Nitrogen fertiliser rate and post-anthesis waterlogging effects on carbohydrate and nitrogen dynamics in wheat

Waterlogging is predicted to increase in both magnitude and frequency along with global warming, and will become one of the most severe adversities for crop production in many regions. Nitrogen is considered to be an effective up-regulatory nutrient for crops grown under stress and non-stress conditions. In this study, we try to evaluate N fertiliser effects on contents of carbohydrate and N dynamics, dry matter accumulation in shoot, yield under post-anthesis waterlogging. Waterlogging after anthesis significantly reduced grain yield due to decrease in thousand-kernel-weight and in grain number per spike. High N fertiliser application aggravated grain yield loss due to post-anthesis waterlogging. These yield losses were related to the decreases in dry matter accumulation, redistribution of stored photosynthate to the grain, and the conversion capacity from carbohydrate to starch in grain. The decrease in dry matter accumulation could be attributed to the reduced activities of Pn (photosynthesis) and SPS (sucrose phosphate synthase) in the flag leaf, while the low capacity in starch synthesis could be explained by the reduced activities of sucrose synthase (SS) and soluble starch synthase (SSS) in grain. Total N uptake in shoot was also reduced, which could contribute to the losses in biomass and yield by waterlogging. The decrease in Pn was inconsistent with the increase in N content in the flag leaf at high N fertiliser application under post-anthesis waterlogging.     

Effects of different water availability at post-anthesis stage on grain nutrition and quality in strong-gluten winter wheat

Wheat (Triticum aestivum L.) is one of the most important agricultural crops worldwide. However, water is the most important limiting factor for wheat production. This study was initiated to test water stress environmental effects on grain quality and nutritional value of wheat by using single different water conditions at post-anthesis stage. Further analyses were conducted to examine variations in concentrations and compositions of the bioactive compounds and nutritions in strong-gluten winter wheat subjected to different levels of water deficit during grain filling. For the experiment on the response to different soil water conditions during post-anthesis stage, effects of soil water environment on protein content and composition in the grains were significant. Soil water conditions in this study greatly affected mineral contents in the grains of winter wheat, particularly with regard to the major minerals (P, K, Ca and Mg). Water deficit during grain filling can result in a decrease in lipid contents in wheat grains, which agrees well with experimental findings elsewhere. Concomitantly, a mild water deficit during grain filling would be beneficial to the grain filling and starch compositions, significantly improved bread-making quality. Therefore, it was concluded that good management of wheat field water at post-anthesis stage was helpful to improving grain quality and nutritions relevant to processing and human nutrition. To cite this article: C.-X. Zhao et al., C. R. Biologies 332 (2009).     

Effects of waterlogging and drought on winter wheat and winter barley grown on a clay and a sandy loam soil

Summary The effects of winter waterlogging and a subsequent drought on the growth of winter barley and winter wheat have been examined. We used lysimeters containing soil monoliths with facilities to control the water table and a mobile shelter to control rainfall. Winter wheat was grown on a clay and on a sandy loam, but winter barley only on the clay soil. Lysimeters were either freely-drained during the winter or waterlogged with the water table 10 cm below the soil surface from 2 December until 31 March (that could occur by rainfall with a return period of 2 to 3 years). The lysimeters then were either irrigated so that the soil moisture deficit did not exceed 84 mm, or subjected to drought by limiting rainfall (equivalent to a 1 in 10 dry year in the driest area of England) so that the deficits reached maximum values of 150 mm in the clay and 159 mm in the sandy loam by harvest.Winter waterlogging restricted tillering and restricted the number of ears for all crops; grain yield of the winter barley was decreased by 219 g/m² (30%), and that of winter wheat by 170 g/m² (24%) and 153 g/m² (21% on the clay and sandy loam respectively.The drought treatment reduced the straw weight of winter barley by 75 g/m² (12%) but did not significantly depress the grain yield. For winter wheat on the clay, where the soil was freely-drained during the winter, drought depressed total shoot weight by 344 g/m² (17%) and grain weight by 137 g/m² (17%), but after winter waterlogging, drought did not further depress total or grain weight. In contrast, the winter wheat on the sandy loam was not significantly affected by drought.From these results, which are discussed in relation to other experiments in the United Kingdom, it seems that winter waterlogging is likely to cause more variation in the yield of winter barley and winter wheat than drought.     

Relationship between mineral coating on roots and yield performance of wheat under waterlogging stress

Waterlogging stress limits the productivity of winter wheat(Triticum aestivum L. emend. Thell.) in many parts of the world.When wheat is grown under waterlogged conditions, a reddish-brownmineral coating can form on the epidermal surface of the roots.In wetland plants such as rice, the amount of mineral coatingformed on root surfaces is positively related to yield. Thisstudy was conducted to determine whether mineral coating onthe roots of wheat is related to yield potential under waterloggedconditions. Root mineral coatings formed under waterlogged conditionswere studied in 12 cultivars and two breeding lines over threeyears of greenhouse pot studies. Soil redox potential in thewaterlogged treatment ranged between - 46 and 171 mV, and grainyield was suppressed by 28–49% compared to well-drainedcontrols. Mineral coating formed on the roots from the waterloggedtreatment was determined to be composed primarily of iron, basedon ICP elemental analysis, iron-specific staining, and ion-mappingby scanning electron microscopy using an X-ray detector. Of11 elements quantified by ICP spectroscopy, six were significantlyaffected by waterlogging treatment, and three of these, Fe,Mn and P, were well-correlated negatively to yield. Aerenchymaformation in the heavily coated waterlogged roots appeared todisrupt the internal root structure, and exceeded 40% of cross-sectionalarea in one cultivar. Unlike rice, which shows a positive relationshipbetween oxygen release from roots, grain yield and mineral coating,in winter wheat, the amount of mineral coating is negativelyrelated to grain yield under waterlogged conditions. Key words: Wheat, mineral coating, waterlogging, aerenchyma, hypoxia     

种植密度和施氮水平对小麦吸收利用土壤氮素的影响

2011-2013小麦季,在大田条件下设置2个氮肥水平(180和240kgN· hm-2)和3个种植密度(135、270和405万·hm-2),并将15N-尿素分别标记在20、60和100 cm土层处,研究种植密度-施氮互作对小麦吸收、利用土壤氮素及硝态氮残留量的影响.结果表明:种植密度从135万·hm-2增加至405万·hm-2,小麦在20、60和100 cm土层的15N吸收量分别增加1.86、2.28和2.51 kg·hm-2,地上部氮素积累量和吸收效率分别提高12.6％和12.6％,氮素利用效率降低5.4％;施氮量由240 kg N·hm-2降至180 kg N·hm-2,小麦在20、60 cm土层的15N吸收量分别降低4.11和1.21 kg·hm-2,在100 cm土层的15N吸收量增加1.02 kg·hm-2,地上部氮素积累量平均降低13.5％,氮素吸收效率和利用效率分别提高9.4％和12.2％.施氮180kg N·hm-2+种植密度为405万·hm-2处理与施氮240 kg N·hm-2+种植密度为270或405万·hm-2处理相比,其籽粒产量无显著差异,深层土壤氮素的吸收量显著提高,氮素吸收效率和利用效率分别提高13.4％和11.9％,O～ 200 cm土层的硝态氮积累量及100～ 200 cm土层硝态氮分布比例降低.在适当降低氮肥用量条件下,通过增加种植密度可以促进小麦吸收深层土壤氮素,减少土壤氮素残留,并保持较高的产量水平.     

Responses of photosynthetic characteristics and antioxidative metabolism in winter wheat to post-anthesis shading

In a field experiment, two winter wheat (Triticum aestivum L.) cultivars, Tainong 18 (a large-spike cultivar) and Jinan 17 (a multiple-spike cultivar), were treated with 78% (S1), 50% (S2), and 10% (S3) of full sunshine (S0, control) from anthesis to maturity to determine the responses of photosynthetic characteristics and antioxidative enzyme activities in a flag leaf. Compared with S0 treatment, the chlorophyll (Chl) content and maximal efficiency of photosystem II (PSII) photochemistry (F_v/F_m) of flag leaves were enhanced in treatments S1 and S2. From 0 to 7 d post flowering, the Chl content and F_v/F_m in S3 were also higher than those in S0, but significantly lower than those in controls, respectively. With the increase of shading intensity, the effective quantum yield of PSII (Φ_PSII) was promoted; whereas, the ratio of Chl a/b declined. Compared with S0, treatments S2 and S3 significantly suppressed the activities of superoxide dismutase (SOD) and peroxidase (POD), net photosynthetic rate (P _N), and contents of total soluble sugar, nevertheless, S1 treatment showed positive effects on the above parameters. Under the same shading condition, Jinan 17 had larger Chl content and higher activities of PSII and antioxidative enzymes, but lower malondialdehyde (MDA) content than Tainong 18. The results indicated that multiple-spike cultivar was more advantageous for the Huang-Huai-Hai Plain, where shading problem occurs later during the growth period, than the large-spike cultivar, because of the lesser damage in a flag leaf and better photosynthetic function of the former one. Wheat plants under S1 shading condition had relatively high activities of antioxidative enzymes and a low degree of membrane lipid peroxidation, which was in favor of stress resistance, maintaining high P _N duration, and accumulation of photosynthates in wheat plants.     

Response of the root system of a winter wheat crop to waterlogging

This study was aimed at analysing and quantifying the response of the root system dynamics of a wheat crop to waterlogging. Two experiments were carried out in parallel: one under controlled conditions with semi-permanent water tables using lysimeters equipped with oxygen measurers, and the other under conditions of artificially drained plots by continually monitoring their hydraulic functioning. The root system was observed frequently using the root mapping method, and this made it possible to measure the growth of the root front, to estimate root densities, and infer growth indices from them. The results showed that the anoxic medium for wheat roots consisted of a water-saturated soil with an oxygen concentration of below a critical threshold estimated to be 0.12 mol m^–3 water. The results also showed that the area which was unfavourable to root growth corresponded to the water table topped with a capillary zone of approximately 6 cm. Once the critical threshold had been reached, it was the water-table duration that explained root behaviour and the first effects were perceptible after approximately 48 h. On the basis of these results, two stress variables were analysed: water-table duration in the root zone (WTD) and proportion of roots in the water table (RPWT). The RPWT variable gave the best results within the two experimental contexts. In the case of the permanent regime, this variable made it possible to consider the root proliferation observations made above the saturated zone. Equations linking the stress variable RPWT to the growth indices are proposed that offer new perspectives to modelling waterlogging effects.     

Proteome analysis of soybean roots under waterlogging stress at an early vegetative stage

To gain better insight into how soybean roots respond to waterlogging stress, we carried out proteomic profiling combined with physiological analysis at two time points for soybean seedlings in their early vegetative stage. Seedlings at the V2 stage were subjected to 3 and 7 days of waterlogging treatments. Waterlogging stress resulted in a gradual increase of lipid peroxidation and in vivo H₂O₂ level in roots. Total proteins were extracted from root samples and separated by two-dimensional gel electrophoresis (2-DE). A total of 24 reproducibly resolved, differentially expressed protein spots visualized by Coomassie brilliant blue (CBB) staining were identified by matrix assisted laser desorption ionization time-of-flight (MALDI-TOF) mass spectrometry or electrospray ionization tandem mass spectrometry (ESI-MS/MS) analysis. Of these, 14 proteins were upregulated; 5 proteins were decreased; and 5 were newly induced in waterlogged roots. The identified proteins include well-known classical anaerobically induced proteins as well as novel waterlogging-responsive proteins that were not known previously as being waterlogging responsive. The novel proteins are involved in several processes, i.e. signal transduction, programmed cell death, RNA processing, redox homeostasis and metabolisms of energy. An increase in abundance of several typical anaerobically induced proteins, such as glycolysis and fermentation pathway enzymes, suggests that plants meet energy requirement via the fermentation pathway due to lack of oxygen. Additionally, the impact of waterlogging on the several programmed cell death- and signal transduction-related proteins suggest that they have a role to play during stress. RNA gel blot analysis for three programmed cell death-related genes also revealed a differential mRNA level but did not correlate well with the protein level. These results demonstrate that the soybean plant can cope with waterlogging through the management of carbohydrate consumption and by regulating programmed cell death. The identification of novel proteins such as a translation initiation factor, apyrase, auxin-amidohydrolase and coproporphyrinogen oxidase in response to waterlogging stress may provide new insight into the molecular basis of the waterlogging-stress response of soybean.     

Productivity of a doubled haploid winter wheat population under heat stress

Breeding of new winter wheat cultivars with good heat tolerance requires better understanding of the genetic background of heat tolerance. In the present work the effect of heat stress on the 6^th day after heading was investigated in a doubled haploid (DH) population arising from a cross between heat-sensitive (Plainsman V) and heat-tolerant (Mv Magma) cultivars. Averaged over the population, heat stress was found to result in a significant reduction in biomass, grain yield and grain number per plant, and in thousand-kernel weight (TKW) and harvest index. High temperature had the greatest effect on the grain yield, with a drop of 36.2% compared with the control. This could be attributed jointly to significant reductions in the TKW of the main ear and in the grain number of the side tillers. The relationship between the yield parameters was confirmed by the positive correlations obtained for the lines in the population. The diverse levels of heat tolerance in the different lines were confirmed by the significant differences in the reduction in the chlorophyll content (SPAD index) of the flag-leaves and in yield parameters. The changes in yield components in stress condition, however, can be still the most effective tools for heat stress evaluation.     

Alterations in photosynthesis and antioxidant enzyme activity in winter wheat subjected to post-anthesis water-logging

Winter wheat (Triticum aestivum L.) cultivars Yangmai 9 (water-logging tolerant) and Yumai 34 (water-logging sensitive) were subjected to water-logging (WL) from 7 d after anthesis to determine the responses of photosynthesis and anti-oxidative enzyme activities in flag leaf. At 15 d after treatment (DAT), net photosynthetic rate under WL was only 3.7 and 8.9 μmol(CO₂) m⁻² s⁻¹ in Yumai 34 and Yangmai 9, respectively, which was much lower than in the control. Ratios of variable to maximum and variable to initial fluorescence, actual photosynthetic efficiency, and photochemical quenching were much lower, while initial fluorescence and non-photochemical quenching were much higher under WL than in control, indicating damage to photosystem 2. WL decreased activities of superoxide dismutase and catalase in both cultivars, and activity of peroxidase (POD) in Yumai 34, while POD activity in Yangmai 9 was mostly increased. The obvious decrease in the amount of post-anthesis accumulated dry matter, which was redistributed to grains, also contributed to the grain yield loss under WL.     

氮、硫肥与灌浆后期高温胁迫对小麦籽粒产量和品质的影响

灌浆期高温胁迫通常导致小麦籽粒产量降低、品质变差.为了探索有效缓解高温胁迫不利影响的有效措施,在大田条件下,以济麦20为试验材料,运用后期高温棚增温的方法,研究了氮、硫肥与灌浆后期高温对小麦籽粒产量和品质的影响.结果表明:高温处理后小麦千粒质量、籽粒产量显著降低(19.7%～28.3%),蛋白质含量升高,蛋白质组分中不溶性谷蛋白含量下降,可溶性谷蛋白含量上升,面团形成时间和稳定时间缩短;氮肥追施比例由50%增加到70%,非高温处理的籽粒产量无显著变化,不溶性谷蛋白质含量和谷蛋白聚合指数(不溶性谷蛋白含量/可溶性谷蛋白含量)升高,面团形成时间和稳定时间延长;高温处理的千粒质量和籽粒产量因氮肥追施比例提高而提高(2.07%～3.58%),面团形成时间和稳定时间缩短;不论高温处理还是非高温处理,与追施硫肥相比,硫肥基施提高籽粒产量、谷蛋白聚合指数,延长面团形成时间和稳定时间;为了有效缓解灌浆后期高温对小麦籽粒产量和品质的不利影响,硫肥基施、氮肥施用基追比50%:50%是较为适宜的施肥模式.     

有机肥对铅胁迫下小麦生长的影响

采用盆栽方法,研究了不同程度铅胁迫条件下施用有机肥对小麦生长的影响.结果表明:无论施肥与否,随着铅胁迫程度的加剧,小麦均表现出株高下降、次生根数减少、总根质量下降、总根长缩短、根系活力和吸收面积降低、根系SOD和POD活性下降、MDA含量增高的趋势,但在不同施肥处理下的下降幅度不同.施用有机肥可以不同程度地缓解铅胁迫对小麦生长的影响,延缓小麦根系衰老,促进根系发育与生长,最终使小麦产量增加,籽粒中的铅含量降低.     

Effects of eyespot on the yield of winter wheat

In two experiments, on Julia and Zephyr spring barley, evidence was obtained that the development of powdery mildew in sprayed plots depended very much on their proximity to unsprayed crop. The effective period of control achieved with one spray was greater where plots were well separated from unsprayed crop than where they were surrounded by or very close to unsprayed crop. There is probably a risk that small plot experiments may not only underestimate the potential effectiveness of fungicides in agricultural practice, but also give misleading information on optimal spray timing.     

种植密度对冬小麦根系时空分布和氮素利用效率的影响

在大田条件下,以大穗型品种泰农18和中穗型品种山农15为材料,研究不同种植密度(泰农18:每公顷135、270、405万株;山农15:每公顷172.5、345、517.5万株)对冬小麦根系时空分布和氮素利用效率的影响.结果表明:在整个生育期,随种植密度的增加,泰农18的根长密度、根系总吸收面积和活跃吸收面积均显著增加;在生育后期,山农15的根长密度、根系总吸收面积和活跃吸收面积在种植密度为每公顷345万株时最大.泰农18的籽粒产量、氮肥吸收利用效率、氮肥偏生产力和氮素利用效率在种植密度为每公顷405万株时最高,山农15在种植密度为每公顷345万株时最高,但与种植密度为每公顷517.5万株的处理差异不显著.随种植密度的增加,冬小麦成熟期土壤硝态氮、铵态氮和无机态氮在不同土层的积累量均降低.泰农18和山农15种植密度分别为每公顷405万株和345万株时,是兼顾高产和高效利用氮素的适宜种植密度.     

Effects of different irrigation modes on winter wheat grain yield and water- and nitrogen use efficiency

Taking the widely planted winter wheat cultivar Tainong 18 as test material, a field experiment was conducted to study the effects of different irrigation modes on the winter wheat grain yield and water- and nitrogen use efficiency in drier year (2009-2010) in Tai' an City of Shandong Province, China. Five treatments were installed, i. e., irrigation before sowing (CK), irrigation before sowing and at jointing stage (W1), irrigation before sowing and at jointing stages and at over-wintering stage with alternative irrigation at milking stage (W2), irrigation before sowing and at jointing and flowering stages (optimized traditional irrigation mode, W3), and irrigation before sowing and at over-wintering, jointing, and milking stages (traditional irrigation mode, W4). The irrigation amount was 600 m3 hm(-2) one time. Under the condition of 119.7 mm precipitation in the winter wheat growth season, no significant difference was observed in the grain yield between treatments W2 and W4, but the water use efficiency was significantly higher in W2 than in W4. Comparing with treatment W3, treatments W2 and W4 had obviously higher grain yield, but the water use efficiency had no significant difference. The partial factor productivity from N fertilization was the highest in W2 and W4, and the NO3(-)-N accumulation amount in 0-100 cm soil layer at harvest was significantly higher in W2 than in W3 and W4, suggesting that W2 could reduce NO3(-)-N leaching loss. Under the conditions of our experiment, irrigation before sowing and jointing stages and at over-wintering stage with alternative irrigation at milking stage was the optimal irrigation mode in considering both the grain yield and the water- and nitrogen use efficiency.     

Photosynthesis and growth of winter wheat in response to waterlogging at different growth stages

A study on photosynthetic and yield effects of waterlogging of winter wheat at four stages of growth was conducted in specially designed experimental tanks during the 2007–2008 and 2008–2009 seasons. Compared with the control, waterlogging treatments at tillering and jointing-booting stages reduced photosynthetic rate (P _N) and transpiration (E) significantly, it also decreased average leaf water-use efficiency (WUE, defined as the ratio of P _N to E) by 3.3% and 3.4% in both years. All parameters returned quickly to the control level after soil was drained. Damage to the photosynthetic apparatus during waterlogging resulted in a lower F_v/F_m ratio, especially at the first two stages. A strong reduction in root length, root mass, root/shoot ratio, total dry mass, and leaf area index were observed. The responses from vegetative plants at tillering and jointing-booting stages were greater than in generative plants at onset of flowering and at milky stages. The number of panicles per hectare at tillering stage and the spikelet per panicle at the stages of jointing-booting and at onset of flowering were also significantly reduced by waterlogging, giving 8.2–11.3% decrease of the grain yield relative to the control in both years. No significant difference in yield components and a grain yield was observed between the control and treatments applied at milky stages. These responses, modulated by the environmental conditions prevailing during and after waterlogging, included negative effects on the growth, photosynthetic apparatus, and the grain yield in winter wheat, but the effect was strongly stage-dependent.     

Effects of water shortage on the yield of winter wheat

The effects of water shortage on the yield of winter wheat (cv. Maris Huntsman) were investigated in covered field plots on silt-loam soil. Plants were subjected to drought at different stages of growth, but none of the treatments significantly decreased yield. Some treatments decreased the number of ears per unit area, and number of grains per ear, but these effects were offset by increases in individual grain weight. We estimate that the soil water deficit (151 mm) in the treatment which had the largest effect on yield, albeit non-significant, would be expected in nature about 15 years in 100 or less. Drought also had little effect on plant water potential and stomatal conductance. The lack of effect of drought on plant water status and yield may be attributable to the large available water content of the silt loam and the deep rooting of winter wheat. On this type of soil, therefore, water shortage appears not to be an important cause of variation in the yield of winter wheat.     

外源脯氨酸对镉胁迫下小麦幼苗生长的影响

以高蛋白小麦品种“北农9549”为试材,研究喷施不同浓度脯氨酸(0、1.0、5.0和10.0 mmol·L-1)对镉胁迫下小麦幼苗生长和重金属吸收的影响.结果表明:以不施镉为对照,1.0 mmol·L-1CdCl2胁迫下,小麦幼苗的根长、株高和干质量分别显著下降24.0％、15.0％和27.5％,叶绿素a、b和类胡萝卜素含量分别显著下降23.3％、6.7％和30.8％,超氧化物歧化酶(SOD)活性降低了18.4％,内源脯氨酸、抗坏血酸和丙二醛(MDA)含量分别显著上升78.6％、31.5％和17.9％,细胞膜相对透性显著升高24.8％,过氧化物酶(POD)活性为对照的2.4倍,并且促进对铜的吸收,抑制锌的吸收.随外源脯氨酸浓度的增加,小麦幼苗的根长、株高、干质量、叶绿素和类胡萝卜素含量均逐渐恢复到对照水平,抗坏血酸、内源游离脯氨酸含量和SOD活性均上升,可溶性蛋白含量先上升后下降,POD活性、MDA含量和细胞膜相对透性下降,而锌积累量升高,镉、铜积累量下降.叶面喷施外源脯氨酸可缓解镉对小麦幼苗生长的胁迫,以喷施5.0～10.0 mmol·L-1外源脯氨酸效果最佳.     

Antioxidative defense system in pigeonpea roots under waterlogging stress

Pigeonpea [Cajanus cajan (L.) Millsp.] is a waterlogging-sensitive legume crop. We studied the effect of waterlogging stress on hydrogen peroxide (H₂O₂) content, lipid peroxidation and antioxidant enzyme activities in two pigeonpea genotypes viz., ICPL-84023 (waterlogging resistant) and MAL-18 (waterlogging susceptible). In a pot experiment, waterlogging stress was imposed for 6 days at early vegetative stage (20 days after sowing). Waterlogging treatment significantly increased hydrogen peroxide accumulation and lipid peroxidation, which indicated the extent of oxidative injury posed by stress conditions. Enzyme activities of peroxidase (POX), catalase (CAT), ascorbate peroxidase (APX), superoxide dismutase (SOD) and polyphenol oxidase (PPO) increased in pigeonpea roots as a consequence of waterlogged conditions, and all the enzyme activities were significantly higher in waterlogged ICPL-84023 than in MAL-18. POX activity was the maximum immediately after imposing stress, therefore, it was suggested to be involved in early scavenging of H₂O₂, while rest of the enzymes (CAT, APX, SOD and PPO) were more important in late responses to waterlogging. Present study revealed that H₂O₂ content is directly related to lipid peroxidation leading to oxidative damage during waterlogging in pigeonpea. Higher antioxidant potential in ICPL-84023 as evidenced by enhanced POX, CAT, APX, SOD and PPO activities increased capacity for reactive oxygen species (ROS) scavenging and indicated relationship between waterlogging resistance and antioxidant defense system in pigeonpea.