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Living trees are the main source of biogenic volatile organic compounds (BVOCs) in forest ecosystems, but substantial emissions originate from leaf and wood litter, the rhizosphere and from microorganisms. This review focuses on temperate and boreal forest ecosystems and the roles of BVOCs in ecosystem function, from the leaf to the forest canopy and from the forest soil to the atmosphere level. Moreover, emphasis is given to the question of how BVOCs will help forests adapt to environmental stress, particularly biotic stress related to climate change. Trees use their vascular system and emissions of BVOCs in internal communication, but emitted BVOCs have extended the communication to tree population and whole community levels and beyond. Future forestry practices should consider the importance of BVOCs in attraction and repulsion of attacking bark beetles, but also take an advantage of herbivore-induced BVOCs to improve the efficiency of natural enemies of herbivores. BVOCs are extensively involved in ecosystem services provided by forests including the positive effects on human health. BVOCs have a key role in ozone formation but also in ozone quenching. Oxidation products form secondary organic aerosols that disperse sunlight deeper into the forest canopy, and affect cloud formation and ultimately the climate. We also discuss the technical side of reliable BVOC sampling of forest trees for future interdisciplinary studies that should bridge the gaps between the forest sciences, health sciences, chemical ecology, conservation biology, tree physiology and atmospheric science.

Plants emit biogenic volatile organic compounds (BVOCs) causing transcriptomic, metabolomic and behavioral responses in receiver organisms. Volatiles involved in such responses are often called "plant language". Arthropods having sensitive chemoreceptors can recognize language released by plants. Insect herbivores, pollinators and natural enemies respond to composition of volatiles from plants with specialized receptors responding to different types of compounds. In contrast, the mechanism of how plants "hear" volatiles has remained obscured. In a plant-plant communication, several individually emitted compounds are known to prime defense response in receiver plants with a specific manner according to the chemical structure of each volatile compound. Further, composition and ratio of volatile compounds in the plant-released plume is important in plant-insect and plant-plant interactions mediated by plant volatiles. Studies on volatile-mediated plant-plant signaling indicate that the signaling distances are rather short, usually not longer than one meter. Volatile communication from plants to insects such as pollinators could be across distances of hundreds of meters. As many of the herbivore induced VOCs have rather short atmospheric life times, we suggest that in long-distant communications with plant volatiles, reaction products in the original emitted compounds may have additional information value of the distance to emission source together with the original plant-emitted compounds.

It is well known that vertical canopy gradients and varying sky conditions influence photosynthesis (Pn), specific leaf area (SLA), leaf thickness (LT) and leaf pigments (lutein, â-carotene and chlorophyll). In contrast, little is known about these effects on monoterpenoid (MT) emissions. Our study examines simultaneously measured Pn, MT emissions and the MT/Pn ratio along the canopy of an adult European beech tree (Fagus sylvatica L.) in natural forest conditions. Dynamic branch enclosure systems were used at four heights in the canopy (7, 14, 21 and 25 m) in order to establish relationships and better understand the interaction between Pn and MT emissions under both sunny and cloudy sky conditions. Clear differences in Pn, MT emissions and the MT/Pn ratio were detected within the canopy. The highest Pn rates were observed in the sun leaves at 25 m due to the higher intercepted light levels, whereas MT emissions (and the MT/Pn ratio) were unexpectedly highest in the semi-shaded leaves at 21 m. The higher Pn rates and, apparently contradictory, lower MT emissions in the sun leaves may be explained by the hypothesis of Owen and Penuelas (2005), stating synthesis of more photo-protective carotenoids may decrease the emissions of volatile isoprenoids (including MTs) because they both share the same biochemical precursors. In addition, leaf traits like SLA, LT and leaf pigments clearly differed with height in the canopy, suggesting that the leaf's physiological status cannot be neglected in future research on biogenic volatile organic compounds (BVOCs) when aiming at developing new and/or improved emission algorithms.

In this study, biogenic volatile organic compound (BVOC) emissions from Abies alba were studied under ambient conditions in Flanders (Belgium). Emission patterns and rates were investigated from April till November 2010 by using the dynamic branch enclosure technique. The present work revealed that A. alba is an isoprene emitter, with isoprene accounting for 86–93% of total BVOC emissions, except during budburst (67%) in May. The emission spectrum of A. alba consisted of 27 compounds. Next to isoprene, the main emitted compounds were α-pinene, β-pinene, camphene and limonene. BVOC emissions showed a peak in June after development of the young needles, followed by a constant emission during summer months and September and a decrease in October. In all the samples isoprene was the most abundant compound with standardized emission rates between 27 μg g(dw)−1 h−1 in June and 4.6 μg g(dw)−1 h−1 in October, while the total standardized terpenoid emission rates ranged from 2.85 μg g(dw)−1 h−1 in June to 0.26 μg g(dw)−1 h−1 in October. The obtained average β coefficients according to the temperature dependent algorithm of Guenther et al. (1993) during April–June, July, August and September–October were as follows: for terpenoids 0.12 ± 0.03, 0.11 ± 0.05, 0.12 ± 0.04, 0.24 ± 0.01 K−1 and sesquiterpenes (SQTs) 0.09 ± 0.02, 0.11 ± 0.01, 0.10 ± 0.05, 0 K−1, respectively. Overall, isoprene detected in this study was never quantified in previous studies on A. alba and this finding could have a significant impact on the regional BVOCs budget. Therefore, the result of this study is very important for modeling and local air quality.

Direct plant stress sensing is the key for a quantitative understanding of drought stress effects on biogenic volatile organic compound (BVOC) emissions. A given level of drought stress might have a fundamentally different effect on the BVOC emissions of different plants. For the first time, we continuously quantified the level of drought stress in a young potted beech (Fagus sylvatica L.) with a linear variable displacement transducer (LVDT) installed at stem level in combination with simultaneous measurements of BVOC emissions and photosynthesis rates at leaf level. This continuous set of measurements allowed us to examine how beech alters its pattern of photosynthesis and carbon allocation to BVOC emissions (mainly monoterpenes, MTs) and radial stem growth during the development of drought stress. We observed an increasing-decreasing trend in the MT emissions as well as in the fraction of assimilated carbon re-emitted back into the atmosphere (ranging between 0.14 and 0.01%). We were able to link these dynamics to pronounced changes in radial stem growth, which served as a direct plant stress indicator. Interestingly, we detected a sudden burst in emission of a non-identified, non-MT BVOC species when drought stress was acute (i.e. pronounced negative stem growth). This burst might have been caused by a certain stress-related green leaf volatile, which disappeared immediately upon re-watering and thus the alleviation of drought stress. These results highlight that direct plant stress sensing creates opportunities to understand the overall complexity of stress-related BVOC emissions.

A dynamic branch enclosure system was used to measure emission rates of biogenic volatile organic compounds (BVOCs) from two common European tree species: Fraxinus excelsior and Quercus robur under ambient conditions in Flanders (Belgium). Both tree species were studied for seasonal variability of BVOC emission rates under natural biotic stress (infestations). Emissions were normalized at standard conditions of temperature and photosynthetic active radiation (PAR) (30°C and 1000 µmol m−2 s−1, respectively). Emission rates from Fraxinus excelsior were highest in May (9.56 µg gDW −1 h−1) and lowest in October (1.17 µg gDW −1 h−1). This tree species emitted (Z)-β-ocimene, (E)-β-ocimene and α-farnesene during the entire measurement period and additionally isoprene only in May. Quercus robur showed isoprene emission variations according to the seasonal cycle with rates of 30, 106 and 29 µg gDW −1 h−1 in May, August and October, respectively. Apart from isoprene, (E)-β-ocimene and β-caryophyllene were emitted t...

Induced volatiles have been a focus of recent research, as not much is known of their emission behavior or atmospheric contribution. BVOC emissions were measured from Pseudotsuga menziesii saplings under natural environmental conditions, using a dynamic branch enclosure system and GC–MS for their analysis. We determined temperature and light dependency of the individual compounds, studied seasonality of the emissions and discuss the effect of heat stress in comparison with two specific biotic stresses that occurred naturally on the trees. A standardized emission rate of 6.8 μg g(dw)−1 h−1 for monoterpenes under stressed conditions was almost a magnitude higher than that obtained for healthy trees (0.8 ± 0.2 μg g(dw)−1 h−1), with higher beta factors characterizing the stressed trees. The response of the emissions to light intensity was different for the individual compounds, suggesting a distinct minimum light intensity to reach saturation. Heat stress changed the relative contribution of specific volatiles, with larger extent of increase of sesquiterpenes, methyl salicylate and linalool emissions compared to monoterpenes. Biotic stress kept low the emissions of sesquiterpenes, (E)-4,8-dimethyl-1,3,7-nonatriene and methylbutenol isomers, and increased the level of methyl salicylate and monoterpenes. The ratio of β-pinene/α-pinene was also found to be significantly enhanced from 1.3 to 2.4 and 3.2 for non-stressed, heat stressed and combined biotic and heat stressed, respectively.

We measured the isoprene and monoterpene fluxes continuously above a mixed forest site at Vielsalm in the eastern part of Belgium, using the disjunct eddy covariance technique with proton transfer reaction-mass spectrometry. Simultaneously, we also measured the carbon dioxide fluxes in order to deduce the gross primary production. The measurements were conducted from July to September 2009. During the day, the seasonal evolution of the isoprene/monoterpene emissions was studied using a monthly temperature and light dependence function deduced from our results to standardize the fluxes. A seasonal decrease in the standard emission factors was observed, probably linked to acclimation or senescence. The standard emission factor for isoprene fluxes (30 °C, 1000 μmol m −2 s −1 ) fell from 0.91 ± 0.01 to 0.56 ± 0.02 μg m −2 s −1 and for monoterpene fluxes from 0.74 ± 0.03 to 0.27 ± 0.03 μg m −2 s −1 . During the night, a slight positive flux of monoterpenes was observed that seemed to be driven by air temperature. The standard emission factor (30°C) for nighttime monoterpene fluxes was equal to 0.093 ± 0.019 μg m −2 s −1 . Finally, we studied the seasonal evolution of the relationship between the gross primary production and the isoprene/monoterpenes fluxes. A linear relationship was observed, highlighting the strong link between carbon assimilation and isoprene/monoterpene emissions.

Although biogenic volatile organic compounds (BVOCs) only represent a very limited fraction of the plant’s carbon (C) budget, they play an important role in atmospheric chemistry for example as a precursor of tropospheric ozone. We performed a study comparing BVOC emissions of European beech (Fagus sylvatica L.) in controlled and natural environmental conditions. A young and adult beech tree was exposed to short-term temperature variations in growth room conditions and in an experimental forest, respectively. This study attempts to clarify how short-term temperature variations between days influenced the ratio between monoterpenoid (MT) emissions and net photosynthesis (Pn). Within a temperature range of 17–27 °C and 13–23 °C, the MT/Pn carbon ratio increased 10–30 fold for the growth room and forest, respectively. An exponential increasing trend between MT/Pn C ratio and air temperature was observed in both conditions. Beech trees re-emitted a low fraction of the assimilated C back into the atmosphere as MT: 0.01–0.12% and 0.01–0.30% with a temperature rise from 17 to 27 °C and 13–23 °C in growth room and forest conditions, respectively. However, the data showed that the MT/Pn C ratio of young and adult beech trees responded significantly to changes in temperature.

Monoterpenoid emissions from Fagus sylvatica L. trees have been measured at light- and temperature-controlled conditions in a growth chamber, using Proton Transfer Reaction Mass Spectrometry (PTR-MS) and the dynamic branch enclosure technique. De novo synthesized monoterpenoid Standard Emission Factors, obtained by applying the G97 algorithm ( Guenther, 1997 ), varied between 2 and 32 μg gDW−1 h−1 and showed a strong decline in late August and September, probably due to senescence. The response of monoterpenoid emissions to temperature variations at a constant daily light pattern could be well reproduced with a modified version of the MEGAN algorithm ( Guenther et al., 2006 ), with a typical dependence on the average temperature over the past five days. The diurnal emissions at constant temperature showed a typical hysteretic behaviour, which could also be adequately described with the modified MEGAN algorithm by taking into account a dependence on the average light levels experienced by the trees during the past 10–13 h. The impact of the past light and temperature conditions on the monoterpenoid emissions from F. sylvatica L. was found to be much stronger than assumed in previous algorithms. Since our experiments were conducted under low light intensity, future studies should aim at confirming and completing the proposed algorithm updates in sunny conditions and natural environments.

The interest in quantitative analysis of biogenic volatile organic compounds (BVOCs) emissions stems from their importance in atmospheric chemistry. In order to compare the most frequently used BVOC measurement techniques, simultaneous on-line PTR-MS and off-line GC-MS data collection was performed on a 3 years old Fagus sylvatica L. tree placed in a growth chamber. Using an internal standard (deuterated toluene) and applying the selective ion mode (SIM) resulted in significant improvements of monoterpene (MT) quantification by TD-GC-MS. PTR-MS quantification of MTs was based on the ion signal at m/z 137. In the course of the experiments the relative contribution of linalool compared to that of MTs was found to be up to 84%. Since this compound has also a PTR-MS signature at m/z 137, quantification of MT emission rates by PTR-MS was disturbed. Comparison of GC-MS and PTR-MS data allowed an estimation of the ratio of the PTR-MS sensitivity for linalool to the one for MTs at m/z 137. This ratio of sensitivities, combined with the information of the relative contribution of linalool to the sum of linalool and MTs obtained by GC-MS, resulted in accurate derivation of the sum of emission rates of linalool and MTs by PTR-MS. The results indicate that fast and on-line PTR-MS measurements of BVOCs are best accompanied by off-line GC measurements to detect possible interferences or to use the additional information for properly quantifying the sum of emission rates of several compounds.

Volatile organic compounds (VOCs) have been the focus of interest to understand atmospheric processes and their consequences in formation of ozone or aerosol particles; therefore, VOCs contribute to climate change. In this study, biogenic VOCs (BVOCs) emitted from Fagus sylvatica L. trees were measured in a dynamic enclosure system. In total 18 compounds were identified: 11 monoterpenes (MT), an oxygenated MT, a homoterpene (C14H18), 3 sesquiterpenes (SQT), isoprene and methyl salicylate. The frequency distribution of the compounds was tested to determine a relation with the presence of the aphid Phyllaphis fagi L. It was found that linalool, (E)-β-ocimene, α-farnesene and a homoterpene identified as (E)-4,8-dimethyl-1,3,7-nonatriene (DMNT), were present in significantly more samples when infection was present on the trees. The observed emission spectrum from F. sylvatica L. shifted from MT to linalool, α-farnesene, (E)-β-ocimene and DMNT due to the aphid infection. Sabinene was quantitatively the most prevalent compound in both, non-infected and infected samples. In the presence of aphids α-farnesene and linalool became the second and third most important BVOC emitted. According to our investigation, the emission fingerprint is expected to be more complex than commonly presumed.