植物对水淹胁迫的响应与适应

水淹是植物遭受的主要的非生物胁迫之一.水淹胁迫使植物处于周期或长期的厌氧或缺氧状态,限制植物的需氧呼吸和维持生命活动所需的能量产生,导致土壤还原势的降低和有毒物质的积累,从而对植物的生存构成严重威胁.在长期的进化过程中,一些植物能够忍耐短期或长期的水淹生境而存活下来.目前分析植物感知水淹胁迫的主要途径为感知体内氧浓度的降低和感知体内乙烯浓度的增加.淹水胁迫下植株的适应策略主要包括:1)茎的伸长生长、不定根和通气组织的形成等形态学方面的适应;2)代谢途径的改变,淹水植物主要通过厌氧代谢获得维持生命的能量;3)通过体内乙烯、赤霉素和脱落酸等激素含量水平的改变来调节生理活动或形态、解剖等方面的变化;4)抗氧化酶系统对厌氧胁迫植株体内有毒的活性氧自由基的清除.运用分子生物学和生物信息学等手段找出由水淹胁迫诱导的相关基因并对其进行克隆,繁殖与培育具有耐水淹能力强的植物种类、品种和生态型,将是从事植物抗水淹胁迫研究的科研人员的目标和方向.     

水体溶氧影响陆生植物喜旱莲子草(Alternanthera philoxeroides)和牛鞭草(Hemarthria altissima)对完全水淹的耐受力

溶氧是水环境中一个重要的环境因子,为了探讨水中的溶氧含量水平是否会对陆生植物的耐淹能力造成影响,研究了陆生植物喜旱莲子草(Alternanthera philoxeroides)和牛鞭草(Hemarthria altissima)在遭受不同溶氧含量水体完全淹没后的生长表现、存活情况和非结构碳水化合物的变化。实验结果表明:(1)水体中的溶氧含量显著影响了处于完全水淹环境中的喜旱莲子草和牛鞭草的存活。受高溶氧水体完全水淹的喜旱莲子草和牛鞭草主茎的完好程度和存活叶的数量均显著高于遭受低溶氧水体完全水淹的喜旱莲子草和牛鞭草,喜旱莲子草和牛鞭草在高溶氧水体完全水淹后的生物量比低溶氧水体完全水淹后要高;(2)水体中的溶氧含量显著影响了处于完全水淹环境中的喜旱莲子草和牛鞭草的生长,受高溶氧水体完全水淹的喜旱莲子草主茎伸长生长和不定根生长显著强于受低溶氧水体完全水淹的喜旱莲子草,在不定根的生长上牛鞭草也具有同样的表现。(3)高溶氧水环境有利于减小被完全淹没的喜旱莲子草和牛鞭草的碳水化合物消耗,两种植物在受高溶氧完全水淹后体内具有的非结构性碳水化合物含量均比受低溶氧完全水淹后高。(4)喜旱莲子草比牛鞭草能更好地耐受完全水淹,当处于低溶氧完全水淹时表现得更为明显,本研究表明入侵物种喜旱莲子草比本地物种牛鞭草具有更强的环境适应能力和水淹耐受能力。     

香附子光合生理对三峡库区消落带陆生环境的响应

植物在消落带水淹结束后的陆生环境中能否合成和储存足够的碳水化合物,对其在消落带长期定居、存活和扩散具有重要影响。香附子是能够适应三峡库区消落带生境的少数多年生草本植物之一。该试验通过对消落带水淹结束后生长在深水(22~27m)、浅水(2~7m)和未淹区段香附子光合生理的定期观测,研究其在陆生环境下的光合生理变化及其形成机制。结果表明:(1)生长在深水区段香附子的净光合速率(P_n)显著高于浅水区段与未淹区段。(2)随着香附子在陆生环境中生活时间的延长,其P_n、气孔导度、表观水分利用效率、表观光能利用效率、表观CO_2利用效率增加。(3)与未淹区段相比,浅水淹没区段香附子的光饱和点显著升高,而光补偿点显著降低。研究发现,淹水胁迫后积极的自我调节能力和光合补偿机制是香附子能够适生于三峡库区消落带的重要原因。     

甲烷胁迫对绿地植物生理生态特性的影响

甲烷胁迫是限制垃圾填埋场绿化植物存活和生长的主要因素。在室内模拟条件下,利用盆栽试验研究了2种绿地植物高羊茅和苜蓿对甲烷胁迫的适应性。结果表明：甲烷胁迫严重影响2种绿化植物的生长,主要表现在生长速度减小、光合作用减弱、干物质积累量减少;2种植物均通过提高体内脯氨酸含量来增加对甲烷胁迫的耐受力,苜蓿还可以通过降低SOD酶的活性来抵御甲烷胁迫;对2种绿化植物而言,脯氨酸含量可以作为评定其耐甲烷胁迫的主要生理指标。     

生长调解物质与植物水分状况的关系

水分是决定植物分布与作物生产力的主要因素之一。水分缺乏可引起植物生命活动的各个方面,包括生长发育、形态解剖与生理代谢等的改变,但是它的最主要而直接的影响是对生长发育的改变。生长调节物质对植物的生长发育有广泛的调节作用。因此关于植物在水分胁迫条件下,内源激素发生什么变化以及激     

完全水淹环境中光照和溶氧对喜旱莲子草表型可塑性的影响

光照和溶氧是水环境和陆地环境间差异显著的两个环境因子,对水淹植物的生长和存活具有重要的意义。以三峡库区常见外来入侵植物喜旱莲子草(Alternanthera philoxeroides)为研究对象,考察了水体中的光照(L)和溶氧(DO)对完全水淹环境中喜旱莲子草的形态特征和生物量分配等表型可塑性的影响。实验设置水淹和非水淹对照两组处理,对水淹组的光照和溶氧两个环境因子再分别设置有(+)、无(-)以及高(+)、低(-)两种水平,共计4个处理。实验结果表明:(1)水淹可促进喜旱莲子草主茎和叶片发生可塑性反应,引发伸长生长。水淹后,其细长的主茎以及长而薄的直立叶更有利于植株早日出露水面。(2)完全水淹条件下,喜旱莲子草主茎和叶片的表型可塑性受光照和溶氧的复合影响,其中主茎的伸长生长主要受溶氧的影响,而叶片的形态变化则主要受光照影响。高溶氧处理下喜旱莲子草的主茎伸长生长显著(P0.05)。在相同光照条件下,高溶氧处理下喜旱莲子草的主茎长、节间数、节间长以及主茎长/主茎直径均明显高于低溶氧处理。不论有光还是无光,高溶氧处理下喜旱莲子草主茎长以及节间数的平均增长率均处于最高水平,分别为61.8%、34.2%。喜旱莲子草叶片的形态变化在有光处理下表现得尤为显著,其平均叶片长宽比、比叶面积以及叶倾角分别较水淹前增加了39.65%、28.3%、45.9°。(3)光照和溶氧对于喜旱莲子草不定根和分枝的发生及发展存在影响差异。有光条件下可促进植株抽枝,而高溶氧处理时更有利于植株生根。这些形态变化有助于喜旱莲子草扩大株型占据有利生境,进一步提高植株的水下存活能力。     

植物低氧胁迫伤害与适应机理的研究进展

不良的通气条件导致了正常生长发育的植物生理性缺氧,低氧胁迫是高等植物主要的非生物胁迫因素之一。本文综述了低氧胁迫对植物生长、植株形态的影响,低氧胁迫对植物内部水分、养分吸收的变化,呼吸代谢途径的变化、激素代谢的变化,氧化系统的变化的影响,以及低氧胁迫过程中植物体内信号的传导、基因的表达、蛋白质的合成等,在不同层面分析了低氧胁迫对植物的伤害及植物对低氧逆境适应机理的最新研究成果。     

克隆整合有助于狗牙根抵御水淹

尽管国内外开展了大量的克隆整合对克隆植物抵御逆境能力影响的研究, 但整合对植物抵御水淹能力的影响研究仍比较缺乏。该文从克隆整合的角度探讨多年生草本植物狗牙根(Cynodon dactylon)对水淹胁迫的响应。试验模拟了先端分株(相对年幼的分株)分别处于0、5和15 cm三种水淹胁迫环境, 并在每个水淹梯度下实施先端分株与基端分株(相对年长的分株)之间匍匐茎连接或切断处理, 调查水淹一个月后基端分株和先端分株以及整个克隆片段在形态和生理上的表现。研究发现: 切断匍匐茎连接显著降低了狗牙根先端分株的生长, 表现在生物量下降、匍匐茎长度减短和分株数减少等方面; 水淹显著抑制了先端分株的生长, 但对基端分株的生长并未造成显著影响; 在5 cm水淹处理下, 匍匐茎保持连接时, 先端分株和整个克隆片段的生长显著增加; 连接或切断处理在不同水淹梯度下对匍匐茎平均节间长没有显著影响, 对先端分株或基端分株在光化学转化效率上也未表现显著性差异。结果表明: 克隆整合效应促进了狗牙根在水淹胁迫下分株的生长, 并有助于整个克隆片段抵御水淹胁迫。     

狗牙根与牛鞭草在三峡库区消落带水淹结束后的抗氧化酶活力

三峡水库周期性水淹结束后,消落带植物积极恢复生长的能力有利于其生存和定居,因此,探究影响植物恢复生长的关键生理生态机制对于消落带生态治理具有重要意义。采用野外采样、对比测定的手段,研究了生长于三峡水库消落带的狗牙根和牛鞭草根系在冬季水淹结束后清除活性氧(ROS)的关键酶活力恢复动态。结果表明:与对照相比,经历水淹的植物恢复初期发生明显氧化胁迫,超氧化歧化酶(SOD)、抗坏血酸过氧化物酶(APX)均维持了水淹诱导增加的活力水平。两个物种的过氧化氢酶(CAT)在水淹中活力均较低;出露之后后狗牙根CAT趋向于对照水平,牛鞭草CAT活力迅速上升但未到显著水平。水淹结束后24 d各个酶活力均回复到对照水平,表明氧化胁迫已基本消失。消落带位点植物不同酶活力之间表现出显著的相关关系,这可能是植株维持ROS代谢内稳态的表现。     

重金属对植物吲哚乙酸合成与分解影响研究进展

生长素在调节植物生长和抗重金属胁迫中具有重要作用。重金属胁迫下植物为维持自身生长,必须维持生长素的内稳态和自身代谢平衡。生长素的内稳态受到生物合成、生长素结合以及水解、代谢失活等生理活动的严格控制。一些涉及生长素合成与分解的相关酶系和基因已被识别或克隆,然而重金属胁迫下与生长素合成与分解有关基因的上调或下调以及相关酶系的激活或失活却研究尚少。揭示植物遭受重金属胁迫后生长素合成与分解变化的机理,可为植物修复实践中合理使用植物生长调节剂提供理论依据。本文以生长素的主要代表物吲哚乙酸(IAA)为例,讨论重金属胁迫下,植物体内IAA合成、分解机制及其赋存形态等方面的研究进展,并从重金属胁迫下植物IAA合成途径的相对重要性、IAA形态变化和作用以及激素间的交互作用等方面探讨了该领域的研究方向。     

Underwater photosynthesis in flooded terrestrial plants: a matter of leaf plasticity

BACKGROUND: Flooding causes substantial stress for terrestrial plants, particularly if the floodwater completely submerges the shoot. The main problems during submergence are shortage of oxygen due to the slow diffusion rates of gases in water, and depletion of carbohydrates, which is the substrate for respiration. These two factors together lead to loss of biomass and eventually death of the submerged plants. Although conditions under water are unfavourable with respect to light and carbon dioxide supply, photosynthesis may provide both oxygen and carbohydrates, resulting in continuation of aerobic respiration. SCOPE: This review focuses on evidence in the literature that photosynthesis contributes to survival of terrestrial plants during complete submergence. Furthermore, we discuss relevant morphological and physiological responses of the shoot of terrestrial plant species that enable the positive effects of light on underwater plant performance. CONCLUSIONS: Light increases the survival of terrestrial plants under water, indicating that photosynthesis commonly occurs under these submerged conditions. Such underwater photosynthesis increases both internal oxygen concentrations and carbohydrate contents, compared with plants submerged in the dark, and thereby alleviates the adverse effects of flooding. Additionally, several terrestrial species show high plasticity with respect to their leaf development. In a number of species, leaf morphology changes in response to submergence, probably to facilitate underwater gas exchange. Such increased gas exchange may result in higher assimilation rates, and lower carbon dioxide compensation points under water, which is particularly important at the low carbon dioxide concentrations observed in the field. As a result of higher internal carbon dioxide concentrations in submergence-acclimated plants, underwater photorespiration rates are expected to be lower than in non-acclimated plants. Furthermore, the regulatory mechanisms that induce the switch from terrestrial to submergence-acclimated leaves may be controlled by the same pathways as described for heterophyllous aquatic plants.     

Resistance to complete submergence in Rumex species with different life histories: the influence of plant size and light

Resistance to complete submergence was tested in three Rumex species that occur in the Dutch river forelands. The species differ in both habitat and life history characteristics. The annual or biennial R. maritimus and the biennial or short lived perennial R. palustris grow on frequently flooded mud flats of low elevation, while the perennial R. thyrsiflorus can be found on dykes and river dunes that are seldom flooded. The flooding characteristics of the habitats of the three species were determined. These data were used to design experiments to determine the survival and biomass development of the three species during submergence and the influence of plant size and light level on these parameters. It was shown in all three species that plants submerged during daytime were much more resistant to flooding than those submerged at night. This is most probably due to the generation of oxygen or carbohydrates by underwater photosynthesis. Mature plants of the three species showed higher survival after submergence than juvenile plants, which might be caused by higher carbohydrate levels in the taproots of mature plants. In addition, the three species clearly differed in survival and biomass development during submergence. Rumex thyrsiflorus , the species least subjected to flooding, is least tolerant to complete submergence. Rumex maritimus , which can avoid the floods by having a short life cycle, is less tolerant to submergence than R. palustris , which has to survive the floods as a vegetative plant. It was noted that some plants that survived the flooding period itself, still died in the following period of drained conditions, possibly due to post-anoxic injury.     

Photosynthetic consequences of phenotypic plasticity in response to submergence: Rumex palustris as a case study

Survival and growth of terrestrial plants is negatively affected by complete submergence. This is mainly the result of hampered gas exchange between plants and their environment, since gas diffusion is severely reduced in water compared with air, resulting in O2 deficits which limit aerobic respiration. The continuation of photosynthesis could probably alleviate submergence-stress in terrestrial plants, but its potential under water will be limited as the availability of CO2 is hampered. Several submerged terrestrial plant species, however, express plastic responses of the shoot which may reduce gas diffusion resistance and enhance benefits from underwater photosynthesis. In particular, the plasticity of the flooding-tolerant terrestrial species Rumex palustris turned out to be remarkable, making it a model species suitable for the study of these responses. During submergence, the morphology and anatomy of newly developed leaves changed: 'aquatic' leaves were thinner and had thinner cuticles. As a consequence, internal O2 concentrations and underwater CO2 assimilation rates were higher at the prevailing low CO2 concentrations in water. Compared with heterophyllous amphibious plant species, underwater photosynthesis rates of terrestrial plants may be very limited, but the effects of underwater photosynthesis on underwater survival are impressive. A combination of recently published data allowed quantification of the magnitude of the acclimation response in this species. Gas diffusion resistance in terrestrial leaves underwater was about 15,000 times higher than in air. Strikingly, acclimation to submergence reduced this factor to 400, indicating that acclimated leaves of R. palustris had an approximately 40 times lower gas diffusion resistance than non-acclimated ones.     

How plants cope with complete submergence

Flooding is a widespread phenomenon that drastically reduces the growth and survival of terrestrial plants. The dramatic decrease of gas diffusion in water compared with in air is a major problem for terrestrial plants and limits the entry of CO(2) for photosynthesis and of O(2) for respiration. Responses to avoid the adverse effects of submergence are the central theme in this review. These include underwater photosynthesis, aerenchyma formation and enhanced shoot elongation. Aerenchyma facilitates gas diffusion inside plants so that shoot-derived O(2) can diffuse to O(2)-deprived plant parts, such as the roots. The underwater gas-exchange capacity of leaves can be greatly enhanced by a thinner cuticle, reorientation of the chloroplasts towards the epidermis and increased specific leaf area (i.e. thinner leaves). At the same time, plants can outgrow the water through increased shoot elongation, which in some species is preceded by an adjustment of leaf angle to a more vertical position. The molecular regulatory networks involved in these responses, including the putative signals to sense submergence, are discussed and suggestions made on how to unravel the mechanistic basis of the induced expression of various adaptations that alleviate O(2) shortage underwater.     

Molecular Genetics of Submergence Tolerance in Rice: QTL Analysis of Key Traits

Flash flooding of young rice plants is a common problem forrice farmers in south and south-east Asia. It severely reducesgrain yield and increases the unpredictability of cropping.The inheritance and expression of traits associated with submergencestress tolerance at the seedling stage are physiologically andgenetically complex. We exploited naturally occurring differencesbetween certain rice lines in their tolerance to submergenceand used quantitative trait loci (QTL) mapping to improve understandingof the genetic and physiological basis of submergence tolerance.Three rice populations, each derived from a single cross betweentwo cultivars differing in their response to submergence, wereused to identify QTL associated with plant survival and variouslinked traits. These included total shoot elongation under water,the extent of stimulation of shoot elongation caused by submergence,a visual submergence tolerance score, and leaf senescence underdifferent field conditions, locations and years. Several majorQTL determining plant survival, plant height, stimulation ofshoot elongation, visual tolerance score and leaf senescenceeach mapped to the same locus on chromosome 9. These QTL weredetected consistently in experiments across all years and inthe genetic backgrounds of all three mapping populations. SecondaryQTL influencing tolerance were also identified and located onchromosomes 1, 2, 5, 7, 10 and 11. These QTL were specific toparticular traits, environments, or genetic backgrounds. Allidentified QTL contributed to increased submergence tolerancethrough their effects on decreased underwater shoot elongationor increased maintenance of chlorophyll levels, or on both.These findings establish the foundations of a marker-assistedscheme for introducing submergence tolerance into agriculturallydesirable cultivars of rice.     

Submergence-induced leaf acclimation in terrestrial species varying in flooding tolerance

Earlier work on the submergence-tolerant species Rumex palustris revealed that leaf anatomical and morphological changes induced by submergence enhance underwater gas exchange considerably. Here, the hypothesis is tested that these plastic responses are typical properties of submergence-tolerant species. Submergence-induced plasticity in leaf mass area (LMA) and leaf, cell wall and cuticle thickness was investigated in nine plant species differing considerably in tolerance to complete submergence. The functionality of the responses for underwater gas exchange was evaluated by recording oxygen partial pressures inside the petioles when plants were submerged. Acclimation to submergence resulted in a decrease in all leaf parameters, including cuticle thickness, in all species irrespective of flooding tolerance. Consequently, internal oxygen partial pressures (pO(2)) increased significantly in all species until values were close to air saturation. Only in nonacclimated leaves in darkness did intolerant species have a significantly lower pO(2) than tolerant species. These results suggest that submergence-induced leaf plasticity, albeit a prerequisite for underwater survival, does not discriminate tolerant from intolerant species. It is hypothesized that these plastic leaf responses may be induced in all species by several signals present during submergence; for example, low LMA may be a response to low photosynthate concentrations and a thin cuticle may be a response to high relative humidity.     

长期水淹条件下香根草(Vetiveria zizanioides)、菖蒲(Acorus calamus)和空心莲子草(Alternanthera philoxeroides)的存活及生长响应

人工构建三峡库区消落区植被是控制消落区水土流失、保护消落区生态环境的重要措施,选择能够耐受长时间完全水淹的植物物种是该措施实施的关键.为了验证香根草、菖蒲、空心莲子草能否用于消落区植被的构建,实验模拟消落区的长期完全水淹条件,设置30d、60d、90d、120d、150d和180d等6个完全水淹时间水平,研究了3种植物在完全水淹条件下生长、生物量积累及存活状况.结果发现:(1)3种植物在经受长时间的完全水淹后有较高的存活率,180d全淹处理后,香根草、菖蒲和空心莲子草的存活率分别为87.5%、100%和50%.(2)这3种植物有不同的水下生长能力.全淹条件下,香根草生长缓慢,几乎没有产生新的叶片,总叶长也没有显著变化;菖蒲能够持续产生较对照植株更为细长的叶片,空心莲子草只在水淹初期(30d内)能够快速伸长地上部分的枝条,并迅速产生新叶片,但随水淹时间的延长,总枝条长及总叶片数没有再显著增加.(3)与对照植株相比,全淹处理抑制了3种植物总生物量的增加,但对3种植物的地上、地下部分生物量抑制程度不同.全淹条件下,香根草的地上部分和地下部分生物量与水淹0d水平(水淹处理开始前一天,下同)相比无显著变化,根冠比高于对照植株;菖蒲的地上部分生物量随水淹时间延长而降低,但却高于对照植株,地下部分生物量始终低于水淹0d水平,根冠比低于对照植株;空心莲子草的地上部分生物量与水淹0d水平相比无显著差异,但地下部分生物量与水淹0d水平相比大幅降低,根冠比低于对照植株.结果表明,这3种植物都有很强的水淹耐受能力,可应用于三峡库区消落区植被的构建.同时,发现植物对长期完全水淹的耐受能力很大程度上与植株在水下的生长情况及植株的营养储备水平相关,剧烈的水下生长会消耗大量的营养储备,进而造成植株存活率降低.植株在全淹条件下有限的生长能力及丰富的营养储备可能是耐淹物种的重要特征.     

Leaf gas films,underwater photosynthesis and plant species distributions in a flood gradient

Traits for survival during flooding of terrestrial plants include stimulation or inhibition of shoot elongation, aerenchyma formation and efficient gas exchange. Leaf gas films form on superhydrophobic cuticles during submergence and enhance underwater gas exchange. The main hypothesis tested was that the presence of leaf gas films influences the distribution of plant species along a natural flood gradient. We conducted laboratory experiments and field observations on species distributed along a natural flood gradient. We measured presence or absence of leaf gas films and specific leaf area of 95 species. We also measured, gas film retention time during submergence and underwater net photosynthesis and dark respiration of 25 target species. The presence of a leaf gas film was inversely correlated to flood frequency and duration and reached a maximum value of 80% of the species in the rarely flooded locations. This relationship was primarily driven by grasses that all, independently of their field location along the flood gradient, possess gas films when submerged. Although the present study and earlier experiments have shown that leaf gas films enhance gas exchange of submerged plants, the ability of species to form leaf gas films did not show the hypothesized relationship with species composition along the flood gradient.     

Morphological and physiological responses of rice seedlings to complete submergence (flash flooding)

Background and Aims

Reducing damage to rice seedlings caused by flash flooding will improve the productivity of rainfed lowland rice in West Africa. Accordingly, the morphological and physiological responses of different forms of rice to complete submergence were examined in field and pot experiments to identify primary causes of damage.

Methods

To characterize the physiological responses, seedlings from a wide genetic base including Oryza sativa, O. glaberrima and interspecific hybrids were compared using principle component analysis.

Key Results

Important factors linked to flash-flood tolerance included minimal shoot elongation underwater, increase in dry matter weight during submergence and post-submergence resistance to lodging. In particular, fast shoot elongation during submergence negatively affected plant growth after de-submergence. Also shoot-elongating cultivars showed a strong negative correlation between dry matter weight of the leaves that developed before submergence and leaves developing during submergence.

Conclusions

Enhancement of shoot elongation during submergence in water that is too deep to permit re-emergence by small seedlings represents a futile escape strategy that takes place at the expense of existing dry matter in circumstances where underwater photosynthetic carbon fixation is negligible. Consequently, it compromises survival or recovery growth once flood water levels recede and plants are re-exposed to the aerial environment. Tolerance is greater in cultivars where acceleration of elongation caused by submergence is minimal.Key words: Africa, flash floods, Oryza glaberrima, rainfed lowland, rice, shoot elongation, stress tolerance, submergence