The Complexity of Factors Driving Volatile Organic Compound Emissions by Plants

The emissions of volatile organic compounds, VOC, from plants have strong relevance for plant physiology, plant ecology, and atmospheric chemistry. We report here on the current knowledge of the many internal (genetic and biochemical) and external (abiotic - temperature, light, water availability, wind, ozone, and biotic - animal, plant and microorganisms interactions) factors that control emission rates of different VOC by altering their synthesis, vapour pressure or diffusion to the atmosphere. The complex net of these factors, their interactions and the different responses of the different VOC produces the large qualitative and quantitative, spatial and temporal variability of emissions and the frequent deviations from current standard emission models. The need for a co-operative multidisciplinary multiscale research to disentangle this complex and important issue of plant VOC emissions is reminded.     

Multiple functions of inducible plant volatiles

A considerable amount of the carbon fixed by plants is emitted back into the atmosphere as volatile organic compounds (VOCs). Novel inducible VOCs released from plants after biotic or abiotic stresses temporarily increase total emissions of carbon substantially. As well as having a role in attracting the natural enemies of herbivores, inducible VOCs are also involved in plant-to-plant signalling, pathogen defence and ozone quenching, as well as tropospheric ozone and fine-particle aerosol formation. To relate these diverse observations to active plant defence, a conceptual framework of four functional levels (plant cellular interspace, leaf boundary layer, ecosystem and atmosphere) of inducible VOCs is proposed to aid understanding of the evolutionary role of inducible plant volatiles.     

Role of phosphorus and nitrogen in photosynthetic and whole plant carbon gain and nutrient use efficiency in eastern white pine

Summary In white pine (Pinus strobus) seedlings grown in five forest soils from New York State, net photosynthetic capacity (Amax) plant^-1 was correlated with total foliar N plant^-1 (r ²=0.57), but was more highly correlated with total foliar P plant^-1 (r ²=0.82). There was no relationship (r ²<0.01) between Amax [g leaf]^-1 and foliar N [g leaf]^-1 for the pooled data set, but there was a significant (P<0.001), but weak (r ²=0.20) positive relationship between Amax [g leaf]^-1 and foliar P [g leaf]^-1 across all soils. However, within two of the five soils leaf N concentration was a significant (P<0.05) determinant of photosynthetic capacity. Due to differences in soil nutrient availabilities a large range in foliar P:N ratio (0.02–0.15) was observed, and the proportion of leaf P:N appeared to control Amax [g leaf N]^-1. Whole plant nitrogen (NUE) and phosphorus (PUE) use efficiencies were well correlated with whole plant P:N ratio. In addition, NUE was well correlated with Amax [g leaf N]^-1 and PUE was well correlated with Amax [g leaf P]^-1. However, NUE was not well correlated with PUE, and Amax [g leaf N]^-1 was not well correlated with Amax [g leaf P]^-1. These results indicated that P and/or N limitations were important components of photosynthetic nutrient relations in white pine grown in these five soils and suggest that both P and N and their proportions should be considered in analyses of photosynthesis-nutrient relations.     

How does deep water rice solve its aeration problem

In partially flooded deep water rice (Oryza sativa L. cv Habiganj Aman II), continuous air layers trapped between the hydrophobic, corrugated surface of the leaf blades and the surrounding water constitute the major path of aeration. The conduction of gases through the internal air spaces of the leaf is negligible compared to the conduction of gases through the external air layers. The total volume of the air layers on both sides of a leaf blade is about 45% of the volume of the leaf blade itself. The size of the air layers around submerged leaf blades of cereals not adapted to conditions of partial flooding, e.g. of oats, barley, and wheat, is considerably smaller than that of rice. Gases move through the air layers not only by diffusion but also by mass flow. In darkness, air is drawn down from the atmosphere through the air layers along a pressure gradient created by solubilization of respiratory CO₂ in the surrounding water. In light, photosynthetic O₂ is expelled through the air layers to the atmosphere because the solubility of O₂ in water is much lower than that of CO₂. Air layers greatly increase the rate of photosynthetic carbon fixation by enlarging the surface of the gas-liquid interface available for CO₂ uptake from the water. Air layers are vital for the survival of the partially submerged rice plant. When leaves are washed with a dilute solution of a surfactant (Triton X-100), no air layers are formed under water. Plants without air layers do not grow in response to submergence, and the submerged parts of the plant deteriorate as evident by rapid loss of chlorophyll and protein. Air layers provide a significant survival advantage even to completely submerged rice plants.     

假眼小绿叶蝉对茶梢挥发物的行为反应

【目的】研究假眼小绿叶蝉Empoasca vitis G?the对健康茶梢和蝉害茶梢挥发物的行为反应,筛选出可引诱假眼小绿叶蝉的茶梢挥发物。【方法】以动态吸附法收集健康茶梢和蝉害茶梢挥发物,用气相色谱–质谱联用仪(GC-MS)进行定性定量分析,借助于Y形嗅觉仪检测多种挥发物引诱假眼小绿叶蝉成虫的活性。【结果】从健康茶梢和蝉害茶梢中共鉴定出30种挥发物组分,其中烯烃类含量较大。健康茶梢和蝉害茶梢挥发物中共有组分有13种,蝉害之后其含量皆上升,其中Z-b-罗勒烯和乙酸叶醇酯的含量分别是健康茶梢中的142.27倍、12.90倍。蝉害茶梢中新出现的组分有12种,其中紫苏烯含量较高。在10-2、10~(-4)、10-6 g/mL浓度下,乙酸叶醇酯表现出极显著的引诱水平(P<0.01);紫苏烯在10~(-2)和10~(-4) g/mL浓度下表现出极显著引诱水平(P<0.01);10~(-4) g/mL浓度下,Z-b-罗勒烯和D-柠檬烯极显著引诱假眼小绿叶蝉(P<0.01);10~(-4) g/mL芳樟醇、10~(-6) g/mL乙酸正丁酯和10~(-6) g/mL D-柠檬烯呈现出显著引诱水平(P<0.05);混合物组分Blend1和Blend2分别表现出极显著和显著引诱水平;而10~(-6) g/mLa-法尼烯显著排斥假眼小绿叶蝉(P<0.05)。【结论】假眼小绿叶蝉成虫对健康茶梢和蝉害茶梢挥发物多种组分具有不同的行为反应,引诱效果较强的单组分或混合组分的选定可为田间引诱效果试验提供参考。     

Tritiated Water Vapour Exchange Method for the Evaluation of Whole Plant Diffusion Resistance

Water vapour exchange between plant and atmosphere was studiedby using THO as a tracer. Pulses of THO vapour were brieflyinjected into the airflow of an open circuit plant chamber.Water vapour exchange was calculated from the difference intotal quantity of THO escaping from the plant chamber with andwithout a plant The amount of THO exchanged was closely dependenton the product of total leaf area x mean leaf conductance. Themethod is suitable for measuring on the same plant short termvariations in leaf conductance following changes in environmentalconditions.     

Root herbivores influence the behaviour of an aboveground parasitoid through changes in plant-volatile signals

It is widely reported that plants emit volatile compounds when they are attacked by herbivorous insects, which may be used by parasitoids and predators to locate their host or prey. The study of herbivore-induced plant volatiles and their role in mediating interactions between plants, herbivores and their natural enemies have been primarily based on aboveground systems, generally ignoring the potential interactions between above and belowground infochemical- and food webs. This study examines whether herbivory by Delia radicum feeding on roots of Brassica nigra (black mustard) affects the behaviour of Cotesia glomerata , a parasitoid of the leaf herbivore Pieris brassicae , mediated by changes in plant volatiles. In a semi-field experiment with root-damaged and root-undamaged plants C. glomerata prefers to oviposit in hosts feeding on root-undamaged plants. In addition, in a flight-cage experiment the parasitoid also prefers to search for hosts on plants without root herbivores. Plants exposed to root herbivory were shown to emit a volatile blend characterized by high levels of specific sulphur volatile compounds, which are reported to be highly toxic for insects, combined with low levels of several compounds, i.e. beta-farnesene, reported to act as attractants for herbivorous and carnivorous insects. Our results provide evidence that the foraging behaviour of a parasitoid of an aboveground herbivore can be influenced by belowground herbivores through changes in the plant volatile blend. Such indirect interactions may have profound consequences for the evolution of host selection behaviour in parasitoids, and may play an important role in the structuring and functioning of communities.     

Role of plant volatiles in host plant location of the leafminer, Liriomyza sativae (Diptera: Agromyzidae)

Abstract The role of plant volatiles in host plant location of the leafminer Liriomyza sativae Blanchard was studied. Four types of antennal sensilla were identified on the funiculus by scanning electron microscopy: trichoid, basiconic, clavate and grooved sensilla. An olfactory pit, containing groups of sensilla, was present on the ventral side of the funiculus. No sexual difference was detected in sensilla diversity and distribution. In behavioural assays, both males and females were attracted by the odour of the bean Phaseolus vulgaris L. They had distinct EAG responses to the bean odour. No significant sexual difference was found in behaviour or EAG responses.
Electroantennograms were recorded from female L. sativae to 14 plant volatile compounds. The most distinct EAG responses were obtained for: (1) the general green leaf volatiles 1-hexanol (E)-2-hexen-1-ol, (E)-3-hexen-1-ol and its isomers, (Z)-3-hexen-1-ol, the acetate (E)-3-hexenylacetate and the aldehyde hexanal; and (2) limonene, a compound associated with tomato, which is a key host plant of this insect. Other volatile compounds associated with host plants, such as α-pinene, myrcene, β-caryophyllene, and eugenol did not elicit responses. The ability of this insect to locate a host plant appears to be augmented by the perception of a combination of host-specific and general green leaf volatiles. A modification of the EAG recording method of Dipteran species was provided.     

顶空进样气相色谱-质谱联用法分析木棉花挥发性成分

为研究木棉Bombax ceiba 鲜花的挥发性成分,采用顶空固相微萃取-气相色谱-质谱联用法(HS-SPME-GC-MS)进行鉴定分析。通过GC-MS,采用面积归一法从木棉花中鉴定了56 个主要组分及其相对含量。结果表明,主要组分为模菲(13.74%)、(1R,3aS,5aS,8aR)-1,3a,4,5a 四甲基-1,2,3,3a,5a,6,7,8-八氢环戊二烯并[c]环戊二烯(10.95%)、2-亚甲基-4,8,8-三甲基-4-乙烯基-双环[5.2.0]壬烷(10.36%)等。其中萜烯类化合物占55.24%,烷类化合物占29.72%,还有少量的醛类、酮类和酯类化合物。     

Shared signals -'alarm calls' from plants increase apparency to herbivores and their enemies in nature

The attraction of natural enemies of herbivores by volatile organic compounds as an induced indirect defence has been studied in several plant systems. The evidence for their defensive function originates mainly from laboratory studies with trained parasitoids and predators; the defensive function of these emissions for plants in natural settings has been rarely demonstrated. In native populations and laboratory Y-tube choice experiments with transgenic Nicotiana attenuata plants unable to release particular volatiles, we demonstrate that predatory bugs use terpenoids and green leaf volatiles (GLVs) to locate their prey on herbivore-attacked plants. By attracting predators with volatile signals, this native plant reduces its herbivore load – demonstrating the defensive function of herbivore-induced volatile emissions. However, plants producing GLVs are also damaged more by flea beetles. The implications of these conflicting ecological effects for the evolution of induced volatile emissions and for the development of sustainable agricultural practices are discussed.     

Why and how terrestrial plants exchange gases with air

This work is intended as a review of gas exchange processes between the atmosphere and the terrestrial vegetation, which have been known for more than two centuries since the discovery of photosynthesis. The physical and biological mechanisms of exchange of carbon dioxide, water vapour, volatile organic compounds emitted by plants and air pollutants taken up by them, is critically reviewed. The role of stomatal physiology is emphasised, as it controls most of these processes. The techniques used for measurement of gas exchange fluxes between the atmosphere and vegetation are outlined.     

The influence of transpiration on the equilibration of leaf water with atmospheric water vapour

Abstract. The authors examine the isotopic composition of leaf water, at natural abundance levels, as influenced by transpiration rate. The isotopic composition of water of wheat leaves ( Triticum aestivum L. var. Aroona) was followed while their transpiration rate adjusted to 'steady-state' environmental conditions. Leaf diffusive resistance was modified by short-term salt treatment and by plant culture in either nutrient solution, free-draining sand, or vermiculite. Resultant changes in ¹⁸O and ²H in leaf water are described and fitted to the model of Leaney et al. (1985). The treatments with lower transpiration rates were found to have a greater fraction of their leaf water equilibrated with water vapour in the atmosphere. Comparable results were obtained with both ¹⁸O and ²H, with some differences being interpreted in terms of turbulence in the vapour diffusion path. The fraction of the leaf water equilibrated with the atmosphere varied between leaves of different ages. However, this may have been due to their different positions in the canopy.     

植物根际促生枯草芽孢杆菌JC01筛选

【目的】筛选高效植物根际促生细菌,阐明产挥发性有机化合物(VOC)菌株JC01的促生机制。【方法】选取从植物根际中分离得到的838株细菌,以固氮、解(溶)磷以及分泌嗜铁素、吲哚乙酸(IAA)活性为指标,对其促生能力进行赋值评估,将赋值在3分以上的107株细菌进行指纹图谱分析,挑选其中不同簇的20株促生潜力细菌进行温室实验,以评价赋值体系与温室促生效果之间的关系,进一步探究具有较好促生效果菌株JC01的作用机理。【结果】共筛选出了来源于指纹图谱中不同簇的4株具有较好促生效果的菌株,细菌的平板活性赋值与促生效果之间的相关系数r大于0.6。其中,菌株JC01分泌的具有促生作用的VOC能够增强番茄植株IAA信号通路关键基因的表达,减弱脱落酸(ABA)、乙烯(ETH)信号通路关键基因的表达。JC01经16S r RNA基因鉴定为枯草芽孢杆菌。【结论】细菌的平板活性赋值与促生效果之间存在较高的正相关性,枯草芽孢杆菌JC01可能通过产生VOC对番茄生长进行调控。     

Volatile Semiochemicals Released from Undamaged Cotton Leaves (A Systemic Response of Living Plants to Caterpillar Damage)

Cotton plants (Gossypium hirsutum L.), attacked by herbivorous insects release volatile semiochemicals (chemical signals) that attract natural enemies of the herbivores to the damaged plants. We found chemical evidence that volatiles are released not only at the damaged site but from the entire cotton plant. The release of volatiles was detected from upper, undamaged leaves after 2 to 3 d of continuous larval damage on lower leaves of the same plant. Compounds released systemically were (Z)-3-hexenyl acetate, (E)-[beta]-ocimene, linalool, (E)-4,8-dimethyl-1,3,7-nonatriene, (E)-[beta]-farnesene, (E,E)-[alpha]-farnesene, and (E,E)-4,8,12-trimethyl-1,3,7,11-tridecatetraene. All systemically released compounds are known to be induced by caterpillar damage and are not released in significant amounts by undamaged plants. Other compounds, specifically indole, isomeric hexenyl butyrates, and 2-methylbutyrates, known to be released by cotton in response to caterpillar damage, were not released systemically. However, when upper, undamaged leaves of a caterpillar-damaged plant were damaged with a razor blade, they released isomeric hexenyl butyrates, 2-methylbutyrates, and large amounts of constitutive compounds in addition to the previously detected induced compounds. Control plants, damaged with a razor blade in the same way, did not release isomeric hexenyl butyrates or 2-methylbutyrates and released significantly smaller amounts of constitutive compounds. Indole was not released systemically, even after artificial damage.     

Volatile organic compound fate in phytoremediation applications: natural and engineered systems

Unique sampling techniques have generated a new understanding regarding the fate of volatile organic compounds (VOCs) in phytoremediation systems. Tissue sampling and diffusion traps were used to determine how VOCs are transported in and diffuse from vegetation, particularly woody species. These techniques were then utilized to observe how plants interact with different contaminated media, showing transport of contaminants occurs from the vadose zone (vapor phase) as well as the saturated zone (aqueous phase). Data was gathered in laboratory studies, in native vegetation, and in engineered phytoremediation systems. The findings reveal that diffusion from the xylem tissues to the atmosphere is a major fate for VOCs in phytoremediation applications. Linking VOCs' fate with groundwater hydraulics, mass removal rates from contaminant plumes can be estimated. These techniques were also utilized to observe the impact of engineered plant/microbe systems, which utilize recombinant, root-colonizing organisms to selectively degrade compounds and subsequently alter the fate of VOCs and other organic compounds. The genetically enhanced rhizoremediation methods pose a novel approach that may allow for biodegradation of compounds that formerly were considered recalcitrant.     

8个茶树品种生化成分分析及抗性成分的初步鉴定

【目的】筛选和鉴定茶树中与假眼小绿叶蝉Empoasca vitis G?the相关的抗性成分。【方法】采用氨基酸分析仪和超高效液相色谱法分别分析8个茶树品种的氨基酸及其组分、咖啡碱和茶多酚及其组分的含量。采用人工饲料喂饲法对可能的抗性物质进行初步鉴定。【结果】通过对假眼小绿叶蝉抗性水平差异较大的茶树品种的30多种生化物质的分析,筛选出抗、感茶树品种间差异较大的物质,包括茶多酚、天冬氨酸、γ-氨基丁酸、绿原酸和茶氨酸。分别将其作为可变因子设置不同浓度梯度,加入到人工饲料饲养假眼小绿叶蝉,统计5种化学成份不同浓度人工饲料上假眼小绿叶蝉的成活率,初步确定了这5种化学成份对假眼小绿叶蝉成活率的影响。其中,γ-氨基丁酸可能是茶树抗虫物质之一。【结论】为阐明茶树抗叶蝉的化学机理奠定基础。     

How do leaf hydraulics limit stomatal conductance at high water vapour pressure deficits?

A reduction in leaf stomatal conductance (g) with increasing leaf-to-air difference in water vapour pressure (D) is nearly ubiquitous. Ecological comparisons of sensitivity have led to the hypothesis that the reduction in g with increasing D serves to maintain leaf water potentials above those that would cause loss of hydraulic conductance. A reduction in leaf water potential is commonly hypothesized to cause stomatal closure at high D. The importance of these particular hydraulic factors was tested by exposing Abutilon theophrasti, Glycine max, Gossypium hirsutum and Xanthium strumarium to D high enough to reduce g and then decreasing ambient carbon dioxide concentration ([CO2]), and observing the resulting changes in g, transpiration rate and leaf water potential, and their reversibility. Reducing the [CO2] at high D increased g and transpiration rate and lowered leaf water potential. The abnormally high transpiration rates did not result in reductions in hydraulic conductance. Results indicate that low water potential effects on g at high D could be overcome by low [CO2], and that even lower leaf water potentials did not cause a reduction in hydraulic conductance in these well-watered plants. Reduced g at high D in these species resulted primarily from increased stomatal sensitivity to [CO2] at high D, and this increased sensitivity may mediate stomatal responses to leaf hydraulics at high D.     

采用多次顶空固相微萃取分析拟南芥绿叶挥发性物质

顶空固相微萃取作为一种新的挥发性和半挥发性物质分析技术,被广泛应用于植物样品的定性分析。由于进行顶空分析时,挥发性组分间的基质效应以及较为复杂的扩散和吸附过程,定量分析一直是SPME分析应用的难题。目标分析物的量看作是达到吸附平衡后单一萃取的物质量的总和,则无需考虑分析样品在顶空、萃取涂层间的分配,同时可以消除基质效应。在利用标准物质进行校正后只需要一次顶空萃取,即可求出分析物质的总量。首先利用DVB/CAR/PDMS定性得到拟南芥挥发性物质的组成,然后采用CAR/PDMS涂层定量,分析了拟南芥的3种绿叶挥发性物质,优化后萃取条件为40℃萃取20min,相对标准偏差小于12%,在3株植物样品中这些挥发性物质的量为78.6~158.4ng.g-1。     

Combined impacts of irradiance and dehydration on leaf hydraulic conductance: insights into vulnerability and stomatal control

The leaf is a hydraulic bottleneck, accounting for a large part of plant resistance. Thus, the leaf hydraulic conductance (K(leaf) ) is of key importance in determining stomatal conductance (g(s) ) and rates of gas exchange. Previous studies showed that K(leaf) is dynamic with leaf water status and irradiance. For four species, we tested the combined impacts of these factors on K(leaf) and on g(s) . We determined responses of K(leaf) and g(s) to declining leaf water potential (Ψ(leaf) ) under low and high irradiance (<6 and >900 μmol photons m(-2) s(-1) photosynthetically active radiation, respectively). We hypothesized greater K(leaf) vulnerability under high irradiance. We also hypothesized that K(leaf) and g(s) would be similar in their responses to either light or dehydration: similar light-responses of K(leaf) and g(s) would stabilize Ψ(leaf) across irradiances for leaves transpiring at a given vapour pressure deficit, and similar dehydration responses would arise from the control of stomata by Ψ(leaf) or a correlated signal. For all four species, the K(leaf) light response declined from full hydration to turgor loss point. The K(leaf) and g(s) differed strongly in their light- and dehydration responses, supporting optimization of hydraulic transport across irradiances, and semi-independent, flexible regulation of liquid and vapour phase water transport with leaf water status.     

In situ translocation of volicitin by beet armyworm larvae to maize and systemic immobility of the herbivore elicitor <Emphasis Type="Italic">in planta</Emphasis>

Volicitin (N-[17-hydroxylinolenoyl]-l glutamine) present in the regurgitant of beet armyworm (Spodoptera exigua) activates the emissions of volatile organic compounds (VOCs) when in contact with damaged corn (Zea mays L.) leaves. VOC emission in turn serves as a signaling defense for the plant by attracting female parasitic wasps that prey on herbivore larvae. Chemical tracking of volicitin within plants has yet to be reported. Here we present biochemical data that beet armyworm regurgitant serves as a vector for the introduction of volicitin to the site of leaf damage under natural feeding conditions. Corn seedlings were ¹⁴CO₂-labeled in situ, and beet armyworm larvae were allowed to feed on the labeled leaves. Herbivore oral secretions collected from late-third-instar larvae contained approximately 120 pmol volicitin (0.05 nCi pmol^–1) per larva. When radiochemically labeled larvae were placed on unlabeled leaves, the amount of volicitin introduced to the damaged site was approximately 5.0 nCi (calc. 100 pmol/larvae). The mobility of volicitin in leaves was examined by allowing radiolabeled beet armyworms to feed on unlabeled plants. In such tracking experiments, radioactivity was not detected in the upper leaves; however, the exogenous application of 5 nCi of [U-¹⁴C]sucrose to the lower leaf did result in subsequent radioactivity being detected in the upper portion of the plant. The detection of labeled sucrose with the same radioactivity as that of administered volicitin indicated that volicitin was not readily transported to undamaged leaves and that volicitin may not directly serve as a mobile messenger in triggering the emissions of VOCs systemically.Abbreviations BAW Beet armyworm (Spodoptera exigua) - dpm Disintegrations per minute - FAA Fatty acid amide - JA Jasmonic acid - VOC Volatile organic compound