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Unpredicted variability in temperature is associated with frequent extreme low-temperature events. Wheat is a leading crop in fulfilling global food requirements. Climate-driven temperature extremes influence the vegetative and reproductive growth of wheat, followed by a decrease in yield. This review describes how low temperature induces a series of modifications in the morphophysiological, biochemical, and molecular makeup of wheat and how it is perceived. To cope with these modifications, crop plants turn on their cold-tolerance mechanisms, characterized by accumulating soluble carbohydrates, signaling molecules, and cold tolerance gene expressions. The review also discusses the integrated management approaches to enhance the performance of wheat plants against cold stress. In this review, we propose strategies for improving the adaptive capacity of wheat besides alleviating risks of cold anticipated with climate change.

Due to global warming, late-spring coldness affecting wheat (LSCW) is one of the major abiotic adversities affecting wheat production. A combination of field and pot trials were conducted in this study. In the field experiment, 20 wheat varieties from the main wheat-producing areas in China were selected as experimental materials. By exploring the effects of LSCW on the spikelet characteristics and yields of different varieties, the evaluation methods and indexes of wheat varieties' resistance to LSCW were established. Three varieties with strong resistance to LSCW (Yannong19, Guomai9, and Shannong17) and five varieties sensitive to LSCW (Zhengmai895, Xinmai26, Zhengmai366, Zhengmai8329, and Fanmai5) were screened out. The wheat varieties Yannong19 (YN19), with a strong resistance to LSCW, and Xinmai26 (XM26), with LSCW sensitivity, were selected as the test materials for the pot experiment. The ultrastructure changes in the wheat in different low-temperature treatments during the anther differentiation period were observed using an ultra-low-temperature artificial climate incubator set to 4 °C and −4 °C for 4 h (1:00–5:00 a.m.). The average temperature of the field during the low-temperature treatment was 10 °C, which was the control temperature (CK). The results showed that the cell morphology and chloroplast and mitochondrial structures of the functional leaves, young ears, and internodes below the ears were damaged, and the degree of damage was related to the cell location, the extent of low-temperature stress, and the resistance of the varieties. The degree of damage to the wheat cells was found to occur in the following order: young ears > internode belove young ears > functional leaves. The degrees of damage to the wheat cell, chloroplast, and mitochondrial structures increased with the intensification of the low-temperature stress. The damage to the XM26 variety was obviously greater than that inflicted on the YN19 variety. The anatomical mechanism of YN19 cells makes the cell structure more stable during late-spring coldness. [ABSTRACT FROM AUTHOR]

In the context of global warming, low temperature (LT) disasters in major crops are also becoming more common. LT stress in the Huang-Huai-Hai Plain, the central wheat region in China, caused a massive reduction in wheat yields. A step towards ensuring wheat yield stability and food security, this study investigated the effects of optimizing phosphorus application on the root physiology, dry matter phosphorus accumulation, translocation, and partitioning of wheat under LT stress, using the representative cultivar Yannong 19 as the test material. The treatments included conventional phosphorus application (R1) and twice-split phosphorus application (R2), followed by −4 °C LT treatment and normal temperature (NT) treatment during the anther interval stage. Analysis of the root physiology (enzymatic activities and acid phosphatase, contents of malondialdehyde, soluble sugar, and soluble protein), phosphorus and dry matter accumulation, translocation, partitioning, and agronomic and yield-related components was carried during this research study. The results showed that the wheat root activity was significantly reduced and the antioxidant enzyme activities were increased to mitigate the damage of LT stress. Moreover, LT treatments damaged root function. The root activity and antioxidant properties were significantly lower than those of the NT treatment at the flowering stage. The dry matter and phosphorus accumulations were reduced by 30.6~33.6% and 15.1~21.3% at the flowering and maturity stages, resulting in final yield losses of 10.3~13.0%. In contrast, root activity increased by 16.1~27.2% in the twice-split phosphorus application treatments, and the root antioxidant characteristics were higher. As a result, dry matter and phosphorus accumulation increased after twice-split phosphorus application and their translocation to the grains was more; the final yield increased by 5.5~7.3%. Overall, the twice-split phosphorus application enhanced the physiological function of the root system and promoted the accumulation of nutrients and their transport to the grain, and alleviated the yield loss of wheat caused by LT stress. [ABSTRACT FROM AUTHOR]

Heat stress during the grain filling is one of most critical factors impacting grain yield in winter wheat. In a two-year field experiment, we studied the effects of 24-Epicastasterone and KH2PO4 on heat-stressed wheat applied during different times. A completely randomized block design was used with three reagents, that is, 24-Epicastasterone, KH2PO4 and H2O and three spraying times, that is, pre-heat, under-heat and post-heat. Two cultivars, that is, Huaimai 33 and Annong 0711 were subjected to a 7 d post-flowering heat stress using plastic tents. Our study suggests that greater grain weight in heat-stressed plants was achieved through sustaining leaf greenness and higher assimilates supplies to grains. When heat-stressed plants sprayed either with 24-Epicastasterone or KH2PO4 could sustain a relatively higher leaf photosynthetic capacity by up-regulating antioxidant enzymes with reduced damage on lipid membranes and leaf chlorophyll. We also found that KH2PO4 was relatively more effective in improving leaf physiology and grain yield of heat-stressed plants than 24-Epicastasterone. Further analysis showed that pre-heat or post-heat was more pronounced in protecting plants from heat injury than spraying the plants under heat stress. Thus, spraying KH2PO4 either prior to or post stress incidence is recommended to protect wheat crops from terminal heat stress. [ABSTRACT FROM AUTHOR]

As one of the most important wheat-producing areas in China, wheat is prone to heat stress during the grain filling period in the Huang-Huai-Hai Plain (3HP), which lowers yields and degrades the grain quality of wheat. To assess the effects of spraying potassium dihydrogen phosphate (KH2PO4) on the physiological traits in flag leaves and grain yield (GY) and quality under heat stress during the filling period, we conducted a two-year field experiment in the winter wheat growing seasons of 2020–2022. In this study, spraying water combined with heat stress (HT), 0.3% KH2PO4 (KDP), and 0.3% KH2PO4 combined with heat stress (PHT) were designed, and spraying water alone was used as a control (CK). The dates for the spraying were the third and eleventh day after anthesis, and a plastic film shed was used to impose heat stress on the wheat plants during the grain filling period. The results showed that spraying KH2PO4 significantly improved the chlorophyll content and net photosynthesis rate (Pn) in flag leaves compared with the non-sprayed treatments. Compared with CK, the Pn in HT decreased by 8.97% after heat stress, while Pn in PHT decreased by 7.44% compared to that of KDP. The activities of superoxide dismutase, catalase, and peroxidase in flag leaves were significantly reduced when the wheat was subjected to heat stress, while malonaldehyde content increased, and the enzyme activities were significantly enhanced when KH2PO4 was sprayed. Heat stress significantly decreased the contribution rate of dry matter accumulation (DM) after anthesis of wheat to grain (CRAA), whereas spraying KH2PO4 significantly increased the CRAA and harvest index. At maturity, the DM in CK was significantly higher than that in HT, KDP was significantly higher than PHT, and KDP had the highest DM. Compared with CK, the GY in KDP significantly increased by 9.85% over the two years, while the GY in HT decreased by 11.44% compared with that of CK, and the GY in PHT decreased by 6.31% compared to that of KDP. Spraying KH2PO4 after anthesis primarily helped GY by maintaining a high thousand grain weight to lessen the negative effects of heat stress on wheat. Moreover, heat stress significantly reduced protein concentration, wet gluten content, dough development time, and hardness index in grains of mature, while spraying KH2PO4 maintained a sufficient grain quality under the conditions of achieving higher yields. Overall, spraying KH2PO4 after anthesis could enhance the heat stress resistance of wheat and maintain the photosynthetic capacity of flag leaves, ensuring the dry matter production and reducing the negative effects on grain yield and quality in the 3HP. [ABSTRACT FROM AUTHOR]

Low-temperature stress has become an important abiotic factor affecting high and stable wheat production. Therefore, it is necessary to take appropriate measures to enhance low-temperature tolerance in wheat. A pot experiment was carried out using Yannong19 (YN19, a cold-tolerant cultivar) and Xinmai26 (XM26, a cold-sensitive cultivar). We employed traditional phosphorus application (TPA, i.e., R1) and optimized phosphorus application (OPA, i.e., R2) methods. Plants undertook chilling (T1 at 4 °C) and freezing treatment (T2 at −4 °C) as well as ambient temperature (CK at 11 °C) during the anther differentiation period to investigate the effects of OPA and TPA on photosynthetic parameters and the accumulation and distribution of dry matter. The net photosynthetic rate (Pn), stomatal conductance (Gs) and transpiration rate (Tr) of flag leaves decreased in low-temperature treatments, whereas intercellular carbon dioxide concentration (Ci) increased. Compared with R1CK, Pn in R1T1 and R1T2 treatments was reduced by 26.8% and 42.2% in YN19 and 34.2% and 54.7% in XM26, respectively. In contrast, it increased by 6.5%, 8.9% and 12.7% in YN19 and 7.7%, 15.6% and 22.6% in XM26 for R2CK, R2T1 and R2T2 treatments, respectively, under OPA compared with TPA at the same temperature treatments. Moreover, low-temperature stress reduced dry matter accumulation at the reproductive growth stage. OPA increased dry matter accumulation of vegetative organs after the flowering stage and promoted the transportation of assimilates to grains. Hence, the grain number per spike (GNPS), 1000-grain weight (TGW) and yield per plant (YPP) increased. The low-temperature treatments of T1 and T2 caused yield losses of 24.1~64.1%, and the yield increased by 8.6~20.5% under OPA treatments among the two wheat cultivars. In brief, OPA enhances low-temperature tolerance in wheat, effectively improves wheat architecture and photosynthesis, increases GNPS and TGW and ultimately lessens yield losses. [ABSTRACT FROM AUTHOR]

The 21st century presents many challenges to mankind, including climate change, fast growing human population, and serious concerns over food security. Wheat is a leading cereal crop that largely fulfills the global food needs. Low temperature stress accompanied by nutrient-starved soils is badly disrupting the source–sink relationship of wheat, thus causing an acute decline in final yield and deteriorating the grain quality. This review paper aimed to understand how low temperature stress affects wheat source–sink organs (i.e., leaves, roots, and spikes) and how phosphorus application reliefs in alleviating its harmful consequences. Also, we discussed mitigation strategies to enhance wheat capacity to adapt to varying temperature extremes and made rational recommendations based on modern agronomic and breeding approaches. Therefore, this study is likely to establish a solid foundation for improving the tolerance to low temperature stress and to improve its phosphorus utilization efficiency in wheat. [ABSTRACT FROM AUTHOR]

Late spring coldness (LSC) is critical for wheat growth and development in the Huang-Huai valleys of China. However, little is known about the molecular mechanisms for young spikes responding to low temperature (LT) stress during anther connective tissue formation phase (ACFP). To elucidate the molecular mechanisms associated with low temperature, we performed a comparative transcriptome analysis of wheat cultivars Xinmai26 (XM26: cold-sensitive) and Yannong19 (YN19: cold-tolerant) using RNA-seq data. Over 4000 differently expressed genes (DEGs) were identified under low temperature conditions (T1: 4°C) and freezing conditions (T2: −4°C) compared with control (CK: 16°C). The number of DEGs associated with two cultivars at two low temperature treatments (T1: 4°C and T2: −4°C) were 834, 1,353, 231, and 1,882 in four comparison groups (Xinmai26-CK vs. Xinmai26-T1, Xinmai26-CK vs. Xinmai26-T2, Yannong19-CK vs. Yannong19-T1, and Yannong19-CK vs. Yannong19-T2), respectively. Furthermore, to validate the accuracy of RNA-seq, 16 DEGs were analyzed using quantitative real-time RT-PCR. Several transcriptome changes were observed through Kyoto Encyclopedia of Genes and Genomes (KEGG) pathway functional enrichment analysis in plant hormone signal transduction, circadian rhythm-plant, and starch and sucrose metabolism under low temperature. In addition, 126 transcription factors (TFs), including AP2-ERF, bHLH, WRKY, MYB, HSF , and members of the bZIP family, were considered as cold-responsive. It is the first study to investigate DEGs associated with low temperature stress at the transcriptome level in two wheat cultivars with different cold resistance capacities. Most likely, the variations in transcription factors (TFs) regulation, and starch and sucrose metabolism contribute to different cold resistance capacities in the two cultivars. Further, physiological activities of superoxide dismutase (SOD), peroxidase (POD), catalase (CAT) enzymes, malondialdehyde (MDA), soluble sugar (SS), and sucrose contents were evaluated to investigate the negative impacts of low temperature in both cultivars. These findings provide new insight into the molecular mechanisms of plant responses to low temperature and potential candidate genes that required for improving wheat's capacity to withstand low temperature stress. [ABSTRACT FROM AUTHOR]

Climate change has caused frequent extreme low-temperature events to threaten global food security. Spring low-temperature stress is one of the major limiting factors for high and stable yields of wheat. We used two wheat varieties differing in spring cold-sensitivity (cold-tolerant variety Yannong 19 and cold-sensitive variety Xinmai 26) to examine the effects of low-temperature stress during the anther differentiation period on wheat photosynthetic performance and spike-setting characteristics. Low-temperature stress was simulated in a climate box at −2 °C, 0 °C or 2 °C (night) and 15 °C (day) for 24 h, 48 h or 72 h. With the extension of the treatment time and the decrease of temperature, the photosynthetic rate, stomatal conductance and transpiration rate of wheat leaves gradually decreased. All treatments except −2 °C for 72 h recovered slowly within 7–15 days after treatment. Low-temperature stress greatly reduced grains per spikelet, 1000-grain weight and yield per plant. By analyzing the spikelets in different stalk locations (upper, middle and lower), we found that the number of upper spikelets was significantly less than lower and middle spikelets after low-temperature stress. The sterile grain of upper spikelets (Xinmai 26, for example) can reach 100% at −2 °C for 48 h and 72 h, and the yield loss rate was 90.52% at 2 °C for 24 h, which was much higher than for the lower spikelets (60.73%) and middle spikelets (50.94%). Overall, these findings suggest that low-temperature stress during the anther differentiation period alters the photosynthetic activity involved in the accumulation of dry matter in wheat, which leads to delaying young spike growth, especially for upper spikelets, and ultimately in a decrease in yield. [ABSTRACT FROM AUTHOR]