Impact of High Night‐Time and High Daytime Temperature Stress on Winter Wheat

High temperature is a major environmental factor that limits wheat (Triticum aestivum L.) productivity. Climate models predict greater increases in night‐time temperature than in daytime temperature. The objective of this research was to compare the effects of high daytime and high night‐time temperatures during anthesis on physiological (chlorophyll fluorescence, chlorophyll concentration, leaf level photosynthesis, and membrane damage), biochemical (reactive oxygen species (ROS) concentration and antioxidant capacity in leaves), growth and yield traits of wheat genotypes. Winter wheat genotypes (Ventnor and Karl 92) were grown at optimum temperatures (25/15 °C, maximum/minimum) until the onset of anthesis. Thereafter, plants were exposed to high night‐time (HN, 25/24 °C), high daytime (HD, 35/15 °C), high daytime and night‐time (HDN, 35/24 °C) or optimum temperatures for 7 days. Compared with optimum temperature, HN, HD and HDN increased ROS concentration and membrane damage and decreased antioxidant capacity, photochemical efficiency, leaf level photosynthesis, seed set, grain number and grain yield per spike. Impact of HN and HD was similar on all traits. Greater impact on seed set, grain number and grain yield per spike was observed at HDN compared with HN and HD. These results suggest that HN and HD during anthesis cause damage of a similar magnitude to winter wheat.     

Growth and Yield Response of Facultative Wheat to Winter Sowing, Freezing Sowing and Spring Sowing at Different Seeding Rates

Growth and yield of wheat are affected by environmental conditions and can be regulated by sowing time and seeding rate. In this study, three sowing times [winter sowing (first week of September), freezing sowing (last week of October) and spring sowing (last week of April)] at seven seeding rates (325, 375, 425, 475, 525, 575 and 625 seeds m^?2) were investigated during the 2002–03 and 2003–04 seasons, in Erzurum (Turkey) dryland conditions, using Kirik facultative wheat. A split‐plot design was used, with sowing times as main plots and seeding rates randomized as subplots. There was a significant year × sowing time interaction for grain yield and kernels per spike. Winter‐sown wheat produced a significantly higher leaf area index, leaf area duration, spikes per square metre, kernel weight and grain yield than freezing‐ and spring‐sown wheat. The optimum time of sowing was winter for the facultative cv. Kirik. Grain yields at freezing and spring sowing were low, which was largely the result of hastened crop development and high temperatures during and after anthesis. Increasing seeding rate up to 525 seeds m^?2 increased the spikes per square metre at harvest, resulting in increased grain yield. Seeding rate, however, was not as important as sowing time in maximizing grain yield. Changes in spikes per square metre were the major contributors to the grain‐yield differences observed among sowing times and seeding rates. Yield increases from higher seeding rates were greater at freezing and spring sowing. We recommended that a seeding rate of 525 seeds m^?2 be chosen for winter sowing, and 575 seeds m^?2 for freezing and spring sowing.     

Wheat Cultivars Adapted to Post-Heading High Temperature Stress

The existence of genetic variation in wheat for tolerance to high temperature stress has been reported but cultivars released for a particular production system often are not characterized. The objective of this study was to identify and describe the characteristics of wheat cultivars adapted to production systems with risks of high temperature during the post‐heading period. Fifteen diverse wheat cultivars and one unreleased genotype were evaluated at the Texas A&M University Agricultural Research and Extension Center, Uvalde, TX, during two seasons characterized by daily maximum temperatures as high as 36 °C. Measurements during both seasons included days to heading, days to physiological maturity and grain yield. Large and significant (P < 0.05) grain yield differences were measured among cultivars within each season. Yield varied between 2979 and 4671 kg ha^?1 in the first season and between 1916 and 5200 kg ha^?1 in the second season. Late planting in the second season delayed heading date resulting in the post‐heading period to coincide with periods of high temperatures. Cultivars that headed early, in general, yielded better than cultivars that headed later within each season with yield reduction of 35.3 kg ha^?1 in the first season and 91.0 kg ha^?1 in the second season for every 1 day delay in heading after mid‐March. Early‐heading cultivars outperformed later‐heading cultivars because of two distinct advantages: the early‐heading cultivars had longer post‐heading and, therefore, longer grain filling period than the later‐heading cultivars. In addition, early‐heading cultivars completed a greater fraction of the grain filling earlier in the season when air temperatures were lower and generally more favourable. The advantage of earlier‐heading cultivars was also manifested in the amount of green leaves retained to anthesis. Earlier‐heading cultivars produced fewer total leaves per tiller but retained more green leaves and lost fewer leaves to senescence at anthesis than later‐heading cultivars. The results suggest that early heading is an important and effective single trait defining wheat cultivars adapted to production systems prone to high temperature stress during the post‐heading period.     

QTL for yield, yield components and canopy temperature depression in wheat under late sown field conditions

A wheat (Triticum aestivum L.) recombinant inbred line (RIL) population was used to identify quantitative trait loci (QTL) associated with yield, yield components, and canopy temperature depression (CTD) under field conditions. The RIL population, consisting of 118 lines derived from a cross between the stress tolerant cultivar ‘Halberd’ and heat stress sensitive cultivar ‘Karl92’, was grown under optimal and late sown conditions to impose heat stress. Yield and yield components including biomass, spikes m^?2, thousand kernel weight, kernel weight and kernel number per spike, as well as single kernel characteristics were determined. In addition, CTD was measured during both moderate (32–33 °C) and extreme heat stress (36–37 °C) during grain-filling. Yield traits showed moderate to high heritability across environments with a large percentage of the variance explained by genetic effects. Composite interval mapping detected 25 stable QTL for the 15 traits measured, with the amount of phenotypic variation explained by individual QTL ranging from 3.5 to 27.1 %. Two QTL for both yield and CTD were co-localized on chromosomes 3BL and 5DL and were independent of phenological QTL. At both loci, the allele from Halberd was associated with both higher yield and a cooler crop canopy. The QTL on 3BL was also pleiotropic for biomass, spikes m^?2, and heat susceptibility index. This region as well as other QTL identified in this study may serve as potential targets for fine mapping and marker assisted selection for improving yield potential and stress adaptation of wheat.     

Traits in Spring Wheat Cultivars Associated with Yield Loss Caused by a Heat Stress Episode after Anthesis

Heat stress resulting from climate change and more frequent weather extremes is expected to negatively affect wheat yield. We evaluated the response of different spring wheat cultivars to a post‐anthesis high temperature episode and studied the relationship between different traits associated with heat tolerance. Fifteen spring wheat (Triticum aestivum L.) cultivars were grown in pots under semifield conditions, and heat stress (35/26 °C) and control treatments (20/12 °C) were applied in growth chambers for 5 days starting 14 days after flowering. The heat stress treatment reduced final yield in all cultivars. Significant variation was observed among cultivars in the reduction in average grain weight and grain dry matter yield under heat stress (up to 36 % and 45 %, respectively). The duration of the grain‐filling period was reduced by 3–12 days by the heat treatment. The reduction in the grain‐filling period was negatively correlated with grain nitrogen yield (r = ?0.60). A positive correlation (r = 0.73) was found between the treatment effect on green leaf area (GLA) and the reduction in yield resulting from heat stress. The amount of stem water‐soluble carbohydrates (WSC) was not related to treatment effects on grain yield or grain weight. However, the treatment effect on stem WSC remobilization was negatively correlated with reduction in grain‐filling duration due to heat stress (r = ?0.74) and positively with treatment effect on grain N yield (r = 0.52). The results suggest that the effect of the heat treatment on GLA was the trait most associated with yield reduction in all cultivars. These findings suggest the importance of ‘stay green’‐associated traits in plant breeding as well as the need for better modelling of GLA in crop models, especially with respect to brief heat episodes during grain filling. There is in particular a need to model how heat and other stresses, including interacting effects of heat and drought, affect duration of GLA after flowering and how this affects source–sink relations during grain filling.     

氮素营养水平对冬小麦氮代谢关键酶活性变化和籽粒蛋白质含量的影响

通过氮素营养水平对籽粒蛋白质含量差异较大的3个小麦品种的氮素代谢关键酶活性变化及籽粒蛋白质含量影响的研究表明: 增加施氮量能够提高氮素同化关键酶硝酸还原酶和谷氨酰胺合成酶的活性, 降低旗叶蛋白质水解酶的活性. 增加施氮量提高籽粒蛋白质含量主要与促进开花后氮素吸收同化能力有关. 不同品种间籽粒蛋白质含量的差异     

Susceptibility of Faba Bean (Vicia faba L.) to Heat Stress During Floral Development and Anthesis

Experiments were conducted over 2 years to quantify the response of faba bean (Vicia faba L.) to heat stress. Potted winter faba bean plants (cv. Wizard) were exposed to temperature treatments (18/10; 22/14; 26/18; 30/22; 34/26 °C day/night) for 5 days during floral development and anthesis. Developmental stages of all flowers were scored prior to stress, plants were grown in exclusion from insect pollinators to prevent pollen movement between flowers, and yield was harvested at an individual pod scale, enabling effects of heat stress to be investigated at a high resolution. Susceptibility to stress differed between floral stages; flowers were most affected during initial green‐bud stages. Yield and pollen germination of flowers present before stress showed threshold relationships to stress, with lethal temperatures (t₅₀) ?28 °C and ~32 °C, while whole plant yield showed a linear negative relationship to stress with high plasticity in yield allocation, such that yield lost at lower nodes was partially compensated at higher nodal positions. Faba bean has many beneficial attributes for sustainable modern cropping systems but these results suggest that yield will be limited by projected climate change, necessitating the development of heat tolerant cultivars, or improved resilience by other mechanisms such as earlier flowering times.     

Improvement in Brazilian wheat breeding: changes in developmental phases and ecophysiological traits

Understanding the processes responsible for genetic progress in grain yield and relating them to the growth and developmental phases of the wheat plant are essential for improving yield potential. This study aimed to evaluate the duration of developmental phases and eco-physiological traits associated to grain yield in Brazilian wheat cultivars released in different decades. Wheat cultivars released from 1940 to 2009 were evaluated during 2010 and 2011 in Pato Branco, Paraná, Brazil. The length of the following periods was compared: sowing-emergence (SW-EM), emergence-double ridge (EM-DR), double ridge-terminal spikelet (DR-TS), terminal spikelet-anthesis (TS-ANT), anthesis-physiological maturity (ANT-PM), sowing-anthesis (SW-ANT) and sowing-physiological maturity (SW-PM). Yield components were also measured. Breeding has reduced the days until anthesis by 14.2%, while it has extended the grain-filling period by 7.6%, compared to the first cultivars released in the country, thereby contributing to a significant increase in 1000-grain weight (12.4 and 9.0% in 2010 and 2011, respectively). The TS-ANT phase was the only phase prior to anthesis exhibiting an extension from old to modern cultivars; this phase increased 1.56 °Cd syear^?1. Spike fertility index (SFI) showed increases of 37.8 and 23.8% in 2010 and 2011, respectively. Shortening of the time to flowering, shifting of critical phases of wheat development to better environmental conditions (i.e., TS-ANT phase), and selection for shorter cultivars have been directly responsible for the increase in spike dry weight at anthesis (SDWa) and SFI [i.e., grain number (GN)].     

Infrared Thermal Imaging as a Rapid Tool for Identifying Water‐Stress Tolerant Maize Genotypes of Different Phenology

The main task of this research was to evaluate canopy temperature and Crop Water Stress Index (CWSI) by assessing genotype variability of maize performance for different water regimes. To that end, three hundred tropical and subtropical maize hybrids with different phenology in terms of date of anthesis were evaluated. The influence of phenology on the change in canopy temperatures and CWSI was not equal during the three dates of measurement. At the end of vegetative growth (82 days after sowing, DAS) and at the blister stage (DAS 97), a high significant difference in temperatures and CWSI (P < 0.001) were obtained between the early‐ and late‐maturity genotypes. During anthesis (DAS 89), phenology had a significant effect (P < 0.01) only for the well‐watered genotypes, while under water‐stress conditions, no differences were found between early and late genotypes in terms of canopy temperature and CWSI. High significant differences (P < 0.001) in stomatal conductance (g_s) between early and late genotypes for different treatments were observed. A relationship (R² = 0.62) between g_s and canopy temperature was obtained. Under a water‐stress canopy, temperature was measured at anthesis, which was negatively correlated with grain yield of the early (r = ?0.55)‐ and late (r = ?0.46)‐maturity genotypes in the water‐stressed condition.     

Investigations on Heat Tolerance of Spring Wheat Varieties of Different Origin Under Growth Chamber Conditions

Winter wheat has replaced spring wheat to a large extent under the climatic conditions of Western and Central Europe. But spring wheat genotypes are still important under the warm climate in large parts of Africa and Central America, since spring wheat needs no vernalization. To reach optimal yield, genotypes must be well adapted to heat stress. In this study 11 spring wheat varieties of Egypt, Sudan, CIMMYT and Germany were exposed to high temperatures at ear emergence and beginning of anthesis. There were reductions in kernel yield and kernel number of the main ear for all genotypes, but the differences in reduction were remarkable. The German variety 'Naxos' was best adapted. Some varieties showed an increased number of florets per spikelet under heat treatment, but this does not prohibit a reduction in kernel number per spikelet.     

Drought Priming at Vegetative Growth Stage Enhances Nitrogen‐Use Efficiency Under Post‐Anthesis Drought and Heat Stress in Wheat

To study the effects of early drought priming at 5th‐leaf stage on grain yield and nitrogen‐use efficiency in wheat (Triticum aestivum L.) under post‐anthesis drought and heat stress, wheat plants were first exposed to moderate drought stress (drought priming; that is, the leaf water potential reached ca. ?0.9 MP a) at the 5th‐leaf stage for 11 days, and leaf water relations and gas exchange rates, grain yield and yield components, and agronomic nitrogen‐use efficiency (ANUE ) of the primed and non‐primed plants under post‐anthesis drought and heat stress were investigated. Compared with the non‐primed plants, the drought‐primed plants possessed higher leaf water potential and chlorophyll content, and consequently a higher photosynthetic rate during post‐anthesis drought and heat stress. Drought priming also resulted in higher grain yield and ANUE in wheat under post‐anthesis drought and heat stress. Drought priming at vegetative stage improves carbon assimilation and ANUE under post‐anthesis drought and heat stress and their combination in wheat, which might be used as a field management tool to enhance stress tolerance of wheat crops to multiple abiotic stresses in a future drier and warmer climate.     

Wheat Yield as Affected by Length of Exposure to Waterlogging During Stem Elongation

Waterlogging, if occurring within the stem elongation period (SE), is particularly critical for yield determination. We quantified for the first time the effect of waterlogging duration during SE on yield and studied whether the effects were only direct on resource capture or whether there were feed‐forward effects as well. We grew wheat (cv. Soissons) outdoors in long tubes (1.25 m deep) forming a normal canopy and imposed different treatments in SE to finish simultaneously around anthesis (0, 4, 8, 12, 16, 20 and 24 days) plus two complementary treatments (8 and 16 days) starting 10 days after the onset of SE. Yield was reduced linearly with the duration of waterlogging c. 2 % d_waterlogging^?1. Treatments mainly affected pre‐anthesis spike growth reducing the number of fertile florets and grains, not affecting fruiting efficiency. The magnitude of grain number loss was inversely proportional to the hierarchy of the spikes and spikelets. Grain weight was more marginally reduced, likely through the effects on the size of the ovaries of the developing florets. This reveals a direct effect of waterlogging on the capture of resources with no major feed‐forward effects. Losses were in agreement with those from other studies for particular durations of waterlogging.     

Frost response in wheat and early detection using proximal sensors

Frost can significantly reduce production of field crops grown in Mediterranean‐type environments, where significant economic losses for Australian wheat occur annually. If non‐destructive sensors could make rapid, spatial assessment of frost damage, this could limit economic losses through timely management decisions. This paper reports on a methodology for imposing frost treatments to wheat under field conditions and the utility of canopy reflectance data for detecting early crop damage. Purpose‐built chambers using stepped additions of dry ice allowed for a range of frost scenarios to be tested when applied at wheat heading and anthesis. For frost treatments applied at anthesis, grain number and yield were reduced by 8.8% and 7.2%, respectively, for every degree Celsius below zero (down to ?4°C). This effect was additive over two consecutive nights. For cold load equivalent, there was a 2.2% and 1.9% reduction in grain number and yield, respectively, per °C hr (below 0°C). For wheat, spectral indices PRI and NDVI (reflectance) and FRF_G and SFR_G (fluorescence) showed significant relationships, with cold load applied for heading treatments. Next steps include targeting frost intensities equivalent to cold loads between 20 and 80°C hr and testing the utility of these proposed indices in a commercial paddock setting.     

Pre-anthesis development and number of fertile florets in wheat as affected by photoperiod sensitivity genes <Emphasis Type="Italic">Ppd-D1</Emphasis>and <Emphasis Type="Italic">Ppd-B1</Emphasis>

Lengthening the late reproductive phase (LRP) of stem elongation in wheat (Triticum aestivumL.), by changing its photoperiod sensitivity independently of the preceding phases, would improve the yield potential through increasing spike weight and the number of fertile florets at anthesis. This paper presents results of a two-year field experiment designed to determine the impact of Ppd-D1and Ppd-B1on (i) the duration of three pre-anthesis developmental phases, and (ii) spike weight and the number of fertile florets at anthesis under two photoperiods during the LRP (natural and an extension of six hours over that). Near isogenic lines of Mercia and single chromosome recombinant lines of Cappelle Desprez were used. Under natural photoperiod, Ppd-D1hastened time to anthesis ca. 500^∘C d in both backgrounds by reducing each of the three pre-anthesis phases. Ppd-B1hastened the time to anthesis under natural photoperiod by 178^∘C d, mainly by reducing the early reproductive phase. The response to photoperiod of the LRP under extended daylength depended on the Ppdlocus present: Ppd-D1was insensitive while Ppd-B1and the recessive controls were sensitive. For all lines, photoperiod treatments and years, the number of fertile florets was associated with spike dry weight at anthesis (R ²≅ 80%, p< 0.01) which, in turn, was positively related to the intercepted radiation accumulated during the LRP (R ² 45%, p< 0.05). Changing the duration of the LRP through extended photoperiod or through Ppd-D1produced similar results in both backgrounds and years. Thus, altering the duration of the LRP by manipulating photoperiod sensitivity may be an alternative to changing the fertile floret number in wheat. Nevertheless, as no particular allele was responsible for the photoperiod sensitivity only during the LRP, new alleles should be studied to identify the control of photoperiod sensitivity of individual phases to fine-tune the pre-anthesis wheat development.     

A strategy of ideotype development for heat-tolerant wheat

Heat stress significantly limits yield in many wheat-growing areas globally including north-western NSW. While various traits linked to high-temperature tolerance have been identified, the combination of traits that optimize the heat tolerance of wheat has not been established in most environments. A total of 554 genotypes were evaluated in the field at different times of sowing in north-western NSW for three consecutive years to develop a heat-tolerant wheat ideotype for this environment. The later sown experiments were exposed to higher temperatures at the critical reproductive and grain-filling stages of development. The impact of high temperature was greatest at anthesis, and eventual grain yield was reduced by between 4% and 7% with every 1°C rise in average maximum temperature above the optimum of 25°C. High temperature reduced yield, plant height, grain weight and days to anthesis and maturity, and increased the percentage of screenings and grain protein content. Genotypes that produced higher yield under heat stress had shorter days to flowering and maturity, higher NDVI during grain filling, greater chlorophyll content at the milk stage of grain fill, taller plants, greater grain weight and number, and lower screenings compared with the benchmark cultivar Suntop. The genotype closest to the predicted heat-tolerant wheat ideotype identified from trait ranges had 79.6% similarity.     

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. This revised version was published online in August 2006 with corrections to the Cover Date.     

Heat,wheat and CO2: The relevance of timing and the mode of temperature stress on biomass and yield

Atmospheric CO₂ enrichment affects C3 crops both directly via increased carbon gain and improved water use efficiency and indirectly via higher temperatures and more frequent climatic extremes. Here we investigated the response of spring wheat (Triticum aestivum L. cv. Triso) to CO₂ enrichment (550 vs. 380 µmol/mol) and heat, applied as a constant +4°C increase or a typical heat wave either before or after anthesis, or as two typical heat waves before and after anthesis. We applied a climate chamber approach closely mimicking ambient conditions. CO₂ enrichment increased above‐ground biomass and yield by c. 7 and 10%, but was not able to compensate for adverse heat stress effects, neither before nor after anthesis, with few exceptions only. Yield depression due to heat stress was most severe when two heat waves were applied (?19%). This adverse effect was, however, compensated by CO₂ enrichment. Applying heat stress before or after anthesis did not exert different effects on yield for both +4°C warming and heat wave application. However, +4°C depressed yield more than a heat wave at ambient CO₂, but not so at elevated CO₂. Thus, the interactive effects were complex and prediction of future wheat yield under CO₂ enrichment and climate extremes deserves more attention.     

Independent and Combined Effects of Soil Warming and Drought Stress During Anthesis on Seed Set and Grain Yield in Two Spring Wheat Varieties

Increase in soil temperature together with decrease in soil moisture during anthesis of spring wheat (Triticum aestivum L) crops is predicted to occur more frequently in a future climate in Denmark. The objective of this study was to investigate the responses of two Danish spring wheat varieties (Trappe and Alora) to soil warming (H), drought (D) and both (HD) during anthesis. The plants were grown in pots in a climate‐controlled glasshouse. In H, the soil temperature was increased by 3 °C compared with the control (C). In both D and HD treatments, the plants were drought‐stressed by withholding irrigation until all of the transpirable soil water had been depleted in the pots. Results showed that, particularly under D treatment, Alora depleted soil water faster than Trappe. In both varieties, flag leaf relative water content (RWC) was significantly lowered, while spikelet abscisic acid (ABA) concentration was significantly increased by D and HD treatments. Compared with the C plants, D and HD treatments significantly reduced ear number, ear to tiller ratio, shoot biomass, grain yield, harvest index and seed set but hardly affected tiller number and 1000‐kernel weight, whereas H treatment alone only decreased shoot biomass and reduced seed set. When analysed across the varieties and the treatments, it was found that the reduction in seed set was closely correlated with the increase in spikelet ABA concentration, indicating that D and HD treatments induced greater spikelet ABA concentrations might have caused seed abortion. It was concluded that the grain yield reduction under D and HD treatments during anthesis in spring wheat is ascribed mainly to a lowered seed set and wheat varieties (i.e. Alora) with more dramatic increase in spikelet ABA concentration are more susceptible to D and HD treatment.     

黄淮麦区冬小麦超高产栽培的理论与实践

根据黄淮麦区的生态特点, 研究了冬小麦超高产(hm2产9000 kg籽粒)栽培的理论与技术. 结果指出, 于土肥水条件良好的高产麦田, 在建立合理群体结构的基础上, 小麦开花至成熟阶段的干物质积累和分配与植株生育后期早衰的矛盾, 是高产(hm2产6000～7500 kg籽粒)向超高产发展的主要限制因素. 揭示了高产条件下小麦的衰老规律, 划分