环境因子对植物释放挥发性化合物的影响

对近年来有关环境因子与植物释放挥发性化合物关系的研究进展进行了综合和概括。本文主要包括3类挥发性化合物。⑴异戊二烯是由叶绿体产生并且直接释放到大气中的C5化合物。⑵单萜类化合物是一类环状或非环状的C10化合物,它在植物体内合成后首先贮存于体内的特殊结构中（如树脂道、油腺）,然后由此通过气孔向大气中释放。⑶含氧挥发性化合物以各种形式释放大大气中。它包括醇、醛、酮、酯和有机酸。本文的重点是前两者,主要阐述了二方面内容：⑴植物军发性化合物的生物合成和释放机理。⑵环境因子（如温度、光照、水分胁迫、营养、CO2浓度、空气湿度）及植物的发育阶段、机械损伤和昆虫取食等对植物挥发性化合物合成与释放的影响机制。     

植物挥发性物质代谢的遗传修饰

植物在生命周期里能合成释放多种多样的挥发性化合物,这些物质在植物生长发育和代谢调控,植物抵御病虫侵害,以及植物与环境信息交流中行使重要功能。在介绍植物挥发性化合物的生理功能、合成途径调控和商业应用的基础上,重点论述应用基因工程调控植物挥发性物质合成的技术策略和研究进展,并讨论了植物挥发性物质遗传修饰存在的问题和发展前景。     

汞在SPAC-人体系统中的转递及主要影响因素

汞（Hg）是环境中一种具有高度毒性的重金属元素,且具有较高挥发性,因而作为一种全球性的污染物而备受关注。汞在SPAC系统中的迁移转化,是全球汞循环的重要环节,与人类的建康密切相关。汞通过食物链进入人体并在体内蓄积受多种因素影响,主要包括3个方面：土壤性质（如土壤汞含量、pH、有机质、粘土矿物等）,植物特性（植物类型、种类和生育期等）,大气状况（如光照强度、气温和大气湿度等）。此外,锌、硒和抗氧化剂等的存在也有较大影响。综述了汞在SPAC-人体系统中的迁移积累及其调控机理。     

转录因子调控植物萜类化合物生物合成研究进展

萜类化合物是植物次生代谢物中结构和数量最多的一类化合物, 它们在植物体内以及植物与环境和其它生命体的相互作用中发挥重要作用。转录因子通过调控代谢通路中基因的转录起始来调节次生代谢物质的产量。目前, 研究发现参与萜类合成的转录因子家族主要有6个, 包括AP2/ERF、bHLH、MYB、NAC、WRKY和bZIP。该文主要对其家族的结构特点、调控模式以及研究进展进行综述, 以期进一步丰富萜烯合成的网络调控, 为植物萜类相关的分子育种、优质栽培和病虫害生物防治等提供新的思路与方法。     

转录因子调控植物萜类化合物生物合成研究进展

萜类化合物是植物次生代谢物中结构和数量最多的一类化合物,它们在植物体内以及植物与环境和其它生命体的相互作用中发挥重要作用。转录因子通过调控代谢通路中基因的转录起始来调节次生代谢物质的产量。目前,研究发现参与萜类合成的转录因子家族主要有6个,包括AP2/ERF、b HLH、MYB、NAC、WRKY和bZIP。该文主要对其家族的结构特点、调控模式以及研究进展进行综述,以期进一步丰富萜烯合成的网络调控,为植物萜类相关的分子育种、优质栽培和病虫害生物防治等提供新的思路与方法。     

植物源挥发性有机物的生态意义(综述)

植物释放的挥发性有机气体(volatile organic compounds, VOCs)在对流层大气中通过一系列氧化还原反应,改变大气的化学组成,对臭氧合成、一氧化碳生成、甲烷氧化等有重要作用,其氧化物质对区域乃至全球的环境和气候都产生一定的影响。本文综述植物释放的VOCs对大气化学、温室效应、光化学烟雾的影响;介绍VOCs释放机制、合成途径及排放速率;对今后研究方向和大面积种植林木、城市绿化提出建议。     

几种海南岛热带雨林优势种植物挥发性有机物排放对模拟氮沉降的短期响应

BVOCs(Biogenic Volatile Organic Compounds)是植物向大气释放的一类重要气态化合物，能参与大气化学过程和陆地生态系统碳素循环。分析环境因子对BVOCs排放的影响，对科学认识未来气候变化具有重要意义。氮素作为植物生长、发育所需的大量营养元素之一，其沉降增加是当前全球气候变化的主要驱动因素之一，但学者对BVOCs如何应对氮沉降增加知之甚少。因此以海南岛热带雨林树种：木荷(Schima superba)、厚壳桂(Cryptocarya chinensis)和线枝蒲桃(Syzygium araiocladum)为研究对象，通过温室盆栽实验模拟氮沉降对3个树种BVOCs释放的短期效应。主要结论如下：(1)自然状态下，从木荷、厚壳桂和线枝蒲桃的枝叶中鉴定出14、34和24种挥发性有机化合物，包括异戊二烯、单萜烯、倍半萜烯和其他挥发性有机化合物(烷烃、羰基、醛、醇、酯、醚和酸),此外三个阔叶树种释放BVOCs的速率呈厚壳桂>木荷>线枝蒲桃；(2)外源施氮均促进了三种植物幼苗VOCs释放，其中总VOCs释放速率和成分数量均随施氮浓度的升高而增加，且叶面...     

植物源异戊二烯及其生态意义

异戊二烯为植物挥的众多有机碳氢化合物中的主要成分,它的合成和释放对全球尺度上的C素平衡、温室效应的大气污染都有极其重要的作用。本文系统论述了影响异戊二烯释放的主要因子,异戊二烯的释放量及其计量方法,以及它可能的生态意义,指出了研究多重环境胁迫与异戊二烯合成和释放的关系的必要性。     

植物为什么释放异戊二烯

自然界中有许多植物具有释放出异戊：1烯（2一甲基一l,3－］一二烯）化合物的能力。据估算,从全球植被中每年释放出的异戊二二烯约为3XIO‘克,与甲烷的释放越相当。由于该化合物能与空气中的羟基离于迅速发生反应,因此它在大气化学中起重要作用。但为什么植物要释放异成二烯,n从本世纪50年代萨纳德兹（Sa。ladze）发现这一现象以来就一直是一个迷,研究发现,植物释放异成二烯的作用有赖于光合作m＿在气温3O℃下,白杨和橡树能将光合作用所同化的碳的2％以拧成二烯的形式释放出来黑暗、光合抑制剂及大气中的纯氮都会阻止异成二烯的释…     

植物挥发性有机物合成与代谢途径及其释放与感知调控机制的研究进展CSCD

植物释放的挥发性有机物(biogenic volatile organic compounds,BVOCs)是具有低沸点、易挥发的低分子量亲脂性化合物,在植物发育的整个过程中,它们通常通过不同底物的几个独立途径由不同的酶催化合成,并受关键基因及转录因子等调控代谢,随后在植物细胞内和细胞之间移动而被释放出来,最终作为植物内部信号被植物感知从而进行传递交流,影响着植物各种生理反应。本文对植物BVOCs参与合成及代谢途径的调控机制、释放的细胞生理和理化性质及植物对BVOCs的感知调控进行分析和解读,以期为BVOCs的合成与代谢、释放与被感知机制的进一步研究提供参考。     

On the induction of volatile organic compound emissions by plants as consequence of wounding or fluctuations of light and temperature

Among the volatile organic compounds (VOCs) emitted by plants, some are characteristic of stress conditions, but their biosynthesis and the metabolic and environmental control over the emission are still unclear. We performed experiments to clarify whether (1) the emission following wounding can occur at distance from the wounding site, from VOC pools subjected to metabolic signals; and (2) the emission of biogenic VOCs generated by membrane damage (e.g. consequent to wounding or ozone exposure) can also be induced by exposure to high light and high temperature, recurrent in nature. In Phragmites australis, leaf cutting caused large and rapid bursts of acetaldehyde both at the cutting site and on parts of the cut leaf distant from the cutting site. This emission was preceded by a transient stomatal opening and did not occur in conditions preventing stomatal opening. This suggests the presence of a large pool of leaf acetaldehyde whose release is under stomatal control. VOCs other than isoprene, particularly acetaldehyde and (E)-2-hexenal, one of the C-6 compounds formed by the denaturation of membrane lipids, were released by leaves exposed to high temperature and high light. The high-temperature treatment (45 degrees C) also caused a rapid stimulation and then a decay of isoprene emission in Phragmites leaves. Isoprene recovered to the original emission level after suspending the high-temperature treatment, suggesting a temporary deficit of photosynthetically formed substrate under high temperature. Emission of C-6 compounds was slowly induced by high temperature, and remained high, indicating that membrane denaturation occurs also after suspending the high-temperature treatment. Conversely, the emission of C-6 compounds was limited to the high-light episode in Phragmites. This suggests that a membrane denaturation may also occur in conditions that do not damage other important plant processes such as the photochemistry of photosynthesis of photoinhibition-insensitive plants. In the photoinhibition-sensitive Arabidopsis thaliana mutant NPQ1, a large but transient emission of (E)-2-hexenal was also observed a few minutes after the high-light treatment, indicating extensive damage to the membranes. However, (E)-2-hexenal emission was not observed in Arabidopsis plants fumigated with isoprene during the high-light treatment. This confirms that isoprene can effectively protect cellular membranes from denaturation. Our study indicates that large, though often transient, VOC emissions by plants occur in nature. In particular, we demonstrate that VOCs can be released by much larger tissues than those wounded and that even fluctuations of light and temperature regularly observed in nature can induce their emissions. This knowledge adds information that is useful for the parameterization of the emissions and for the estimate of biogenic VOC load in the atmosphere.     

Detection of plant volatiles after leaf wounding and darkening by proton transfer reaction "time-of-flight" mass spectrometry (PTR-TOF)

Proton transfer reaction-time of flight (PTR-TOF) mass spectrometry was used to improve detection of biogenic volatiles organic compounds (BVOCs) induced by leaf wounding and darkening. PTR-TOF measurements unambiguously captured the kinetic of the large emissions of green leaf volatiles (GLVs) and acetaldehyde after wounding and darkening. GLVs emission correlated with the extent of wounding, thus confirming to be an excellent indicator of mechanical damage. Transient emissions of methanol, C5 compounds and isoprene from plant species that do not emit isoprene constitutively were also detected after wounding. In the strong isoprene-emitter Populus alba, light-dependent isoprene emission was sustained and even enhanced for hours after photosynthesis inhibition due to leaf cutting. Thus isoprene emission can uncouple from photosynthesis and may occur even after cutting leaves or branches, e.g., by agricultural practices or because of abiotic and biotic stresses. This observation may have important implications for assessments of isoprene sources and budget in the atmosphere, and consequences for tropospheric chemistry.     

Isoprene emission and primary metabolism in Phragmites australis grown under different phosphorus levels

Aquatic plants are generally used for wastewater purification and phytoremediation, but some of them also emit large amounts of isoprene, the most abundant biogenic volatile organic compound. Since isoprenoid biosynthesis requires high amounts of phosphorylated intermediates, the emission may also be controlled by inorganic phosphorus concentration (Pi) in leaves. We carried out experiments to determine the emission of isoprene from Phragmites australis plants used in reconstructed wetlands to phytoremediate elevated levels of phosphorus contributed by urban wastes. Four groups of plants were grown hydroponically in water containing different levels of KH(2)PO(4). High levels of phosphorus in the water resulted in high Pi in the leaves. High Pi stimulated photosynthesis at intercellular CO(2) concentrations lower and higher than ambient, implying higher ribulose 1,5-bisphosphate carboxylase (Rubisco) activity and higher ribulose 1,5-bisphosphate regeneration rates, respectively. However, isoprene emission was substantially lower at high Pi than at low Pi, and was not associated to photosynthesis rates at high Pi. This surprising result suggests that isoprene is limited by processes other than photosynthetic intermediate availability or by energetic (ATP) requirements under high Pi levels. Irrespective of the mechanism responsible for the observed reduction of isoprene emission, our results show that Phragmites plants may effectively remove phosphorus from water without concurrently increase isoprene emission, at least on a leaf area basis. Thus, Phragmites used in reconstructed wetlands for phytoremediation of urban wastes rich of phosphates will not contribute high loads of hydrocarbons which may influence air quality over urban and peri-urban areas.     

Isoprene is more affected by climate drivers than monoterpenes: A meta‐analytic review on plant isoprenoid emissions

Isoprene and monoterpenes (MTs) are among the most abundant and reactive volatile organic compounds produced by plants (biogenic volatile organic compounds). We conducted a meta‐analysis to quantify the mean effect of environmental factors associated to climate change (warming, drought, elevated CO₂, and O₃) on the emission of isoprene and MTs. Results indicated that all single factors except warming inhibited isoprene emission. When subsets of data collected in experiments run under similar change of a given environmental factor were compared, isoprene and photosynthesis responded negatively to elevated O₃ (?8% and ?10%, respectively) and drought (?15% and ?42%), and in opposite ways to elevated CO₂ (?23% and +55%) and warming (+53% and ?23%, respectively). Effects on MTs emission were usually not significant, with the exceptions of a significant stimulation caused by warming (+39%) and by elevated O₃ (limited to O₃‐insensitive plants, and evergreen species with storage organs). Our results clearly highlight individual effects of environmental factors on isoprene and MT emissions, and an overall uncoupling between these secondary metabolites produced by the same methylerythritol 4‐phosphate pathway. Future results from manipulative experiments and long‐term observations may help untangling the interactive effects of these factors and filling gaps featured in the current meta‐analysis.     

陆地生态系统植物挥发性有机化合物的排放及其生态学功能研究进展

综述了国内外生物源挥发性有机化合物 (Biologicalvolatileorganiccompounds, BVOCs) 研究现状及未来的研究方向, 侧重介绍了陆地生态系统中植物排放BVOCs的种类、生物学功能及其对大气化学过程的影响。BVOCs按其化学结构以及在大气中的滞留时间可以分为 4类 :异戊二烯、单萜、其它活性BVOCs和其它次活性BVOCs。不同的植物类群排放不同的BVOCs种类并具有不同的排放特性, 环境条件对植物不同BVOCs的排放影响也不同。BVOCs作为有机物质被排放到体外, 从植物能量代谢的角度来讲要消耗一部分植物光合作用产物从而降低植物的生产力, 因此推测植物排放BVOCs具有一定的生理学或者生态学的功能。其中比较成熟的假说是抗热胁迫假说, 其次是抗氧化假说, 也有一些其它假说例如促氮同化假说等。但这些假说都还缺乏直接的有力证据, 有待更多的研究来支持。BVOCs被排放到大气中对大气化学过程的影响更是科学家关注的问题, BVOCs对大气的影响一方面是在大气对流层中促进臭氧 (O3 ) 的形成, 造成环境污染, 另一方面BVOCs通过对大气中的OH自由基和臭氧等氧化物浓度的调整而影响到大气中甲烷等温室气体的平衡, 对大气温室效应具有间接的贡献。我国在BVOCs的研究上也做了大量的工作, 包括分析鉴定了一些植物排放的BVOCs, 探讨了环境因子对植物BVOCs排放速率的影响, 从不同尺度估测了BVOCs的排放量等等。今后对BVOCs的研究将会集中在以下几个方面 :1) 进一步研究不同植物类群释放的BVOCs种类及其它们在大气中的理化性质 ;2 ) 继续探讨植物排放BVOCs的合成与代谢途径及其生物学功能 ;3) 研究BVOCs对大气化学过程的作用, 以及区域植被变化对BVOCs排放格局进而对区域乃至全球环境变化的影响 ;4 ) 加强对一些研究比较薄弱的生态系统例如在热带地区所进行的BVOCs研究工作 ;5 ) 进一步建立和完善BVOCs排放的理论模型, 以模拟不同陆地生态系统BVOCs排放的时空动态。     

Volatile emissions and phenolic compound concentrations along a vertical profile of <Emphasis Type="Italic">Populus nigra</Emphasis> leaves exposed to realistic ozone concentrations

Plants are exposed to increasing levels of tropospheric ozone concentrations. This pollutant penetrates in leaves through stomata and quickly reacts inside leaves, thus making plants valuable ozone sinks, but at the same time triggers oxidation processes which lead to leaf injuries. To counteract these negative effects, plants produce an array of antioxidants which react with ozone and reactive molecules which ozone generates in the leaf tissues. In this study, we measured the effect of an ozone concentration which is likely to be attained in many areas of the world in the near future (80 ppb) on leaves of the vertical profile of the widespread agroforestry species Populus nigra. Changes in (1) physiological parameters (photosynthesis and stomatal conductance), (2) ozone uptake, (3) emission of volatile organic compounds (VOCs, i.e. isoprene, methanol and other oxygenated compounds), (4) concentration of antioxidant surface compounds, and (5) concentration of phenolic compounds were assessed. The aim was to assess whether the defensive pathways leading to isoprenoids and phenolics formation were induced when a moderate and chronic increment of ozone is not able to damage photosynthesis. No visual injuries and minor changes in physiology and ozone uptake were observed. The emission of isoprene and oxygenated six-carbon (C6) volatiles were inhibited by ozone, whereas methanol emission was increased, especially in developing leaves. We interpret these results as suggesting an ontogenetic shift in ozone-treated leaves, leading to a slower development and a faster senescence. Most surface and phenolic compounds showed a declining trend in concentration from the youngest to the fully expanded leaves. Ozone reduced the concentrations of chlorogenic acid derivatives at the leaf surface, whereas in total leaf extracts a metabolic shift towards few phenolics with higher antioxidant capacity was observed.     

Rapid regulation of the methylerythritol 4-phosphate pathway during isoprene synthesis

More volatile organic carbon is lost from plants as isoprene than any other molecule. This flux of carbon to the atmosphere affects atmospheric chemistry and can serve as a substrate for ozone production in polluted air. Isoprene synthesis may help leaves cope with heatflecks and active oxygen species. Isoprene synthase, an enzyme related to monoterpene synthases, converts dimethylallyl diphosphate derived from the methylerythritol 4-phosphate pathway to isoprene. We used dideuterated deoxyxylulose (DOX-d(2)) to study the regulation of the isoprene biosynthetic pathway. Exogenous DOX-d(2) displaced endogenous sources of carbon for isoprene synthesis without increasing the overall rate of isoprene synthesis. However, at higher concentrations, DOX-d(2) completely suppressed isoprene synthesis from endogenous sources and increased the overall rate of isoprene synthesis. We interpret these results to indicate strong feedback control of deoxyxylulose-5-phosphate synthase. We related the emission of labeled isoprene to the concentration of labeled dimethylallyl diphosphate in order to estimate the in situ K(m) of isoprene synthase. The results confirm that isoprene synthase has a K(m) 10- to 100-fold higher for its allylic diphosphate substrate than related monoterpene synthases for geranyl diphosphate.     

Reduced levels of volatile emissions in jasmonate-deficient spr2 tomato mutants favour oviposition by insect herbivores

Tomato plants release volatile organic compounds (VOCs) following insect or mechanical damage. In this study, the constitutive and wound-induced emission levels of VOCs in suppressor of prosystemin-mediated responses2 (spr2) mutant plants, compromised in linolenic acid (LA) and jasmonic acid (JA) synthesis, and in 35S::prosystemin (35S::prosys) plants, having upregulated direct defence responses, were compared. The spr2 mutants produced constitutively lower levels of VOCs, which were nonetheless increased in response to (a)biotic damage, although at lower levels than wild-type (Wt) and 35S::prosys plants. No significant differences in VOC emissions were detected between the latter two genotypes, thereby suggesting that systemin does not regulate indirect defence responses, whereas differences in fatty acid composition in spr2 plants led to the predominant emission of saturated C6 volatiles in response to wounding. The expression of 1-deoxy-D-xylulose 5-phosphate synthase (DXS2), a key gene involved in VOC synthesis in the chloroplast, was only upregulated in Manduca sexta L.-damaged Wt and 35S::prosys plants. However, its expression was restored in spr2 plants by exogenous LA or JA, suggesting that abated VOC emissions in spr2 plants are correlated with lowered DXS2 expression. Bioassays with two different insects showed that adult females significantly preferred spr2 plants, indicating that lowered levels of VOCs in tomato influence plant selection by insects during oviposition.     

Health monitoring of plants by their emitted volatiles: trichome damage and cell membrane damage are detectable at greenhouse scale

Pathogen attack and herbivore infestation have a major impact on plant health. In a model study, these two plant health issues were simulated to study whether plant health can be monitored at greenhouse scale through the analysis of volatile organic compounds (VOCs) in greenhouse atmosphere. To simulate pathogen attack and herbivore infestation, we repeatedly stroked the stems of tomato plants ( Lycopersicon esculentum ) and repeatedly removed their side shoots. In addition, we studied the effect of fruit picking on the concentration of plant-emitted VOCs in greenhouse atmosphere. Analysis of air samples obtained before these treatments revealed up to 17 VOCs that are known to be released from tomato plants, of which the most dominant one was the monoterpene β-phellandrene. When plants were 7 weeks old, the concentration of this VOC was approximately 0.06 ppbv before treatment. When plants were 12 weeks old, this concentration was raised to approximately 0.14 ppbv. Stroking of the stems, removing the side shoots and fruit picking resulted in an increase in the concentrations of all mono- and most sesquiterpenes up to 60-fold, which was expected because these VOCs are well-known constituents of trichomes. The treatments did not result in substantially increased concentrations of the stress-related compounds α-copaene, methyl salicylate and ( E,E )-4,8,12-trimethyl-1,3,7,11-tridecatetraene. In contrast to stroking and fruit picking, shoot removal resulted in the emission of the lipoxygenase-derived product ( Z )-3-hexenol in greenhouse atmosphere expressing cell membrane degradation. The findings presented in this paper focus on the feasibility of monitoring plant health through the analysis of VOCs in greenhouse air, but findings might also be relevant for atmospheric chemistry.