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.

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.

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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.

In a pot experiment, we explored the regulatory pathways through which melatonin (MT) protects wheat growth and grain yield loss from waterlogging injury. Two wheat cultivars, Yangmai 18 and Yannong 19, were exposed to seven days of soil waterlogging at flowering. Melatonin (100 μmol·L−1) was sprayed before and after waterlogging to explore its regulation on root growth, photosynthetic characteristics, dry matter accumulation, and grain yield. Soil waterlogging intensified malondialdehyde (MDA) and O2− production rates in wheat tissues, impairing leaf photosynthesis, biomass accumulation, and final grain yield formation. In this study, the roots waterlogged at 7 days after anthesis (DAA) accumulated 20.9%, 76.2%, 17.6%, 28.5%, and 5.6% higher MDA content, O2− production rate, pyruvate decarboxylase (PDC), lactate dehydrogenase (LDH), and alcohol dehydrogenase (ADH) activities, respectively, in Yangmai 18, and 25.7%, 74.8%, 35.8%, 70.8%, and 30.7% higher in Yannong 19, respectively, compared with their respective non-waterlogged controls. Further, Yangmai 18 achieved a maximum net photosynthetic rate (Pn) reduction of 22.1% at 7 DAA, while the maximum Pn reduction of Yannong 19 was 27.4% at 14 DAA, respectively, compared with their respective non-waterlogged plants. Thus, waterlogging decreased total dry matter accumulation, 1000-grain weight (TGW), and total grain yield by 14.0%, 13.8%, and 16.2%, respectively, in Yangmai 18, and 16.0%, 8.1%, and 25.1%, respectively, in Yannong 19. Our study also suggests that exogenously applied melatonin can protect wheat root tissues from waterlogging-induced oxidative injury by upregulating antioxidant enzymes and sustaining leaf photosynthesis. The plants treated with melatonin showed better water status and less oxidative damage, which was conducive to maintaining a higher photosynthetic capacity, thereby improving the waterlogging tolerance of wheat. For example, compared with waterlogged plants, melatonin treatments significantly reduced MDA content, O2− production rate, PDC, LDH, and ADH activities by 7.7%, 25.4%, 2.6%, 32.1%, and 3.2%, respectively, in Yangmai 18, and 6.7%, 17.9%, 4.1%, 22.0%, and 15.3%, respectively, in Yannong 19. MT treatments significantly increased total dry matter accumulation, TGW, and yield by 5.9%, 8.7%, and 14.9%, respectively, in Yangmai 18, and 3.2%, 7.3%, and 26.0%, respectively, in Yannong 19.

High temperature stress during grain filling substantially decreases wheat productivity; thus, to ensure food security, heat tolerance in wheat must be developed. It remains unclear whether exogenous salicylic acid (SA) can induce tolerance to high temperatures in wheat at the grain-filling stage. In this study, a two-year pot culture experiment using the wheat cultivar ‘Yangmai 18’ was conducted from 2018 to 2020. The plants were pre-sprayed with SA from the heading stage (SAH), anthesis stage (SAA), 5 days after anthesis (DAA; SA5), and 10 DAA (SA10). After that, the wheat plants were subjected to high temperature stress (G) simulated using a passive warming method during the period between 15 and 19 DAA. The results showed that, compared with the normal temperature control group (NN), high temperature stress at the grain-filling stage significantly reduced the yield and photosynthetic capacity of wheat. The application of SA at different stages reduced the yield loss and the damage to the photosynthetic capacity caused by high temperature stress; the effectiveness of the treatments in descending order was SAAG > SA5G > SA10G > SAHG. Exogenous SA treatment increased the amount and proportion of dry matter distributed in the stem sheaths and leaves and grains, and decreased the amount and proportion of dry matter distributed in the rachises and glumes at the maturity stage, thereby reducing the yield loss under high temperature stress. The application of SA significantly increased the leaf area, stomatal density, chlorophyll content, soluble protein content, maximum photochemical efficiency (Fv/Fm), actual photochemical efficiency (ΦPSII), and activity of sucrose phosphate synthase (SPS) of the wheat flag leaves under high temperature stress at the grain-filling stage, thereby improving the photosynthetic performance of the flag leaves under stress. In summary, exogenous SA significantly restored the photosynthetic capacity of wheat flag leaves injured by post-anthesis high temperature stress, which effectively alleviated the inhibition of wheat growth caused by the stress and ultimately reduced the yield loss. Spraying SA at the anthesis stage had the greatest effect abating the loss of yield and reduced photosynthetic performance under high temperature stress.

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