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Journal of Integrative Agriculture  2024, Vol. 23 Issue (2): 536-550    DOI: 10.1016/j.jia.2023.06.024
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Night warming increases wheat yield by improving pre-anthesis plant growth and post-anthesis grain starch biosynthesis

Yonghui Fan1, Boya Qin1, Jinhao Yang1, 2, Liangliang Ma1, Guoji Cui1, Wei He1, Yu Tang1, Wenjing Zhang1, Shangyu Ma1, Chuanxi Ma1#, Zhenglai Huang1# 

1 College of Agronomy, Anhui Agricultural University/Key Laboratory of Wheat Biology and Genetic Improvement in South Yellow &     Huai River Valley, Ministry of Agriculture and Rural Affairs, Hefei 230036, China

2 Xianyang Central Station for Agricultural Technology Extension, Xianyang 712000, China

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摘要  

全球变暖的主要特征是非对称性增温,即冬春季和夜间增温幅度大于夏秋季和白天的增温幅度。于2019~20202020~2021年两个小麦生长季,以扬麦18YM18)、苏麦188SM188)、烟农19YN19)和安农0711AN0711)为试验材料,采用被动式夜间增温方法,对小麦生育前期进行不同阶段夜间增温处理,即分蘖期至拔节期夜间增温处理(NWT-J)、拔节期至孕穗期夜间增温处理(NWJ-B)、孕穗期至开花期夜间增温处理(NWB-A),以不增温为对照(NN通过小麦干物质积累转运特性,籽粒蔗糖和淀粉积累特性,研究不阶段夜间增温对小麦产量形成的影响。结果表明,不同阶段夜间增温通过提高小麦的千粒重以及可孕小穗数从而提高小麦产量,NWT-J处理4个品种小麦产量均显著高于NN,半冬性小麦品种YN19和AN0711受增温的影响大于春性小麦品种YM18和SM188。NWT-J处理通过提高小麦营养生长阶段的生长速率从而增加了小麦开花期和成熟期各器官干物质积累量,且以旗叶和穗部的干物质积累量提升比例较大。NWT-J处理还提高了小麦灌浆前期和中期的籽粒蔗糖和淀粉含量,从而促进产量的形成。综上所述,分蘖到拔节期夜间增温通过促进小麦花前的生长速率从而提高了小麦的干物质生产能力进而有利于产量的提高。



Abstract  

Global climate change is characterized by asymmetric warming, i.e., greater temperature increases in winter, spring, and nighttime than in summer, autumn, and daytime.  Field experiments were conducted using four wheat cultivars, namely ‘Yangmai 18’ (YM18), ‘Sumai 188’ (SM188), ‘Yannong 19’ (YN19), and ‘Annong 0711’ (AN0711), in the two growing seasons of 2019–2020 and 2020–2021, with passive night warming during different periods in the early growth stage.  The treatments were night warming during the tillering–jointing (NWT–J), jointing–booting (NWJ–B), and booting–anthesis (NWB–A) stages, with ambient temperature (NN) as the control.  The effects of night warming during different stages on wheat yield formation were investigated by determining the characteristics of dry matter accumulation and translocation, as well as sucrose and starch accumulation in wheat grains.  The wheat yields of all four cultivars were significantly higher in NWT–J than in NN in the 2-year experiment.  The yield increases of semi-winter cultivars YN19 and AN0711 were greater than those of spring cultivars YM18 and SM188.  Treatment NWT–J increased wheat yield mainly by increasing the 1,000-grain weight and the number of fertile spikelets, and it increased dry matter accumulation in various organs of wheat at the anthesis and maturity stages by increasing the growth rate at the vegetative growth stage.  The flag leaf and spike showed the largest increases in dry matter accumulation.  NWT–J also increased the grain sucrose and starch contents in the early and middle grain-filling stages, promoting yield formation.  Overall, night warming between the tillering and jointing stages increased the pre-anthesis growth rate, and thus, wheat dry matter production, which contributed to an increase in wheat yield.

Keywords:  wheat        asymmetric warming        dry matter accumulation and translocation        starch        yield  
Received: 06 January 2023   Accepted: 05 May 2023
Fund: 

This work was supported by the Project of Natural Science Foundation of Anhui Province, China (2008085qc118), the National Natural Science Foundation of China (U19A2021), the Major Science and Technology Special Project of Anhui Province, China (S202003a06020035), and the Jiangsu Collaborative Innovation Center for Modern Crop Production, China (JCIC-MCP).

About author:  Yonghui Fan, E-mail: yonghuifan66@163.com; #Correspondence Zhenglai Huang, E-mail: ahauhzl@163.com; Chuanxi Ma, E-mail: machuanxi@ahau.edu.cn

Cite this article: 

Yonghui Fan, Boya Qin, Jinhao Yang, Liangliang Ma, Guoji Cui, Wei He, Yu Tang, Wenjing Zhang, Shangyu Ma, Chuanxi Ma, Zhenglai Huang. 2024.

Night warming increases wheat yield by improving pre-anthesis plant growth and post-anthesis grain starch biosynthesis . Journal of Integrative Agriculture, 23(2): 536-550.

Aneela U, Ansar M, Shafique A K, Sami U A. 2021. Elevated carbon dioxide offers promise for wheat adaptation to heat stress by adjusting carbohydrate metabolism. Physiology and Molecular Biology of Plants, 27, 2345–2355.

Asseng S, Ewert F, Martre P, Rötter R P, Lobell D B, Cammarano D, Kimball B A, Ottman M J, Wall G, White J W. 2015. Rising temperatures reduce global wheat production. Nature Climate Change, 5, 143–147.

Bala S, Asthir B, Taggar M S, Goyal M, Bains N S, Sharma A, Chhuneja P. 2021. Grain carbon metabolism and stem reserve mobilization compensate high temperature stress in wheat. Agrochimica, 65, 39–52.

Chavas D R, Izaurralde R C, Thomson A M, Gao X J. 2009. Long-term climate change impacts on agricultural productivity in eastern China. Agricultural and Forest Meteorology, 149, 1118–1128.

Chen J, Tian Y L, Zhang X, Zheng C Y, Song Z W, Deng A X, Zhang W J. 2014. Nighttime warming will increase winter wheat yield through improving plant development and grain growth in North China. Journal of Plant Growth Regulation, 33, 397–407.

Chen W, Li H B, Deng X P. 2018. The characteristics of sucrose metabolism in flag leaf and grain starch accumulation during grain filling period under different cultivation modes. Acta Agriculturae Boreali-Occidentalis Sinica, 27, 641–649. (in Chinese)

Chu J P, Guo X H, Zheng F N, Zhang X, Dai X L, He M R. 2023. Effect of delayed sowing on grain number, grain weight, and protein concentration of wheat grains at specific positions within spikes. Journal of Integrative Agriculture, 22, 2359–2369.

Djanaguiraman M, Narayanan S, Erdayani E, Prasad P V V. 2020. Effects of high temperature stress during anthesis and grain filling periods on photosynthesis, lipids and grain yield in wheat. BMC Plant Biology, 20, 2–12.

Fan Y H, Tian M Y, Jing Q, Tian Z W, Han H M, Jiang D, Cao W X, Dai T B. 2015. Winter night warming improves pre-anthesis crop growth and post-anthesis photosynthesis involved in grain yield of winter wheat (Triticum aestivum L.). Field Crops Research, 178, 100–108.

Fan Y H, Tian Z W, Yan Y Y, Hu C X, Muhammad A, Jiang D, Ma C X, Huang Z L, Dai T B. 2017. Winter night-warming improves post-anthesis physiological activities and sink strength in relation to grain filling in winter wheat (Triticum aestivum L.). Frontiers in Plant Science, 8, 1–14.

Grant R F, Kimball B A, Conley M M, White J W, Wall G W, Ottman M J. 2011. Controlled warming effects on wheat growth and yield: field measurements and modeling. Agronomy Journal, 103, 1742–1754.

Gomez D, Salvador P, Sanz J, Luis C J. 2021. Modelling wheat yield with antecedent information, satellite and climate data using machine learning methods in Mexico. Agricultural and Forest Meteorology, 300, 108317–108322.

Hansen J, Moller I. 1975. Percolation of starch and soluble carbohydrates from plant tissue for quantitative determination with anthrone. Analytical Biochemistry, 68, 87–94.

He D, Fang S B, Liang H Y, Wang E L, Wu D. 2020. Contrasting yield responses of winter and spring wheat to temperature rise in China. Environmental Research Letters, 15, 1–10.

Hunt L A, Poorten G, Parajasingham S. 1991. Postanthesis temperature effects on duration and rate of grain filling in some winter and spring wheats. Canadian Journal of Plant Science, 71, 609–617.

IPCC (Intergovernmental Panel on Climate Change). 2007. Climate change 2007: The physical science basis. In: Cntribution of Working Group I to the Fourth Assessment Report of the Intergovernmental Panel on Vlimate Change. Cambridge University Press, Cambridge, United Kingdom.

Jenner C. 1991. Effects of exposure of wheat ears to high temperature on dry matter accumulation and carbohydrate metabolism in the grain of two cultivars. I. immediate responses. Functional Plant Biology, 18, 165–177.

Jia W, Zhang J H. 2008. Stomatal movements and long-distance signaling in plants. Plant Signaling & Behavior, 3, 772–777.

Kakar K, Xuan T D, Noori Z, Aryan S, Gulab G. 2020. Effects of organic and onorganic fertilizer application on growth, yield, and grain quality of rice. Agriculture, 10, 544–555.

Kanno K, Mae T, Makino A. 2009. High night temperature stimulates photosynthesis, biomass production and growth during the vegetative stage of rice plants. Soil Science & Plant Nutrition, 55, 124–131.

Lawas L M F, Shi W, Yoshimoto M, Hasegawa T, Hincha D K, Zuther E, Jagadishi S V K. 2018. Combined drought and heat stress impact during flowering and grain filling in contrasting rice cultivars grown under field conditions. Field Crops Research, 229, 66–77.

Lazar C, Lazar D A. 2010. Simulation of temperature increase influence on winter wheat yields and development in South-Eastern Romania. Romania Agricultural Research, 27, 7–15.

Li T Y, Ullah S, Liang H, Ali I, Zhao Q, Iqbal A, Wei S Q, Shah T, Luo Y Q, Jiang L G. 2020. The enhancement of soil frtility, dry matter transport and accumulation, nitrogen uptake and yield in rice via green manuring. Phyton-International Journal of Experimental Botany, 90, 223–243.

Liu J J, Xiao Y G, Zhu F B, Cheng D G, Li H S, Liu A F, Song J M. 2009. Effect of canopy temperature on yield traits of different genotypes of winter wheat. Journal of Triticeae Crops, 29, 283–288. (in Chinese)

Liu L T, Hu C S, Olesen J E, Ju Z Q, Yang P P, Zhang Y M. 2013. Warming and nitrogen fertilization effects on winter wheat yields in northern China varied between four years. Field Crops Research, 151, 56–64.

Lobell D B. 2007. Changes in diurnal temperature rand and national cereal yields. Agricultural and Forest Meteorology, 145, 229–238.

Lv L H, Liang S B, Zhang L H, Jia X L, Dong Z Q, Yao Y R. 2016. Yield in response to accumulated temperature before winter in winter wheat. Acta Agronomica Sinica, 37, 1047–1055. (in Chinese)

Madhukar A, Dashora K, Kumar V. 2021. Climate trends in temperature and water variables during wheat growing season and impact on yield. Environmental Processes, 8, 1047–1072.

Mamrutha H M, Rinki K, Venkatesh K, Gopalareddy K, Khan H, Mishra C N, Kumar S, Kumar Y, Singh G, Singh G P. 2020. Impact of high night temperature stress on different growth stages of wheat. Plant Physiology Reports, 25, 707–715.

Mirosavljevic M, Mikic S, Zupunski V, Spika A K, Trkulja D, Ottosen C, Zhou R, Abdelhakim L. 2021. Effects of high temperature during anthesis and grain filling on physiological characteristics of winter wheat cultivars. Journal of Agronomy and Crop Science, 207, 823–832.

Nijs I, Kockelbergh H, Blum H, Hendrey G, Impens I. 2010. Free air temperature increase (FATI) A new tool to study global warming effects on plants in the field. Plant Cell & Environment, 19, 495–502.

Pepin N, Bradley R S, Diaz H F, Baraer M, Caceres B, Forsythe N, Fowler H J, Greenwood G, Hashmi M Z R, Liu X D, Miller J R, Ning L, Ohmura A, Palazzi E, Rangwala I, Schner W, Severskiy I, Shahgedanova M, Wang M B, Williamson S N, et al. 2015. Elevation-dependent warming in mountain regions of the world. Nature Climate Change, 5, 424–430.

Plaut Z, Butow B J, Blumenthal C S, Wrigley C W. 2004. Transport of dry matter into developing wheat kernels and its contribution to grain yield under post-anthesis water deficit and elevated temperature. Field Crop Research, 86, 185–198.

Qian J X, Xi Y X. 2008. Analysis of characteristics and its influencing facters of winter wheat main growth stages. Chinese Agricultural Science Bulletin, 11, 438–443. (in Chinese)

Qu C H, Li X X, Ju H, Liu Q. 2019. The impacts of climate change on wheat yield in the Huang-Huai-Hai Plain of China using DSSAT-CERES-Wheat model under different climate scenarios. Journal of Integrative Agriculture, 18, 1379–1391.

Shi J J, Jiang X D, Shi H W, Chen Y J, Yuan J K, Jiang M. 2015. Effect of winter warming treatments on photosynthesis and yield of wheat. Journal of Triticeae Crops, 35, 352–356. (in Chinese)

Skinner D Z. 2014. Time and temperature interactions in freezing tolerance of winter wheat. Crop Science, 54, 1523–1529.

Subedi K D, Gregory P J, Summertfield R J, Gooding M J. 1998. Cold temperatures and boron deficiency caused grain set failure in spring wheat (Triticum aestivum L.). Field Crops Research, 57, 277–288.

Takai T, Matsuura S, Nishio T, Ohsumi A, Shiraiwa T, Horie T. 2006. Rice yield potential is closely related to crop growth rate during late reptoductive period. Field Crops Research, 96, 328–335.

Tashiro T, Wardlaw I F. 1990. The effect of high temperature at different stages of ripening on grain set, grain weight and grain dimensions in the semi-dwarf wheat ‘Banks’. Annals of Botany, 65, 51–61.

Tian Y L, Chen J, Chen C Q, Deng A X, Song Z W, Zheng C Y, Hoogmoed W, Zhang W J. 2012. Warming impacts on winter wheat phenophase and grain yield under field conditions in Yangtze Delta Plain, China. Field Crops Research, 134, 193–199.

Tian Y L, Chen J, Dong W J, Deng A X, Zhang W J. 2011. Effects of asymmetric warming on key enzyme activities of starch synthesis in superior and inferior grains of winter wheat under FATI facility. Acta Agronomica Sinica, 37, 1031–1038. (in Chinese)

Venkatesh K, Senthilkumar K M, Mamrutha H M, Singh G, Singh G P. 2022. High-temperature stress in wheat under climate change scenario, effects and mitigation strategies. Climate Change and Crop Stress, 8, 209–229.

Villegas D, Aparicio N, Blanco R, Royo C. 2001. Biomass accumulation and main stem elongation of durum wheat grown under mediterranean conditions. Annals of Botany, 88, 617–627.

Way D A, Yamori W. 2014. Thermal acclimation of photosynthesis: On the importance of adjusting our definitions and accounting for thermal acclimation of respiration. Photosynthesis Research, 119, 89–100.

Wrigley C W, Blumenthal C, Gras P W, Barlow E W R. 1994. Temperature variation during grain filling and changes in wheat-grain quality. Functional Plant Biology, 21, 875–885.

Xiao G J, Zhang Q, Li Y, Wang R Y, Yao Y B, Zhao H, Bai H Z. 2010. Impact of temperature increase on the yield of winter wheat at low and high altitudes in semiarid northwestern China. Agricultural Water Management, 97, 1360–1364.

Xiao G J, Zhang Q, Zhang F J, Lou C K, Wang R Y. 2011. The impact of rising temperature on spring wheat production in the Yellow River irrigation region of Ningxia. Acta Ecologica Sinica, 21, 6588–6593. (in Chinese)

Xu L, Zhao D Y, Wang Y Q, Zhang C Z. 2017. Determination of D-tagatose content by resorcinol method. Journal of Dalian Polytechnic University, 36, 168–170. (in Chinese)

Yamamori M, Quynh N. 2000. Differential effects of Wx-A1, -B1 and -D1 protein deficiencies on apparent amylose content and starch pasting properties in common wheat. Theoretical and Applied Genetics, 100, 32–38.

Yu X R, Hao D, Yang J Q, Ran L P, Zang Y, Xiong F. 2020. Effects of low temperature at stem elongation stage on the development, morphology, and physicochemical properties of wheat starch. PeerJ, 8, 2–22.

Zhang C H, Jiang D, Liu F L, Cai J, Dai T B, Cao W C. 2010. Starch granules size distribution in superior and inferior grains of wheat is related to enzyme activities and their gene expressions during grain filling. Journal of Cereal Science, 51, 226–233.

Zhao H, Dai T B, Jing Q, Cao W X. 2007. Leaf senescence and grain filling affected by post-anthesis high temperatures in two different wheat cultivars. Plant Growth Regulation, 51, 149–158.

Zhao H, Dai T B, Qi J, Cao W X. 2005. Effects of temperature during grain filling on the contents of grain protein components and free amino acid in two different wheat cultivars. Acta Agronomica Sinica, 31, 1466–1472. (in Chinese)

Zheng C, Zhang J, Chen J, Chen C Q, Tian Y L, Deng A X, Song Z W, Nawaz M M, Van G K J, Zhang W J. 2017. Nighttime warming increases winter-sown wheat yield across major Chinese cropping regions. Field Crops Research, 214, 202–210.

Zhong M, Yoshida H, Zhao P, Wang X, Han L, Hu X, Huang S, Huang J, Sun Z. 2008. Changes in frost resistance of wheat young ears with development during jointing stage. Journal of Agronomy and Crop Science, 194, 343–349.

Aneela U, Ansar M, Shafique A K, Sami U A. 2021. Elevated carbon dioxide offers promise for wheat adaptation to heat stress by adjusting carbohydrate metabolism. Physiology and Molecular Biology of Plants, 27, 2345–2355.

Asseng S, Ewert F, Martre P, Rötter R P, Lobell D B, Cammarano D, Kimball B A, Ottman M J, Wall G, White J W. 2015. Rising temperatures reduce global wheat production. Nature Climate Change, 5, 143–147.

Bala S, Asthir B, Taggar M S, Goyal M, Bains N S, Sharma A, Chhuneja P. 2021. Grain carbon metabolism and stem reserve mobilization compensate high temperature stress in wheat. Agrochimica, 65, 39–52.

Chavas D R, Izaurralde R C, Thomson A M, Gao X J. 2009. Long-term climate change impacts on agricultural productivity in eastern China. Agricultural and Forest Meteorology, 149, 1118–1128.

Chen J, Tian Y L, Zhang X, Zheng C Y, Song Z W, Deng A X, Zhang W J. 2014. Nighttime warming will increase winter wheat yield through improving plant development and grain growth in North China. Journal of Plant Growth Regulation, 33, 397–407.

Chen W, Li H B, Deng X P. 2018. The characteristics of sucrose metabolism in flag leaf and grain starch accumulation during grain filling period under different cultivation modes. Acta Agriculturae Boreali-Occidentalis Sinica, 27, 641–649. (in Chinese)

Chu J P, Guo X H, Zheng F N, Zhang X, Dai X L, He M R. 2023. Effect of delayed sowing on grain number, grain weight, and protein concentration of wheat grains at specific positions within spikes. Journal of Integrative Agriculture, 22, 2359–2369.

Djanaguiraman M, Narayanan S, Erdayani E, Prasad P V V. 2020. Effects of high temperature stress during anthesis and grain filling periods on photosynthesis, lipids and grain yield in wheat. BMC Plant Biology, 20, 2–12.

Fan Y H, Tian M Y, Jing Q, Tian Z W, Han H M, Jiang D, Cao W X, Dai T B. 2015. Winter night warming improves pre-anthesis crop growth and post-anthesis photosynthesis involved in grain yield of winter wheat (Triticum aestivum L.). Field Crops Research, 178, 100–108.

Fan Y H, Tian Z W, Yan Y Y, Hu C X, Muhammad A, Jiang D, Ma C X, Huang Z L, Dai T B. 2017. Winter night-warming improves post-anthesis physiological activities and sink strength in relation to grain filling in winter wheat (Triticum aestivum L.). Frontiers in Plant Science, 8, 1–14.

Grant R F, Kimball B A, Conley M M, White J W, Wall G W, Ottman M J. 2011. Controlled warming effects on wheat growth and yield: field measurements and modeling. Agronomy Journal, 103, 1742–1754.

Gomez D, Salvador P, Sanz J, Luis C J. 2021. Modelling wheat yield with antecedent information, satellite and climate data using machine learning methods in Mexico. Agricultural and Forest Meteorology, 300, 108317–108322.

Hansen J, Moller I. 1975. Percolation of starch and soluble carbohydrates from plant tissue for quantitative determination with anthrone. Analytical Biochemistry, 68, 87–94.

He D, Fang S B, Liang H Y, Wang E L, Wu D. 2020. Contrasting yield responses of winter and spring wheat to temperature rise in China. Environmental Research Letters, 15, 1–10.

Hunt L A, Poorten G, Parajasingham S. 1991. Postanthesis temperature effects on duration and rate of grain filling in some winter and spring wheats. Canadian Journal of Plant Science, 71, 609–617.

IPCC (Intergovernmental Panel on Climate Change). 2007. Climate change 2007: The physical science basis. In: Cntribution of Working Group I to the Fourth Assessment Report of the Intergovernmental Panel on Vlimate Change. Cambridge University Press, Cambridge, United Kingdom.

Jenner C. 1991. Effects of exposure of wheat ears to high temperature on dry matter accumulation and carbohydrate metabolism in the grain of two cultivars. I. immediate responses. Functional Plant Biology, 18, 165–177.

Jia W, Zhang J H. 2008. Stomatal movements and long-distance signaling in plants. Plant Signaling & Behavior, 3, 772–777.

Kakar K, Xuan T D, Noori Z, Aryan S, Gulab G. 2020. Effects of organic and onorganic fertilizer application on growth, yield, and grain quality of rice. Agriculture, 10, 544–555.

Kanno K, Mae T, Makino A. 2009. High night temperature stimulates photosynthesis, biomass production and growth during the vegetative stage of rice plants. Soil Science & Plant Nutrition, 55, 124–131.

Lawas L M F, Shi W, Yoshimoto M, Hasegawa T, Hincha D K, Zuther E, Jagadishi S V K. 2018. Combined drought and heat stress impact during flowering and grain filling in contrasting rice cultivars grown under field conditions. Field Crops Research, 229, 66–77.

Lazar C, Lazar D A. 2010. Simulation of temperature increase influence on winter wheat yields and development in South-Eastern Romania. Romania Agricultural Research, 27, 7–15.

Li T Y, Ullah S, Liang H, Ali I, Zhao Q, Iqbal A, Wei S Q, Shah T, Luo Y Q, Jiang L G. 2020. The enhancement of soil frtility, dry matter transport and accumulation, nitrogen uptake and yield in rice via green manuring. Phyton-International Journal of Experimental Botany, 90, 223–243.

Liu J J, Xiao Y G, Zhu F B, Cheng D G, Li H S, Liu A F, Song J M. 2009. Effect of canopy temperature on yield traits of different genotypes of winter wheat. Journal of Triticeae Crops, 29, 283–288. (in Chinese)

Liu L T, Hu C S, Olesen J E, Ju Z Q, Yang P P, Zhang Y M. 2013. Warming and nitrogen fertilization effects on winter wheat yields in northern China varied between four years. Field Crops Research, 151, 56–64.

Lobell D B. 2007. Changes in diurnal temperature rand and national cereal yields. Agricultural and Forest Meteorology, 145, 229–238.

Lv L H, Liang S B, Zhang L H, Jia X L, Dong Z Q, Yao Y R. 2016. Yield in response to accumulated temperature before winter in winter wheat. Acta Agronomica Sinica, 37, 1047–1055. (in Chinese)

Madhukar A, Dashora K, Kumar V. 2021. Climate trends in temperature and water variables during wheat growing season and impact on yield. Environmental Processes, 8, 1047–1072.

Mamrutha H M, Rinki K, Venkatesh K, Gopalareddy K, Khan H, Mishra C N, Kumar S, Kumar Y, Singh G, Singh G P. 2020. Impact of high night temperature stress on different growth stages of wheat. Plant Physiology Reports, 25, 707–715.

Mirosavljevic M, Mikic S, Zupunski V, Spika A K, Trkulja D, Ottosen C, Zhou R, Abdelhakim L. 2021. Effects of high temperature during anthesis and grain filling on physiological characteristics of winter wheat cultivars. Journal of Agronomy and Crop Science, 207, 823–832.

Nijs I, Kockelbergh H, Blum H, Hendrey G, Impens I. 2010. Free air temperature increase (FATI) A new tool to study global warming effects on plants in the field. Plant Cell & Environment, 19, 495–502.

Pepin N, Bradley R S, Diaz H F, Baraer M, Caceres B, Forsythe N, Fowler H J, Greenwood G, Hashmi M Z R, Liu X D, Miller J R, Ning L, Ohmura A, Palazzi E, Rangwala I, Schner W, Severskiy I, Shahgedanova M, Wang M B, Williamson S N, et al. 2015. Elevation-dependent warming in mountain regions of the world. Nature Climate Change, 5, 424–430.

Plaut Z, Butow B J, Blumenthal C S, Wrigley C W. 2004. Transport of dry matter into developing wheat kernels and its contribution to grain yield under post-anthesis water deficit and elevated temperature. Field Crop Research, 86, 185–198.

Qian J X, Xi Y X. 2008. Analysis of characteristics and its influencing facters of winter wheat main growth stages. Chinese Agricultural Science Bulletin, 11, 438–443. (in Chinese)

Qu C H, Li X X, Ju H, Liu Q. 2019. The impacts of climate change on wheat yield in the Huang-Huai-Hai Plain of China using DSSAT-CERES-Wheat model under different climate scenarios. Journal of Integrative Agriculture, 18, 1379–1391.

Shi J J, Jiang X D, Shi H W, Chen Y J, Yuan J K, Jiang M. 2015. Effect of winter warming treatments on photosynthesis and yield of wheat. Journal of Triticeae Crops, 35, 352–356. (in Chinese)

Skinner D Z. 2014. Time and temperature interactions in freezing tolerance of winter wheat. Crop Science, 54, 1523–1529.

Subedi K D, Gregory P J, Summertfield R J, Gooding M J. 1998. Cold temperatures and boron deficiency caused grain set failure in spring wheat (Triticum aestivum L.). Field Crops Research, 57, 277–288.

Takai T, Matsuura S, Nishio T, Ohsumi A, Shiraiwa T, Horie T. 2006. Rice yield potential is closely related to crop growth rate during late reptoductive period. Field Crops Research, 96, 328–335.

Tashiro T, Wardlaw I F. 1990. The effect of high temperature at different stages of ripening on grain set, grain weight and grain dimensions in the semi-dwarf wheat ‘Banks’. Annals of Botany, 65, 51–61.

Tian Y L, Chen J, Chen C Q, Deng A X, Song Z W, Zheng C Y, Hoogmoed W, Zhang W J. 2012. Warming impacts on winter wheat phenophase and grain yield under field conditions in Yangtze Delta Plain, China. Field Crops Research, 134, 193–199.

Tian Y L, Chen J, Dong W J, Deng A X, Zhang W J. 2011. Effects of asymmetric warming on key enzyme activities of starch synthesis in superior and inferior grains of winter wheat under FATI facility. Acta Agronomica Sinica, 37, 1031–1038. (in Chinese)

Venkatesh K, Senthilkumar K M, Mamrutha H M, Singh G, Singh G P. 2022. High-temperature stress in wheat under climate change scenario, effects and mitigation strategies. Climate Change and Crop Stress, 8, 209–229.

Villegas D, Aparicio N, Blanco R, Royo C. 2001. Biomass accumulation and main stem elongation of durum wheat grown under mediterranean conditions. Annals of Botany, 88, 617–627.

Way D A, Yamori W. 2014. Thermal acclimation of photosynthesis: On the importance of adjusting our definitions and accounting for thermal acclimation of respiration. Photosynthesis Research, 119, 89–100.

Wrigley C W, Blumenthal C, Gras P W, Barlow E W R. 1994. Temperature variation during grain filling and changes in wheat-grain quality. Functional Plant Biology, 21, 875–885.

Xiao G J, Zhang Q, Li Y, Wang R Y, Yao Y B, Zhao H, Bai H Z. 2010. Impact of temperature increase on the yield of winter wheat at low and high altitudes in semiarid northwestern China. Agricultural Water Management, 97, 1360–1364.

Xiao G J, Zhang Q, Zhang F J, Lou C K, Wang R Y. 2011. The impact of rising temperature on spring wheat production in the Yellow River irrigation region of Ningxia. Acta Ecologica Sinica, 21, 6588–6593. (in Chinese)

Xu L, Zhao D Y, Wang Y Q, Zhang C Z. 2017. Determination of D-tagatose content by resorcinol method. Journal of Dalian Polytechnic University, 36, 168–170. (in Chinese)

Yamamori M, Quynh N. 2000. Differential effects of Wx-A1, -B1 and -D1 protein deficiencies on apparent amylose content and starch pasting properties in common wheat. Theoretical and Applied Genetics, 100, 32–38.

Yu X R, Hao D, Yang J Q, Ran L P, Zang Y, Xiong F. 2020. Effects of low temperature at stem elongation stage on the development, morphology, and physicochemical properties of wheat starch. PeerJ, 8, 2–22.

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