To ensure the food security of future generations and to address the challenge of the 'no hunger zone' proposed by the FAO (Food and Agriculture Organization), crop production must be doubled by 2050, but environmental stresses are counteracting this goal. Heat stress in particular is affecting agricultural crops more frequently and more severely. Since the discovery of the physiological, molecular, and genetic bases of heat stress responses, cultivated plants have become the subject of intense research on how they may avoid or tolerate heat stress by either using natural genetic variation or creating new variation with DNA technologies, mutational breeding, or genome editing. This review reports current understanding of the genetic and molecular bases of heat stress in crops together with recent approaches to creating heat-tolerant varieties. Research is close to a breakthrough of global relevance, breeding plants fitter to face the biggest challenge of our time.© The Author(s) 2020. Published by Oxford University Press on behalf of the Society for Experimental Biology.

构建考虑高温天气过程(包括发生时间、持续日数)的水稻高温热害指标,可以实现对水稻高温热害等级的动态判识,对精准监测、预警与评估水稻高温热害意义重大。以江西早稻为对象,利用气象资料、早稻高温热害灾情史料和生育期资料,反演历史早稻高温热害,采用K-S分布拟合检验和置信区间方法,构建基于高温天气过程的早稻高温热害动态指标,并采用预留的独立的早稻高温热害样本进行检验验证。在此基础上,计算江西各站点早稻高温热害指数(M),并分析水稻热害。结果表明: 高温天气过程起始时间、持续日数是影响早稻高温热害发生程度的关键因子,其中,起始时间的影响大于持续日数。3～5 d早稻轻、中、重度高温热害的起始时间阈值分别为抽穗后第10～12、5～9、2～4天;6～8 d早稻轻、中、重度高温热害的起始时间阈值为抽穗后第11～18、8～10和1～7天;>8 d早稻轻、中、重度高温热害的起始时间阈值为抽穗后第12～18、8～11和0～7天。指标验证完全一致的吻合率为73.7%,完全一致及相差1级的吻合率为89.5%。1981—2015年,M的线性倾向率为0.04·a^-1,1999年左右发生由低到高突变;M高值区域主要位于江西中部和东北部,M>0.18;江西中部、东北部和南部地区M值呈显著增加趋势,线性倾向率均大于0.04·a^-1。总体来说,本文构建的指标实现了基于高温天气过程的早稻高温热害动态判识,江西中部和东北部是早稻高温热害的高风险区域。

【目的】研究水稻产量对不同时期不同类型高温的响应差异,为耐热水稻品种的筛选与培育提供依据。【方法】在桶栽条件下,于人工气候室内,以耐热水稻品种Nagina 22、汕优63和热敏感品种两优培九为材料,设置三个处理时期,即穗分化期、开花期和灌浆期;四种处理温度,即白天高温、夜间高温、全天高温和适宜温度,研究水稻产量和产量构成因子对不同时期高温的响应。【结果】汕优63在三个时期均表现出较强的耐热性;Nagina 22在花期和灌浆期对高温耐性较强,但易受穗分化期高温影响;两优培九对不同时期高温均较敏感。汕优63在不同时期高温下均表现出稳定的伤流强度,而高温敏感品种两优培九的伤流强度受高温影响均显著下降。【结论】水稻对不同时期的高温耐性并不具有相关性,筛选或培育耐热品种时应考虑其对不同时期高温的综合耐受性。

为探究脱落酸(ABA)对水稻(Oryza sativa)抽穗开花期高温胁迫的诱抗效应, 以江西省主推水稻品种黄华占为材料, 于孕穗期用蒸馏水、ABA溶液(10、50和100 µmol∙L^-1)、氟啶酮(FLU)和原花青素(PC) 6种溶液进行叶面喷施, 然后移入对照(CK)和高温胁迫(HS)环境处理8天, 考查籽粒活性氧(ROS)积累、抗氧化防御能力、产量构成及相关基因的表达。结果表明, 高温胁迫下, 水稻的穗长、穗重、结实率、千粒重和产量与超氧阴离子和过氧化氢含量呈显著负相关。高温胁迫下, 喷施ABA显著上调了ABA应答和抗氧化防御基因的表达, 籽粒中活性氧含量下降了8.24%-31.35%; 喷施ABA显著增加了水稻的穗长、穗重、结实率和千粒重, 显著上调了产量形成基因的表达, 增产12.73%-20.77%。高温胁迫下, 喷施FLU可抑制ABA的生物合成, 导致ROS过量积累和水稻减产; 喷施抗氧化剂PC则抑制ROS过量积累, 使产量增加。以上结果表明, 高温胁迫下, 孕穗期喷施ABA不仅能够激发ABA信号通路, 而且上调抗氧化防御能力和产量形成基因的表达, 进而提高水稻在抽穗开花期的耐热性, 达到增产目的。

Global warming has caused frequent occurrence of heat stress at the flowering stage of single-season rice in the Yangtze River region of China, which results in declines of spikelet fertility and yield in rice. Rice flowering stage is the most sensitive period to high temperatures, and therefore, the key for heat stress happening is the flowering stage coinciding with high temperature, which causes spikelet fertility decreasing in heat-sensitive varieties, and is the major factor for heat injury differences among various rice planting regions. With the development of rice breeding, temperature indexes for heat stress has been converted from daily maximum temperature of 35 ºC to 38 ºC with the stress duration of more than 3 d. During the flowering stage, anther dehiscence inhibition and low pollen shedding onto the stigma are two main reasons for spikelet fertility reduction under high temperatures. At panicle initiation stage, high temperatures aggravate spikelet degeneration, and destroy floral organ development. Various types of rice varieties coexist in production, and indica-japonica hybrid rice demonstrates the highest heat resistance in general, followed by indica and japonica rice varieties. In production, avoiding high temperature is the main strategy of preventing heat stress, and planting suitable cultivars and adjustment of sowing date are the most effective measures. Irrigation is an effective real-time cultivation measure to decline the canopy temperature during the rice flowering stage. We suggested that further study should be focused on exploring heat injury differences among different rice variety types, and innovating rice-planting methods according to planting system changes in rice planting regions with extreme heat stress. Meanwhile, high temperature monitor and warning systems should be improved to achieve optimal heat stress management efficiencies.

【目的】为了明确灌浆结实前中期不同温度对晚粳稻籽粒灌浆、总淀粉及其组分含量的影响特征及其机理,【方法】以优质食味粳稻南粳9108为材料,利用植物培养箱控温,于水稻灌浆结实期设置高温(HT,日均温度31℃)、中温(MT,日均温度26℃)和低温(LT,日均温度21℃)三个处理,以自然温度为对照(CK),研究灌浆结实前中期持续高低温对晚粳稻籽粒灌浆、内源激素含量、淀粉及其及合成关键酶活性的影响。【结果】随温度上升(21℃~31℃),晚粳稻结实率和千粒重降低;灌浆前中期籽粒总淀粉积累速率加快,但持续时间短,总含量降低,而直链淀粉含量及其所占比例在整个灌浆期均呈降低趋势;灌浆前中期脱落酸(ABA)含量显著增加,赤霉素(GA₃)含量降低,玉米素核苷(ZR)含量呈“先增先降”趋势;MT处理有利于提高灌浆结实期籽粒生长素(IAA)含量,HT和LT处理均降低籽粒IAA含量,LT处理下,IAA含量先降后升,而HT处理籽粒IAA含量一直处于较低水平;籽粒可溶性淀粉合成酶(SSS)活性与ZR变化规律相似,也呈“先增先降”趋势,但二者在灌浆结实期并不同步;淀粉分支酶(SBE)活性增加而淀粉脱分支酶(DBE)活性降低。【结论】不同温度显著影响晚粳稻籽粒内源激素含量和淀粉合成关键酶活性的变化,进而影响晚粳稻籽粒灌浆、总淀粉及其组分的积累。

高温和干旱是影响水稻生长、发育、产量和品质的两个最重要的环境因子,全面理解高温和干旱胁迫对评价气候变化对水稻生产的影响至关重要。概述了高温、干旱及其复合胁迫对水稻生长发育、产量形成和稻米品质的影响;从光合作用、抗氧化系统、内源激素、蔗糖-淀粉代谢途径关键酶活性、分子机制等方面阐述其生理机制;提出减轻水稻高温干旱胁迫的调控措施;对未来深入开展水稻高温干旱逆境的研究提出建议。

全球气候变暖环境下,水稻高温热害频发,孕穗-开花期和灌浆期高温已成为制约水稻产量和品质的主要因素之一。本文综述了水稻高温热害的发生特点(鉴定与分级、区域和时间)和高温对水稻生长发育(生理、产量和品质)的影响,总结了水稻高温热害的数量性状座位定位、转录组和蛋白质组分析等分子生物学研究及监测预警与风险评估,重点阐述了高温热害的防御措施,包括选用耐热品种、改善田间管理和喷施外源物质,并对今后水稻高温热害研究进行了展望,以期为水稻高温热害防御和农业减灾增效提供科学支持。

为研究杂交稻对开花结实期高温胁迫的生理生态适应性，选用具代表性的主栽杂交稻组合国稻6号、协优46，人工设计极值高温40~42℃，自始穗期至此后15 d每天6 h行热害处理，以自然气候条件为对照，比较研究高温胁迫对国稻6号与协优46结实的生理影响。结果表明，国稻6号具较高的受孕小穗成粒效应，即不仅不孕小穗率低，而且秕谷率也低。国稻6号对开花结实期高温较协优46钝感，两者在小穗育性和热害指数上的差异达到显著水平(P<0.05)。究其生理原因, (1)成熟期间稻株剑叶光能转化效率及光合效率较高，茎鞘储藏物质较丰；(2)成熟期间稻株仍具较强根系吸水能力与叶片蒸腾水平，维持蒸腾流利于光合物质运输；(3)成熟灌浆期稻株自动调节粒间顶端优势，增进迟开花的弱势粒充实成实粒。

High temperature (HT) at meiosis stage is one of most important environment constraint affecting spikelet fertility and rice yield. In this paper, the effects of HT exposure at meiosis stage on the ROS (reactive oxygen species) accumulation, various superoxide dismutase (SOD, EC1.15.1.11) isozymes in developing anther, and its relationship with HT-induced decline in pollen viability and floret fertility were investigated by using four rice cultivars differing in heat tolerance under well-controlled climatic condition. Results showed that HT exposure significantly increased ROS level and malondialdehyde (MDA) content in rice anther, and this occurrence was strongly responsible for the HT-induced decline in pollen viability and harmful effect of HT adversity on floret fertility. However, the increased extent of ROS concentration in rice anther under HT exposure was greatly variable, depending on both the intensity and duration of HT exposure and different rice cultivars used. The SOD and CAT activities of HT-sensitive cultivars decreased more profoundly than those of HT-tolerant cultivars under the same HT regimes. Among various types of SOD enzymes, Cu/Zn-SODa expressed highly in rice anther and responded sensitively to HT exposure, while Cu/Zn-SODb expressed weakly in rice anther and preferentially in rice leaves. HT exposure suppressed the expression of Cu/Zn-SODa in developing anther, which was closely associated with the down-regulated transcripts of cCu/Zn-SOD1 gene. Hence, Cu/Zn-SODa may play a central role in the regulation of total SOD activity and ROS detoxification in rice anther as affected by HT exposure at meiosis stage.Copyright © 2017. Published by Elsevier Masson SAS.

Around the intensity, frequency, duration, accumulated temperature, and even extremes of high-temperature events, nine selected temperature-related indices were used to explore the space and time changes of rice heat stress in Jiangxi province, southeast China. Several statistical methods including Mann-Kendall trend test (M-K test) and principal component analysis (PCA) were used in this study, and main results were listed as follows: (1) The changes in the intensity indices for high-temperature events were more significant, it was mainly embodied in that more than 80 % of stations had positive trends. (2) R-mode PCA was applied to the multiannual average values of nine selected indices of whole stations, and the results showed that the higher hazard for rice heat stress could be mainly detected in the middle and northeast area of Jiangxi. (3) S-mode PCA was applied to the integrated heat stress index series, and the results demonstrated that Jiangxi could be divided into four sub-regions with different variability in rice heat stress. However, all the sub-regions are dominated by increasing tendencies in rice heat stress since 1990. (4) Further analysis indicated that the western north Pacific sub-tropical high (WPSH) had the significant dominant influence on the rice heat stress in Jiangxi province.

Background: Pollen tube elongation in the pistil is a key step for pollination success in plants, and auxins play an important role in this process. However, the function of auxins in pollen tube elongation in the pistil of rice under heat stress has seldom been previously reported.Results: Two rice genotypes differing in heat tolerance were subjected to heat stress of 40 degrees C for 2 h after flowering. A sharp decrease in spikelet fertility was found in the Nipponbare (NPB) and its mutant High temperature susceptible (HTS) under heat stress, but the stress-induced spikelet sterility was reversed by 1-naphthaleneacetic acid (NAA), especially the HTS. Under heat stress, the pollen tubes of NPB were visible in ovule, while those of HTS were invisible. However, we found the pollen tubes in ovule when sprayed with NAA. During this process, a significant increase in indole-3-acetic acid (IAA) and reactive oxygen species (ROS) levels was found in the pistil of heat-stressed NPB, while in heat-stressed HTS they were obviously decreased. Additionally, the peroxidase (POD) activity in pistil of NPB was significantly decreased by heat stress, whereas there was no difference between the heat-stressed and non-heat-stressed pistils of HTS.Conclusion: It was concluded that the enhancement of heat tolerance in plants by NAA was achieved through the increase of the levels of auxins, which prevented the inhibition of pollen tube elongation in pistil, and the crosstalk between auxins and ROS, which might be involved in this process. In addition, POD might be a negative mediator in pollen tube elongation under heat stress due to its ability to scavenge ROS and degrade auxin.

【目的】 穗粒数是水稻产量构成因子之一。为明确减数分裂期高温对水稻穗粒数的可能影响,构建其定量估算模型,【方法】于2014–2015年以两优培九和南粳45为供试材料,于减数分裂期设置不同强度和持续时间的高温处理,以自然无高温环境为对照,分析了减数分裂期不同高温强度和高温持续时间对穗粒数的影响,建立了减数分裂期高温与穗粒数的定量关系,最后根据温度日变化特征,构建了自然环境下的减数分裂期高温对穗粒数的定量影响算法。【结果】 减数分裂期高温对穗粒数的影响程度与高温强度、持续时间有关。同样高温条件下,穗粒数随高温持续天数增加呈指数减少;日相对穗粒数随温度增加而减少,可用二次曲线进行描述。穗粒数对高温敏感的终止时间大约为抽穗前5 d。利用2015年试验资料对上述模型进行了检验,两优培九和南粳45穗粒数模拟值的相对均方根差为0.094和0.085,说明模型可很好地模拟减数期高温对穗粒数的影响。【结论】 对完善高温对水稻生长过程的定量影响具有重要作用。

【目的】随着全球气候变暖的加剧,灌浆期高温逼熟对水稻产量和品质的威胁日趋严重,明确水稻不同耐热性品种对灌浆期高温逼熟的反应及其与气象因子间的关系,对提高优质稻栽培和育种水平意义重大。【方法】对48个不同类型的水稻品种分4期进行大田播种,使各品种在灌浆期经历不同气象条件,探讨其主要产量、品质性状表现及其与气象因子的关系。【结果】多数品种粒重和品质指标在播期间差异显著;根据各指标变异系数和指标间相关性,确定千粒重、整精米率和垩白度等作为水稻品种灌浆期响应气象条件的主要农艺学指标;而气象因子中,日平均温度、相对湿度和日照时数为主要影响指标。采用隶属函数值对品种进行分类,感性品种千粒重、整精米率和垩白度的平均差异分别比耐性品种大1.98 g、30.02%和19.81%。进一步分析隶属函数值随日平均气温、相对湿度和日照时数的分布发现,齐穗后1–15 d日平均气温≥28℃、相对湿度≤84%、日照时数6~10 h时品种间耐性差异最大。【结论】垩白度、整精米率和千粒重可作为品种对高温逼熟响应的代表性农艺学指标;田间形成高温逼熟危害的气象条件是以温湿度为主导的综合气象条件,齐穗后1–15 d日平均气温≥28℃、平均相对湿度≤84%和日照时数≥6.5 h时,品种间耐性差异最大,是田间高温逼熟耐性筛选和鉴定的适宜气象条件。

【目的】研究抽穗结实期不同时段高温处理对稻米品质的影响，阐明高温对稻米品质的影响机理及其时段效应。【方法】选用耐热性不同的两个水稻品系，利用人工气候室进行抽穗结实期不同时段温度试验，研究抽穗结实期高温对稻米外观品质、加工品质、直链淀粉含量、胶稠度、蛋白质含量、RVA特征谱、Mg/ K及籽粒的淀粉粒结构的影响。【结果】抽穗结实期高温使两品系稻米长宽比变小，垩白粒率和垩白度增加，糙米率、精米率、整精米率降低，直链淀粉含量和蛋白质含量增加，胶稠度降低；淀粉RVA谱特征值崩解值降低，冷胶粘度、消减值和糊化温度升高，峰值时间延长，Mg、K含量增加，Mg/K比值降低；淀粉粒扫描电镜显示，高温条件下，淀粉粒间结合较为疏松，籽粒内大多以单个淀粉粒的形式存在，折光率下降，胚乳透明度降低，导致垩白形成。相同处理条件下，耐热品系996稻米品质受高温影响低于热敏感品系4628。抽穗结实期不同时段高温对稻米品质影响存在显著差异，以始穗后8—14 d对稻米品质影响最大，其次为始穗后15—21 d、1—7 d、始穗后22—28 d影响较小。【结论】始穗后8—21 d是温度对稻米品质影响的关键时期，对稻米品质形成具有决定作用，而结实后期（始穗22 d后）环境温度对稻米品质影响较弱。

Heat stress often results in the generation of reactive oxygen species, such as hydrogen peroxide, which plays a vital role as a secondary messenger in the process of abscisic acid (ABA)-mediated stomatal closure. Here, we characterized the rice (Oryza sativa) HEAT TOLERANCE AT SEEDLING STAGE (OsHTAS) gene, which plays a positive role in heat tolerance at the seedling stage. OsHTAS encodes a ubiquitin ligase localized to the nucleus and cytoplasm. OsHTAS expression was detected in all tissues surveyed and peaked in leaf blade, in which the expression was concentrated in mesophyll cells. OsHTAS was responsive to multiple stresses and was strongly induced by exogenous ABA. In yeast two-hybrid assays, OsHTAS interacted with components of the ubiquitin/26S proteasome system and an isoform of rice ascorbate peroxidase. OsHTAS modulated hydrogen peroxide accumulation in shoots, altered the stomatal aperture status of rice leaves, and promoted ABA biosynthesis. The results suggested that the RING finger ubiquitin E3 ligase OsHTAS functions in leaf blade to enhance heat tolerance through modulation of hydrogen peroxide-induced stomatal closure and is involved in both ABA-dependent and DROUGHT AND SALT TOLERANCE-mediated pathways.

Episodes of high temperature at anthesis, which in rice is the most sensitive stage to temperature, are expected to occur more frequently in future climates. The morphology of the reproductive organs and pollen number, and changes in anther protein expression, were studied in response to high temperature at anthesis in three rice (Oryza sativa L.) genotypes. Plants were exposed to 6 h of high (38 degrees C) and control (29 degrees C) temperature at anthesis and spikelets collected for morphological and proteomic analysis. Moroberekan was the most heat-sensitive genotype (18% spikelet fertility at 38 degrees C), while IR64 (48%) and N22 (71%) were moderately and highly heat tolerant, respectively. There were significant differences among the genotypes in anther length and width, apical and basal pore lengths, apical pore area, and stigma and pistil length. Temperature also affected some of these traits, increasing anther pore size and reducing stigma length. Nonetheless, variation in the number of pollen on the stigma could not be related to measured morphological traits. Variation in spikelet fertility was highly correlated (r=0.97, n=6) with the proportion of spikelets with > or = 20 germinated pollen grains on the stigma. A 2D-gel electrophoresis showed 46 protein spots changing in abundance, of which 13 differentially expressed protein spots were analysed by MS/MALDI-TOF. A cold and a heat shock protein were found significantly up-regulated in N22, and this may have contributed to the greater heat tolerance of N22. The role of differentially expressed proteins and morphology during anther dehiscence and pollination in shaping heat tolerance and susceptibility is discussed.

Rice grain yield and quality are predicted to be highly vulnerable to global warming. Five genotypes including heat-tolerant and susceptible checks, a heat-tolerant near-isogenic line and two hybrids were exposed to control (31 °C/23 °C, day/night), high night-time temperature (HNT; 31 °C/30 °C), high day-time temperature (HDT; 38 °C/23 °C) and high day- and night-time temperature (HNDT; 38 °C/30 °C) treatments for 20 consecutive days during the grain-filling stage. Grain-filling dynamics, starch metabolism enzymes, temporal starch accumulation patterns and the process of chalk formation were quantified. Compensation between the rate and duration of grain filling minimized the impact of HNT, but irreversible impacts on seed-set, grain filling and ultimately grain weight were recorded with HDT and HNDT. Scanning electron microscopy demonstrated irregular and smaller starch granule formation affecting amyloplast build-up with HDT and HNDT, while a quicker but normal amylopast build-up was recorded with HNT. Our findings revealed temporal variation in the starch metabolism enzymes in all three stress treatments. Changes in the enzymatic activity did not derail starch accumulation under HNT when assimilates were sufficiently available, while both sucrose supply and the conversion of sucrose into starch were affected by HDT and HNDT. The findings indicate differential mechanisms leading to high day and high night temperature stress-induced loss in yield and quality. Additional genetic improvement is needed to sustain rice productivity and quality under future climates.© Society for Experimental Biology 2017.