Differences in Sugar Accumulation and Mobilization between Sequential and Non-Sequential Senescence Wheat Cultivars under Natural and Drought Conditions.
ABSTRACT: Wheat leaf non-sequential senescence at the late grain-filling stage involves the early senescence of younger flag leaves compared to that observed in older second leaves. On the other hand, sequential senescence involves leaf senescence that follows an age-related pattern, in which flag leaves are the latest to undergo senescence. The characteristics of sugar metabolism in two sequential senescence cultivars and two non-sequential senescence cultivars under both natural and drought conditions were studied to elucidate the underlying mechanism of drought tolerance in two different senescence modes. The results showed that compared to sequential senescence wheat cultivars, under natural and drought conditions, non-sequential senescence wheat cultivars showed a higher leaf net photosynthetic rate, higher soluble sugar levels in leaves, leaf sheaths, and internodes, higher leaf sucrose synthase (SS) and sucrose phosphate synthase (SPS) activity, and higher grain SS activity, thereby suggesting that non-sequential senescence wheat cultivars had stronger source activity. Spike weight, grain weight per spike, and 100-grain weight of non-sequential senescence cultivars at maturity were significantly higher than those of sequential senescence cultivars under both natural and drought conditions. These findings indicate that the higher rate of accumulation and the higher mobilization of soluble sugar in the leaves, leaf sheaths and internodes of non-sequential senescence cultivars improve grain weight and drought tolerance. At the late grain-filling stage, drought conditions adversely affected leaf chlorophyll content, net photosynthetic rate, soluble sugar and sucrose content, SS and SPS activity, gain SS activity, and weight. This study showed that higher rates of soluble sugar accumulation in the source was one of the reasons of triggering leaf non-sequential senescence, and higher rates of soluble sugar mobilization during leaf non-sequential senescence promoted high and stable wheat yield and drought tolerance.
Project description:We report a case study of natural variations and correlations of some photosynthetic parameters, green biomass and grain yield in Cappelle Desprez and Plainsman V winter wheat (Triticum aestivum L.) cultivars, which are classified as being drought sensitive and tolerant, respectively. We monitored biomass accumulation from secondary leaves in the vegetative phase and grain yield from flag leaves in the grain filling period. Interestingly, we observed higher biomass production, but lower grain yield stability in the sensitive Cappelle cultivar, as compared to the tolerant Plainsman cv. Higher biomass production in the sensitive variety was correlated with enhanced water-use efficiency. Increased cyclic electron flow around PSI was also observed in the Cappelle cv. under drought stress as shown by light intensity dependence of the ratio of maximal quantum yields of Photosystem I and Photosystem II, as well by the plot of the Photosystem I electron transport rate as a function of Photosystem II electron transport rate. Higher CO2 uptake rate in flag leaves of the drought-stressed Plainsman cv. during grain filling period correlates well with its higher grain yield and prolonged transpiration rate through spikes. The increase in drought factor (DFI) and performance (PI) indices calculated from variable chlorophyll fluorescence parameters of secondary leaves also showed correlation with higher biomass in the Cappelle cultivar during the biomass accumulation period. However, during the grain filling period, DFI and PI parameters of the flag leaves were higher in the tolerant Plainsman V cultivar and showed correlation with grain yield stability. Our results suggest that overall biomass and grain yield may respond differentially to drought stress in different wheat cultivars and therefore phenotyping for green biomass cannot be used as a general approach to predict grain yield. We also conclude that photosynthetic efficiency of flag and secondary leaves is correlated with grain yield and green biomass, respectively. In addition, secondary trait associated mechanisms like delayed senescence and higher water-use efficiency also contribute to biomass stability. Our studies further prove that photosynthetic parameters could be used to characterize environmental stress responses.
Project description:Drought is a major environmental stress that severely restricts plant growth and crop productivity. A previous study showed that TaWRKY2 from wheat (Triticum aestivum) plays an important role in drought stress tolerance. In the present study, we isolated the promoter of TaWRKY2 and identified multiple regulatory cis-elements in the promoter region. The activity of the TaWRKY2 promoter was induced by drought, salt, heat, and abscisic acid (ABA). We also generated TaWRKY2-overexpressing transgenic wheat, and found that the transgenic seedlings exhibited significantly enhanced tolerance to drought stress, as evidenced by a higher survival rate and lower water loss rate of detached leaves compared with wild type (WT) plants. In addition, the transgenic lines had higher contents of free proline, soluble sugar, and chlorophyll. During a prolonged period of drought stress before the heading stage, the growth of WT plants was inhibited, whereas the TaWRKY2-overexpressing lines progressed to the heading stage. The increased grain yield of the transgenic wheat lines reflected the cumulative effects of longer panicle length, more kernels per spike, and greater aboveground biomass. Our findings show that TaWRKY2 can enhance drought tolerance and increase grain yield in wheat, thus providing a promising candidate target for improving the drought tolerance of wheat cultivars through genetic engineering.
Project description:In the Mediterranean-type environment of Australia and other parts of the world, end-of- season or terminal drought is the most significant abiotic stress affecting wheat grain yields. This study examined the response of two wheat cultivars with contrasting root system size to terminal drought and the effect of terminal drought on grain yield and yield components. The cultivars were grown in 1.0 m deep PVC columns filled with soil in a glasshouse under well-watered conditions until the onset of ear emergence (Z51) when well-watered and terminal drought treatments were imposed. Terminal drought reduced stomatal conductance, leaf photosynthesis, and transpiration rates faster in Bahatans-87 (larger root system size) than Tincurrin (smaller root system size). Terminal drought reduced grain yield in both cultivars, more so in Bahatans-87 (80%) with the large root system than Tincurrin (67%) with the small root system, which was mainly due to a reduction in grain number and grain size in Bahatans-87 and grain size in Tincurrin. In the terminal drought treatment, Bahatans-87 had 59% lower water use efficiency than Tincurrin, as Bahatans-87 used 39% more water and reduced grain yield more than Tincurrin. The lesser reduction in grain yield in Tincurrin was associated with slower water extraction by the small root system and slower decline in stomatal conductance, leaf photosynthesis, and transpiration rates, but more importantly to faster phenological development, which enabled grain filling to be completed before the severe effects of water stress.
Project description:Nitrogen (N) fertilizer represents a significant cost for the grower and may also have environmental impacts through nitrate leaching and N2O (a greenhouse gas) emissions associated with denitrification. The objectives of this study were to quantify the genetic variability in N partitioning and N remobilization in Indian spring wheat cultivars and identify traits for improved grain yield and grain protein content for application in breeding N-efficient cultivars. Twenty-eight bread wheat cultivars and two durum wheat cultivars were tested in field experiments in two years in Maharashtra, India. Growth analysis was conducted at anthesis and harvest to assess above-ground dry matter (DM) and dry matter and N partitioning. Flag-leaf photosynthesis rate (A max ), flag-leaf senescence rate and canopy normalized difference vegetation index (NDVI) were also assessed. Significant N?×?genotype level interaction was observed for grain yield and N-use efficiency. There was a positive linear association between post-anthesis flag-leaf A max and grain yield amongst the 30 genotypes under high N (HN) conditions. Flag-leaf A max was positively associated with N uptake at anthesis (AGNA). Under both HN and low N (LN) conditions, higher N uptake at anthesis was associated with delayed onset of flag-leaf senescence and higher grain yield. Under N limitation, there was a genetic negative correlation between grain yield and grain protein concentration. Deviation from this negative relationship (grain protein deviation or GPD) was related to genotypic differences in post-anthesis N uptake. It is concluded that N uptake at anthesis was an important determinant of flag-leaf photosynthesis rate and grain yield under high N conditions; while post-anthesis N uptake was an important determinant of GPD of wheat grown under low to moderate N conditions in India.
Project description:Active and passive sensors are available for ground-based, high-throughput phenotyping in the field. However, these sensor systems have seldom been compared with respect to their determination of plant water status and water use efficiency related parameters under drought conditions. In this study, five passive and active reflectance sensors, including a hyperspectral passive sensor, an active flash sensor (AFS), the Crop Circle, and the GreenSeeker, were evaluated to assess drought-related destructive and non-destructive morphophysiological parameters (ground cover, relative leaf water content, leaf temperature, and carbon isotope discrimination of leaves and grain) and grain yield of twenty wheat (Triticum aestivum L.) cultivars. Measurements were conducted in a 2-year study, including a drought stress and a control environment under field conditions. A comparison of the active sensors at the heading, anthesis and grain-filling stages indicated that the Crop Circle provided the most significant and robust relationships with drought-related parameters (relative leaf water content and leaf and grain carbon isotope discrimination). In comparison with the passive sensor, the five water and normalized water indices (WI and NWI-1 to 4), which are only provided by the passive sensor, showed the strongest relationships with the drought stress-related parameters (r = -0.49 to -0.86) and grain yield (r = -0.88) at anthesis. This paper indicates that precision phenotyping allows the integration of water indices in breeding programs to rapidly and cost-effectively identify drought-tolerant genotypes. This is supported by the fact that grain yield and the water indices showed the same heritability under drought conditions.
Project description:Drought stress induced pollen sterility is a detrimental factor reducing grain number in wheat. Exploring the mechanisms underlying pollen fertility under drought conditions could assist breeding high-yielding wheat cultivars with stress tolerance. Here, by using two Chinese wheat cultivars subjected to different levels of polyethylene glycol (PEG)-induced drought stress, possible links between pollen fertility and stress tolerance were analyzed under different levels of drought stress at the young microspore stage. In both cultivars, higher grain number reduction was observed under condition of lower water availability. Overall, the drought tolerant cultivar (Jinmai47) exhibited less grain number reduction than the drought sensitive cultivar (Shiluan02-1) under all stress conditions. Compared with Shiluan02-1, Jinmai47 exhibited superior physiological performance in terms of leaf photosynthetic rate, ear carbohydrate accumulation, pollen sink strength, pollen development and fertility under stress. Moreover, Jinmai47 showed a lower increase in endogenous abscisic acid in ears than Shiluan02-1. Furthermore, higher levels of superoxide dismutase (SOD) and peroxidase (POD) activities were also found in the drought tolerant cultivar Jinmai47 under PEG stress, compared with the drought sensitive cultivar Shiluan02-1. Changes in these physiological traits could contribute to better pollen development and male fertility, ultimately leading to the maintenance of grain number under drought stress.
Project description:Previous studies have shown that wheat grain yield is seriously affected by drought stress, and leaf cuticular wax is reportedly associated with drought tolerance. However, most studies have focused on cuticular wax biosynthesis and model species. The effects of cuticular wax on wheat drought tolerance have rarely been studied. The aims of the current study were to study the effects of leaf cuticular wax on wheat grain yield under drought stress using the above-mentioned wheat NILs and to discuss the possible physiological mechanism of cuticular wax on high grain yield under drought stress. Compared to water-irrigated (WI) conditions, the cuticular wax content (CWC) in glaucous and non-glaucous NILs under drought-stress (DS) conditions both increased; mean increase values were 151.1 and 114.4%, respectively, which was corroborated by scanning electronic microscopy images of large wax particles loaded on the surfaces of flag leaves. The average yield of glaucous NILs was higher than that of non-glaucous NILs under DS conditions in 2014 and 2015; mean values were 7368.37 kg·ha<sup>-1</sup> and 7103.51 kg·ha<sup>-1</sup>. This suggested that glaucous NILs were more drought-tolerant than non-glaucous NILs (<i>P</i> = 0.05), which was supported by the findings of drought tolerance indices TOL and SSI in both years, the relatively high water potential and relative water content, and the low ELWL. Furthermore, the photosynthesis rate (<i>P<sub>n</sub></i> ) of glaucous and non-glaucous wheat NILs under DS conditions decreased by 7.5 and 9.8%, respectively; however, glaucous NILs still had higher mean values of <i>P<sub>n</sub></i> than those of non-glaucous NILs, which perhaps resulted in the higher yield of glaucous NILs. This could be explained by the fact that glaucous NILs had a smaller <i>F<sub>v</sub>/F<sub>m</sub></i> reduction, a smaller <i>PI</i> reduction and a greater <i>ABS/RC</i> increase than non-glaucous NILs under DS conditions. This is the first report to show that wheat cuticular wax accumulation is associated with drought tolerance. Moreover, the leaf CWC can be an effective selection criterion in the development of drought-tolerant wheat cultivars.
Project description:In order to clarify the transcriptional regulatory network and physiological mechanisms governing leaf senescence response to drought stress in wheat, experiments were performed using two wheat varieties with contrasting drought tolerance: Fu287 (F287, a drought-sensitive genotype) and Shannong20 (SN20, a drought-resistant genotype). The latter has higher SPAD values, salicylic acid (SA), jasmonic acid (JA), zeatin (Z), zeatin riboside (ZR), and gibberellin (GA 3) content as well as higher expression levels of Cu/Zn-SOD, Mn-SOD, Fe-SOD,POD,CAT, and APX under various water deficit conditions. Conjoint analysis of physiological and biochemical indicators and transcriptome data by weighted gene co-expression network analysis (WGCNA) in the present study provides a useful genomic and molecular resource for studying drought adaptation in wheat. The flag leaf senescence process was changed by altering the concentration of phytohormones. SA, JA, abscisic acid (ABA), Z, ZR, and GA 3 coordinate with each other to control leaf senescence and plant adaptation under drought stress. Further, the leaf senescence process was divided into two phases: the persistence phase and the rapid loss phase. Shorter Chltotal (duration of the flag leaf being photosynthetically active), shorter Chlper (persistence phase), reduced M (inflection point cumulative temperature when senescence rate is the maximum), decreased r max (the maximum senescence rate), larger r 0 (the initial senescence rate), and increased r aver (the average senescence rate) were slightly associated with low grain mass. We speculated that extending the period of the persistence phase by cultivation or chemical control measures could further increase the drought survivability and productivity of wheat.
Project description:Drought stress triggers mature leaf senescence, which supports plant survival and remobilization of nutrients; yet leaf senescence also critically decreases post-drought crop yield. Drought generally results in carbon/nitrogen imbalance, which is reflected in the increased carbon:nitrogen (C:N) ratio in mature leaves, and which has been shown to be involved in inducing leaf senescence under normal growth conditions. Yet the involvement of the carbon/nitrogen balance in regulation of drought-induced leaf senescence is unclear. To investigate the role of carbon/nitrogen balance in drought-induced senescence, sorghum seedlings were subjected to a gradual soil drought treatment. Leaf senescence symptoms and the C:N ratio, which was indicated by the ratio of non-structural carbohydrate to total N content, were monitored during drought progression. In this study, leaf senescence developed about 12 days after the start of drought treatment, as indicated by various senescence symptoms including decreasing photosynthesis, photosystem II photochemistry efficiency (Fv/Fm) and chlorophyll content, and by the differential expression of senescence marker genes. The C:N ratio was significantly enhanced 10 to 12 days into drought treatment. Leaf senescence occurred in the older (lower) leaves, which had higher C:N ratios, but not in the younger (upper) leaves, which had lower C:N ratios. In addition, a detached leaf assay was conducted to investigate the effect of carbon/nitrogen availability on drought-induced senescence. Exogenous application of excess sugar combined with limited nitrogen promoted drought-induced leaf senescence. Thus our results suggest that the carbon/nitrogen balance may be involved in the regulation of drought-induced leaf senescence.
Project description:In the present study we analyzed the responses of wheat to mild salinity and drought with special emphasis on the so far unclarified interaction of these important stress factors by using high-throughput phenotyping approaches. Measurements were performed on 14 genotypes of different geographic origin (Austria, Azerbaijan, and Serbia). The data obtained by non-invasive digital RGB imaging of leaf/shoot area reflect well the differences in total biomass measured at the end of the cultivation period demonstrating that leaf/shoot imaging can be reliably used to predict biomass differences among different cultivars and stress conditions. On the other hand, the leaf/shoot area has only a limited potential to predict grain yield. Comparison of gas exchange parameters with biomass accumulation showed that suppression of CO2 fixation due to stomatal closure is the principal cause behind decreased biomass accumulation under drought, salt and drought plus salt stresses. Correlation between grain yield and dry biomass is tighter when salt- and drought stress occur simultaneously than in the well-watered control, or in the presence of only salinity or drought, showing that natural variation of biomass partitioning to grains is suppressed by severe stress conditions. Comparison of yield data show that higher biomass and grain yield can be expected under salt (and salt plus drought) stress from those cultivars which have high yield parameters when exposed to drought stress alone. However, relative yield tolerance under drought stress is not a good indicator of yield tolerance under salt (and salt plus drought) drought stress. Harvest index of the studied cultivars ranged between 0.38 and 0.57 under well watered conditions and decreased only to a small extent (0.37-0.55) even when total biomass was decreased by 90% under the combined salt plus drought stress. It is concluded that the co-occurrence of mild salinity and drought can induce large biomass and grain yield losses in wheat due to synergistic interaction of these important stress factors. We could also identify wheat cultivars, which show high yield parameters under the combined effects of salinity and drought demonstrating the potential of complex plant phenotyping in breeding for drought and salinity stress tolerance in crop plants.