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.

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.

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 (P_n), stomatal conductance (G_s), and transpiration rate (T_r) 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.

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.