effects of drought, high temperature, and their interactions on photosynthesis and grain-growth of wheat (Triticum aestivum L.). Plants (cv. Len) were grown uniformly in well-watered soil at 25/20 ± 2 °C day/night until anthesis, when they were subjected to regimes of no drought (soil at field capacity) and drought (plant water potential of −.0 to −2.4 MPa) at 15/10, 25/20, and 35/30 °C in controlled environments until physiological maturity. Drought decreased photosynthesis, stomatal conductance, viable leaf area, shoot and grain mass, and weight and soluble sugar content of kernels but increased plant water-use efficiency. High temperature hastened the decline in photosynthesis and leaf area, decreased shoot and grain mass as well as weight and sugar content of kernels, and reduced water-use efficiency. Interactions between the two stresses were pronounced, and consequences of drought on all physiological parameters were more severe at high temperature than low temperature. The synergistic interactions indicated that productivity of wheat is reduced considerably more by the combined stresses than by either stress alone, and that much of the effect is on photosynthetic processes.

Combined effects of drought and high temperature on water relations of wheat and sorghum

Drought and high temperature are major factors limiting crop production. The two stresses occur together in many regions, but they usually are investigated separately. This study tested the hypothesis that high temperature interacts with drought to affect water relations, and the effect is greater in heat-sensitive wheat (Triticum aestivum L.) than in sorghum (Sorghum bicolor L. Moench). Wheat and sorghum were grown in soil that was well watered or not watered in controlled chambers at 15/10, 25/20, 35/30 and 40/35 °C day/night. Soil water content (SWC), leaf relative water content (RWC), leaf water potential (Ψ), leaf osmotic potential (π), leaf turgor potential (P) and osmotic adjustment (OA) were determined at 2-d intervals. All values held nearly constant at all temperatures when soil was well watered but were affected strongly by high temperature when water was withheld. The combined stresses reduced SWC, RWC, Psi and π, and unevenly raised P over time, particularly in sorghum. Sorghum also exhibited marked OA at high temperature, which was usually lethal to wheat. High temperature appeared to interact with drought to affect water relations by altering SWC and not by influencing OA. The results demonstrated that crops maintain nearly stable water relations regardless of temperature when moisture is ample, but high temperature strongly affects water relations when water is limiting. Increasing the thermotolerance of wheat might improve its potential to acclimate to both high temperature and drought.

Evaluation of selection strategies for wheat adaptation across water regimes

Drought frequently constrains production of wheat (Triticum aestivum L.), but development of tolerant cultivars is hampered by low heritability for drought tolerance and a lack of effective selection strategies. Our objective was to identify an optimum selection regime for wheat in drought-prone environments. Six-hundred entries derived from 10 crosses were developed by selection under continuous high moisture, alternating high with low moisture, alternating low with high moisture, and continuous low moisture conditions for five generations. The selections were evaluated in two low-yield, a medium-yield, and a high-yield environment in the Yaqui Valley, Sonora, Mexico. The mean performance of entries derived from a particular selection regime was dependant on the stress level of the evaluation environment. Lines developed and selected under continuous high moisture and continuous low-moisture regimes produced the highest mean yields in the low moisture evaluation environment. There was no relationship between continuous selection under either high yielding conditions or low yielding conditions and the mean performance of the resultant lines in their respective high and low yielding evaluation environments. The mean yield of lines selected using the alternating high/low moisture regime as well as the five highest yielding lines were superior in the HY environment, and had similar performance with other regimes under the low yielding evaluation environment. Our results indicate that alternating selection between high and low yielding environments is the most effective way to develop wheat germplasm adapted to environments where intermittent drought occurs.

EDTA-assisted heavy-metal uptake by poplar and sunflower grown at a long-term sewage-sludge farm

Little information is available concerning the efficacy of chelates applied to biosolids (sewage-sludge)-treated soil for heavy-metal removal. The purpose of the experiment was to determine the availability to sunflower (Helianthus annuus L.) and hybrid poplar (Populus deltoides Marsh. × P. nigra L.) seedlings, of non-essential (Cd, Ni, Pb) and essential heavy metals (Cu, Fe, Mn, Zn) in field soil injected with biosolids since 1976 and treated with ethylenediamine-tetraacetic acid (EDTA) in 2001. Sunflower was grown at two densities, 20 000 and 60 000 plants/ha, and poplar at 10 000 plants/ha. The tetrasodium salt of EDTA was applied at rates of 0, 0.5, 1, and 2 g EDTA salt per kg surface (25-cm depth) soil. The EDTA did not affect uptake by poplar of the three non-essential (Cd, Ni, Pb) and four essential (Cu, Fe, Mn, Zn) heavy metals. For sunflower, the 1.0 g/kg rate of chelate addition resulted in maximal removal of the three non-essential heavy metals (Cd, Ni, Pb). Uptake of the essential heavy metals by sunflower was little affected by the EDTA. At the 20 000 plants/ha density, leaves of sunflower grown with 1.0 g EDTA Na4ċ2H2O per kg soil accumulated more Cd, Ni, and Pb than leaves of sunflower grown without the EDTA salt. At this density, concentrations of Cd in leaves of sunflower without EDTA and with 1.0 g/kg EDTA salt were 2.2 and 6.5 μg/g, respectively; for Ni, they were 6.7 and 19.2 μg/g, respectively; and for Pb, they were 15.6 and 46.9 μg/g, respectively. At the 60 000 plants/ha density, stems of sunflower grown with 1.0 g EDTA Na4ċ2H2O per kg soil accumulated more Cd, Ni, and Pb than stems of sunflower grown without the EDTA salt. At this density, concentrations of Cd in stems of sunflower without EDTA and with 1.0 g/kg EDTA salt were 0.6 and 4.6 μg/g, respectively; for Ni, they were 1.7 and 17.6 μg/g, respectively; and for Pb, they were 5.2 and 42.8 μg/g, respectively. Removal of the non-essential heavy metals by sunflower was greater at the higher plant density (60 000 plants/ha) compared to the lower one (20 000 plants/ha).

Quantitative and molecular characterization of heat tolerance in hexaploid wheat

Understanding the genetic basis of tolerance to high temperature is important for improving the productivity of wheat (Triticum aestivum L.) in regions where the stress occurs. The objective of this study was to estimate inheritance of heat tolerance and the minimum number of genes for the trait in bread wheat by combining quantitative genetic estimates and molecular marker analyses. Two cultivars, Ventnor (heat-tolerant) and Karl92 (heat-susceptible), were crossed to produce F1, F2, and F3populations, and their grain-filling duration (GFD) at 30/25 °C 16/8 h day/night was determined as a measure of heat tolerance. Distribution of GFD in the F1 and F2 populations followed the normal model (χ2, p > 0.10). A minimum of 1.4 genes with both additive and dominance effects, broad-sense heritability of 80%, and realized heritability of 96%for GFD were determined from F2 and F3 populations. Products from 59primer pairs among 232 simple sequence repeat (SSR) pairs were polymorphic between the parents. Two markers, Xgwm11 andXgwm293, were linked to GFD by quantitative trait loci (QTL) analysis of the F2 population. The Xgwm11-linked QTL had only additive gene action and contributed 11% to the total phenotypic variation in GFD in the F2population, whereas the Xgwm293-linked QTL had both additive and dominance action and contributed 12% to the total variation in GFD. The results demonstrated that heat tolerance of common wheat is controlled by multiple genes and suggested that marker-assisted selection with microsatellite primers might be useful for developing improved cultivars.

Genotypic differences in utilization of assimilate sources during maturation of wheat under chronic heat and heat shock stresses

Heat stress from chronic, prolonged exposure up to 32 °C or heat shock from brief exposure to 33 °C and above alters the source of assimilates for grain growth of wheat (Triticum asetivum L.). Our objectives were to identify genotypes that resist chronic heat stress and heat shock and to determine the relative contributions of photosynthesis and stem reserves to grain filling under both conditions. Twenty-eight genotypes were grown in controlled enviroments at 20/15 and 30/25 °C day/night in light and darkness during maturation in the first experiment, and six genotypes were grown in light at the same temperatures and at 40/35 °C followed by 20/15 or 30/25 °C in the second experimnet. Heat susceptibility indices (HSI) were calculated from grain yields of the genotypes in both experiments. The ratio of chlorophyll variable fluorescence to maximum fluorescence (Fv/Fm), a measure of the stability of photosynthesis, and carbohydrate reserves in the stems were measured in the second experiment. Photosynthesis provided 63 and 65% of assimilates in the grain at 20/15 and 30/25 °C, respectively, but both stable photosynthesis in some genotypes and high content of reserves in other genotypes were associated with low susceptibility to stress. The Fv/Fm ratio was decreased by heat shock and returned to normal values intolerant genotypes when the treatment was followed by 20/15 °C but not 30/25 °C. Grain yield was highly correlated among 20/15, 30/25, and 40/35 °C followed by 20/15 °C treatments, suggesting that similar plant traits were involved. We conclude that assimilates from either stable photosynthesis or high reserve levels provided for high grain yields during heat stress. Combining the two traits could improve heat tolerance of wheat but might not be feasible if other traits are impeded.

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.

Growth and senescence characteristics associated with tolerance of wheat-alien amphiploids to high temperature under controlled conditions

Tolerance of wheat (Triticum aestivumL.) to high temperature might be improved by introducing alien genes from amphiploids. Our objectives were to determine responses of synthetic hexaploid and octaploid amphiploid wheats to high temperature and evaluate their potential usefulness for developing improved cultivars. Thirty synthetic hexaploids from durum wheat (T. turgidum L.) × Aegilops tauschii Cos. Accessions and four octaploid amphiploids from Chinese Spring wheat × different grasses were grown at 20/15 and 30/25 °C day/night during maturation. Tolerance was ascertained by two measures of senescence, leaf chlorophyll content and grain filling duration, plus grain yield and its components. Leaf chlorophyll was measured after 10 and 15 days of treatment, and grain yield was determined at maturity to calculate the heat susceptibility index(HSI), a gauge of the reduction in yield at high temperature of each line relative to all other lines. Chlorophyll content, grain filling duration, yield, and kernel weight were highly negatively correlated with HIS of the hexaploid amphiploids at30/25 °C, but grain yield was positively correlated with HSI at20/15 °C. The hexaploid lines might be useful for improving wheat for regions where stress from high temperature occurs frequently. Chlorophyll content and grain filling duration also were highly negatively correlated with HSI of the octaploid lines, but they would be less directly useful for improving wheat because the kernel number was reduced greatly due to unbalanced meiotic chromosomal segregation.

Functional and ultrastructural injury to photosynthesis in wheat by high temperature during maturation

Abstract Wheat (Triticum aestivum L. cv. Len) plants were exposed to 15/10, 25/20, and 35/30°C day/night temperature regimes after anthesis until plants grown at 25/20°C reached physiological maturity. Increasing temperature during maturation enhanced leaf senescence, accentuated the loss of chloroplast integrity, increased thylakoid luminal volume and decreased extent of appressed thylakoid membrane area, and accelerated the decline of PSII-mediated electron transport. Stomatal conductance declined slower than other photosynthetic processes at high temperature. Lability of PSII and stability of PSI activities to prolonged high temperature contrasted with measurements of Chl-binding proteins, such as LHCII, type-II LHCI and the PsaA-PsaB polypeptides. While the activity of PSII declined rapidly, the LHCII polypeptides remained as prominent thylakoid membrane components. Activity of PSI, in contrast, was sustained despite a diminished 735:685 nm fluorescence emission ratio and severe damage to the Chl-binding proteins, such as PsaA-PsaB and type-II LHCI. The dissimilar responses of LHCII and type-II LHCI to high temperature injury may be related to their localization in the thylakoid membrane; LHCII, predominantly in the appressed membrane regions, may be relatively shielded from proteolytic activity as compared with type-II LHCI in the exposed membrane regions.

Auxin-Enhanced Root Growth for Phytoremediation of Sewage-Sludge Amended Soil

A technology to increase root growth would be advantageous for phytoremediation of trace metal polluted soil, because more roots would be available for metal uptake. The objective of this study was to determine if the auxin, indole-3-acetic acid (IAA), would increase root growth in soil with metals from sewage sludge, when the tetrasodium salt of the chelate EDTA (ethylenediamine-tetraacetic acid) was added to solubilize the metals. Sunflower (Helianthus annuus L.) plants grew in large pots containing either soil from a sludge farm or composted sludge. The EDTA salt was added at a rate of 1 g kg−1 soil 37 days after planting. IAA at the rate of 3 or 6 mg l−1 was sprayed on the leaves (500 ml) and added to the soil (500 ml) three times: 41, 50, and 74 days after planting. At harvest 98 days after planting, oven-dry weights were measured, and plant organs were analyzed for Cd, Cu, Fe, Mn, Ni, Pb, and Zn. Metal uptake was determined as the product of metal concentration in an organ and weight. IAA increased root growth of plants grown in the soil with sludge when no EDTA was present. With no EDTA, Mn and Ni in leaves of plants grown in the soil were higher at 3 and 6 mg l−1 IAA compared to 0 mg l−1 IAA. With and without EDTA, Cd and Pb in leaves of plants grown in the compost were higher with 3 and 6 mg l−1 IAA compared to 0 mg l−1 IAA.