Hybridization of European oaks (Quercus ilex × Q. robur) results in a mixed isoprenoid emitter type

European oaks have been reported to emit isoprene or monoterpenes derived from recently fixed photosynthetic carbon. The emission type is plant species specific and can be used as chemo‐taxonomic marker. In the present article the isoprenoid biochemical properties of mature Quercus × turneri‘Pseudoturneri’ hybrids resulting from a crossing of a Mediterranean evergreen monoterpene‐emitting species (subgenus Sclerophyllodrys; Quercus ilex L.) and an isoprene‐emitting deciduous oak species (subgenus Lepidobalanus; Quercus robur L.) are described. Both species are compared with respect to the capacity for isoprenoid synthesis and the actual isoprenoid emission pattern of different tree‐types. The analysis showed that the oak hybrid combines properties of both parental species. Furthermore, it could be shown that the enzyme activities of isoprene synthase and monoterpene synthases are reflected in the isoprenoid emission pattern of the hybrids as well as in the observed emission rates.     

Monoterpene emission and monoterpene synthase activities in the Mediterranean evergreen oak Quercus ilex L. grown at elevated CO2 concentrations

Monoterpene emissions, monoterpene synthase activities, photosynthesis, fluorescence yield in the dark and drought stress indicators (stomatal conductance and mid‐day water potential) were concurrently measured under similar temperature and illumination in current‐year leaves of Quercus ilex L. of plants grown in open‐top chambers at ambient (350 ppm) and elevated (700 ppm) CO₂. The study was undertaken to understand the effect of CO₂ on monoterpene biosynthesis, and to predict and parameterize the biogenic emissions at growing CO₂ concentrations. The results of the 1998 and 1999 studies show that at elevated CO₂, and in the absence of persistent environmental stresses, photosynthesis was stimulated with respect to ambient CO₂, but that the emission of the three most abundantly emitted monoterpenes (α‐pinene, sabinene and β‐pinene) was inhibited by approximately 68%. The enzyme activities of the monoterpene synthases catalysing the formation of the three monoterpenes were also inhibited at elevated CO₂ and an excellent relationship was found between monoterpene emission and activity of the corresponding enzyme both at ambient and elevated CO₂. Interestingly, however, limonene emission was enhanced in conditions of elevated CO₂ as it was also the corresponding synthase. The ratio between enzyme activity and emission of the three main monoterpenes was high (above 20) at ambient CO₂ but it was below 10 at elevated CO₂ and, for limonene, on both treatments. Our results indicate that the overall emission of monoterpenes at elevated CO₂ will be inhibited because of a concurrent, strong down‐regulation of monoterpene synthase activities. When the enzyme activity does not change, as for limonene, the high photosynthetic carbon availability at elevated CO₂ conditions may even stimulate emission. The results of the 1997 study show that severe and persistent drought, as commonly occurs in the Mediterranean, may inhibit both photosynthesis and monoterpene (α‐pinene) emission, particularly at ambient CO₂. Thus, emission is probably limited by photosynthetic carbon availability; the effect of elevated CO₂per se is not apparent if drought, and perhaps other environmental stresses, are also present.     

Interactive effects of elevated CO2 and soil fertility on isoprene emissions from Quercus robur

The effects of global change on the emission rates of isoprene from plants are not clear. A factor that can influence the response of isoprene emission to elevated CO₂ concentrations is the availability of nutrients. Isoprene emission rate under standard conditions (leaf temperature: 30°C, photosynthetically active radiation (PAR): 1000 μmol photons m^?2 s^?1), photosynthesis, photosynthetic capacity, and leaf nitrogen (N) content were measured in Quercus robur grown in well‐ventilated greenhouses at ambient and elevated CO₂ (ambient plus 300 ppm) and two different soil fertilities. The results show that elevated CO₂ enhanced photosynthesis but leaf respiration rates were not affected by either the CO₂ or nutrient treatments. Isoprene emission rates and photosynthetic capacity were found to decrease with elevated CO₂, but an increase in nutrient availability had the converse effect. Leaf N content was significantly greater with increased nutrient availability, but unaffected by CO₂. Isoprene emission rates measured under these conditions were strongly correlated with photosynthetic capacity across the range of different treatments. This suggests that the effects of CO₂ and nutrient levels on allocation of carbon to isoprene production and emission under near‐saturating light largely depend on the effects on photosynthetic electron transport capacity.     

Enhanced isoprene emission capacity and altered light responsiveness in aspen grown under elevated atmospheric CO2 concentration

Controversial evidence of CO₂‐responsiveness of isoprene emission has been reported in the literature with the response ranging from inhibition to enhancement, but the reasons for such differences are not understood. We studied isoprene emission characteristics of hybrid aspen (Populus tremula x P. tremuloides) grown under ambient (380 μmol mol^?1) and elevated (780 μmol mol^?1) [CO₂] to test the hypothesis that growth [CO₂] effects on isoprene emission are driven by modifications in substrate pool size, reflecting altered light use efficiency for isoprene synthesis. A novel in vivo method for estimation of the pool size of the immediate isoprene precursor, dimethylallyldiphosphate (DMADP) and the activity of isoprene synthase was used. Growth at elevated [CO₂] resulted in greater leaf thickness, more advanced development of mesophyll and moderately increased photosynthetic capacity due to morphological “upregulation”, but isoprene emission rate under growth light and temperature was not significantly different among ambient‐ and elevated‐[CO₂]‐grown plants independent of whether measured at 380 μmol mol^?1 or 780 μmol mol^?1 CO₂. However, DMADP pool size was significantly less in elevated‐[CO₂]‐grown plants, but this was compensated by increased isoprene synthase activity. Analysis of CO₂ and light response curves of isoprene emission demonstrated that the [CO₂] for maximum isoprene emission was shifted to lower [CO₂] in elevated‐[CO₂]‐grown plants. The light‐saturated isoprene emission rate (I_max,Q) was greater, but the quantum efficiency at given I_max,Q was less in elevated‐[CO₂]‐grown plants, especially at higher CO₂ measurement concentration, reflecting stronger DMADP limitation at lower light and higher [CO₂]. These results collectively demonstrate important shifts in light and CO₂‐responsiveness of isoprene emission in elevated‐[CO₂]‐acclimated plants that need consideration in modeling isoprene emissions in future climates.     

Light and temperature dependence of the emission of cyclic and acyclic monoterpenes from holm oak (Quercus ilex L.) leaves

In a laboratory study, we investigated the monoterpene emissions from Quercus ilex, an evergreen sclerophyllous Mediterranean oak species whose emissions are light dependent. We examined the light and temperature responses of individual monoterpenes emitted from leaves under various conditions, the effect of heat stress on emissions, and the emission-onset during leaf development. Emission rate increased 10-fold during leaf growth, with slight changes in the composition. At 30 °C and saturating light, the monoterpene emission rate from mature leaves averaged 4·1 nmol m^–2 s^–1, of which α-pinene, sabinene and β-pinene accounted for 85%. The light dependence of emission was similar for all monoterpenes: it resembled the light saturation curve of CO₂ assimilation, although monoterpene emission continued in the dark. Temperature dependence differed among emitted compounds: most of them exhibited an exponential increase up to 35 °C, a maximum at 42 °C, and a slight decline at higher temperatures. However, the two acyclic isomers cis-β-ocimene and trans-β-ocimene were hardly detected below 35 °C, but their emission rates increased above this temperature as the emission rates of other compounds fell, so that total emission of monoterpenes exponentially increased from 5 to 45 °C. The ratio between ocimene isomers and other compounds increased with both absolute temperature and time of heat exposure. The light dependence of emission was insensitive to the temperature at which it was measured, and vice versa the temperature dependence was insensitive to the light regime. The results demonstrated that none of the models currently applied to simulate isoprene or monoterpene emissions correctly predicts the short-term effects of light and temperature on Q. ilex emissions. The percentage of fixed carbon lost immediately as monoterpenes ranged between 0·1 and 6·0% depending on temperature, but rose up to 20% when leaves were continuously exposed to temperatures between 40 and 45 °C.     

Impact of rising CO2 on emissions of volatile organic compounds: isoprene emission from Phragmites australis growing at elevated CO2 in a natural carbon dioxide spring†

Isoprene basal emission (the emission of isoprene from leaves exposed to a light intensity of 1000 µmol m^?2 s^?1 and maintained at a temperature of 30 °C) was measured in Phragmites australis plants growing under elevated CO₂ in the Bossoleto CO₂ spring at Rapolano Terme, Italy, and under ambient CO₂ at a nearby control site. Gas exchange and biochemical measurements were concurrently taken. Isoprene emission was lower in the plants growing at elevated CO₂ than in those growing at ambient CO₂. Isoprene emission and isoprene synthase activity (IsoS) were very low in plants growing at the bottom of the spring under very rich CO₂ and increased at increasing distance from the spring (and decreasing CO₂ concentration). Distance from the spring did not significantly affect photosynthesis making it therefore unlikely that there is carbon limitation to isoprene formation. The isoprene emission rate was very quickly reduced after rapid switches from elevated to ambient CO₂ in the gas‐exchange cuvette, whereas it increased when switching from ambient to elevated CO₂. The rapidity of the response may be consistent with post‐translational modifications of enzymes in the biosynthetic pathway of isoprene formation. Reduction of IsoS activity is interpreted as a long‐term response. Basal emission of isoprene was not constant over the day but showed a diurnal course opposite to photosynthesis, with a peak during the hottest hours of the day, independent of stomatal conductance and probably dependent on external air temperature or temporary reduction of CO₂ concentration. The present experiments show that basal emission rate of isoprene is likely to be reduced under future elevated CO₂ levels and allow improvement in the modelling of future isoprene emission rates.     

Profiles of isoprene emission and photosynthetic parameters in hybrid poplars exposed to free‐air CO2 enrichment†

Poplar (Populus × euroamericana) saplings were grown in the field to study the changes of photosynthesis and isoprene emission with leaf ontogeny in response to free air carbon dioxide enrichment (FACE) and soil nutrient availability. Plants growing in elevated [CO₂] produced more leaves than those in ambient [CO₂]. The rate of leaf expansion was measured by comparing leaves along the plant profile. Leaf expansion and nitrogen concentration per unit of leaf area was similar between nutrient treatment, and this led to similar source–sink functional balance. Consequently, soil nutrient availability did not cause downward acclimation of photosynthetic capacity in elevated [CO₂] and did not affect isoprene synthesis. Photosynthesis assessed in growth [CO₂] was higher in plants growing in elevated than in ambient [CO₂]. After normalizing for the different number of leaves over the profile, maximal photosynthesis was reached and started to decline earlier in elevated than in ambient [CO₂]. This may indicate a [CO₂]‐driven acceleration of leaf maturity and senescence. Isoprene emission was adversely affected by elevated [CO₂]. When measured on the different leaves of the profile, isoprene peak emission was higher and was reached earlier in ambient than in elevated [CO₂]. However, a larger number of leaves was emitting isoprene in plant growing in elevated [CO₂]. When integrating over the plant profile, emissions in the two [CO₂] levels were not different. Normalization as for photosynthesis showed that profiles of isoprene emission were remarkably similar in the two [CO₂] levels, with peak emissions at the centre of the profile. Only the rate of increase of the emission of young leaves may have been faster in elevated than in ambient [CO₂]. Our results indicate that elevated [CO₂] may overall have a limited effect on isoprene emission from young seedlings and that plants generally regulate the emission to reach the maximum at the centre of the leaf profile, irrespective of the total leaf number. In comparison with leaf expansion and photosynthesis, isoprene showed marked and repeatable differences among leaves of the profile and may therefore be a useful trait to accurately monitor changes of leaf ontogeny as a consequence of elevated [CO₂].     

Modelling the drought impact on monoterpene fluxes from an evergreen Mediterranean forest canopy

In many ecosystems drought cycles are common during the growing season but their impact on volatile monoterpene emissions is unclear. Therefore, we aimed to develop and evaluate a process-based modelling approach to explore the explanatory power of likely mechanisms. The biochemically based isoprene and monoterpene emission model SIM-BIM2 has been modified and linked to a canopy model and a soil water balance model. Simulations are carried out for Quercus ilex forest sites and results are compared to measured soil water, photosynthesis, terpene-synthase activity, and monoterpene emission rates. Finally, the coupled model system is used to estimate the annual drought impact on photosynthesis and emission. The combined and adjusted vegetation model was able to simulate photosynthesis and monoterpene emission under dry and irrigated conditions with an R ² of 0.74 and 0.52, respectively. We estimated an annual reduction of monoterpene emission of 67% for the extended and severe drought period in 2006 in the investigated Mediterranean ecosystem. It is concluded that process-based ecosystem models can provide a useful tool to investigate the involved mechanisms and to quantify the importance of specific environmental constraints.     

Effect of water stress on monoterpene emissions from young potted holm oak (Quercus ilex L.) trees

We investigated the effects of a short period of water stress on monoterpene emissions from Quercus ilex, a common oak species of the Mediterranean vegetation and a strong emitter of monoterpenes. The experiment was carried out on two young saplings with a branch enclosure system under semi-controlled conditions. Under unstressed conditions, small qualitative (cis--ocimene, trans--ocimene, -caryophyllene and 1,8-cineol) and large quantitative (as much as 40% for the main compounds emitted) differences were observed between the two apparently similar trees. Nevertheless these differences did not affect the short-and long-term responses to temperature and water stress. Daily courses of emissions and gas exchanges were similar before and after the stress. During the most severe stress, emissions were reduced by a factor of two orders of magnitude and the log-linear relationship between emissions and temperature no longer existed. Photosynthesis and transpiration rates decreased as soon as the soil started to dry, whereas monoterpene emissions slightly increased for few days and then dropped when the daily CO₂ balance approached zero. We concluded that under water stress monoterpene emissions were highly limited by monoterpene synthesis resulting from a lack of carbon substrate and/or ATP. After rewatering, both emissions and gas exchanges recovered immediately, but to a level lower than the pre-stress level. These results have many implications for monoterpene emission modelling in the Mediterranean area, since the dry period generally extends from May to August. If our results are confirmed by field experiments, water stress could lead to a large overestimation of the emissions under summer conditions, when the algorithms based on light and temperature would give high emission rates.     

Effects of environmental conditions on isoprene emission from live oak

Live-oak plants (Quercus virginiana Mill.) were subjected to various levels of CO₂, water stress or photosynthetic photon flux density to test the hypothesis that isoprene biosynthesis occurred only under conditions of restricted CO₂ availability. Isoprene emission increases as the ambient CO₂ concentration decreased, independent of the amount of time that plants had photosynthesized at ambient CO₂ levels. When plants were water-stressed over a 4-d period photosynthesis and leaf conductance decreased 98 and 94%, respectively, while isoprene emissions remained constant. Significant isoprene emissions occurred when plants were saturated with CO₂, i.e., below the light compensation level for net photosynthesis (100 mol m^-2 s^-1). Isoprene emission rates increased with photosynthetic photon flux density and at 25 and 50 mol m^-2 s^-1 were 7 and 18 times greater than emissions in the dark. These data indicate that isoprene is a normal plant metabolite and not — as has been suggested — formed exclusively in response to restricted CO₂ or various stresses.Abbreviation PPFD photosynthetic photon flux density     

Impact of global warming on the tree species composition of boreal forests in Finland and effects on emissions of isoprenoids

This study aims to identify how climate change may influence total emissions of monoterpene and isoprene from boreal forest canopies. The whole of Finland is assumed to experience an annual mean temperature (T) increase of 4 °C and a precipitation increase of 10% by the year 2100. This will increase forest resources throughout the country. At the same time, the proportions of Scots pine (Pinus sylvestris) and Norway spruce (Picea abies) in southern Finland (60°≤ latitude < 65°N) will be reduced from the current 40–50% to less than 10–20%, with increased dominance of birches (Betula pendula and Betula pubescens). In northern Finland (65°≤ latitude < 70°N), the proportions of Norway spruce and Scots pine will be balanced at a level of about 40% as the result of an increase in Norway spruce from the current 21% to 37% and a concurrent reduction in Scots pine from 63% to 40%. The proportion of birches is predicted to increase from the current 17% to 23%, but these will become the dominant species only on the most fertile sites. Total mean emissions of monoterpene by Scots pine will be reduced by 80% in southern Finland, but will increase by 62% in the north. Emissions from Norway spruce canopies will increase by 4% in the south but by 428% in the north, while those from birch canopies will increase by about 300% and 113%, respectively. Overall emissions of monoterpene over the whole country amount to about 950 kg km^?2 y^?1 under current temperature conditions and will increase by 17% to 1100 kg km^?2 y^?1 with elevated temperature and precipitation, mainly because of an increase at northern latitudes. Under current conditions, emissions of isoprene follow the spatial distribution of spruce canopies (the only isoprene‐emitting tree species that forms forests in Finland) with four times higher emissions in the south than in the north. The elevated temperature and the changes in the areal distribution of Norway spruce will result in increases in isoprene emissions of about 37% in southern Finland and 435% in northern Finland. Annual mean isoprene emissions from Norway spruce canopies over the whole country will increase by about 60% up to the year 2100.     

Isoprene emissions from plants are mediated by atmospheric CO2 concentrations

The tropical African tree species Acacia nigrescens Oliv. was grown in environmentally controlled growth chambers at three CO₂ concentrations representative of the Last Glacial Maximum (～180 ppmv), the present day (～380 ppmv), and likely mid‐21st century (～600 ppmv) CO₂ concentrations. Isoprene (C₅H₈) emissions, per unit leaf area, were greater at lower‐than‐current CO₂ levels and lower at higher‐than‐current CO₂ levels relative to controls grown at 380 ppmv CO₂. Changes in substrate availability and isoprene synthase (IspS) activity were identified as the mechanisms behind the observed leaf‐level emission response. In contrast, canopy‐scale emissions remained unaltered between the treatments as changes in leaf‐level emissions were offset by changes in biomass and leaf area. Substrate concentration and IspS activity‐CO₂ responses were used in a biochemical model, coupled to existing isoprene emission algorithms, to model isoprene emissions from A. nigrescens grown for over 2 years at three different CO₂ concentrations. The addition of the biochemical model allowed for the use of emission factors measured under present day CO₂ concentrations across all three CO₂ treatments. When isoprene emissions were measured from A. nigrescens in response to instantaneous changes in CO₂ concentration, the biochemical model satisfactorily represented the observed response. Therefore, the effect of changes in atmospheric CO₂ concentration on isoprene emission at any timescale can be modelled and predicted.     

Coupling isoprene and monoterpene emissions from Amazonian tree species with physiological and environmental parameters using a neural network approach

The ability to predict isoprene emissions from plants is important for predicting atmospheric chemistry. To improve the basis for prediction capability, data obtained from continuous field measurements of isoprene and monoterpene emissions from three Amazonian tree species were related to observed environmental and leaf physiological parameters using a new neural network approach. The environmental parameters included leaf temperature, light, relative humidity, water vapour pressure deficit, and the history of ambient temperature and ozone concentration, whereas the physiological parameters included stomatal conductance, assimilation and intercellular CO₂ concentration. The neural approach with 24 different combinations of these parameters was applied to predict the emission variability observed during short time periods (2–3 d) with individual tree branches and, on a longer-term scale, in aggregated data sets from different seasons, leaf developmental stage, and light environment. The results were compared to the quasi standard emission algorithm for isoprene. On the short-term scale, good agreement (r²≈ 0.9) was obtained between observations and predictions of the standard algorithm as well as predictions of the neural network using the same input parameters (leaf temperature and light). When these predictors were used to model the long-term emission variability, r² was reduced to < 0.5 for both approaches. Remarkably, for the neural technique, more than 50% of the unexplained variance could be explained by the mean temperature of the preceding 36 h. An even better network performance was obtained with physiological parameter combinations (r² > 0.9) suggesting a strong and applicable link between isoprenoid emission and leaf primary metabolism.     

Response of isoprene emission to ambient CO2 changes and implications for global budgets

We explore the potential role of atmospheric carbon dioxide (CO₂) on isoprene emissions using a global coupled land–atmosphere model [Community Atmospheric Model–Community Land Model (CAM–CLM)] for recent (year 2000, 365 ppm CO₂) and future (year 2100, 717 ppm CO₂) conditions. We incorporate an empirical model of observed isoprene emissions response to both ambient CO₂ concentrations in the long‐term growth environment and short‐term changes in intercellular CO₂ concentrations into the MEGAN biogenic emission model embedded within the CLM. Accounting for CO₂ inhibition has little impact on predictions of present‐day global isoprene emission (increase from 508 to 523 Tg C yr^?1). However, the large increases in future isoprene emissions typically predicted in models, which are due to a projected warmer climate, are entirely offset by including the CO₂ effects. Projected global isoprene emissions in 2100 drop from 696 to 479 Tg C yr^?1 when this effect is included, maintaining future isoprene sources at levels similar to present day. The isoprene emission response to CO₂ is dominated by the long‐term growth environment effect, with modulations of 10% or less due to the variability in intercellular CO₂ concentration. As a result, perturbations to isoprene emissions associated with changes in ambient CO₂ are largely aseasonal, with little diurnal variability. Future isoprene emissions increase by more than a factor of two in 2100 (to 1242 Tg C yr^?1) when projected changes in vegetation distribution and leaf area density are included. Changing land cover and the role of nutrient limitation on CO₂ fertilization therefore remain the largest source of uncertainty in isoprene emission prediction. Although future projections suggest a compensatory balance between the effects of temperature and CO₂ on isoprene emission, the enhancement of isoprene emission due to lower ambient CO₂ concentrations did not compensate for the effect of cooler temperatures over the last 400 thousand years of the geologic record (including the Last Glacial Maximum).     

Monoterpene ‘thermometer’ of tropical forest‐atmosphere response to climate warming

Tropical forests absorb large amounts of atmospheric CO₂ through photosynthesis but elevated temperatures suppress this absorption and promote monoterpene emissions. Using ¹³CO₂ labeling, here we show that monoterpene emissions from tropical leaves derive from recent photosynthesis and demonstrate distinct temperature optima for five groups (Groups 1–5), potentially corresponding to different enzymatic temperature‐dependent reaction mechanisms within β‐ocimene synthases. As diurnal and seasonal leaf temperatures increased during the Amazonian 2015 El Niño event, leaf and landscape monoterpene emissions showed strong linear enrichments of β‐ocimenes (+4.4% °C^?1) at the expense of other monoterpene isomers. The observed inverse temperature response of α‐pinene (?0.8% °C^?1), typically assumed to be the dominant monoterpene with moderate reactivity, was not accurately simulated by current global emission models. Given that β‐ocimenes are highly reactive with respect to both atmospheric and biological oxidants, the results suggest that highly reactive β‐ocimenes may play important roles in the thermotolerance of photosynthesis by functioning as effective antioxidants within plants and as efficient atmospheric precursors of secondary organic aerosols. Thus, monoterpene composition may represent a new sensitive ‘thermometer’ of leaf oxidative stress and atmospheric reactivity, and therefore a new tool in future studies of warming impacts on tropical biosphere‐atmosphere carbon‐cycle feedbacks.     

Monoterpene emission from coniferous trees in response to elevated CO2 concentration and climate warming

It was hypothesized that high CO₂ availability would increase monoterpene emission to the atmosphere. This hypothesis was based on resource allocation theory which predicts increased production of plant secondary compounds when carbon is in excess of that required for growth. Monoterpene emission rates were measured from needles of (a) Ponderosa pine grown at different CO₂ concentrations and soil nitrogen levels, and (b) Douglas fir grown at different CO₂ concentrations. Ponderosa pine grown at 700 μmol mol^–1 CO₂ exhibited increased photosynthetic rates and needle starch to nitrogen (N) ratios when compared to trees grown at 350 μmol mol^–1 CO₂. Nitrogen availability had no consistent effect on photosynthesis. Douglas fir grown at 550 μmol mol^–1 CO₂ exhibited increased photosynthetic rates as compared to growth at 350 μmol mol^–1 CO₂ in old, but not young needles, and there was no influence on the starch/N ratio. In neither species was there a significant effect of elevated growth CO₂ on needle monoterpene concentration or emission rate. The influence of climate warming and leaf area index (LAI) on monoterpene emission were also investigated. Douglas fir grown at elevated CO₂ plus a 4 °C increase in growth temperature exhibited no change in needle monoterpene concentration, despite a predicted 50% increase in emission rate. At elevated CO₂ concentration the LAI increased in Ponderosa pine, but not Douglas fir. The combination of increased LAI and climate warming are predicted to cause an 80% increase in monoterpene emissions from Ponderosa pine forests and a 50% increase in emissions from Douglas fir forests. This study demonstrates that although growth at elevated CO₂ may not affect the rate of monoterpene emission per unit biomass, the effect of elevated CO₂ on LAI, and the effect of climate warming on monoterpene biosynthesis and volatilization, could increase canopy monoterpene emission rate.     

The effects of glacial atmospheric CO2 concentrations and climate on isoprene emissions by vascular plants

Isoprene (C₅H₈) emissions by terrestrial vegetation vary with temperature and light intensity, and play an important role in biosphere–chemistry–climate interactions. Such interactions were probably substantially modified by Pleistocene climate and CO₂ cycles. Central to understanding the nature of these modifications is assessment and analysis of how emissions changed under glacial environmental conditions. Currently, even the net direction of change is difficult to predict because a CO₂‐depleted atmosphere may have stimulated emissions compensating for the negative impacts of a cooler climate. Here, we address this issue and attempt to determine the direction of change from an experimental standpoint by investigating the interaction between isoprene emissions and plant growth of two known isoprene‐emitting herbaceous species (Mucuna pruriens and Arundo donax) grown at glacial (180 ppm) to present (366 ppm) CO₂ levels. We found a significant enhancement of isoprene emissions per unit leaf area in M. pruriens under subambient CO₂ concentrations relative to ambient controls but not for A. donax. In contrast, canopy emissions remained unaltered for both plant species because enhanced leaf emissions were offset by reductions in biomass and leaf area. Separate growth experiments with M. pruriens revealed that lowering day/night temperatures by 5°C decreased canopy isoprene emissions irrespective of the CO₂ level. Incorporation of these results into a simple canopy emissions model highlights their potential to attenuate reductions in the total isoprene flux from forests under glacial conditions predicted by standard models.     

Why only some plants emit isoprene

Isoprene (2‐methyl‐1,3‐butadiene) is emitted from many plants and it appears to have an adaptive role in protecting leaves from abiotic stress. However, only some species emit isoprene. Isoprene emission has appeared and been lost many times independently during the evolution of plants. As an example, our phylogenetic analysis shows that isoprene emission is likely ancestral within the family Fabaceae (= Leguminosae), but that it has been lost at least 16 times and secondarily gained at least 10 times through independent evolutionary events. Within the division Pteridophyta (ferns), we conservatively estimate that isoprene emissions have been gained five times and lost two times through independent evolutionary events. Within the genus Quercus (oaks), isoprene emissions have been lost from one clade, but replaced by a novel type of light‐dependent monoterpene emissions that uses the same metabolic pathways and substrates as isoprene emissions. This novel type of monoterpene emissions has appeared at least twice independently within Quercus, and has been lost from 9% of the individuals within a single population of Quercus suber. Gain and loss of gene function for isoprene synthase is possible through relatively few mutations. Thus, this trait appears frequently in lineages; but, once it appears, the time available for evolutionary radiation into environments that select for the trait is short relative to the time required for mutations capable of producing a non‐functional isoprene synthase gene. The high frequency of gains and losses of the trait and its heterogeneous taxonomic distribution in plants may be explained by the relatively few mutations necessary to produce or lose the isoprene synthase gene combined with the assumption that isoprene emission is advantageous in a narrow range of environments and phenotypes.     

Strong correlation between isoprene emission and gross photosynthetic capacity during leaf phenology of the tropical tree species Hymenaea courbaril with fundamental changes in volatile organic compounds emission composition during early leaf development

Changes of the volatile organic compounds (VOC) emission capacity and composition of different developmental stages of the tropical tree species Hymenaea courbaril were investigated under field conditions at a remote Amazonian rainforest site. The basal emission capacity of isoprene changed considerably over the course of leaf development, from young to mature and to senescent leaves, ultimately spanning a wide range of observed isoprene basal emission capacities from 0.7 to 111.5 µg C g^?1 h^?1 during the course of the year. By adjusting the standard emission factors for individual days, the diel courses of instantaneous isoprene emission rates could nevertheless adequately be modelled by a current isoprene algorithm. The results demonstrate the inadequacy of using one single standard emission factor to represent the VOC emission capacity of tropical vegetation for an entire seasonal cycle. A strong linear correlation between the isoprene emission capacity and the gross photosynthetic capacity (GP_max) covering all developmental stages and seasons was observed. The present results provide evidence that leaf photosynthetic properties may confer a valuable basis to model the seasonal variation of isoprenoid emission capacity; especially in tropical regions where the environmental conditions vary less than in temperate regions. In addition to induction and variability of isoprene emission during early leaf development, considerable amounts of monoterpenes were emitted in a light‐dependent manner exclusively in the period between bud break and leaf maturity. The fundamental change in emission composition during this stage as a consequence of resource availability (supply side control) or as a plant's response to the higher defence demand of young emerging leaves (demand‐side control) is discussed. The finding of a temporary emergence of monoterpene emission may be of general interest in understanding both the ecological functions of isoprenoid production and the regulatory processes involved.     

Isoprenoid emissions,photosynthesis and mesophyll diffusion conductance in response to blue light

The effects of blue light (BL) on leaf gas exchange of Populus × canadensis, a strong isoprene emitter, and Quercus ilex and Citrus reticulata, two monoterpene emitters with respectively small and large storage pools for monoterpenes, were studied. Leaves were initially exposed to a saturating photosynthetic photon flux density (PPFD) of white light (WL), which was then progressively reduced to perform WL-response curves. Leaves acclimated to saturating WL were then quickly exposed to equivalent BL levels to perform BL-response curves. Blue light did not significantly affect photosynthetic parameters in the light-limited portion of the PPFD-response curves in both P. × canadensis and Q. ilex. Whereas photosynthesis (A), stomatal conductance (g_s), and mesophyll conductance (g_m) were significantly decreased at high PPFDs of BL. A was similarly inhibited by BL in C. reticulata, but there was no significant effect of light quality on g_s. Overall these results show that the negative effect of BL on photosynthesis is widespread in tree species with different leaf characteristics, and that this involves coordinated reductions in g_s and g_m. BL negatively affected isoprene emission and, to a lesser extent monoterpene emissions, in concert with photosynthetic inhibition. Interesting, both isoprene and monoterpene emissions were shown to be inversely dependent upon intercellular [CO₂]. These results indicate that a change in light spectral quality, which can vary during the day, between days and within seasons, can alter photosynthesis and isoprenoid emissions, depending on the PPFD intensity. Such effects should be strongly considered in photosynthesis and volatile isoprenoid emission models.