水稻高温胁迫的生理响应及耐热机理研究进展

自国际水稻所最先开展耐热水稻品种选育至今近三十几年来,国内外大批学者对于高温灾害评估与预测,耐热水稻品种选育及高温胁迫抗性鉴定,高温胁迫水稻形态发育、产量、品质、生理生化响应及其防御与调控,耐高温胁迫水稻品种遗传改良等方面开展了大量研究,并取得了很大的进展。该综述简要回顾了国内外水稻高温胁迫研究进展,重点评述了高温胁迫对水稻生长的影响,水稻高温胁迫下的生理响应,高温肋迫下其他环境因素对水稻生长的综合效应,以及水稻耐热/热响应的可能分子机制,以期为水稻耐热性研究提供参考。     

小麦对高温胁迫的响应机制研究进展

小麦是中国主要粮食作物之一,为了应对全球气候变暖对农作物的影响,明确小麦对高温环境胁迫的响应,本文归纳了最新研究高温对小麦生长发育、产量品质的影响。从生理生化方面归纳了小麦光合作用、相关酶活性、氧化系统、植物激素对高温胁迫的响应。根据文献分析提出小麦未来研究发展的建议：（1）选育抗逆性强的品种,（2）合理栽培管理的调控措施来减轻高温对小麦的影响。     

高温胁迫对水稻的影响及其对策的研究进展

在全球气候变暖的大环境下,极端天气尤其是高温热害频繁发生,研究高温对水稻影响的机理与对策对促进水稻生产具有重要意义。综述了高温胁迫对水稻生长发育、产量形成和稻米品质的影响,并从光合特性及干物质积累、代谢酶、激素和抗逆物质等方面阐述了其生理机制,提出减轻水稻高温胁迫的具体防御措施,并对未来全球气温升高环境下的水稻生产研究提出建议。     

水稻生长对干旱胁迫的响应及抗旱性研究进展

水稻是我国主要粮食作物,也是用水量最多的作物.研究水稻抗旱性,培育抗旱水稻品种,实现水资源不足背景下水稻的高产和稳产,有助于缓和粮食生产与水资源短缺之间的矛盾.国内外学者对水稻抗旱机理、抗旱鉴定指标以及抗旱相关基因的分析等做了大量研究并取得了突破性进展.本文从形态特征、生理生化特性以及分子水平等方面综述了干旱胁迫对水稻生长的影响,全面了解水稻在干旱胁迫下的形态变化及其生理生化抗旱机制,以期为鉴定筛选抗旱种质、选育抗旱品种提供参考.     

植物有性生殖对高温胁迫的响应机制

极端高温天气频发,严重威胁农作物的生产。高温胁迫对有性生殖过程的影响与作物减产密切相关,解析其中的分子机制对于指导作物耐高温遗传改良具有重要意义。然而,与模式植物拟南芥相比,目前有关作物有性生殖过程中耐高温的相关研究十分有限。本文从植物有性生殖过程出发,概述了在减数分裂、花粉绒毡层降解、小孢子发育、花粉管萌发与授精以及籽粒发育等一系列生殖发育过程中响应高温胁迫的分子机制。据此,我们提出了作物耐高温改良的可行策略,以期为耐高温品种的遗传改良提供理论依据。     

根系对土壤环境胁迫响应的研究进展

根系是土壤和植物的动态界面,植物根系具有锚定植株、吸收和运输土壤中的水分及养分、合成和贮藏营养物质等重要功能。近年来,随着研究方法的改进和现代根系定量分析技术的发展,对根系的研究日益受到重视和关注。因而,根系的研究在多个新的研究方向上展开。本文简要介绍了国内外近年来在根系对土壤环境胁迫响应的研究进展,以供根系研究参考。     

湿地松优良半同胞家系蛋白质及糖类对水分胁迫的生理响应

为研究湿地松优良半同胞家系蛋白质及糖类对水分逆境的生理响应，采用盆栽试验和吸光度测定结合的方法，以湿地松普通种子为对照（CK），测定了七个湿地松半同胞家系在正常水分状态（土壤相对含水量为70%）、弱度水分胁迫（土壤相对含水量为55%~60%）、中度水分胁迫（土壤相对含水量为35%~40%）和强度水分胁迫（土壤相对含水量为20%~25%）条件下的可溶性蛋白质、还原性糖、可溶性糖的含量，研究结果表明：无论是不同水分梯度还是不同家系水平，蛋白质、还原性糖、可溶性糖含量的差异均达极显著性水平。随着水分胁迫程度的加剧，各家系蛋白质含量呈先升后降的趋势，在水分胁迫初期，各家系蛋白质含量增加，水分胁迫中、后期各家系蛋白含量呈直线下降；随着水分胁迫的加剧，各家系还原性糖、可溶性糖含量呈直线上升。家系1027、101、464在水分胁迫条件下具有较高的可溶性糖含量，其抗旱性较其它家系要强，是这三个家系速生、高产的生理原因之一。     

高温胁迫下水稻叶片的蛋白响应及其基因型和生育期差异

高温已经成为水稻产量的主要限制因素,且其影响由于全球温室效应的加剧而呈扩大趋势。本研究在鉴定耐热水稻基因型的基础上,从生理学和蛋白质组学上进行耐性机理研究。结果表明,苗期或抽穗期高温处理导致结实率、SPAD值、株高、根长和生物量下降,丙二醛、过氧化氢、超氧阴离子含量增加和超氧化物歧化酶活性提高。同时,高温胁迫对热敏感品种明恢63的影响大于对耐热品种密阳46的影响。蛋白质组学分析表明,高温使光合作用相关蛋白、能量类蛋白、代谢类蛋白表达量下降,抗逆相关蛋白表达量上升。另外,蛋白试验结果佐证了密阳46的耐热性以及水稻抽穗期对高温的敏感性。本研究还首次发现抗逆相关蛋白2-cys过氧化物酶BAS1的表达量在高温下上升。     

转PuP5CS基因烟草对低温胁迫的生理响应

以朝鲜碱茅PuP5CS基因(HQ637435)为目的基因,构建了PuP5CS基因的正义表达载体和反义表达载体,通过农杆菌介导的叶盘法转化烟草,研究了PuP5CS基因对烟草耐寒性的影响。结果显示,低温胁迫下,正义转基因烟草体内可溶性糖、脯氨酸、叶绿素a和叶绿素b的含量均显著高于野生型烟草,丙二醛含量增幅显著低于野生型烟草,提高了转基因烟草的抗寒性,而反义转基因烟草体内各项生理指标变化不明显。     

水分胁迫对干热河谷乡土树种清香木的光合影响

为了研究水分胁迫对清香木光合特性的影响,根据土壤田间持水量设置不同的土壤水分含量梯度（W1~W6,土壤田间持水量分别为100%、80%、50%、30%、10%、5%）。以清香木为材料,利用Li-6400测量6个不同土壤水分含量梯度下清香木叶片的净光速率(Pn)、胞间CO₂浓度(Ci)、气孔导度(Gs)、蒸腾速率(Tr)和水分利用率(WUE)。了解清香木对水分过多或过少胁迫的响应,旨在为云南干热河谷地区的植被恢复、清香木的培育种植提供一定的理论指导和技术参考。结果表明：（1）不同土壤水分含量梯度下清香木净光合速率Pn值显著差异,总体Pn呈现“降—升—降”的态势,日变化曲线呈“双峰”型,具明显的午休现象。Pn下降主要受到气孔限制和非气孔限制因素的影响。W1、W2受到气孔的限制,W3~W6受非气孔限制的因素影响,随着土壤含水量的下降,气孔限制向非气孔限制转变。（2）18:00以后,除了W3以外,其他Pn出现负值,呼吸作用大于光合作用,植物的光合生产力下降,植物死亡;W3则为水分适度的最佳选择,其田间持水量和土壤含水量分别在50%和33.6%左右。（3）气孔导度随着土壤含水量的减小而下降,土壤含水量在W3~W6时,气孔导度Gs日变化曲线无明显,基本呈现平缓状态。W1~W6的Gs值在8:00时最大,主要是空气湿度引起。（4）水分利用率WUE随着土壤含水量的下降而增加,适当的干旱胁迫有利于提高清香木光合生产力和水分利用率。     

杂草稻种子芽期对盐碱胁迫的响应

盐碱胁迫对未吸胀种子发芽率、发芽势、芽长和主根长伤害率大于吸胀种子.WR 03-12的吸胀种子在各浓度盐碱胁迫下发芽率和发芽势受影响较小,未吸胀种子发芽率和发芽势只在高中性盐和高碱性盐处理下与对照相比达显著差异.无论是吸胀种子还是未吸胀种子,WR 03-12发芽率及发芽势相对伤害率均小于其他两个材料.     

Physiological Response to Heat Stress During Seedling and Anthesis Stage in Tomato Genotypes Differing in Heat Tolerance

Tomato cultivars differ in their sensitivity to heat stress, and the sensitivity depends on the developmental stage of the plants. It is less known how heat stress affects tomato at the anthesis stage in terms of leaf physiology and fruit set and whether the ability of tomato to tolerate heat at different developmental stages is linked. To investigate photosynthetic gas exchange characteristics, carbohydrate content and fruit set during heat stress, a thermo‐tolerant cultivar (‘LA1994’) and a thermo‐sensitive cultivar (‘Aromata’) were studied at the seedling and anthesis stage. The photosynthetic parameters, maximum quantum efficiency of photosystem II (F_v/F_m), chlorophyll content, carbohydrate content and fruit set were determined in plants grown at 26/18 °C (control) and 36/28 °C (heat stress). The physiological responses including net photosynthetic rate (P_N), chlorophyll content and F_v/F_m decreased in ‘Aromata’ at both developmental stages during heat stress, whereas they were unaltered in ‘LA1994’ during heat stress as compared to the respective control. This was accompanied by lower contents of glucose and fructose in mature leaves of ‘Aromata’ at the seedling stage under heat stress. In contrast, the glucose content increased while the fructose content was unaltered in mature leaves of ‘LA1994’ at the seedling stage under heat stress. High temperature induced a similar change in carbohydrate content in the young leaves of both cultivars at anthesis. The fructose and sucrose content were unaffected in the mature leaves of ‘Aromata’ but significantly increased in ‘LA1994’ under heat stress at anthesis. The heat stress treatment decreased pollen viability and inhibited fruit set due to flower wilting and abnormal abscission in ‘Aromata’, whereas fruit set was not inhibited in ‘LA1994’. A decrease in chlorophyll content, photosynthesis and carbohydrate content in the mature leaves of tomato could be related to fruit set failure at high temperature. The results show that physiological responses to heat stress at the seedling stage correspond with the responses at the anthesis stage, demonstrating that screening for heat stress sensitivity can be carried out in young plants.     

Heat Stress Effects are Stronger on Spikelets Than on Flag Leaves in Rice Due to Differences in Dissipation Capacity

Rice is most sensitive to heat stress at the flowering stage, with different degrees of heat damage in spikelets and leaves. To investigate the heat damage in spikelets and flag leaves, two rice genotypes, N22 (heat‐tolerant) and GT937 (heat‐sensitive), were subjected to a heat‐stress treatment (40 °C for 15 days). The results showed that more damaging was found in spikelets than in flag leaves and the heat stress significantly decreased the seed‐setting rate by 12.41 % in N22 and by 65.02 % in GT937. However, the difference in the net photosynthetic rate of the flag leaf between heat‐stressed and control was not significant. Moreover, the difference of temperatures in spikelets and flag leaves was attributed to the differences in heat dissipating. Under heat stress, the transpiration rate was significantly higher in flag leaves than in spikelets, and the temperature in flag leaves were at least 4 °C cooler than in spikelets. Although the spikelet temperatures did not differ significantly between the two genotypes under heat stress, spikelets of GT937 were more severely damaged than those of N22, which might result from the differences in the antioxidant capacities between genotypes. Results showed that little difference of superoxide dismutase, peroxidase and catalase activities of spikelets was found in N22, while significant reduction was found in GT937 under heat stress, compared with control. These results suggest that organ temperature is controlled mainly by transpirational cooling, and that heat stress is an indirect result of oxidative stress, rather than a direct result of heat damage.     

Evaluation of Cold Tolerance in NERICAs Compared with Japanese Standard Rice Varieties at the Reproductive Stage

New Rice for Africa (NERICA) is a general name for interspecific rice varieties derived from a cross between the high‐yielding Asian rice (Oryza sativa L.) between locally adapted African rice (Oryza glaberrima Steud.). Eight NERICAs were evaluated for cold tolerance (CT) at the reproductive stage and compared with their O. sativa parents and three Japanese standard rice varieties over 3 years. Cold tolerance was evaluated based on the filled grain ratio (FGR) after cold water irrigation. The FGR was greatly reduced by cold water irrigation. NERICA 1, 2 and 7 had higher FGR (51.9–57.9 %), while NERICA 6, 15 and 16 had lower FGR (6.2–14.5 %). NERICA 1, 2 and 7 were less affected by cold stress, with a 31 % mean reduction in FGR, while NERICA 6, 15 and 16 were greatly affected, with their FGRs being reduced by more than 80 %. NERICA 3 and 4 were moderately affected by cold stress, with about 45 % reduction rate in FGR. FGR significantly influenced the grain weights of the varieties with strong positive correlations (r = 0.83–0.91; P < 0.001), and thus, similar trends in grain weights were observed. Grain weights were reduced by 61.7–96.4 % under cold stress. NERICA 1, 2 and 7 showed significantly better performance than NERICA 3 and 4, while NERICA 6, 15 and 16 performed poorly under cold water irrigation. The Japanese varieties Koshihikari (very tolerant) and Ozora (moderately tolerant) were more affected by cold water irrigation than NERICA 1, 2 and 7. On the basis of the mean reduction rate (%) in FGR under cold stress, the varieties were classified as follows: NERICA 1, 2 and 7 as tolerant; NERICA 3 and 4 as moderately tolerant; and NERICA 6, 15 and 16 as susceptible to cold stress. However, NERICA 7 grain yields were lower under cold stress due to both greatly reduced number of panicles per plant and number of spikelets per panicle. Therefore, NERICA 1 and 2 are suitable candidates for production in the highland regions of East Africa and should be promoted for production.     

北方粳稻灌浆期水分胁迫条件下叶片内源激素变化的研究

为了阐明北方超级粳稻‘沈农265’的抗旱性生理机制,采用盆栽人工控制水分的方法,以北方超级粳稻‘沈农265’和米质优良的常规粳稻品种‘丰锦’为材料,在灌浆期进行不同程度的水分处理,比较两者生理反应上的差异。结果表明,与‘丰锦’相比,中度胁迫使‘沈农265’剑叶中叶绿素含量比‘丰锦’高5.03%,可溶性蛋白、可溶性糖含量及内源激素(IAA+GA3+ZR)/ABA比值分别比‘丰锦’低32.4%、11.41%和33.33%;重度胁迫下,‘沈农265’剑叶中叶绿素含量和可溶性蛋白分别比‘丰锦’高6.38%和20.2%,可溶性糖含量及(IAA+GA3+ZR)/ABA比值分别比‘丰锦’低51.5%和25%。上述结果说明,‘沈农265’具有较强的渗透调节能力,重度胁迫下,可溶性蛋白可能起主要作用;同时,‘沈农265’在相对高的(IAA+GA3+ZR)水平基础上增加ABA含量,不仅提高了干旱适应能力,也延缓了衰老,使其具有比丰锦更强的抗旱能力。     

中国水稻热害研究历史、进展与展望

水稻热害的研究正引起世界范围内的高度关注。近50年中国长江流域共发生重大水稻热害事件6次。最近一次发生在2003年,保守估计全流域受害面积达3×107 hm^2,损失稻谷达5.18×107 t。长江流域水稻热害的严重发生,主要与其特殊的气候环境有关,也与大面积推广的部分杂交稻品种对高温较敏感有关。2003年以后,长江流域水稻热害渐成为中国农业研究的一个热点,目前主要研究集中在热害形成的田间小气候环境、持续高温天气的形成机制与预测、水稻抗性材料和品种的鉴定方法、水稻抗性性状的确认、基因定位与利用和水稻高温反应机理等方面。     

水稻幼苗生长对弱光胁迫的响应及相关分析

耐弱光性不同的水稻品种对光照条件反应不同,确定水稻耐弱光鉴定指标,可以快速有效地从大量种质资源中筛选耐弱光的水稻品种,提高品种耐弱光选择效率。试验对不同基因型的61份水稻品种（品系）于苗期进行遮光处理,分别研究各品种在不同光照下的叶龄、苗高、根长、地上部干重、根干重、根冠比、根体积、叶绿素（SPAD值）、单株重的变化,并对各性状的相对指数进行相关及通径分析。试验结果表明,在55%遮光条件下,不同水稻品种的叶绿素含量均增加,苗高变化不一,多数品种的根长、叶龄、根体积、根干重有不同程度的下降,少数品种的根长、叶龄、根体积、根干重增加,各品种地上部干重和单株重均有不同程度的下降;而且,弱光下地上部干重、根干重、根长的相对指数与单株重相对指数直接通径系数分别为0.7592、0.2105、0.0346,相关系数为0.9034、0.7634、0.6862,可见,弱光下的地上部干重相对指数是鉴定水稻耐弱光性的可靠指标,弱光下的根干重和根长相对指数可作为耐弱光性鉴定的重要辅助指标。     

化控措施对直播晚稻抽穗扬花期低温胁迫的缓解效应

直播晚稻抽穗扬花期易遭受寒露风危害,影响产量与品质。利用自然低温条件,以陵两优722和中早35为试验材料,设置T1（芸苔素内酯+赤霉素）、T2（芸苔素内酯+赤霉素+磷酸二氢钾）、T3（赤霉素+磷酸二氢钾）和对照（CK,清水）4个处理,研究化学调控剂对直播晚稻遭遇寒露风危害的缓解作用。结果表明,在寒露风危害下,喷施不同化学调控剂均可有效缩短水稻抽穗时间,与CK相比,陵两优722和中早35生育期平均缩短了2.5d,其中,T1生育期均缩短1d,T2分别缩短3d和5d,T3分别缩短2d和4d,喷施化学调控剂还可有效减缓叶绿素相对值（SPAD值）与净光合速率（P_n）的下降,CK的SPAD值较第1次测定最大下降幅度平均为6.55%,T1、T2和T3分别为3.50%、4.22%和3.77%;CK的P_n较第1次测定最大下降幅度平均为10.59%,T1、T2和T3分别为5.96%、1.22%和3.36%。喷施化学调控剂不仅能稳定结实率与千粒重,且产量、稻米加工品质及外观品质均优于CK,与CK相比,陵两优722与中早35在T1处理下结实率分别提高8.17%与2.31%,T2分别提高6.20%与5.36%,T3分别提高7.88%与6.31%;千粒重则仅T2有促进作用,陵两优722与中早35较CK分别提高了1.67%与2.92%;与CK相比,陵两优722和中早35在T1处理产量平均提升9.01%,T2平均提升20.69%,T3平均提升19.01%,出糙率、精米率与整精米率较CK均明显提高,垩白度与垩白粒率明显降低。总体而言,磷酸二氢钾、赤霉素和0.01%芸苔素内酯混合液（T2）对水稻低温胁迫的缓解效果最明显。     

皖浙地区早稻高温热害发生规律及其对产量结构的影响研究

探讨高温与水稻产量的关系,在水稻高温热害防御中有重要意义。利用皖浙地区气象数据与早稻生育状况观测数据,分析了皖浙地区早稻高温热害规律及高温为害与早稻产量结构的关系。结果表明：皖浙2省早稻受高温热害危害的敏感期常年为6月中旬至7月中旬,且高温热害发生在时间分布上存在着周期震荡,其中20世纪80年代中期以前高温热害在发生日数和次数上呈减少趋势,其后则呈增多趋势;在空间分布上,安徽省沿江、江淮早稻主产区的高温热害重于沿淮地区;浙江省的金华—丽水一带为早稻高温热害重发区。在高温热害对早稻产量结构因素的影响中,高温热害与穗结实粒数、千粒重之间为负相关,与空壳率和秕谷率之间呈正相关,且扬花灌浆期高温热害对水稻空壳率和穗粒数影响较为显著。研究认为早稻抽穗扬花期遭遇高温热害,会对水稻结实率、千粒重、空壳率、秕谷率造成不等程度危害,其中穗粒数减少、空壳率增加是高温热害影响产量的主要原因。     

Mapping of Quantitative Trait Loci Associated with Reproductive‐Stage Salt Tolerance in Rice

Salinity tolerance in rice varies with the state of growth, with the seedling and reproductive stages being the most sensitive. However, association between tolerances at the two stages is poor, suggesting that they are regulated by different processes and genes. Tolerance at the reproductive stage is the most crucial as it determines grain yield. An F₂ mapping population was developed from two rice genotypes contrasting in tolerance: Cheriviruppu and Pusa Basmati 1 (PB1). Cheriviruppu is highly tolerant at the reproductive stage, while PB1 is highly sensitive at both seedling and reproductive stages. One hundred and thirty‐one microsatellite markers polymorphic between the parents were used to construct a linkage map of 1458.5 cM (Kosambi), with a mean intermarker distance of 11.1 cM. Sixteen QTLs with LOD values ranging from 3.2 to 22.3 were identified on chromosomes 1, 7, 8 and 10, explaining 4–47 % of the phenotypic variation. The maximum number of QTL clusters for different component traits was colocalized on the long arm of chromosome 1 and chromosome 7. We identified several significant epistatic interactions, including three inter‐QTL interactions, using MapManager. The results suggest that pollen fertility, Na⁺ concentration and Na/K ratio in the flag leaf are the most important mechanisms controlling salt tolerance at the reproductive stage in rice. The study reports the construction of a genetic map for reproductive‐stage salt tolerance in rice and demonstrates its utility for molecular mapping of QTLs controlling salinity tolerance‐related traits, which will be useful in marker‐assisted selection in the future.