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。（1）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。（2）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。（3）含氧挥发性化合物以各种形式释放到大气中。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：(1)植物挥发性化合物的生物合成和释放机理。(2)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。     

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。     

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。     

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。     

山海螺的化学成分研究

从药用植物册海螺（Ｃｏｄｏｎｏｐｓｉｓ Ｉａｎｃｅｏｌａｔａ）中分离得到４个化合物,分别鉴定为：莽草酸（ｓｈｉｋｉｍｉｃ ａｃｉｄ）⑴、顺丁烯二酸（ｓｕｃｃｉｎｉｃ ａｃｉｄ）⑵、丁香脂素（ｓｙｒｉｎｇａｒｓ８ｉｎｏｌ）⑶、鸢尾甙（ｔｅｃｔｏｒｉｄｉｎ）⑷、化合物１－４均为首次从该植物分得。     

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。     

植物释放挥发性有机物(VOCs)的研究进展

植物释放的有机物(VOCs)对大气圈臭氧的动态、一氧化碳的产生和甲烷的氧化起重要作用。本文主要综述了异戊二烯、单萜这两种重要的植物释放物,总结了国内外的研究进展,概括了它们的合成、释放以及环境因子的影响。同时简要介绍了其它几种植物释放的有机物,并就有机物的释放对大气化学、温室效应和全球变化的影响也作了简要分析。     

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.     

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.     

Modelling changes in VOC emission in response to climate change in the continental United States

The alteration of climate is driven not only by anthropogenic activities, but also by biosphere processes that change in conjunction with climate. Emission of volatile organic compounds (VOCs) from vegetation may be particularly sensitive to changes in climate and may play an important role in climate forcing through their influence on the atmospheric oxidative balance, greenhouse gas concentration, and the formation of aerosols. Using the VEMAP vegetation database and associated vegetation responses to climate change, this study examined the independent and combined effects of simulated changes in temperature, CO₂ concentration, and vegetation distribution on annual emissions of isoprene, monoterpenes, and other reactive VOCs (ORVOCs) from potential vegetation of the continental United States. Temperature effects were modelled according to the direct influence of temperature on enzymatic isoprene production and the vapour pressure of monoterpenes and ORVOCs. The effect of elevated CO₂ concentration was modelled according to increases in foliar biomass per unit of emitting surface area. The effects of vegetation distribution reflects simulated changes in species spatial distribution and areal coverage by 21 different vegetation classes. Simulated climate warming associated with a doubled atmospheric CO₂ concentration enhanced total modelled VOC emission by 81.8% (isoprene + 82.1%, monoterpenes + 81.6%, ORVOC + 81.1%), whereas a simulated doubled CO₂ alone enhanced total modelled VOC emission by only + 11.8% (isoprene + 13.7%, monoterpenes + 4.1%, ORVOC + 11.7%). A simulated redistribution of vegetation in response to altered temperatures and precipitation patterns caused total modelled VOC emission to decline by 10.4% (isoprene – 11.7%, monoterpenes – 18.6%, ORVOC 0.0%) driven by a decline in area covered by vegetation classes emitting VOCs at high rates. Thus, the positive effect of leaf-level adjustments to elevated CO₂ (i.e. increases in foliar biomass) is balanced by the negative effect of ecosystem-level adjustments to climate (i.e. decreases in areal coverage of species emitting VOC at high rates).     

Volatile terpenes – mediators of plant-to-plant communication

Plants interact with other organisms employing volatile organic compounds (VOCs). The largest group of plant-released VOCs are terpenes, comprised of isoprene, monoterpenes, and sesquiterpenes. Mono- and sesquiterpenes are well-known communication compounds in plant–insect interactions, whereas the smallest, most commonly emitted terpene, isoprene, is rather assigned a function in combating abiotic stresses. Recently, it has become evident that different volatile terpenes also act as plant-to-plant signaling cues. Upon being perceived, specific volatile terpenes can sensitize distinct signaling pathways in receiver plant cells, which in turn trigger plant innate immune responses. This vastly extends the range of action of volatile terpenes, which not only protect plants from various biotic and abiotic stresses, but also convey information about environmental constraints within and between plants. As a result, plant–insect and plant–pathogen interactions, which are believed to influence each other through phytohormone crosstalk, are likely equally sensitive to reciprocal regulation via volatile terpene cues. Here, we review the current knowledge of terpenes as volatile semiochemicals and discuss why and how volatile terpenes make good signaling cues. We discuss how volatile terpenes may be perceived by plants, what are possible downstream signaling events in receiver plants, and how responses to different terpene cues might interact to orchestrate the net plant response to multiple stresses. Finally, we discuss how the signal can be further transmitted to the community level leading to a mutually beneficial community-scale response or distinct signaling with near kin.     

Isoprene emission from plants: why and how

BACKGROUND: Some, but not all, plants emit isoprene. Emission of the related monoterpenes is more universal among plants, but the amount of isoprene emitted from plants dominates the biosphere-atmosphere hydrocarbon exchange. SCOPE: The emission of isoprene from plants affects atmospheric chemistry. Isoprene reacts very rapidly with hydroxyl radicals in the atmosphere making hydroperoxides that can enhance ozone formation. Aerosol formation in the atmosphere may also be influenced by biogenic isoprene. Plants that emit isoprene are better able to tolerate sunlight-induced rapid heating of leaves (heat flecks). They also tolerate ozone and other reactive oxygen species better than non-emitting plants. Expression of the isoprene synthase gene can account for control of isoprene emission capacity as leaves expand. The emission capacity of fully expanded leaves varies through the season but the biochemical control of capacity of mature leaves appears to be at several different points in isoprene metabolism. CONCLUSIONS: The capacity for isoprene emission evolved many times in plants, probably as a mechanism for coping with heat flecks. It also confers tolerance of reactive oxygen species. It is an example of isoprenoids enhancing membrane function, although the mechanism is likely to be different from that of sterols. Understanding the regulation of isoprene emission is advancing rapidly now that the pathway that provides the substrate is known.     

The atmospheric chemistry of trace gases and particulate matter emitted by different land uses in Borneo

We report measurements of atmospheric composition over a tropical rainforest and over a nearby oil palm plantation in Sabah, Borneo. The primary vegetation in each of the two landscapes emits very different amounts and kinds of volatile organic compounds (VOCs), resulting in distinctive VOC fingerprints in the atmospheric boundary layer for both landscapes. VOCs over the Borneo rainforest are dominated by isoprene and its oxidation products, with a significant additional contribution from monoterpenes. Rather than consuming the main atmospheric oxidant, OH, these high concentrations of VOCs appear to maintain OH, as has been observed previously over Amazonia. The boundary-layer characteristics and mixing ratios of VOCs observed over the Borneo rainforest are different to those measured previously over Amazonia. Compared with the Bornean rainforest, air over the oil palm plantation contains much more isoprene, monoterpenes are relatively less important, and the flower scent, estragole, is prominent. Concentrations of nitrogen oxides are greater above the agro-industrial oil palm landscape than over the rainforest, and this leads to changes in some secondary pollutant mixing ratios (but not, currently, differences in ozone). Secondary organic aerosol over both landscapes shows a significant contribution from isoprene. Primary biological aerosol dominates the super-micrometre aerosol over the rainforest and is likely to be sensitive to land-use change, since the fungal source of the bioaerosol is closely linked to above-ground biodiversity.     

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.     

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。     

亚热带城乡复合系统BVOC排放清单——以台州地区为例

挥发性有机复合物(VOC)对大气的化学组成有重要的影响。植物排放的VOC(BVOC)比人为源VOC(AVOC)具有更高的反应活性。通过实地调查台州地区的植被构成和分布,综合BVOC排放速率的测定值和文献数值、当地气象资料等,研究了台州城市和周边地区的BVOC排放强度、时空格局及种类特征。结果表明,2009年台州地区的BVOC年排放量为4.6×1010 g C,其中异戊二烯、单萜和其他VOC所占的比重分别为93.8%、3.5%和2.7%。野外森林中毛竹林排放强度(133.8 t C.km-.2a-1)高出马尾松林、杉木林和常绿阔叶林(分别为0.9、0.8、0.6 t C.km-.2a-1)2个数量级,因此从改善大气质量角度考虑应当抑制毛竹林的扩张。城市行道树及河道树的BVOC排放强度为2.4 t C.km-.2a-1,公园及居民区绿地为1.6 t C.km-.2a-1,均高于野外除毛竹林外的其他植被类型。台州市区的主要树种中,垂柳、合欢、龙爪槐和枫香的单株排放强度较高,而香樟、广玉兰、银杏等单株排放强度相对较低。可优先选择这些排放强度低的树种用作城市绿化。研究结果也可为处在相近气候带的城市中绿化树种的选择和大气质量的改善提供科学依据。