Range size and growth temperature influence Eucalyptus species responses to an experimental heatwave

Understanding forest tree responses to climate warming and heatwaves is important for predicting changes in tree species diversity, forest C uptake, and vegetation–climate interactions. Yet, tree species differences in heatwave tolerance and their plasticity to growth temperature remain poorly understood. In this study, populations of four Eucalyptus species, two with large range sizes and two with comparatively small range sizes, were grown under two temperature treatments (cool and warm) before being exposed to an equivalent experimental heatwave. We tested whether the species with large and small range sizes differed in heatwave tolerance, and whether trees grown under warmer temperatures were more tolerant of heatwave conditions than trees grown under cooler temperatures. Visible heatwave damage was more common and severe in the species with small rather than large range sizes. In general, species that showed less tissue damage maintained higher stomatal conductance, lower leaf temperatures, larger increases in isoprene emissions, and less photosynthetic inhibition than species that showed more damage. Species exhibiting more severe visible damage had larger increases in heat shock proteins (HSPs) and respiratory thermotolerance (T_max). Thus, across species, increases in HSPs and T_max were positively correlated, but inversely related to increases in isoprene emissions. Integration of leaf gas‐exchange, isoprene emissions, proteomics, and respiratory thermotolerance measurements provided new insight into mechanisms underlying variability in tree species heatwave tolerance. Importantly, warm‐grown seedlings were, surprisingly, more susceptible to heatwave damage than cool‐grown seedlings, which could be associated with reduced enzyme concentrations in leaves. We conclude that species with restricted range sizes, along with trees growing under climate warming, may be more vulnerable to heatwaves of the future.     

Drought effect on isoprene production and consumption in Biosphere 2 tropical rainforest

Isoprene is the most abundant of the hydrocarbon compounds emitted from vegetation and plays a major role in tropospheric chemistry. Models predict that future climate change scenarios may lead to an increase in global isoprene emissions as a consequence of higher temperatures and extended drought periods. Tropical rainforests are responsible for more than 80% of global isoprene emissions, so it is important to obtain experimental data on isoprene production and consumption in these ecosystems under control of environmental variables. We explored isoprene emission and consumption in the tropical rainforest model ecosystem of Biosphere 2 laboratory during a mild water stress, and the relationship with light and temperature. Gross isoprene production (GIP) was not significantly affected by mild water stress in this experiment because the isoprene emitters were mainly distributed among the large, canopy layer trees with deep roots in the lower soil profile where water content decreased much less than the top 30 cm. However, as found in previous leaf level and whole canopy studies, the ecosystem gross primary production was reduced by (32%) during drought, and as a consequence the percentage of fixed C lost as isoprene tended to increase during drought, from ca. 1% in wet conditions to ca. 2% when soil water content reached its minimum. GIP correlated very well with both light and temperature. Notably, soil isoprene uptake decreased dramatically during the drought, leading to a large increase in daytime net isoprene fluxes.     

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.     

Within‐plant isoprene oxidation confirmed by direct emissions of oxidation products methyl vinyl ketone and methacrolein

Isoprene is emitted from many terrestrial plants at high rates, accounting for an estimated 1/3 of annual global volatile organic compound emissions from all anthropogenic and biogenic sources combined. Through rapid photooxidation reactions in the atmosphere, isoprene is converted to a variety of oxidized hydrocarbons, providing higher order reactants for the production of organic nitrates and tropospheric ozone, reducing the availability of oxidants for the breakdown of radiatively active trace gases such as methane, and potentially producing hygroscopic particles that act as effective cloud condensation nuclei. However, the functional basis for plant production of isoprene remains elusive. It has been hypothesized that in the cell isoprene mitigates oxidative damage during the stress‐induced accumulation of reactive oxygen species (ROS), but the products of isoprene‐ROS reactions in plants have not been detected. Using pyruvate‐2‐¹³C leaf and branch feeding and individual branch and whole mesocosm flux studies, we present evidence that isoprene (i) is oxidized to methyl vinyl ketone and methacrolein (i_ox) in leaves and that i_ox/i emission ratios increase with temperature, possibly due to an increase in ROS production under high temperature and light stress. In a primary rainforest in Amazonia, we inferred significant in plant isoprene oxidation (despite the strong masking effect of simultaneous atmospheric oxidation), from its influence on the vertical distribution of i_ox uptake fluxes, which were shifted to low isoprene emitting regions of the canopy. These observations suggest that carbon investment in isoprene production is larger than that inferred from emissions alone and that models of tropospheric chemistry and biota–chemistry–climate interactions should incorporate isoprene oxidation within both the biosphere and the atmosphere with potential implications for better understanding both the oxidizing power of the troposphere and forest response to climate change.     

Variation in potential for isoprene emissions among Neotropical forest sites

As part of the Large Scale Biosphere–Atmosphere Experiment in Amazônia (LBA), we have developed a bottom‐up approach for estimating canopy‐scale fluxes of isoprene. Estimating isoprene fluxes for a given forest ecosystem requires knowledge of foliar biomass, segregated by species, and the isoprene emission characteristics of the individual tree species comprising the forest. In this study, approximately 38% of 125 tree species examined at six sites in the Brazilian Amazon emitted isoprene. Given logistical difficulties and extremely high species diversity, it was possible to screen only a small percentage of tree species, and we propose a protocol for estimating the emission capacity of unmeasured taxa using a taxonomic approach, in which we assign to an unmeasured genus a value based on the percentage of genera within its plant family which have been shown to emit isoprene. Combining this information with data obtained from 14 tree censuses at four Neotropical forest sites, we have estimated the percentage of isoprene‐emitting biomass at each site. The relative contribution of each genus of tree is estimated as the basal area of all trees of that genus divided by the total basal area of the plot. Using this technique, the percentage of isoprene‐emitting biomass varied from 20% to 42% (mean=31%; SD=8%). Responses of isoprene emission to varying light and temperature, measured on a sun‐adapted leaf of mango (Mangifera indica L.), suggest that existing algorithms developed for temperate species are adequate for tropical species as well. Incorporating these algorithms, estimates of isoprene‐emitting biomass, isoprene emission capacity, and site foliar biomass into a canopy flux model, canopy‐scale fluxes of isoprene were predicted and compared with the above‐canopy fluxes measured at two sites. Our bottom‐up approach overestimates fluxes by about 50%, but variations in measured fluxes between the two sites are largely explained by observed variation in the amount of isoprene‐emitting biomass.     

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.     

Seasonal differences in isoprene and light-dependent monoterpene emission by Amazonian tree species

Whereas for extra‐tropical regions model estimates of the emission of volatile organic compounds (VOC) predict strong responses to the strong annual cycles of foliar biomass, light intensity and temperature, the tropical regions stand out as a dominant source year round, with only little variability mainly due to the annual cycle of foliar biomass of drought‐deciduous trees. As part of the Large Scale Biosphere Atmosphere Experiment in Amazônia (LBA‐EUSTACH), a remote secondary tropical forest site was visited in the dry‐to‐wet season transition campaign, and the trace gas exchange of a strong isoprene emitter and a monoterpene emitter are compared to the wet‐to‐dry season transition investigations reported earlier. Strong seasonal differences of the emission capacity were observed. The standard emission factor for isoprene emission of young mature leaves of Hymenaea courbaril was about twofold in the end of the dry season (111.5 μgC g^?1 h^?1 or 41.2 nmol m^?2 s^?1) compared to old mature leaves investigated in the end of the wet season (45.4 μgC g^?1 h^?1 or 24.9 nmol m^?2 s^?1). Standardized monoterpene emission rate of Apeiba tibourbou were 2.1 and 3.6 μgC g^?1 h^?1 (or 0.3 and 0.8 nmol m^?2 s^‐1), respectively. This change in species‐specific VOC emission capacity was mirrored by a concurrent change in the ambient mixing ratios. The growth conditions vary less in tropical areas than in temperate regions of the world, and the seasonal differences in emission strength could not be reconciled solely with meteorological data of instantaneous light intensity and temperature. Hence the inadequacy of using a single standard emission factor to represent an entire seasonal cycle is apparent. Among a host of other potential factors, including the leaf developmental stage, water and nutrient status, and abiotic stresses like the oxidative capacity of the ambient air, predominantly the long‐term growth temperature may be applied to predict the seasonal variability of the isoprene emission capacity. The dry season isoprene emission rates of H. courbaril measured at the canopy top were also compared to isoprene emissions of the shade‐adapted species Sorocea guilleminiana growing in the understory. Despite the difference in VOC emission composition and canopy position, one common algorithm was able to predict the diel emission pattern of all three tree species.     

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).     

Ecosystem‐scale volatile organic compound fluxes during an extreme drought in a broadleaf temperate forest of the Missouri Ozarks (central USA)

Considerable amounts and varieties of biogenic volatile organic compounds (BVOCs) are exchanged between vegetation and the surrounding air. These BVOCs play key ecological and atmospheric roles that must be adequately represented for accurately modeling the coupled biosphere–atmosphere–climate earth system. One key uncertainty in existing models is the response of BVOC fluxes to an important global change process: drought. We describe the diurnal and seasonal variation in isoprene, monoterpene, and methanol fluxes from a temperate forest ecosystem before, during, and after an extreme 2012 drought event in the Ozark region of the central USA. BVOC fluxes were dominated by isoprene, which attained high emission rates of up to 35.4 mg m^?2 h^?1 at midday. Methanol fluxes were characterized by net deposition in the morning, changing to a net emission flux through the rest of the daylight hours. Net flux of CO₂ reached its seasonal maximum approximately a month earlier than isoprenoid fluxes, which highlights the differential response of photosynthesis and isoprenoid emissions to progressing drought conditions. Nevertheless, both processes were strongly suppressed under extreme drought, although isoprene fluxes remained relatively high compared to reported fluxes from other ecosystems. Methanol exchange was less affected by drought throughout the season, confirming the complex processes driving biogenic methanol fluxes. The fraction of daytime (7–17 h) assimilated carbon released back to the atmosphere combining the three BVOCs measured was 2% of gross primary productivity (GPP) and 4.9% of net ecosystem exchange (NEE) on average for our whole measurement campaign, while exceeding 5% of GPP and 10% of NEE just before the strongest drought phase. The megan v2.1 model correctly predicted diurnal variations in fluxes driven mainly by light and temperature, although further research is needed to address model BVOC fluxes during drought events.     

The regulation of isoprene emission responses to rapid leaf temperature fluctuations

Isoprene emission from leaves is temperature dependent and may protect leaves from damage at high temperatures. We measured the temperature of white oak ( Quercus alba L.) leaves at the top of the canopy. The largest short-term changes in leaf temperature were associated with changes in solar radiation. During these episodes, leaf temperature changed with a 1 min time constant, a measure of the rate of temperature change. We imposed rapid temperature fluctuations on leaves to study the effect of temperature change rate on isoprene emission. Leaf temperature changed with a 16 s time constant; isoprene responded more slowly with a 37 s time constant. This time constant was slow enough to cause a lag in isoprene emission when leaf temperature fluctuated rapidly but isoprene emission changed quickly enough to follow the large temperature changes observed in the oak canopy. This is consistent with the theory that isoprene functions to protect leaves from short periods of high temperature. Time constant analysis also revealed that there are two processes that cause isoprene emission to increase with leaf temperature. The fastest process likely reflects the influence of temperature on reaction kinetics, while the slower process may reflect the activation of an enzyme.     

The Influence of Light and Temperature on Isoprene Emission Rates from Live Oak

There is a growing awareness of the role of vegetation as a source of reactive hydrocarbons that may serve as photochemical oxidant precursors. A study was designed to assess independently the influence of variable light and temperature on isoprene emissions from live oak (Quercus virginiana Mill.). Plants were conditioned in a growth chamber and then transferred to an environmentally controlled gas-exchange chamber. Samples of the chamber atmosphere were collected; isoprene was concentrated cryogenically and measured by gas chromatography. A logistic function was used to model isoprene emission rates. Under regimes of low temperature (20°C) or darkness, isoprene emissions were lowest. With increasing temperature or light intensity, the rate of isoprene emission increased, reaching maxima at 800 μE m^-2 s^-1 and 40–44°C, respectively. Higher temperatures caused a large decrease in emissions. Since the emissions of isoprene were light-saturated at moderate intensities, temperature appeared to be the main factor controlling emissions during most of the day. Carbon lost through isoprene emissions accounted for 0.1 to 2% of the carbon fixed during photosynthesis depending on light intensity and temperature.     

Process-based modelling of isoprene emission by oak leaves

The emission rate of the volatile reactive compound isoprene, emitted predominantly by trees, must be known before the level of photo‐oxidants produced during summer smog can be predicted reliably. The emission is dependent on plant species and local conditions, and these dependencies must be quantified to be included in any empirical algorithm for the calculation of isoprene production. Experimental measurements of isoprene emission rates are expensive, however, and existing data are scarce and fragmentary. To overcome these difficulties, it is promising to develop a numerical model capable of precisely calculating the isoprene emission by trees for diverse ecosystems, even under changing environmental conditions. A basic process‐based biochemical isoprene emission model (BIM) has therefore been developed, which describes the enzymatic reactions in leaf chloroplasts leading to the formation of isoprene under varying environmental conditions (e.g. light intensity, temperature). Concentrations of the precursors of isoprene formation, 3‐phosphoglyceric acid and glyceraldehyde 3‐phosphate, are provided by a published light fleck photosynthesis model. Specific leaf and enzyme parameters were determined for the pedunculate oak (Quercus robur L.), so that the BIM is capable of calculating oak‐specific isoprene emission rates as influenced by the leaf temperature and light intensity. High correlation was observed between isoprene emission rates calculated by the BIM and the diurnal isoprene emission rates of leaves measured under controlled environmental conditions. The BIM was even capable of describing changes in isoprene emission caused by midday depression of net photosynthesis.     

Isoprenoid emission in trees of Quercus pubescens and Quercus ilex with lifetime exposure to naturally high CO2 environment†

The long‐term effect of elevated atmospheric CO₂ on isoprenoid emissions from adult trees of two Mediterranean oak species (the monoterpene‐emitting Quercus ilex L. and the isoprene‐emitting Quercus pubescens Willd.) native to a high‐CO₂ environment was investigated. During two consecutive years, isoprenoid emission was monitored both at branch level, measuring the actual emissions under natural conditions, and at leaf level, measuring the basal emissions under the standard conditions of 30 °C and at light intensity of 1000 µmol m^?2 s^?1. Long‐term exposure to high atmospheric levels of CO₂ did not significantly affect the actual isoprenoid emissions. However, when leaves of plants grown in the control site were exposed for a short period to an elevated CO₂ level by rapidly switching the CO₂ concentration in the gas‐exchange cuvette, both isoprene and monoterpene basal emissions were clearly inhibited. These results generally confirm the inhibitory effect of elevated CO₂ on isoprenoid emission. The absence of a CO₂ effect on actual emissions might indicate higher leaf temperature at elevated CO₂, or an interaction with multiple stresses some of which (e.g. recurrent droughts) may compensate for the CO₂ effect in Mediterranean ecosystems. Under elevated CO₂, isoprene emission by Q. pubescens was also uncoupled from the previous day's air temperature. In addition, pronounced daily and seasonal variations of basal emission were observed under elevated CO₂ underlining that correction factors may be necessary to improve the realistic estimation of isoprene emissions with empirical algorithms in the future. A positive linear correlation of isoprenoid emission with the photosynthetic electron transport and in particular with its calculated fraction used for isoprenoid synthesis was found. The slope of this relationship was different for isoprene and monoterpenes, but did not change when plants were grown in either ambient or elevated CO₂. This suggests that physiological algorithms may usefully predict isoprenoid emission also under rising CO₂ levels.     

Comparison of Isoprene Emission, Intercellular Isoprene Concentration and Photosynthetic Performance in Water-Limited Oak (Quercus pubescens Willd. and Quercus robur L.) Saplings

Abstract: The influence of prolonged water limitation on leaf gas exchange, isoprene emission, isoprene synthase activities and intercellular isoprene concentrations was investigated under standard conditions (30 °C leaf temperature and 1000 μmol photons m^-2 s^-1 PPFD) in greenhouse experiments with five-year-old pubescent oak ( Quercus pubescens Willd.) and four-year-old pedunculate oak ( Quercus robur L.) saplings. Net assimilation rates proved to be highly sensitive to moderate drought in both oak species, and were virtually zero at water potentials (Ψ_pd) below - 1.3 MPa in Q. robur and below - 2.5 MPa in Q. pubescens . The response of stomatal conductance to water stress was slightly less distinct. Isoprene emission was much more resistant to drought and declined significantly only at Ψ_pd below - 2 MPa in Q. robur and below - 3.5 MPa in Q. pubescens . Even during the most severe water stress, isoprene emission of drought-stressed saplings was still approximately one-third of the control in Q. robur and one-fifth in Q. pubescens . Isoprene synthase activities were virtually unaffected by drought stress. Re-watering led to partial recovery of leaf gas exchange and isoprene emission. Intercellular isoprene concentrations were remarkably enhanced in water-limited saplings of both oak species during the first half of the respective drought periods with maximum mean values up to ca. 16 μl l^-1 isoprene for Q. pubescens and ca. 11 μl l^-1 isoprene for pedunculate oak, supporting the hypothesis that isoprene serves as a short-term thermoprotective agent in isoprene-emitting plant species.     

Circadian control of isoprene emissions from oil palm (Elaeis guineensis)

The emission of isoprene from the biosphere to the atmosphere has a profound effect on the Earth's atmospheric system. Until now, it has been assumed that the primary short-term controls on isoprene emission are photosynthetically active radiation and temperature. Here we show that isoprene emissions from a tropical tree (oil palm, Elaeis guineensis) are under strong circadian control, and that the circadian clock is potentially able to gate light-induced isoprene emissions. These rhythms are robustly temperature compensated with isoprene emissions still under circadian control at 38 degrees C. This is well beyond the acknowledged temperature range of all previously described circadian phenomena in plants. Furthermore, rhythmic expression of LHY/CCA1, a genetic component of the central clock in Arabidopsis thaliana, is still maintained at these elevated temperatures in oil palm. Maintenance of the CCA1/LHY-TOC1 molecular oscillator at these temperatures in oil palm allows for the possibility that this system is involved in the control of isoprene emission rhythms. This study contradicts the accepted theory that isoprene emissions are primarily light-induced.     

Effects of heat and drought stress on post‐illumination bursts of volatile organic compounds in isoprene‐emitting and non‐emitting poplar

Over the last decades, post‐illumination bursts (PIBs) of isoprene, acetaldehyde and green leaf volatiles (GLVs) following rapid light‐to‐dark transitions have been reported for a variety of different plant species. However, the mechanisms triggering their release still remain unclear. Here we measured PIBs of isoprene‐emitting (IE) and isoprene non‐emitting (NE) grey poplar plants grown under different climate scenarios (ambient control and three scenarios with elevated CO₂ concentrations: elevated control, periodic heat and temperature stress, chronic heat and temperature stress, followed by recovery periods). PIBs of isoprene were unaffected by elevated CO₂ and heat and drought stress in IE, while they were absent in NE plants. On the other hand, PIBs of acetaldehyde and also GLVs were strongly reduced in stress‐affected plants of all genotypes. After recovery from stress, distinct differences in PIB emissions in both genotypes confirmed different precursor pools for acetaldehyde and GLV emissions. Changes in PIBs of GLVs, almost absent in stressed plants and enhanced after recovery, could be mainly attributed to changes in lipoxygenase activity. Our results indicate that acetaldehyde PIBs, which recovered only partly, derive from a new mechanism in which acetaldehyde is produced from methylerythritol phosphate pathway intermediates, driven by deoxyxylulose phosphate synthase activity.     

Stimulation of isoprene emissions and electron transport rates as key mechanisms of thermal tolerance in the tropical species Vismia guianensis

Tropical forests absorb large amounts of atmospheric CO₂ through photosynthesis, but high surface temperatures suppress this absorption while promoting isoprene emissions. While mechanistic isoprene emission models predict a tight coupling to photosynthetic electron transport (ETR) as a function of temperature, direct field observations of this phenomenon are lacking in the tropics and are necessary to assess the impact of a warming climate on global isoprene emissions. Here we demonstrate that in the early successional species Vismia guianensis in the central Amazon, ETR rates increased with temperature in concert with isoprene emissions, even as stomatal conductance (g_s) and net photosynthetic carbon fixation (P_n) declined. We observed the highest temperatures of continually increasing isoprene emissions yet reported (50°C). While P_n showed an optimum value of 32.6 ± 0.4°C, isoprene emissions, ETR, and the oxidation state of PSII reaction centers (q_L) increased with leaf temperature with strong linear correlations for ETR (? = 0.98) and q_L (? = 0.99) with leaf isoprene emissions. In contrast, other photoprotective mechanisms, such as non‐photochemical quenching, were not activated at elevated temperatures. Inhibition of isoprenoid biosynthesis repressed P_n at high temperatures through a mechanism that was independent of stomatal closure. While extreme warming will decrease g_s and P_n in tropical species, our observations support a thermal tolerance mechanism where the maintenance of high photosynthetic capacity under extreme warming is assisted by the simultaneous stimulation of ETR and metabolic pathways that consume the direct products of ETR including photorespiration and the biosynthesis of thermoprotective isoprenoids. Our results confirm that models which link isoprene emissions to the rate of ETR hold true in tropical species and provide necessary “ground‐truthing” for simulations of the large predicted increases in tropical isoprene emissions with climate warming.     

Human-induced changes in US biogenic volatile organic compound emissions: evidence from long-term forest inventory data

Volatile organic compounds (VOCs) emitted by woody vegetation influence global climate forcing and the formation of tropospheric ozone. We use data from over 250 000 re‐surveyed forest plots in the eastern US to estimate emission rates for the two most important biogenic VOCs (isoprene and monoterpenes) in the 1980s and 1990s, and then compare these estimates to give a decadal change in emission rate. Over much of the region, particularly the southeast, we estimate that there were large changes in biogenic VOC emissions: half of the grid cells (1°× 1°) had decadal changes in emission rate outside the range ?2.3% to +16.8% for isoprene, and outside the range 0.2–17.1% for monoterpenes. For an average grid cell the estimated decadal change in heatwave biogenic VOC emissions (usually an increase) was three times greater than the decadal change in heatwave anthropogenic VOC emissions (usually a decrease, caused by legislation). Leaf‐area increases in forests, caused by anthropogenic disturbance, were the most important process increasing biogenic VOC emissions. However, in the southeast, which had the largest estimated changes, there were substantial effects of ecological succession (which decreased monoterpene emissions and had location‐specific effects on isoprene emissions), harvesting (which decreased monoterpene emissions and increased isoprene emissions) and plantation management (which increased isoprene emissions, and decreased monoterpene emissions in some states but increased monoterpene emissions in others). In any given region, changes in a very few tree species caused most of the changes in emissions: the rapid changes in the southeast were caused almost entirely by increases in sweetgum (Liquidambar styraciflua) and a few pine species. Therefore, in these regions, a more detailed ecological understanding of just a few species could greatly improve our understanding of the relationship between natural ecological processes, forest management, and biogenic VOC emissions.     

Controls of the quantum yield and saturation light of isoprene emission in different‐aged aspen leaves

Leaf age alters the balance between the use of end‐product of plastidic isoprenoid synthesis pathway, dimethylallyl diphosphate (DMADP), in prenyltransferase reactions leading to synthesis of pigments of photosynthetic machinery and in isoprene synthesis, but the implications of such changes on environmental responses of isoprene emission have not been studied. Because under light‐limited conditions, isoprene emission rate is controlled by DMADP pool size (S_DMADP), shifts in the share of different processes are expected to particularly strongly alter the light dependency of isoprene emission. We examined light responses of isoprene emission in young fully expanded, mature and old non‐senescent leaves of hybrid aspen (Populus tremula x P. tremuloides) and estimated in vivo S_DMADP and isoprene synthase activity from post‐illumination isoprene release. Isoprene emission capacity was 1.5‐fold larger in mature than in young and old leaves. The initial quantum yield of isoprene emission (α_I) increased by 2.5‐fold with increasing leaf age primarily as the result of increasing S_DMADP. The saturating light intensity (Q_I90) decreased by 2.3‐fold with increasing leaf age, and this mainly reflected limited light‐dependent increase of S_DMADP possibly due to feedback inhibition by DMADP. These major age‐dependent changes in the shape of the light response need consideration in modelling canopy isoprene emission.