Effects of exogenous trehalose on filling characteristics and sugar component content of wheat under high temperature stress during the filling period.

Taking the heat-sensitive wheat variety 'Fanmai 5' (FM5) and the heat-tolerant variety 'Huaimai 33' (HM33), which were screened out in the previous experiments, as experimental materials, we conducted a field experiment with passive heat-enhancing shelters to simulate post-flowering high-temperature environment (average temperature increase of 5.13 ℃) during 2021-2022. During the filling period, we analyzed the effects of exogenous trehalose (10, 15 and 20 mmol·L-1) on the filling characteristics and sugar fraction under high temperature, with no spraying at ordinary temperature as control (CK). The results showed that treating without spraying exogenous trehalose at high temperature (H) significantly reduced wheat grain yield and grain weight during the filling period, and spraying exogenous trehalose alleviated the reduction of grain yield and grain weight at the filling stage under high temperature stress. Compared with the H treatment, grain yield and grain weight of HM33 and FM5 wheat varie-ties increased by 3.5%, 6.7% and 4.2%, 5.4%, respectively. High temperature stress significantly increased the trehalose content and trehalase (THL) activity in flag leaves of both wheat varieties, and decreased the fructose and glucose contents. Spraying exogenous trehalose increased the contents of trehalose, fructose, and glucose in flag leaves, and decreased the trehalase activity in flag leaves compared with H treatment, which could improve the glucose metabolism capacity of wheat at filling stage. The increasing effect of FM5 was higher than that of HM33. High temperature stress significantly reduced starch content of flag leaves and grains, while spraying exogenous trehalose alleviated the decrease of starch content of flag leaves and grains under high temperature stress, which was profit able for the substance accumulation of wheat grains under high temperature stress. Under the conditions of this experiment, spraying 15 mmol·L-1 trehalose at flowering stage was the best treatment for the two wheat varieties.

Low-temperature at booting reduces starch content and yield of wheat by affecting dry matter transportation and starch synthesis.

With the continuous change of global climate, the frequency of low-temperature stress (LTS) in spring increased greatly, which led to the increase of wheat yield decline. The effects of LTS at booting on grain starch synthesis and yield were examined in two wheat varieties with differing low-temperature sensitivities (insensitive variety Yannong 19 and sensitive variety Wanmai 52). A combination of potted and field planting was employed. For LTS treatment at booting, the wheat plants were placed in a climate chamber for 24 h at -2°C, 0°C or 2°C from 19:00 to 07:00 then 5°C from 07:00 to 19:00. They were then returned to the experimental field. The effects of flag leaf photosynthetic characteristics, the accumulation and distribution of photosynthetic products, enzyme activity related to starch synthesis and relative expression, the starch content, and grain yield were determined. LTS at booting caused a significant reduction in the net photosynthetic rate (Pn), stomatal conductance (Gs), and transpiration rate (Tr) of the flag leaves at filling. The development of starch grains in the endosperm is also hindere, there are obvious equatorial grooves observed on the surface of the A-type starch granules, and a reduction in the number of B-type starch granules. The abundance of 13C in the flag leaves and grains decreased significantly. LTS also caused a significant reduction in translocation amount of pre-anthesis stored dry matte from vegetative organs to grains and amount of post-anthesis transfer of accumulated dry matte into grains, and the distribution rate of dry matter in the grains at maturity. The grain filling time was shortened, and the grain filling rate decreased. A decrease in the activity and relative expression of enzymes related to starch synthesis was also observed, with a decrease in the total starch content. As a result, a decrease in the grain number per panicle and 1000-grain weight were also observed. These findings highlight the underlying physiological cause of decreased starch content and grain weight after LTS in wheat.

Exogenous 6-benzylaminopurine enhances waterlogging and shading tolerance after anthesis by improving grain starch accumulation and grain filling.

Due to the frequent occurrence of extreme weather events, the area of wheat affected by continuous cloudy rainfall is increasing, with waterlogging becoming a major limiting factor of wheat yield. To alleviate the effect, spraying exogenous plant growth regulators is often used. In this study, two wheat cultivars, waterlogging-tolerant Yangmai 18 and waterlogging-sensitive Sumai 188, were selected for waterlogging and shading (WS) after anthesis for 7, 11, and 15 days respectively. Three concentrations of 6-benzylaminoadenine (6-BA) solution (15, 25, and 35 mg·L-1) were sprayed after WS treatment and water was sprayed as the control. Then, the effect of spraying 6-BA on photosynthetic characteristics, starch content, grain filling characteristics, and yield was explored under artificially stimulated continuous cloudy rainfall during anthesis. Compared with the control, the application of 6-BA caused a significant increase in grain plumpness throughout grain filling, as well as increases in the net photosynthetic rate (P n), stomatal conductance (G s), and transpiration rate (T r), and a significant decrease in the intercellular CO2 concentration (C i) of the flag leaves, all of which enhanced the photosynthetic capacity. The content of total starch, amylose, and amylopectin in the grains also increased significantly compared with the control. After WS for 15 days, the starch content increased by 3.81%-11.41% compared with the control. Spraying 6-BA also prolonged grain filling, increased the average grain filling rate, and significantly increased the 1000-grain weight and yield. The thousand-grain weight increased by 5.06%-43.28%, and wheat yield increased by 8.93%-64.27% after spraying 25 mg·L-1 of the 6-BA solution. These findings suggest that the application of 6-BA after WS stress could significantly improve the photosynthetic performance, which is propitious to the accumulation and transport of photosynthetic products after anthesis. Besides, spraying 6-BA can also increase the duration and rate of grain filling and starch accumulation content and improve grain weight, thereby alleviating the adverse effects of WS on wheat yield. Overall, spraying 25 mg·L-1 of the 6-BA solution had an optimal effect. These findings provide a theoretical basis for the exploration of cultivation techniques and measures aimed at alleviating damage caused by continuous rainfall during wheat anthesis.

Post-flowering Soil Waterlogging Curtails Grain Yield Formation by Restricting Assimilates Supplies to Developing Grains.

Soil waterlogging is among the major factors limiting the grain yield of winter wheat crops in many parts of the world, including the middle and lower reaches of the Yangtze River China. In a field study, we investigated the relationship between leaf physiology and grain development under a varying duration of post-flowering waterlogging. A winter wheat cultivar Ningmai 13 was exposed to soil waterlogging for 0 (W0), 3 (W3), 6 (W6), and 9 d (W9) at anthesis. Increasing waterlogging duration significantly reduced flag leaf SPAD (soil plant analysis development) values and net photosynthetic rate (Pn). There was a linear reduction in flag leaf Pn and SPAD as plant growth progressed under all treatments; however, the speed of damage was greater in the waterlogged leaves. For example, compared with their respective control (W0), flag leaves of W9 treatment have experienced 46% more reduction in Pn at 21 d after anthesis (DAA) than at 7 DAA. Increasing waterlogging duration also induced oxidative damage in flag leaves, measured as malondialdehyde (MDA) contents. The capacity to overcome this oxidative damage was limited by the poor performance of antioxidant enzymes in wheat leaves. Inhibited leaf Pn and capacity to sustain assimilate synthesis under waterlogged environments reduced grain development. Compared with W0, W6 and W9 plants experienced a 20 and 22% reduction in thousand grain weight (TGW) in response to W6 and W9, respectively at 7 DAA and 11 and 19%, respectively at 28 DAA. Sustained waterlogging also significantly reduced grain number per spike and final grain yield. Averaged across two years of study, W9 plants produced 28% lesser final grain yield than W0 plants. Our study suggested that wheat crops are highly sensitive to soil waterlogging during reproductive and grain filling phases due to their poor capacity to recover from oxidative injury to photosynthesis. Management strategies such as planting time, fertilization and genotype selection should be considered for the areas experiencing frequent waterlogging problems.

Night warming at the vegetative stage improves pre-anthesis photosynthesis and plant productivity involved in grain yield of winter wheat.

We conducted pot experiments during the 2018-2020 growing seasons to study the effects of night warming at different growth stages of wheat on the photosynthetic performance; accumulation, transportation, and distribution of dry matter; and grain yield of winter wheat. Night warming at all the different growth stages resulted in an elevation of wheat yield by increasing the 1000-grain weight and the number of grains per ear. Night warming during the period from jointing to booting stage resulted in the greatest increase in wheat yield. It also increased the amount of overall dry matter and transferrable amount of dry matter in plants and increased the distribution of dry matter to grains to increase grain weight. Night warming treatments at three different growth stages enhanced pre-anthesis photosynthetic capacity by increasing flag leaf net photosynthetic rate, chlorophyll content, and photochemical efficiency of winter wheat at the early stage of grain filling, especially in the night warming treatment from jointing to booting stage. Night warming not only increased the stomatal density and stomatal index of wheat leaves but also increased stomatal conductance and transpiration rate in the early stage of grain filling, thus being conducive to the smooth progress of photosynthesis. In conclusion, night warming treatment at different growth stages increased the photosynthesis of flag leaves at the early stage of grain filling, and promoted the accumulation of dry matter in plants after anthesis, which was conducive to the grain yield of winter wheat.

Night-Warming Priming at the Vegetative Stage Alleviates Damage to the Flag Leaf Caused by Post-anthesis Warming in Winter Wheat (Triticum aestivum L.).

The asymmetric warming in diurnal and seasonal temperature patterns plays an important role in crop distribution and productivity. Asymmetric warming during the early growth periods of winter wheat profoundly affects its vegetative growth and post-anthesis grain productivity. Field experiments were conducted on winter wheat to explore the impact of night warming treatment in winter (Winter warming treatment, WT) or spring (Spring warming treatment, ST) on the senescence of flag leaves and yield of wheat plants later treated with night warming during grain filling (Warming treatment during grain filling, FT). The results showed that FT decreased wheat yield by reducing the number of grains per panicle and per 1,000-grain weight and that the yield of wheat plants treated with FT declined to a greater extent than that of wheat plants treated with WT + FT or ST + FT. The net photosynthetic rate, chlorophyll content, and chlorophyll fluorescence parameters of the flag leaves of wheat plants treated with WT + FT or ST + FT were higher than those under the control treatment from 0 to 7 days after anthesis (DAA) but were lower than those under the control treatment and higher than those of wheat plants treated with FT alone from 14 to 28 DAA. The soluble protein and Rubisco contents in the flag leaves of wheat plants treated with WT + FT or ST + FT were high in the early grain-filling period and then gradually decreased to below those of the control treatment. These contents were greater in wheat plants treated with WT + FT than in wheat plants treated with ST + FT from 0 to 14 DAA, whereas the opposite was true from 21 to 28 DAA. Furthermore, WT + FT and ST + FT inhibited membrane lipid peroxidation by increasing superoxide dismutase and peroxidase activities and lowering phospholipase D (PLD), phosphatidic acid (PA), lipoxygenase (LOX), and free fatty acid levels in the early grain-filling period, but their inhibitory effects on membrane lipid peroxidation gradually weakened during the late grain-filling period. Night-warming priming alleviated the adverse effect of post-anthesis warming on yield by delaying the post-anthesis senescence of flag leaves.

The Effects of Short-Term Exposure to Low Temperatures During the Booting Stage on Starch Synthesis and Yields in Wheat Grain.

Low temperatures (LT) in spring can have a major impact on the yields of wheat in winter. Wheat varieties with different cold sensitivities (the cold-tolerant Yannong 19 variety and the cold-sensitive Yangmai 18 variety) were used to study the responses of the wheat grain starch synthesis and dry material accumulation to short-term LT during the booting stage. The effects of short-term LT on the activities of key wheat grain starch synthesis enzymes, starch content and grain dry-matter accumulation were determined by exposing the wheat to simulated LT of from -2 to 2°C. Short-term LT stress caused a decrease in the fullness of the wheat grains along with decreased activities of adenosine diphosphate glucose pyrophosphorylase (AGPase, EC2.7.7.27), soluble starch synthase (SSS, EC2.4.1.21), granule-bound starch synthase (GBSS, EC2.4.1.21), and starch branching enzyme (SBE, EC2.4.1.18) at different spike positions during the filling stage. The rate of grain starch accumulation and starch content decreased with decreasing temperatures. Also, the duration of grain filling increased, the mean and the maximum filling rates were reduced and the quality of the grain dry-matter decreased. The number of grains per spike and the thousand-grain weight of the mature grains also decreased. Our data showed that short-term LT stress at the booting stage caused a decrease in the activities of key starch synthesis enzymes at the grain-filling stage. These changes reduced the accumulation of starch, decreased the filling rate, and lowered the accumulation of grain dry matter to ultimately decrease grain yields.

Effects of Low Temperature at Booting Stage on Sucrose Metabolism and Endogenous Hormone Contents in Winter Wheat Spikelet.

Low spring temperatures often occur during the winter wheat booting stage, when the young ears are very sensitive to cold. In this study, we used two wheat varieties differing in cold sensitivity (sensitive variety Yangmai 18 and tolerant variety Yannong 19) to examine the effect of low temperature on wheat grain number at booting stage. Low temperature stress was simulated in an artificial climate chamber at 4°C for 60 h in 2016 and at 2, 0, or -2°C for 24 h in morphological assays, showing that the development of wheat spikelets was inhibited and floret growth was delayed following low temperature stress. However, an increase in the sucrose content of young panicles was also observed, and the activity of enzymes involved in sucrose metabolism was dynamically altered. Sucrose phosphate synthase activity was enhanced, and sucrose synthase activity significantly increased after treatment at 4 and 2°C, respectively. However, activities of sucrose synthase and invertase decreased with a reduction in temperature. Gene expression assays further revealed downregulation of TaSuS1 expression and upregulation of TaSuS2, while expression of CWINV was inhibited. Moreover, phytohormone content assays showed an increase in the content of abscisic acid in young wheat ears, but a decrease in the content of auxin and gibberellins. The grain number per spike and 1000-grain weight also showed a downward trend following low temperature stress. Overall, these findings suggest that low temperature at booting induces abscisic acid accumulation in winter wheat, altering the activity of the enzymes involved in sucrose metabolism, which leads to an accumulation of sucrose in the young ears, thereby having a negative effect on wheat production.

Heat Priming During Early Reproductive Stages Enhances Thermo-Tolerance to Post-anthesis Heat Stress via Improving Photosynthesis and Plant Productivity in Winter Wheat (Triticum aestivum L.).

Heat stress during grain filling substantially decreases wheat productivity; thus, to ensure food security, heat tolerance in wheat needs to be developed. In this study, we evaluated the effect of heat priming applied during the stem-elongation stage, booting and anthesis, followed by 5 days of severe heat stress (a 7.86°C rise in temperature) during the grain-filling stage on physiological activities and grain yield of winter wheat in pot experiments during the 2015-2017 growing seasons using the winter wheat cultivars Yangmai 18 (a vernal type) and Yannong 19 (a facultative type). Compared with the damage observed in non-primed plants, heat priming during the stem-elongation stage and booting significantly prevented the grain-yield damage caused by heat stress during grain filling. Heat-primed plants displayed higher sucrose contents and sucrose-phosphate activity in leaves and greater above-ground dry matter than non-primed plants. Priming during stem elongation and booting led to increased photosynthetic capacity, stomatal conductance and chlorophyll contents in comparison with non-priming. Improved tolerance to heat stress due to the enhanced activities of antioxidant enzymes superoxide dismutase and peroxidase and reductions in reactive oxygen species and malondialdehyde production was observed in primed plants compared with non-primed plants of both cultivars. The positive effect of heat priming on the response to heat stress during grain filling was more pronounced in plants primed at the booting stage than in those primed at the stem-elongation or anthesis stage. Moreover, the vernal-type Yangmai 18 benefited more from heat priming than did Yannong 19, as evidenced by its higher productivity. We conclude that heat priming during early reproductive-stage growth can improve post-anthesis heat tolerance in winter wheat.

Winter Night-Warming Improves Post-anthesis Physiological Activities and Sink Strength in Relation to Grain Filling in Winter Wheat (Triticum aestivum L.).

The diurnal and seasonal temperature rising patterns "asymmetric warming," plays an important role in crop distribution and productivity. Asymmetric warming during the early growth periods of winter wheat (Triticum aestivum L.) profoundly affects vegetative growth and post-anthesis grain productivity, but the underlying physiological mechanism is still unclear. We conducted field experiments from 2012 to 2014 using two wheat cultivars, namely, Yangmai-13 (vernal type) and Yannong-19 (semi-winter type), to investigate the influences of night-warming during the winter (warming by 1.56-1.67°C from tillering to jointing) or during the spring (warming by 1.78-1.92°C from jointing to booting) on post-anthesis physiological activities and grain-filling processes. Both night-warming treatments enhanced the source activity by increasing flag leaf area, chlorophyll content, and photosynthetic capability in both cultivars compared with those of the control. The night-warming treatments caused an increase in the antioxidant activities of superoxide dismutase (SOD), peroxidase, and catalase (CAT) in the flag leaves of both cultivars, while ROS contents such as superoxide anion radical ([Formula: see text]) and hydrogen peroxide (H2O2) decreased. Moreover, the expression levels of Rubisco activase B (RcaB), major chlorophyll a/b-binding protein (Cab), chloroplast Cu/Zn superoxide dismutase (Cu/Zn-SOD), mitochondrial manganese superoxide dismutase (Mn-SOD), and CAT genes were upregulated at anthesis and were associated with higher photosynthetic capacity and antioxidant activities. Furthermore, night-warming improved sink activities by increasing the concentrations of grain indole-3-acetic acid and cytokinins as well as the sucrose synthase activity for both cultivars. Winter night-warming showed greater potential for improving source strength and grain filling, with consistent performance in both cultivars compared with that of spring night-warming. We concluded form these results that night-warming can improve source and sink capacities in winter wheat, and winter night-warming has greater advantages in this respect than does spring warming.

[Effects of low temperature stress on photosynthetic performance of different genotypes wheat cultivars].

Different genotypes wheat cultivars (spring wheat Yangmai 18, semi-spring wheat Zhengmai 9023, and semi-winter wheat Yannong 19) were chosen to study their photosynthetic and chlorophyll fluorescence characteristics at tillering and stem elongation stages under low temperature stress. After treated with low temperature (-10 degrees C at night) at tillering stage for 2 days, the net photosynthetic rate (Pn), stomata conductance (g(s)), maximum quantum yield of PSII (Fv/Fm), photochemical quenching (q(P)), non-photochemical quenching coefficient (NPQ), and acyclic photosynthetic electron transfer rate of PSII (ETR) of Yannong 19 were significantly higher than those of Yangmai 18 and Zhengmai 9023, the g(s), Fv/Fm, qP, and NPQ of Zhengmai 9023 were significantly higher than those of Yangmai 18, while the intercellular CO2 concentration (Ci) of Yannong 19 was significantly lower than that of Zhengmai 9023 and Yangmai 18. In addition, the minimal fluorescence (Fo) of Yangmai 18 was significantly higher than that of Zhengmai 9023 and Yannong 19. Similarly, after treated with low temperature (0 degrees C at night) at stem elongation stage for 3 days, the Pn, g(s), qP, and Fv/Fm of Yannong 19 were significantly higher than those of Yangmai 18 and Zhengmai 9023, and the NPQ and ETR of Yannong 19 were significantly higher than those of Yangmai 18. In addition, the Pn, g(s), Fv/Fm, and qP of Zhengmai 9023 were significantly higher than those of Yangmai 18, the Fo of Zhengmai 9023 was significantly higher than that of Yannong 19, and the Ci and Fo of Yangmai 18 were significantly higher than those of Zhengmai 9023 and Yannong 19. It was suggested that under low temperature stress at tillering and stem elongation stages, semi-winter wheat Yannong 19 had the highest photosynthetic activity and the best self-protection mechanism, followed by semi-spring wheat Zhengmai 9023, and spring wheat Yangmai 18.