高山植物圆锥南芥的光合系统耐热性及其修复机制(英文)

高温胁迫包括极端高温和中高温,严重影响了植物的一系列生理活动,尤其是光合作用,而植物应对极端高温和中高温胁迫具有不同的策略。高山植物因长期生长于相对寒冷的环境中,相比而言应缺少对高温胁迫的适应机制。本文以圆锥南芥作为一种高山模式植物来探索其在中高温下是否表现出耐热能力,如果具有耐热能力,那么在光合方面与拟南芥存在怎样的差异。研究发现,圆锥南芥在中高温处理后具有更高的光化学效率及快速可逆的恢复过程,表现出了较强的耐热能力。两物种的F0没有明显的差异,而圆锥南芥在热处理后及恢复过程中具有更高的Fm,促进其快速光合修复。在热处理后,非光化学能量耗散快速瞬时上升,及时保护光系统II免受光损伤和热伤害,另外,HSP101蛋白迅速诱导可能启动了光化学修复。最后,圆锥南芥在严重高温处理后具有更高的存活率再次验证了它在中高温下的耐热能力。结果表明,圆锥南芥具有更耐热的光合系统以及有效的光合修复机制来耐受中高温胁迫。     

高山植物圆锥南芥的光合系统耐热性及其修复机制?

高温胁迫包括极端高温和中高温,严重影响了植物的一系列生理活动,尤其是光合作用,而植物应对极端高温和中高温胁迫具有不同的策略。高山植物因长期生长于相对寒冷的环境中,相比而言应缺少对高温胁迫的适应机制。本文以圆锥南芥作为一种高山模式植物来探索其在中高温下是否表现出耐热能力,如果具有耐热能力,那么在光合方面与拟南芥存在怎样的差异。研究发现,圆锥南芥在中高温处理后具有更高的光化学效率及快速可逆的恢复过程,表现出了较强的耐热能力。两物种的F0没有明显的差异,而圆锥南芥在热处理后及恢复过程中具有更高的Fm,促进其快速光合修复。在热处理后,非光化学能量耗散快速瞬时上升,及时保护光系统II免受光损伤和热伤害,另外, HSP101蛋白迅速诱导可能启动了光化学修复。最后,圆锥南芥在严重高温处理后具有更高的存活率再次验证了它在中高温下的耐热能力。结果表明,圆锥南芥具有更耐热的光合系统以及有效的光合修复机制来耐受中高温胁迫。     

植物在缺钾胁迫中膜稳定性的变化

植物维持膜的功能是其抵御胁迫的关键问题,而维持膜功能必须要保持膜的稳定性和合适的流动性。我们前期的研究发现植物主要是通过积累叶片膜脂和保持根部膜脂基本不变来适应长期缺钾。在本研究中,以拟南芥和其具有耐受缺钾胁迫特性的近缘种须弥芥为对象,研究了与膜的流动性密切相关的双键指数（double bond index,DBI）的变化,发现长期缺钾条件下,两种植物叶片中总的DBI保持不变,根部总的DBI略有降低。同时研究了与膜稳定性密切相关的溶血磷脂的含量和DGDG/MGDG以及PC/PE这两个比值的变化,发现长期缺钾后拟南芥和须弥芥叶片中溶血磷脂的总量呈上升趋势,根部溶血磷脂总量基本保持不变;无论在对照还是缺钾条件下,拟南芥溶血磷脂的总含量要高于须弥芥。须弥芥叶片具有更高的DGDG/MGDG值,根部具有更高的PC/PE值,说明长期缺钾条件下须弥芥膜的稳定性可能更好。这可能是须弥芥耐缺钾的原因之一。     

植物耐热相关基因研究进展

植物受到高温胁迫时,会激活某些特定基因的表达,从而增强植物的耐热性。近年来,随着生物技术的不断发展,植物耐热相关基因被相继克隆并转化植物体。本文对植物耐热的分子机制、相关基因的克隆、耐热性基因工程研究进展进行了综述,并提出了植物耐热基因工程的研究方向。     

青藏高原高山区和泛北极地区种子植物多倍体比较

多倍化是植物快速适应极端环境胁迫的一种重要机制。青藏高原高山区和泛北极地区具有相似的极端低温环境, 且两地的植物曾有密切的交流和联系。然而, 多倍体物种对两地植物区系生物多样性的贡献是否相同仍不清楚。我们系统地收集两地已有染色体数目和倍性报道的种子植物物种名录, 共计1,770种, 其中青藏高原高山区774种, 泛北极地区996种; 同时也相应地收集了每个物种的生活型信息。分析显示青藏高原高山区多倍体植物的比例为20.9%, 泛北极地区多倍体植物比例为61.5%; 青藏高原高山区一年生草本、多年生草本和木本植物中多倍体的比例分别为20.7%、21.6%和12.8%, 泛北极地区一年生草本、多年生草本和木本植物中多倍体的比例分别为60.2%、65.5%和38.3%。这些结果表明泛北极地区比青藏高原高山区具有较高比例的多倍体物种。青藏高原高山植物区系在渐新世‒中新世之交开始兴起, 此时高原已达到一定高度, 而后的高寒环境相对稳定, 致使多倍体物种相对较少; 而泛北极地区植物区系在3-4 Ma兴起, 此后经历了冰期‒间冰期、海平面波动等反复剧烈的气候环境变化, 可能促进了大量的多倍化事件发生。本研究通过比较青藏高原高山区和泛北极地区植物多倍体物种的比例, 揭示了两地多倍体比例差异的可能原因, 将提高对多倍体适应极端环境的理解。     

植物耐热性及热激信号转导机制研究进展

随着温室效应的加剧,全球气候变暖已经成为现代农业生产体系所面临的严峻挑战．高温灾害性气候是影响作物产量的一种主要的非生物胁迫．因此,对于农作物生产而言,研究植物耐热信号转导机制不仅有重要的科学意义,而且有现实的紧迫性．最近几年,在阐明植物耐热信号转导机制的研究方面取得了很多重要的进展,这些进展涵盖植物高温胁迫的感受机制、热激转录因子和热激蛋白的表达调控、热激转录因子结合蛋白参与耐热性调控的分子机制等几个主要的方面．热胁迫影响细胞膜系统、RNA、蛋白质的稳定性,同时改变酶的活性和细胞骨架系统．当热胁迫来临时,植物的转录组会发生显著变化,所涉及的基因大约占基因组的2％．这些高温胁迫响应基因构成了热激响应网络,是植物抵御热胁迫的第一道防线．植物的耐热性分为基础耐热性和获得性耐热性．基础耐热性是植物固有的耐热性．获得性耐热性是温和的热驯化诱导的耐热性．获得性耐热性状的形成反映了植物在自然生长环境下适应高温胁迫的生理机制．     

低温胁迫下脱落酸对线粒体膜磷脂酶D活性的影响

高山离子芥(Choraspora bungeana)是一种稀有高山冰缘植物,其生活环境具有低温、强紫外线等胁迫因子。PLD在膜磷脂降解及磷脂信号转导过程中发挥着重要作用,但其活性往往受到多种因素的影响。该研究以高山离子芥试管苗为材料,研究了4℃、0℃和-4℃胁迫下,ABA对高山离子芥试管苗叶中线粒体膜结合态PLD活性的影响。结果表明:10,50和100μmol·L~(-1)脱落酸(ABA)处理高山离子芥后,线粒体膜结合态PLD活性均较未添加ABA的处理组线粒体膜结合态PLD活性高,其中以50μmol·L~(-1) ABA对离子芥叶中线粒体膜结合态PLD活性的促进作用最为显著;外施0.3 mmol·L~(-1)的ABA合成抑制剂钨酸钠处理高山离子芥后,线粒体膜结合态PLD活性较对照组线粒体膜结合态PLD活性降低;在50μmol·L~(-1) ABA+5 mmol·L~(-1) EGTA处理组中,高山离子芥叶中线粒体膜结合态PLD活性低于未添加EGTA处理组线粒体膜结合态PLD活性;在0.3 mmol·L~(-1)钨酸钠+10 mmol·L~(-1)CaCl_2处理组中,高山离子芥叶中线粒体膜结合态PLD活性高于未添加CaCl_2处理组线粒体膜结合态PLD活性。由此推测,低温胁迫下ABA可能通过Ca~(2+)介导影响高山离子芥叶中线粒体膜结合态PLD的活性。     

抽穗扬花期高温胁迫对水稻主要品质指标的影响

高温热害对水稻生产造成了巨大的损失。利用国际水稻研究所耐热圃材料,在抽穗扬花期进行高温处理,研究了该时期高温胁迫对水稻主要品质评价指标的影响,结果如下：水稻耐热圃材料的耐热性有较大的差别,依然存在高温相对敏感材料;抽穗扬花期高温胁迫对水稻主要品质评价指标影响较大;高温胁迫对水稻籽粒的透明度、长宽比、垩白率、糊化温度和蛋白质含量等品质指标与品种的耐热性相关关系不大;高温胁迫下直链淀粉含量变化与品种耐热系数表现出显著的正相关关系（r=0.659 78）;高温胁迫下胶稠度变化与不同水稻品种的耐热系数呈显著的负相关关系（r=-0.516 70）。     

膜脂组成与植物抗冷性的关系及其分子生物学研究进展

植物的抗冷性与膜脂的组分和结构密切相关,与质膜中脂肪酸的不饱和度关系更为密切。膜脂不饱和脂肪酸含量越高,膜脂相变温度越低,植物的抗冷性提高。植物体内存在一些降低膜脂脂肪酸饱和程度的酶,如甘油3磷酸酰基转移酶,ω3脂肪酸去饱和酶等,它们能够催化膜中脂肪酸的去饱和反应,生成不饱和脂肪酸,从而提高植物的抗冷性。本文就低温对膜脂的影响、膜脂组成与植物抗寒性的关系及其分子生物学研究进展作一简单综述。     

植物适应长期缺钾——积累叶片膜脂、维持根膜脂组成不变

环境对植物的胁迫可能是短期快速的、也可能是长期而缓慢的,而植物应对这两种胁迫的策略可能不同。膜脂组成变化是植物响应环境胁迫的主要手段之一,其响应长期胁迫和短期胁迫的样式也可能不同。植物膜脂组成对短期缺钾胁迫的响应已经有报道,但是对长期缺钾的响应如何尚且未知。我们设置了4种（51,051,0051和0mmol·L-1）不同的钾浓度,比较了拟南芥（Arabidopsis thaliana）及其生长于贫钾生境中的近缘种须弥芥（Crucihimalaya himalaica）长期缺钾后（18天）的生理和生化变化,发现须弥芥具有耐受贫钾的能力。我们进一步运用脂类组学的方法检测比较了拟南芥和须弥芥在长期缺钾胁迫下脂类组成的变化,发现：(1) 两种植物叶片中总脂以及几乎所有脂类的含量明显上升;(2) 两种植物都是地上部分膜脂的变化幅度大于根部膜脂的变化幅度;(3) 地上部分膜脂变化幅度,须弥芥的大于拟南芥的;地下部分的膜脂变化幅度,须弥芥的小于拟南芥的;(4) 拟南芥叶片和根中PA的含量显著上升,与PA相对应的是PE含量的显著下降,由此我们推测拟南芥中PA的积累主要来自于PE的水解。上述结果提示,在细胞水平上,植物主要通过积累叶片膜脂和维持根部膜脂组成基本不变来适应长期缺钾。     

Arabidopsis mutants reveal that short- and long-term thermotolerance have different requirements for trienoic fatty acids

The photosynthetic thylakoid has the highest level of lipid unsaturation of any membrane. In Arabidopsis thaliana plants grown at 22°C, approximately 70% of the thylakoid fatty acids are trienoic - they have three double bonds. In Arabidopsis, and other species, the levels of trienoic fatty acids decline substantially at higher temperatures. Several genetic studies indicate that reduced unsaturation improves photosynthetic function and plant survival at high temperatures. Here, these studies are extended using the Arabidopsis triple mutant, fad3-2 fad7-2 fad8 that contains no detectable trienoic fatty acids. In the short-term, fluorescence analyses and electron-transport assays indicated that photosynthetic functions in this mutant are more thermotolerant than the wild type. However, long-term photosynthesis, growth, and survival of plants were all compromised in the triple mutant at high temperature. The fad3-2 fad7-2 fad8 mutant is deficient in jasmonate synthesis and this hormone has been shown to mediate some aspects of thermotolerance; however, additional experiments demonstrated that a lack of jasmonate was not a major factor in the death of triple-mutant plants at high temperature. The results indicate that long-term thermotolerance requires a basal level of trienoic fatty acids. Thus, the success of genetic and molecular approaches to increase thermotolerance by reducing membrane unsaturation will be limited by countervailing effects that compromise essential plant functions at elevated temperatures.     

Not changes in membrane fluidity but proteotoxic stress triggers heat shock protein expression in Chlamydomonas reinhardtii

A conserved reaction of all organisms exposed to heat stress is an increased expression of heat shock proteins (HSPs). Several studies have proposed that HSP expression in heat‐stressed plant cells is triggered by an increased fluidity of the plasma membrane. Among the main lines of evidence in support of this model are as follows: (a) the degree of membrane lipid saturation was higher in cells grown at elevated temperatures and correlated with a lower amplitude of HSP expression upon a temperature upshift, (b) membrane fluidizers induce HSP expression at physiological temperatures, and (c) membrane rigidifier dimethylsulfoxide dampens heat‐induced HSP expression. Here, we tested whether this holds also for Chlamydomonas reinhardtii. We show that heat‐induced HSP expression in cells grown at elevated temperatures was reduced because they already contained elevated levels of cytosolic HSP70A/90A that apparently act as negative regulators of heat shock factor 1. We find that membrane rigidifier dimethylsulfoxide impaired translation under heat stress conditions and that membrane fluidizer benzyl alcohol not only induced HSP expression but also caused protein aggregation. These findings support the classical model for the cytosolic unfolded protein response, according to which HSP expression is induced by the accumulation of unfolded proteins. Hence, the membrane fluidity model should be reconsidered.     

Effect of growth temperature on membrane fatty acid composition and susceptibility to cold shock of Bacillus amyloliquefaciens.

We investigated the fatty acid composition of the membrane of Bacillus amyloliquefaciens grown at different temperatures. A decrease in growth temperature was accompanied by an increase in the ratio of branched- to straight-chain fatty acids and a marked increase in the level of unsaturation of branched-chain fatty acids. When cells of this organism grown at 30 degrees C were cold shocked, viability and ability to secrete extracellular protease were lost. Growth of this organism at lower temperatures or addition of Tween 80 to cells caused the critical temperature zone for cold shocking to be lowered significantly. These results suggest a direct correlation between membrane fluidity and the susceptibility to cold shock.     

Identification of genetic diversity and mutations in higher plant acquired thermotolerance

Plants experience high air and soil temperatures during periods of drought and when fields receive limited irrigation. Elevated plant temperatures that occur under these conditions negatively impact plant health and productivity. Plants, like all organisms, respond to an elevation in temperature by the synthesis of heat shock proteins (HSP). The appearance of plant HSP is strongly correlated to the development of a condition termed 'acquired thermotolerance'. Acquired thermotolerance is induced by pre-exposure to elevated but non-lethal temperatures and leads to enhanced protection of plant cells from subsequent heat induced injury. Although the correlation between the development of acquired thermotolerance and the appearance of HSP is strong, a cause-and-effect relationship between the two has been difficult to demonstrate. To understand the relationship between HSP and acquired thermotolerance, mutations would be required that result in a coordinate change in the expressions of HSP. This paper describes research efforts leading to the development of a screening procedure for the isolation and characterization of acquired thermotolerance mutants. This method for identifying mutants is based on the inhibition of chlorophyll accumulation in etiolated tissue following challenges at lethal temperatures and the prevention of this inhibition by pre-incubation at a non-lethal elevated temperature; i.e. acquired thermotolerance. Arabidopsis thaliana mutants deficient in varying levels of acquired thermotolerance have been identified from both the RLD and Columbia ecotypes and these mutants are currently undergoing a detailed characterization at both the protein and molecular levels.     

Regulatory role of membrane fluidity in gene expression and physiological functions

Plants, algae, and photosynthetic bacteria experience frequent changes in environment. The ability to survive depends on their capacity to acclimate to such changes. In particular, fluctuations in temperature affect the fluidity of cytoplasmic and thylakoid membranes. The molecular mechanisms responsible for the perception of changes in membrane fluidity have not been fully characterized. However, the understanding of the functions of the individual genes for fatty acid desaturases in cyanobacteria and plants led to the directed mutagenesis of such genes that altered the membrane fluidity of cytoplasmic and thylakoid membranes. Characterization of the photosynthetic properties of the transformed cyanobacteria and higher plants revealed that lipid unsaturation is essential for protection of the photosynthetic machinery against environmental stresses, such as strong light, salt stress, and high and low temperatures. The unsaturation of fatty acids enhances the repair of the damaged photosystem II complex under stress conditions. In this review, we summarize the knowledge on the mechanisms that regulate membrane fluidity, on putative sensors that perceive changes in membrane fluidity, on genes that are involved in acclimation to new sets of environmental conditions, and on the influence of membrane properties on photosynthetic functions.     

Changes of Membrane Stability in Potassium Stressed Plants

The maintenance of membrane function is critical to the ability of plants to resist environmental stresses; specifically, the stability and appropriate fluidity of membranes are crucial to their normal function. We previously demonstrated that plants adapt to long term potassium (K+) deficiency by accumulation of membrane lipids in leaves and maintenance of the lipid composition in roots. In this study, which involved Arabidopsis thaliana and its K+ deficiency tolerant relative Crucihimalaya himalaica, we first calculated the double bond index (DBI) as an indicator of membrane fluidity. After exposure to long term K+ deficiency stress, the DBI of the total lipids in leaves of Athaliana and Chimalaica showed no significant changes, whereas the DBI of the total lipids in the roots of these species showed slight increases. Changes in lysophospholipids (lysoPLs) levels, and digalactosyldiacylglycerol/monogalactosyldiacylglycerol (DGDG/MGDG) and phosphatidylcholine/phosphatidylethanolamine (PC/PE) ratios, all of which strongly reflect membrane stability, were also studied in K+ stressed Athaliana and Chimalaica. After long term K+ deficiency, total lysoPLs levels increased in Athaliana and Chimalaica leaves, but showed no significant changes in roots. DGDG/MGDG and PC/PE ratios were higher in Chimalaica leaves and roots than in those of Athaliana. These results indicate that Chimalaica exhibits superior membrane stability compared with Athaliana. This may explain its superior growth and tolerance under K+ deficient conditions.     

Heat‐stress Memory is Responsible for Acquired Thermotolerance in Bangia fuscopurpurea

The environmental stresses that sessile organisms experience usually fluctuate dramatically and are often recurrent. Terrestrial plants can acquire memory of exposure to sublethal heat stress to acquire thermotolerance and survive subsequent lethal high‐temperature stress; however, little is known concerning whether seaweeds acquire thermotolerance via heat‐stress memory. We have demonstrated that the red seaweed Bangia fuscopurpurea can indeed acquire memory of sublethal high‐temperature stress, resulting in the acquisition of thermotolerance that protects against subsequent lethal high‐temperature stress. Moreover, the maintenance of heat‐stress memory was associated with a slight increase in the saturation level of membrane fatty acids. This suggests that the modification of membrane fluidity via changes in membrane fatty acid composition is involved in the establishment and maintenance of heat‐stress memory in B. fuscopurpurea. These findings provide insights into the physiological survival and growth strategies of sessile red seaweeds to cope with recurrent changes in environmental conditions.     

Regulation of fatty acid unsaturation in encystingAcanthamoeba cells

The degree of fatty acid unsaturation and average chain length are closely similar for microsomal membranes from exponential-phase trophozoites and cysts ofAcanthamoeba castellanii despite significant differences in fatty acid composition. The same trend was apparent for total fatty acids extracted from whole cells. The observations suggest that the organism regulates these lipid parameters during differentiation in order to maintain optimum membrane lipid viscosity, and are consistent with previous electron spin resonance measurements indicating that the fluidity of microsomal membranes does not change during encystment. About 75% of the microsomal fatty acids are unsaturated for both cysts and amoebae. Wide-angle X-ray diffraction of phospholipid liposomes prepared from lipid extracts of the membranes has indicted that this high level of unsaturation renders the phospholipid exclusively liquid-crystalline at temperatures as low as 9°C for rough microsomes and-1.5°C for smooth microsomes. Thus, by retaining a high proportion of unsaturated fatty acids throughout its differentiation cycle, the organism gains some protection in its natural soil habitat against lateral phase separation of membrane lipids.