植物对UV-B辐射的响应与调控机制

UV-B(ultraviolet-B)辐射会在多个方面对植物产生影响,随着研究的不断深入,人们发现UV-B除了作为一个胁迫因子外,还会作为信号调节因子来调节植物的生长发育.本文把UV-B辐射对植物形态建成和生理代谢的影响、植物响应UV-B辐射的信号通路以及UV-B与其他因子复合作用对植物的影响等进行了论述,并对植物响应UV-B辐射的研究做了展望.     

植物响应UV-B辐射的研究进展

地表UV-B辐射的增强对植物的生长生理产生了多方面影响。随着研究的不断深入, 人们认识到UV-B辐射不仅是一种胁迫因子, 而且是一个重要的信号调节分子。该文论述了近年来植物响应UV-B辐射研究的一系列成果, 包括UV-B辐射对植物形态建成、生理代谢、UV-B光受体UVR8蛋白、细胞程序性死亡、细胞骨架和细胞周期的影响, 及其它因素与UV-B复合处理对植物的作用; 并对植物响应UV-B辐射研究进行了望。     

UV-B辐射对植物影响的分子水平研究进展

大气平流层中的臭氧层被破坏,导致到达地球表面太阳的UV-B辐射增强,对植物产生了多方面影响,其中包括对DNA、抗氧化酶系统、光系统Ⅱ（PSⅡ）的作用以及植物体内类黄酮等保护物质的生物合成等。UV-B辐射也是环境中重要的非生物因子,植物在长期的进化演替过程中,形成了对它的适应机制,可能被作为一种环境信号调节植物体内一系列的基因表达过程。本文论述了近些年来在分子水平上UV-B辐射对植物DNA损伤的修复、抗氧化酶和光系统Ⅱ的基因表达影响,以及有关UV-B信号传导,并对UV-B辐射的植物分子生物学研究作了展望。     

盐生植物胀果甘草和光果甘草对UV-B以及盐胁迫的不同响应

盐胁迫及增强的UV-B辐射(280~320 nm)是影响干旱地区植物生长及生存的重要环境因子。在本研究中,研究单独的UV-B辐射和不同浓度盐胁迫及其复合作用对两种甘草属盐生植物(胀果甘草Glycyrrhiza inflata Bat.和光果甘草Glycyrrhiza glabra L.)的种子萌发、幼苗的抗氧化酶活性、丙二醛、脯氨酸、酚类物质含量以及生长相关指标的影响进行了分析。结果表明:UV-B辐射对两种植物的种子萌发都没有显著性影响,但是高盐胁迫的存在明显降低了它们的3种萌发相关参数,而胀果甘草表现出比光果甘草更快的萌发速率;两种甘草属植物除了地上及地下部分的干重外,其根长、株高、地上、地下部分鲜重都明显受高盐胁迫及其与UV-B复合作用的影响而降低。在本研究中所有的胁迫条件下,尤其是UV-B强辐射,都造成了胀果甘草的脂质过氧化,而光果甘草的脂质过氧化仅受到UV-B和高盐复合胁迫的影响。所有的胁迫条件都增加了两种植物的脯氨酸含量,而最高含量出现在两种植物的高盐胁迫组和UV-B处理的胀果甘草组中。不同胁迫处理对两种甘草的抗氧化酶系统有不同的效应,盐胁迫明显提高两种甘草的POD活性及光果甘草的SOD活性,但使光果甘草CAT活性明显降低。所有胁迫条件下植物的花青素及类黄酮类物质含量都得到了提升,而增强的UV-B辐射条件下最明显。从以上结果可以看出,两种甘草的生物量都没有受到胁迫环境的影响,这可能和脯氨酸、酚类物质及抗氧化酶所发挥的保护效应有关。此外,胀果甘草对盐胁迫以及UV-B胁迫的耐受性要高于光果甘草。     

非生物胁迫下植物细胞壁组分变化

植物细胞壁主要由纤维素、半纤维素、果胶、木质素和糖蛋白组成,其在植物生长中主要起结构支持、物质运输和抵御逆境的作用.植物生长在受到各种环境信号影响后,细胞壁特性会发生很大改变.这些环境信号也会改变细胞壁组分的含量和结构,从而改变细胞壁机械特性.这种细胞壁的改变可以认为是植物对环境胁迫的响应.本文主要综述在非生物环境胁迫下,包括水分亏缺、低温胁迫、重金属胁迫和增强UV-B辐射下细胞壁多糖含量和结构,细胞壁结构蛋白和细胞壁相关酶活性,以及分布在细胞间隙的小分子物质的响应和机制,结合近年来细胞壁相关基因水平、基因组水平和蛋白组水平方面的研究结果,讨论了今后该领域的研究方向.     

UV-B辐射胁迫下不同起源时期的3种木本植物幼苗的生长及光合特性

平流层臭氧的减薄导致到达地表UV-B辐射增强是全球所面临的环境问题之一。UV-B辐射胁迫对植物的生物学效应研究成为继全球大气二氧化碳浓度升高对植物影响研究之后的又一热点领域。设置了UV-B滤光减弱组、UV-B辐射增强组和自然光对照组3组大田实验,选择不同起源时期的乐东拟单性木兰(Parakmeria lotungensi)、青冈(Cyclobalanopsis glauca)、山核桃(Carya cathayensis)幼苗为实验材料,测定每组中3种植物的生长量与光合特征参数,通过对比组间和种间差异,研究不同起源时期的3种木本植物对UV-B辐射胁迫的响应模式,分析3种植物对于UV-B辐射胁迫的适应性与自身起源和进化时间的关系,为"起源时间越早的木本植物生长发育和光合生理能否更好地适应UV-B辐射胁迫"这一科学命题的探讨提供一定实验参考。得到如下结果:(1)相对于自然光照条件,增强UV-B辐射胁迫对3种木本植物的地径和株高都有抑制作用;对乐东拟单性木兰、青冈的Pn和Amax有一定的抑制作用,对山核桃Pn和Amax则具有一定的促进作用。减弱UV-B辐射胁迫对3种木本植物的地径起到抑制作用,对乐东拟单性木兰、青冈幼苗的株高生长有促进作用,但对山核桃的株高却具有抑制作用;对乐东拟单性木兰以及山核桃的Pn和Amax有一定的抑制作用,而对青冈Pn和Amax则有促进作用。(2)对比种间差异,发现3种不同的UV-B光照条件下青冈的地径生长量都最大,乐东拟单性木兰次之,山核桃最小;株高生长量种间大小排序不一致;相对于自然光照条件,增强UV-B辐射强度下山核桃Pn、Amax的比值都最大,青冈次之,乐东拟单性木兰最小;减弱UV-B辐射强度下青冈的Pn、Amax的比值都最大,乐东拟单性木兰次之,山核桃最小;表明不同起源时间对植物抗UV-B辐射胁迫能力有一定的影响,但不是决定性因素。UV-B辐射增强和过滤减弱胁迫对3个树种幼苗的生长发育、光合作用、叶绿素均有影响,但不同起源时期3种木本植物幼苗光合特征参数的响应模式不一,其机制尚待进一步开展实验进行求证。本研究结果可丰富和补充UV-B辐射胁迫对木本植物的影响研究,为从进化角度筛选UV-B胁迫抗性较强的植物提供了一定的依据。     

UV-B辐射方向对白三叶克隆整合的影响

增强UV-B辐射会对植物生长和生理生化过程产生有害效应。克隆植物中,相连的克隆分株对经常共享资源和激素,然而鲜有关于异质性UV-B辐射下UV-B辐射方向对克隆整合的影响及克隆植物形态结构变化的报道。模拟同质(克隆分株片段均处于自然背景辐射)和异质(克隆分株一端处于自然背景辐射,另一端处于补加的UV-B辐射)UV-B辐射,以克隆植物白三叶为材料,进行连接和隔断处理,研究UV-B辐射方向对克隆整合强度变化、叶片形态结构特化及生理可塑性的影响。结果表明:异质性UV-B辐射下,15N同位素标记端保留的15N百分比高于同质UV-B辐射处理,转移到无标记相连端的15N含量则降低,紫外辐射处理和同位素标记是否处于同一分株端对结果无显著性影响,说明克隆植物白三叶生理整合存在但整合强度降低,辐射方向与克隆整合强度无关;隔断处理组气孔长度增加,栅栏组织增厚,但连接处理组却无此变化,表明生理整合在白三叶叶片形态结构特化中发挥作用。UV-B辐射下,最小荧光、电子传递速率及光化学淬灭系数降低但非光化学淬灭系数升高,而生理整合却使结果相反;叶绿素和紫外吸收物可在异质性UV-B辐射相连的两端运输分享。以上均表明异质UV-B辐射环境中,UV-B辐射胁迫端克隆分株通过生理整合从非胁迫端获益,并以此提高胁迫环境中克隆植物对资源的利用效率。     

克隆整合提高异质性UV-B辐射下活血丹抗氧化酶活力

增强UV-B辐射会对植物生长和生理生化过程产生有害效应。克隆植物中,相连的克隆分株对经常共享资源和激素,然而鲜有关于异质性UV-B辐射下抗氧化酶活力变化的报道。本研究模拟同质(克隆分株片段均处于自然背景辐射)和异质(克隆分株一端处于自然背景辐射,另一端处于补加的UV-B辐射)UV-B辐射,以克隆植物活血丹为材料,进行连接和隔断处理,研究异质性UV-B辐射下,克隆整合对活血丹抗氧化酶(超氧化物歧化酶(SOD),过氧化物酶(POD)和过氧化氢酶(CAT))活力的影响。结果表明:与处于同质UV-B辐射环境相比,异质UV-B辐射下连接处理中的活血丹UV-B辐射端抗氧化酶活力显著增加,说明克隆植物生理整合存在,且克隆整合提高了活血丹抗氧化酶活力。表明异质UV-B辐射环境中,UV-B辐射胁迫端克隆分株通过生理整合从非胁迫端获益,最大化地利用资源。     

植物对UV-B辐射增强应答的分子机制及信号级联研究进展

地表的UV-B辐射量伴随着大气平流层臭氧层的变薄而不断增强,给地球生态系统带来严重影响.UV-B主要通过抑制植物光合作用、伤害生物膜及DNA等生物大分子来影响其生长发育,最终导致生物量及产量降低,甚至致死.植物在进化过程中形成了自我防护及防御机制,如DNA损伤的自我修复,活性氧自由基的酶促及非酶促清除机制,以及紫外吸收物质的诱导合成等;同时,在植物中也有许多物质及不同途径来感受和应答UV-B胁迫.本文从UV-B辐射增强对植物造成损伤的主要途径、植物对UV-B辐射增强的应答机制及信号级联过程等方面的研究进展进行综述.     

光受体UVR8对UV-B响应机制研究进展

来自太阳光谱中的UV-B辐射被认为是一种重要的环境信号,可以被植物感受并诱导植物调整自身生长和发育状态以适应环境。人们对植物中光敏色素、隐花色素和蓝光受体向光素的研究已非常深入,但对植物响应UV-B的机制仅在最近才取得一些突破性进展。这些研究发现,植物中存在着UV-B受体UVR8(UV Resistance Locus 8)。目前认为,UVR8二聚体感应UV-B后瞬间解聚为单体,并与E3泛素连接酶COP1(constitutively photomorphogenic 1)相互作用,从而激活UV-B响应基因的表达。该文从UVR8的发现、UVR8的结构和感受UV-B机制、UVR8二聚体重新形成以及UV-B信号传导与可见光信号传导途径间的差异等方面综述了关于UV-B受体UVR8的最新研究成果。     

Photosynthetic and biochemical characterization of pigments and UV-absorbing compounds in <Emphasis Type="Italic">Phormidium tenue</Emphasis> due to UV-B radiation

We report the effect of UV-B radiation (0.8 ± 0.1 mW cm⁻²) and UV-B radiation supplemented with low-intensity PAR (∼80 μmol photons m⁻² s⁻¹) on the photosynthesis, photosynthetic pigments, phosphoglycolipids, oxidative damage, enzymatic antioxidants, and UV-absorbing compounds in Phormidium tenue, a marine cyanobacterium. UV-B radiation resulted in a decline in photosynthesis and photosynthetic pigments leading to lower biomass. P. tenue synthesized UV-absorbing compounds like mycosporine-like amino acids (MAAs) and scytonemin in response to UV-B radiation. Quantity of MAAs and scytonemin was higher when UV-B was supplemented with low-level PAR. UV-B treatment also resulted in quantitative changes in phosphoglycolipids of the membrane. The UV-B treatment resulted in a slight increase in the level of peroxidation of cell membrane and very little increase in the activity of superoxide dismutase (SOD). Results indicate that UV-B affected photosynthesis and that the main protective system was the synthesis of MAAs and scytonemin-like compounds rather than antioxidant enzymes such as SOD.     

水稻对UV-B辐射增强的抗性遗传及其生理生化特性研究

本研究结果表明紫外线B(UVB,280～330nm)能严重影响水稻秧苗生长发育,从而引起植株矮化,叶片干枯,叶绿素含量降低,单株干物质减少甚至死亡。不同水稻基因型对紫外线的反应存在明显差异。在3个供试品种(HR、M63、Hr)中,HR最抗UVB危害。遗传分析表明水稻抗UVB的性状为数量遗传性状且受多对隐性基因控制,生化分析表明水稻秧苗经UVB处理后,在0～3d内,随着时间推移,秧苗保护酶活性明显增强,之后,保护酶活性逐惭下降,MDA含量相应增加。抗UVB品种HR比感UVB品种Hr保护酶活性一直较高,膜质过氧化程度较低,反映了保护酶对抵抗UVB危害的重要作用。     

植物UV-B生理效应的分子机制研究进展

中波紫外线UV-B（280～320nm）是植物必需的太阳光线的组成部分,具有明显的双重效应：一方面UV-B在强度较高时,就触发产生大量活性氧对DNA、蛋白质以及生物膜等造成伤害,同时植物通过抗氧化系统对其作出防御反应以减轻伤害;另一方面,低强度的UV-B是植物生长发育的光信号因子之一,经由UVR8等光受体介导中、低、极低强度的UV-B信号,可能通过几个分子途径控制相关基因的表达,分别对植物的UV-B保护基因表达、形态建成、昼夜节律、生长发育等进行调控。目前对UVR8介导的低强度UV-B信号转导的分子机制研究相对深入。在本文中,将对UV-B生理效应分子机制的最新研究进展作一个比较全面的介绍。     

连续多代UV-B照射对麦长管蚜保护酶活性的影响

【目的】为探明中波紫外线（UV-B）连续多代处理对麦长管蚜 Sitobion avenae (Fabricius ) 的作用机制,发现在UV-B环境压力下害虫进化动力。【方法】用强度为0.25 W/m²的UV-B对每代麦长管蚜进行6 h照射,以处理后成蚜当天产的仔蚜为继代饲养材料,连续饲养18代,提取第1,4,8,12,15和18代的对照组和处理组无翅孤雌成蚜的保护酶液,利用分光光度计测定其超氧化物歧化酶（SOD）、过氧化氢酶（CAT）和过氧化物酶（POD）3种保护酶活性。【结果】不同世代的对照组间3种保护酶活性均无差异（P>0.05）,处理组间有显著差异（P<0.05）。随着UV-B胁迫世代增加,SOD活性呈先升高后下降的趋势,活性高峰出现在第15 代。POD活性呈下降趋势,处理组活性均高于对照组,且在第1和4代处理组与对照组间差异极显著（P<0.01）。CAT活性先下降后升高,在第8, 12和15代处理组与对照组间无显著差异（P>0.05）,而至第18代处理组与对照组间差异又恢复为极显著（P<0.01）。【结论】麦长管蚜受到短期UV-B胁迫时,其体内SOD,POD和CAT 3种保护酶活性均升高,以抵御UV-B胁迫引起的自由基伤害;而经过到长期UV-B胁迫后,主要依靠SOD和CAT 2种保护酶清除活性氧自由基。     

Ultraviolet-B radiation effects on the Mediterranean ruderal Dittrichia viscosa

Young seedlings of Dittrichia viscosa L. (syn. Inula viscosa (L.) Aiton) (Asteraceae) were extensively treated with artificial rain in order to remove the water soluble component of their epicuticular UV-B absorbing compounds. As a result, 75% of the epicuticular absorbing capacity at 300 nm was lost. The seedlings were subsequently grown in a naturaly lit glasshouse for 80 days under 0.06, 6.41 and 10.14 kJ m^-2 day ^-1 biologicaly effective UV-B radiation doses. The initial, pre-rain values of the water soluble, epicuticular UV-B absorbing potential was restored in about three weeks. During this transient period the plants were exposed to the enhanced UV-B radiation doses with part of their UV-B radiation screen removed. Although a trend for increased accumulation of epicuticular UV-B absorbing capacity was observed with increasing UV-B radiation doses, the allelopathic potential of the epicuticular material remained unchanged. Internal (cellular) UV-B absorbing compounds and chlorophylls were unaffected, but total carotenoids were increased, indicating a possible protective role against UV-B radiation damage. Leaf, stem and root dry mass were the same under all treatments but UV-B radiation caused a reduction in the dry mass invested per unit leaf area with a concomitant increase in leaf area. The importance of this UV-B radiation induced selective allocation of photosynthate to the production of assimilative surfaces is discussed.     

Effects of ultraviolet-B radiation on the growth and yield of crop plants

This paper reviews growth chamber, greenhouse, and field studies on the effects of ultraviolet-B (UV-B. between 280 and 320 nm) radiation on agricultural crop plants. Our understanding of the physiological effects of UV-B radiation comes primarily from growth chamber studies, where UV-B is artificially supplied via filtered lamps. Both photosystems I and II, as well as carboxylating enzymes, are sensitive to UV-B radiation. Ultraviolet-B radiation also affects stomatal resistance, chlorophyll concentration, soluble leaf proteins, lipids, and carbohydrate pools. In general, the effects of UV-B radiation are accentuated by the low levels of visible radiation typically found inside growth chambers. Ultraviolet-B radiation has also been shown to affect anatomical and morphological plant characteristics. Commonly observed UV-B induced changes include plant stunting, reductions in leaf area and total biomass, and alterations in the pattern of biomass partitioning into various plant organs. In sensitive plants, evidence of cell and tissue damage often appears on the upper leaf epidermis as bronzing, glazing, and chlorosis. Epidermal transmission in the UV region decreases in irradiated leaves. This decrease is primarily associated with a stimulation in flavonoid biosynthesis and is thought to be a protective, screening response to the deleterious effects of UV-B. A considerable degree of variability exists in sensitivity to UV-B radiation between different species. Approximately 30% of the species tested were resistant, another 20% were extremely sensitive, and the remainder were of intermediate sensitivity, in terms of reductions in total dry weight. In addition to this sizable interspecific variability, there appears to be a similarly wide intraspecific variability in UV-B response. The effects of UV-B radiation on crop yield have only been examined in a limited number of field studies, with ambient levels of UV-B radiation being supplemented with fluorescent sun lamps. Due to various deficiencies, all these field experiments to date have only limited utility for assessing the potential impact of enhanced levels of UV-B on crop productivity.     

Impact of solar ultraviolet-B radiation (290-320 nm) upon marine microalgae

For years scientists and laymen alike have casually noted the impact of solar ultraviolet radiation upon the non-human component of the biosphere. It was not until recently, when human activities were thought to threaten the protective stratospheric ozone shield, that researchers undertook intensive studies into the biological stress caused by the previously neglected short-wavelength edge of the global solar spectrum. Stratospheric ozone functions effectively as an ultraviolet screen by filtering out solar radiation in the 220–320 nm waveband as it penetrates through the atmosphere, thus allowing only small amounts of the longer wavelengths of radiation in this waveband to leak through to the surface of the earth. Although this ultraviolet radiation (UV-B radiation, 290–320 nm) comprises only a small fraction (less than 1%) of the total solar spectrum, it can have a major impact on biological systems due to its actinic nature. Many organic molecules, most notably DNA and proteins, absorb UV-B radiation which can initiate photochemical reactions. It is life's ability, or lack thereof, to cope with enhanced levels of solar UV-B radiation that has generated the concern over the potential depletion of stratospheric ozone. The defense mechanisms that serve to protect both plants and animals from current levels of UV-B radiation are quite varied. Whether these mechanisms will suffice for marine microalgae under conditions of enhanced levels of UV-B radiation is the subject of this review.     

Responses of aquatic algae and cyanobacteria to solar UV-B

Continuous depletion of the stratospheric ozone layer has resulted in an increase in solar ultraviolet-B (UV-B; 280–315 nm) radiation reaching the Earth's surface. The consequences for aquatic phototrophic organisms of this small change in the solar spectrum are currently uncertain. UV radiation has been shown to adversely affect a number of photochemical and photobiological processes in a wide variety of aquatic organisms, such as cyanobacteria, phytoplankton and macroalgae. However, a number of photosynthetic organisms counteract the damaging effects of UV-B by synthesizing UV protective compounds such as mycosporine-like amino acids (MAAs) and the cyanobacterial sheath pigment, scytonemin. The aim of this contribution is to discuss the responses of algae and cyanobacteria to solar UV-B radiation and the role of photoprotective compounds in mitigating UV-B damage.     

Outdoor Studies on the Effects of Solar UV-B on bryophytes: Overview and Methodology

In this review all recent field studies on the effects of UV-B radiation on bryophytes are discussed. In most of the studies fluorescent UV-B tubes are used to expose the vegetation to enhanced levels of UV-B radiation to simulate stratospheric ozone depletion. Other studies use screens to filter the UV-B part of the solar spectrum, thereby comparing ambient levels of UV-B with reduced UV-B levels, or analyse effects of natural variations in UV-B arising from stratospheric ozone depletion. Nearly all studies show that mosses are well adapted to ambient levels of UV-B radiation since UV-B hardly affects growth parameters. In contrast with outdoor studies on higher plants, soluble UV-B absorbing compounds in bryophytes are typically not induced by enhanced levels of UV-B radiation. A few studies have demonstrated that UV-B radiation can influence plant morphology, photosynthetic capacity, photosynthetic pigments or levels of DNA damage. However, there is only a limited number of outdoor studies presented in the literature. More additional, especially long-term, experiments are needed to provide better data for statistical meta-analyses. A mini UV-B supplementation system is described, especially designed to study effects of UV-B radiation at remote field locations under harsh conditions, and which is therefore suited to perform long-term studies in the Arctic or Antarctic. The first results are presented from a long-term UV-B supplementation experiment at Signy Island in the Maritime Antarctic.     

Effects of Enhanced UV-B Radiation on the Activity and Expression of Alternative Oxidase in Red Kidney Bean Leaves

An increase in ultraviolet （UV） B radiation on the earth＇s surface is a feature of current global climate changes. It has been reported that alternative oxidase （AOX） may have a protective role against oxidative stress induced by environmental stresses, such as UV-B. To better understand the characteristic tolerance of plants to UV-B radiation, the effects of enhanced UV-B radiation on the activity and expression of AOX in red kidney bean （Phaseolus vulgaris） leaves were investigated in the present study. The results show that the total respiration rate and AOX activity in red kidney bean leaves increased significantly during treatment with enhanced UV-B. However, cytochrome oxidase （COX） activity did not change significantly. The H2O2 content was also markedly increased and reached a maximum of 4.45 mmol·L^-1·g^-1 DW （dry weight） at 24 h of UV-B treatment, before dropping rapidly. Both alternative pathway content and alternative pathway activity were increased in the presence of exogenous H2O2. Immunoblotting analysis with anti-AOX monoclonal antibody revealed that expression of the AOX protein increased in red kidney bean leaves under enhanced UV-B radiation, reaching a peak at 72 h. In addition, AOX expression in red kidney bean leaves was induced by exogenous H2O2. These data indicate that the increase in AOX activity in red kidney bean leaves under enhanced UV-B radiation was mainly due to H2O2-induced AOX expression.