Do cities simulate climate change? A comparison of herbivore response to urban and global warming

Cities experience elevated temperature, CO₂, and nitrogen deposition decades ahead of the global average, such that biological response to urbanization may predict response to future climate change. This hypothesis remains untested due to a lack of complementary urban and long‐term observations. Here, we examine the response of an herbivore, the scale insect Melanaspis tenebricosa, to temperature in the context of an urban heat island, a series of historical temperature fluctuations, and recent climate warming. We survey M. tenebricosa on 55 urban street trees in Raleigh, NC, 342 herbarium specimens collected in the rural southeastern United States from 1895 to 2011, and at 20 rural forest sites represented by both modern (2013) and historical samples. We relate scale insect abundance to August temperatures and find that M. tenebricosa is most common in the hottest parts of the city, on historical specimens collected during warm time periods, and in present‐day rural forests compared to the same sites when they were cooler. Scale insects reached their highest densities in the city, but abundance peaked at similar temperatures in urban and historical datasets and tracked temperature on a decadal scale. Although urban habitats are highly modified, species response to a key abiotic factor, temperature, was consistent across urban and rural‐forest ecosystems. Cities may be an appropriate but underused system for developing and testing hypotheses about biological effects of climate change. Future work should test the applicability of this model to other groups of organisms.     

CO2 exchange in an organic field growing barley or grass in eastern Finland

The CO₂ dynamics were measured in an organic soil in eastern Finland during the growing season and wintertime, and the annual CO₂ balance was calculated for plots where barley or grass was grown. During the summer, the CO₂ dynamics were measured by transparent and opaque chambers using a portable infrared gas analyser for the CO₂ analyses. During the winter, the CO₂ release was measured by opaque chambers analysing the samples in the laboratory with a gas chromatograph. Statistical response functions for CO₂ dynamics were constructed to evaluate the annual CO₂ exchange from the climatic data. The net CO₂ exchange was calculated for every hour in the snow‐free season. The carbon balance varied extensively depending on the weather conditions, and type and phenology of vegetation. During the growing season, the grassland was a net source while the barley field was a net sink for CO₂. However, both soils were net sources for CO₂ when autumn, winter and spring were included also. The annual CO₂ emissions from the grassland and barley soil were 750 g CO₂‐C m^?2 and 400 g CO₂‐C m^?2, respectively. The carbon accumulated in root and shoot biomass during the growing season was 330 g m^?2 for grass and 520 g m^?2 for barley. The C in the aboveground plant biomass ranged from 43 to 47% of the carbon fixed in photosynthesis (P_G) and the proportion of C in the root biomass was 10% of the carbon fixed in photosynthesis. The bare soils had 10–60% higher net CO₂ emission than the vegetated soils. These results indicate that the carbon balance of organic soils is affected by the characteristics of the prevailing plant cover. The dry summer of 1997 may have limited the growth of grass in the late summer thus reducing photosynthesis, which could be one reason for the high CO₂ release from this grass field.     

Future wood productivity of Pinus radiata in New Zealand under expected climatic changes

The physiologically based growth model CenW was used to simulate wood‐productivity responses of Pinus radiata forests to climate change in New Zealand. The model was tested under current climatic conditions against a comprehensive set of observations from growth plots located throughout the country. Climate change simulations were based on monthly climate change fields of 12 GCMs forced by the SRES B1, A1B and A2 emission scenarios for 2040 and 2090. Simulations used either constant or increasing CO₂ concentrations corresponding to the different emission scenarios. With constant CO₂, there were only slight growth responses to climate change across the country as a whole. More specifically, there were slight growth reductions in the warmer north but gains in the cooler south, especially at higher altitudes. For sites where P. radiata is currently grown, and across the full suite of GCMs and emission scenarios, changes in wood productivity averaged +3% for both 2040 and 2090. When increasing CO₂ concentration was also included, responses of wood productivity were generally positive, with average increases of 19% by 2040 and 37% by 2090. These responses varied regionally, ranging from relatively minor changes in the north of the country to very significant increases in the south, where the beneficial effect of increasing CO₂ combined with the beneficial effect of increasing temperatures. These relatively large responses to CO₂ depend on maintenance of the current adequate fertility levels in most commercial plantations. Productivity enhancements came at the expense of some soil‐carbon losses. Average losses for the country were simulated to average 3.5% under constant CO₂ and 1.5% with increasing CO₂ concentration. Again, there were regional differences, with larger losses for regions with lesser growth enhancements, and lesser reductions in regions where greater productivity enhancements could partly balance the effect of faster decomposition activity.     

Contrasting acclimation responses to elevated CO2 and warming between an evergreen and a deciduous boreal conifer

Rising atmospheric carbon dioxide (CO₂) concentrations may warm northern latitudes up to 8°C by the end of the century. Boreal forests play a large role in the global carbon cycle, and the responses of northern trees to climate change will thus impact the trajectory of future CO₂ increases. We grew two North American boreal tree species at a range of future climate conditions to assess how growth and carbon fluxes were altered by high CO₂ and warming. Black spruce (Picea mariana, an evergreen conifer) and tamarack (Larix laricina, a deciduous conifer) were grown under ambient (407 ppm) or elevated CO₂ (750 ppm) and either ambient temperatures, a 4°C warming, or an 8°C warming. In both species, the thermal optimum of net photosynthesis (T_optA) increased and maximum photosynthetic rates declined in warm‐grown seedlings, but the strength of these changes varied between species. Photosynthetic capacity (maximum rates of Rubisco carboxylation, V_cmax, and of electron transport, J_max) was reduced in warm‐grown seedlings, correlating with reductions in leaf N and chlorophyll concentrations. Warming increased the activation energy for V_cmax and J_max (E_aV and E_aJ, respectively) and the thermal optimum for J_max. In both species, the T_optA was positively correlated with both E_aV and E_aJ, but negatively correlated with the ratio of J_max/V_cmax. Respiration acclimated to elevated temperatures, but there were no treatment effects on the Q₁₀ of respiration (the increase in respiration for a 10°C increase in leaf temperature). A warming of 4°C increased biomass in tamarack, while warming reduced biomass in spruce. We show that climate change is likely to negatively affect photosynthesis and growth in black spruce more than in tamarack, and that parameters used to model photosynthesis in dynamic global vegetation models (E_aV and E_aJ) show no response to elevated CO₂.     

Trees tolerate an extreme heatwave via sustained transpirational cooling and increased leaf thermal tolerance

Heatwaves are likely to increase in frequency and intensity with climate change, which may impair tree function and forest C uptake. However, we have little information regarding the impact of extreme heatwaves on the physiological performance of large trees in the field. Here, we grew Eucalyptus parramattensis trees for 1 year with experimental warming (+3°C) in a field setting, until they were greater than 6 m tall. We withheld irrigation for 1 month to dry the surface soils and then implemented an extreme heatwave treatment of 4 consecutive days with air temperatures exceeding 43°C, while monitoring whole‐canopy exchange of CO₂ and H₂O, leaf temperatures, leaf thermal tolerance, and leaf and branch hydraulic status. The heatwave reduced midday canopy photosynthesis to near zero but transpiration persisted, maintaining canopy cooling. A standard photosynthetic model was unable to capture the observed decoupling between photosynthesis and transpiration at high temperatures, suggesting that climate models may underestimate a moderating feedback of vegetation on heatwave intensity. The heatwave also triggered a rapid increase in leaf thermal tolerance, such that leaf temperatures observed during the heatwave were maintained within the thermal limits of leaf function. All responses were equivalent for trees with a prior history of ambient and warmed (+3°C) temperatures, indicating that climate warming conferred no added tolerance of heatwaves expected in the future. This coordinated physiological response utilizing latent cooling and adjustment of thermal thresholds has implications for tree tolerance of future climate extremes as well as model predictions of future heatwave intensity at landscape and global scales.     

Risky future for Mediterranean forests unless they undergo extreme carbon fertilization

Forest performance is challenged by climate change but higher atmospheric [CO₂] (c_a) could help trees mitigate the negative effect of enhanced water stress. Forest projections using data assimilation with mechanistic models are a valuable tool to assess forest performance. Firstly, we used dendrochronological data from 12 Mediterranean tree species (six conifers and six broadleaves) to calibrate a process‐based vegetation model at 77 sites. Secondly, we conducted simulations of gross primary production (GPP) and radial growth using an ensemble of climate projections for the period 2010–2100 for the high‐emission RCP8.5 and low‐emission RCP2.6 scenarios. GPP and growth projections were simulated using climatic data from the two RCPs combined with (i) expected c_a; (ii) constant c_a = 390 ppm, to test a purely climate‐driven performance excluding compensation from carbon fertilization. The model accurately mimicked the growth trends since the 1950s when, despite increasing c_a, enhanced evaporative demands precluded a global net positive effect on growth. Modeled annual growth and GPP showed similar long‐term trends. Under RCP2.6 (i.e., temperatures below +2 °C with respect to preindustrial values), the forests showed resistance to future climate (as expressed by non‐negative trends in growth and GPP) except for some coniferous sites. Using exponentially growing c_a and climate as from RCP8.5, carbon fertilization overrode the negative effect of the highly constraining climatic conditions under that scenario. This effect was particularly evident above 500 ppm (which is already over +2 °C), which seems unrealistic and likely reflects model miss‐performance at high c_a above the calibration range. Thus, forest projections under RCP8.5 preventing carbon fertilization displayed very negative forest performance at the regional scale. This suggests that most of western Mediterranean forests would successfully acclimate to the coldest climate change scenario but be vulnerable to a climate warmer than +2 °C unless the trees developed an exaggerated fertilization response to [CO₂].     

Distinct seasonal dynamics of responses to elevated CO2 in two understorey grass species differing in shade‐tolerance

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Modelling the influence of predicted future climate change on the risk of wind damage within New Zealand's planted forests

Wind is the major abiotic disturbance in New Zealand's planted forests, but little is known about how the risk of wind damage may be affected by future climate change. We linked a mechanistic wind damage model (ForestGALES) to an empirical growth model for radiata pine (Pinus radiata D. Don) and a process‐based growth model (cenw ) to predict the risk of wind damage under different future emissions scenarios and assumptions about the future wind climate. The cenw model was used to estimate site productivity for constant CO₂ concentration at 1990 values and for assumed increases in CO₂ concentration from current values to those expected during 2040 and 2090 under the B1 (low), A1B (mid‐range) and A2 (high) emission scenarios. Stand development was modelled for different levels of site productivity, contrasting silvicultural regimes and sites across New Zealand. The risk of wind damage was predicted for each regime and emission scenario combination using the ForestGALES model. The sensitivity to changes in the intensity of the future wind climate was also examined. Results showed that increased tree growth rates under the different emissions scenarios had the greatest impact on the risk of wind damage. The increase in risk was greatest for stands growing at high stand density under the A2 emissions scenario with increased CO₂ concentration. The increased productivity under this scenario resulted in increased tree height, without a corresponding increase in diameter, leading to more slender trees that were predicted to be at greater risk from wind damage. The risk of wind damage was further increased by the modest increases in the extreme wind climate that are predicted to occur. These results have implications for the development of silvicultural regimes that are resilient to climate change and also indicate that future productivity gains may be offset by greater losses from disturbances.     

Environmental and developmental controls over the seasonal pattern of isoprene emission from aspen leaves

Isoprene emission from plants represents one of the principal biospheric controls over the oxidative capacity of the continental troposphere. In the study reported here, the seasonal pattern of isoprene emission, and its underlying determinants, were studied for aspen trees growing in the Rocky Mountains of Colorado. The springtime onset of isoprene emission was delayed for up to 4 weeks following leaf emergence, despite the presence of positive net photosynthesis rates. Maximum isoprene emission rates were reached approximately 6 weeks following leaf emergence. During this initial developmental phase, isoprene emission rates were negatively correlated with leaf nitrogen concentrations. During the autumnal decline in isoprene emission, rates were positively correlated with leaf nitrogen concentration. Given past studies that demonstrate a correlation between leaf nitrogen concentration and isoprene emission rate, we conclude that factors other than the amount of leaf nitrogen determine the early-season initiation of isoprene emission. The late-season decline in isoprene emission rate is interpreted as due to the autumnal breakdown of metabolic machinery and loss of leaf nitrogen. In potted aspen trees, leaves that emerged in February and developed under cool, springtime temperatures did not emit isoprene until 23 days after leaf emergence. Leaves that emrged in July and developed in hot, midsummer temperatures emitted isoprene within 6 days. Leaves that had emerged during the cool spring, and had grown for several weeks without emitting isoprene, could be induced to emit isoprene within 2 h of exposure to 32°C. Continued exposure to warm temperatures resulted in a progressive increase in the isoprene emission rate. Thus, temperature appears to be an important determinant of the early season induction of isoprene emission. The seasonal pattern of isoprene emission was examined in trees growing along an elevational gradient in the Colorado Front Range (1829–2896 m). Trees at different elevations exhibited staggered patterns of bud-break and initiation of photosynthesis and isoprene emission in concert with the staggered onset of warm, springtime temperatures. The springtime induction of isoprene emission could be predicted at each of the three sites as the time after bud break required for cumulative temperatures above 0°C to reach approximately 400 degree days. Seasonal temperature acclimation of isoprene emission rate and photosynthesis rate was not observed. The temperature dependence of isoprene emission rate between 20 and 35°C could be accurately predicted during spring and summer using a single algorithm that describes the Arrhenius relationship of enzyme activity. From these results, it is concluded that the early season pattern of isoprene emission is controlled by prevailing temperature and its interaction with developmental processes. The late-season pattern is determined by controls over leaf nitrogen concentration, especially the depletion of leaf nitrogen during senescence. Following early-season induction, isoprene emission rates correlate with photosynthesis rates. During the season there is little acclimation to temperature, so that seasonal modeling simplifies to a single temperature-response algorithm.     

Effects of climate warming on Sphagnum photosynthesis in peatlands depend on peat moisture and species‐specific anatomical traits

Climate change will influence plant photosynthesis by altering patterns of temperature and precipitation, including their variability and seasonality. Both effects may be important for peatlands as the carbon (C) sink potential of these ecosystems depends on the balance between plant C uptake through photosynthesis and microbial decomposition. Here, we show that the effect of climate warming on Sphagnum community photosynthesis toggles from positive to negative as the peatland goes from rainy to dry periods during summer. More particularly, we show that mechanisms of compensation among the dominant Sphagnum species (Sphagnum fallax and Sphagnum medium) stabilize the average photosynthesis and productivity of the Sphagnum community during summer despite rising temperatures and frequent droughts. While warming had a negligible effect on S. medium photosynthetic capacity (A_max) during rainy periods, A_max of S. fallax increased by 40%. On the opposite, warming exacerbated the negative effects of droughts on S. fallax with an even sharper decrease of its A_max while S. medium A_max remained unchanged. S. medium showed a remarkable resistance to droughts due to anatomical traits favouring its water holding capacity. Our results show that different phenotypic plasticity among dominant Sphagnum species allow the community to cope with rising temperatures and repeated droughts, maintaining similar photosynthesis and productivity over summer in warmed and control conditions. These results are important because they provide information on how soil water content may modulate the effects of climate warming on Sphagnum productivity in boreal peatlands. It further confirms the transitory nature of warming‐induced photosynthesis benefits in boreal systems and highlights the vulnerability of the ecosystem to excess warming and drying.     

Assessing forest vulnerability to climate warming using a process‐based model of tree growth: bad prospects for rear‐edges

Growth models can be used to assess forest vulnerability to climate warming. If global warming amplifies water deficit in drought‐prone areas, tree populations located at the driest and southernmost distribution limits (rear‐edges) should be particularly threatened. Here, we address these statements by analyzing and projecting growth responses to climate of three major tree species (silver fir, Abies alba; Scots pine, Pinus sylvestris; and mountain pine, Pinus uncinata) in mountainous areas of NE Spain. This region is subjected to Mediterranean continental conditions, it encompasses wide climatic, topographic and environmental gradients, and, more importantly, it includes rear‐edges of the continuous distributions of these tree species. We used tree‐ring width data from a network of 110 forests in combination with the process‐based Vaganov–Shashkin‐Lite growth model and climate–growth analyses to forecast changes in tree growth during the 21st century. Climatic projections were based on four ensembles CO₂ emission scenarios. Warm and dry conditions during the growing season constrain silver fir and Scots pine growth, particularly at the species rear‐edge. By contrast, growth of high‐elevation mountain pine forests is enhanced by climate warming. The emission scenario (RCP 8.5) corresponding to the most pronounced warming (+1.4 to 4.8 °C) forecasted mean growth reductions of ?10.7% and ?16.4% in silver fir and Scots pine, respectively, after 2050. This indicates that rising temperatures could amplify drought stress and thus constrain the growth of silver fir and Scots pine rear‐edge populations growing at xeric sites. Contrastingly, mountain pine growth is expected to increase by +12.5% due to a longer and warmer growing season. The projections of growth reduction in silver fir and Scots pine portend dieback and a contraction of their species distribution areas through potential local extinctions of the most vulnerable driest rear‐edge stands. Our modeling approach provides accessible tools to evaluate forest vulnerability to warmer conditions.     

Large‐scale variations in the vegetation growing season and annual cycle of atmospheric CO2 at high northern latitudes from 1950 to 2011

We combine satellite and ground observations during 1950–2011 to study the long‐term links between multiple climate (air temperature and cryospheric dynamics) and vegetation (greenness and atmospheric CO₂ concentrations) indicators of the growing season of northern ecosystems (>45°N) and their connection with the carbon cycle. During the last three decades, the thermal potential growing season has lengthened by about 10.5 days (P < 0.01, 1982–2011), which is unprecedented in the context of the past 60 years. The overall lengthening has been stronger and more significant in Eurasia (12.6 days, P < 0.01) than North America (6.2 days, P > 0.05). The photosynthetic growing season has closely tracked the pace of warming and extension of the potential growing season in spring, but not in autumn when factors such as light and moisture limitation may constrain photosynthesis. The autumnal extension of the photosynthetic growing season since 1982 appears to be about half that of the thermal potential growing season, yielding a smaller lengthening of the photosynthetic growing season (6.7 days at the circumpolar scale, P < 0.01). Nevertheless, when integrated over the growing season, photosynthetic activity has closely followed the interannual variations and warming trend in cumulative growing season temperatures. This lengthening and intensification of the photosynthetic growing season, manifested principally over Eurasia rather than North America, is associated with a long‐term increase (22.2% since 1972, P < 0.01) in the amplitude of the CO₂ annual cycle at northern latitudes. The springtime extension of the photosynthetic and potential growing seasons has apparently stimulated earlier and stronger net CO₂ uptake by northern ecosystems, while the autumnal extension is associated with an earlier net release of CO₂ to the atmosphere. These contrasting responses may be critical in determining the impact of continued warming on northern terrestrial ecosystems and the carbon cycle.     

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].     

Tree mineral nutrition is deteriorating in Europe

The response of forest ecosystems to increased atmospheric CO₂ is constrained by nutrient availability. It is thus crucial to account for nutrient limitation when studying the forest response to climate change. The objectives of this study were to describe the nutritional status of the main European tree species, to identify growth‐limiting nutrients and to assess changes in tree nutrition during the past two decades. We analysed the foliar nutrition data collected during 1992–2009 on the intensive forest monitoring plots of the ICP Forests programme. Of the 22 significant temporal trends that were observed in foliar nutrient concentrations, 20 were decreasing and two were increasing. Some of these trends were alarming, among which the foliar P concentration in F. sylvatica, Q. Petraea and P. sylvestris that significantly deteriorated during 1992–2009. In Q. Petraea and P. sylvestris, the decrease in foliar P concentration was more pronounced on plots with low foliar P status, meaning that trees with latent P deficiency could become deficient in the near future. Increased tree productivity, possibly resulting from high N deposition and from the global increase in atmospheric CO₂, has led to higher nutrient demand by trees. As the soil nutrient supply was not always sufficient to meet the demands of faster growing trees, this could partly explain the deterioration of tree mineral nutrition. The results suggest that when evaluating forest carbon storage capacity and when planning to reduce CO₂ emissions by increasing use of wood biomass for bioenergy, it is crucial that nutrient limitations for forest growth are considered.     

Recent CO2 rise has modified the sensitivity of tropical tree growth to rainfall and temperature

Atmospheric CO₂ (c_a) rise changes the physiology and possibly growth of tropical trees, but these effects are likely modified by climate. Such c_a × climate interactions importantly drive CO₂ fertilization effects of tropical forests predicted by global vegetation models, but have not been tested empirically. Here we use tree‐ring analyses to quantify how c_a rise has shifted the sensitivity of tree stem growth to annual fluctuations in rainfall and temperature. We hypothesized that c_a rise reduces drought sensitivity and increases temperature sensitivity of growth, by reducing transpiration and increasing leaf temperature. These responses were expected for cooler sites. At warmer sites, c_a rise may cause leaf temperatures to frequently exceed the optimum for photosynthesis, and thus induce increased drought sensitivity and stronger negative effects of temperature. We tested these hypotheses using measurements of 5,318 annual rings from 129 trees of the widely distributed (sub‐)tropical tree species, Toona ciliata. We studied growth responses during 1950–2014, a period during which c_a rose by 28%. Tree‐ring data were obtained from two cooler (mean annual temperature: 20.5–20.7°C) and two warmer (23.5–24.8°C) sites. We tested c_a × climate interactions, using mixed‐effect models of ring‐width measurements. Our statistical models revealed several significant and robust c_a × climate interactions. At cooler sites (and seasons), c_a × climate interactions showed good agreement with hypothesized growth responses of reduced drought sensitivity and increased temperature sensitivity. At warmer sites, drought sensitivity increased with increasing c_a, as predicted, and hot years caused stronger growth reduction at high c_a. Overall, c_a rise has significantly modified sensitivity of Toona stem growth to climatic variation, but these changes depended on mean climate. Our study suggests that effects of c_a rise on tropical tree growth may be more complex and less stimulatory than commonly assumed and require a better representation in global vegetation models.     

Isoprene emission from aspen leaves : influence of environment and relation to photosynthesis and photorespiration

Isoprene emission rates from quaking aspen (Populus tremuloides Michx.) leaves were measured simultaneously with photosynthesis rate, stomatal conductance, and intercellular CO₂ partial pressure. Isoprene emission required the presence of CO₂ or O₂, but not both. The light response of isoprene emission rate paralleled that of photosynthesis. Isoprene emission was inhibited by decreasing ambient O₂ from 21% to 2%, only when there was oxygen insensitive photosynthesis. Mannose (10 millimolar) fed through cut stems resulted in strong inhibition of isoprene emission rate and is interpreted as evidence that isoprene biosynthesis requires either the export of triose phosphates from the chloroplast, or the continued synthesis of ATP. Light response experiments suggest that photosynthetically generated reductant or ATP is required for isoprene biosynthesis. Isoprene biosynthesis and emission are not directly linked to glycolate production through photorespiration, contrary to previous reports. Isoprene emission rate was inhibited by above-ambient CO₂ partial pressures (640 microbar outside and 425 microbar inside the leaf). The inhibition was not due to stomatal closure. This was established by varying ambient humidity at normal and elevated CO₂ partial pressures to measure isoprene emission rates over a range of stomatal conductances. Isoprene emission rates were inhibited at elevated CO₂ despite no change in stomatal conductance. Addition of abscisic acid to the transpiration stream dramatically inhibited stomatal conductance and photosynthesis rate, with a slight increase in isoprene emission rate. Thus, isoprene emission is independent of stomatal conductance, and may occur through the cuticle. Temperature had an influence on isoprene emission rate, with the Q₁₀ being 1.8 to 2.4 between 35 and 45°C. At these high temperatures the amount of carbon lost through isoprene emission was between 2.5 and 8% of that assimilated through photosynthesis. This represents a significant carbon cost that should be taken into account in determining midsummer carbon budgets for plants that are isoprene emitters.     

Sap‐feeding insects on forest trees along latitudinal gradients in northern Europe: a climate‐driven patterns

Knowledge of the latitudinal patterns in biotic interactions, and especially in herbivory, is crucial for understanding the mechanisms that govern ecosystem functioning and for predicting their responses to climate change. We used sap‐feeding insects as a model group to test the hypotheses that the strength of plant–herbivore interactions in boreal forests decreases with latitude and that this latitudinal pattern is driven primarily by midsummer temperatures. We used a replicated sampling design and quantitatively collected and identified all sap‐feeding insects from four species of forest trees along five latitudinal gradients (750–1300 km in length, ten sites in each gradient) in northern Europe (59 to 70°N and 10 to 60°E) during 2008–2011. Similar decreases in diversity of sap‐feeding insects with latitude were observed in all gradients during all study years. The sap‐feeder load (i.e. insect biomass per unit of foliar biomass) decreased with latitude in typical summers, but increased in an exceptionally hot summer and was independent of latitude during a warm summer. Analysis of combined data from all sites and years revealed dome‐shaped relationships between the loads of sap‐feeders and midsummer temperatures, peaking at 17 °C in Picea abies, at 19.5 °C in Pinus sylvestris and Betula pubescens and at 22 °C in B. pendula. From these relationships, we predict that the losses of forest trees to sap‐feeders will increase by 0–45% of the current level in southern boreal forests and by 65–210% in subarctic forests with a 1 °C increase in summer temperatures. The observed relationships between temperatures and the loads of sap‐feeders differ between the coniferous and deciduous tree species. We conclude that climate warming will not only increase plant losses to sap‐feeding insects, especially in subarctic forests, but can also alter plant‐plant interactions, thereby affecting both the productivity and the structure of future forest ecosystems.     

Scaling net ecosystem CO2 exchange from the community to landscape‐level at a subarctic fen

Landscape‐ and community‐level CO₂ measurements were made at a subarctic sedge fen near Churchill Manitoba during the 1997 growing season. Climatic conditions were warmer and drier than the 30‐y normal. Landscape‐scale micrometeorological measurements indicated that the wetland gained 49 g CO₂ m^?2 during the growing season. Chamber‐scale measurements from the main vegetation community types showed that small hummocks (Carex spp. sites) dominated the CO₂ exchange, yielding an effective scaling factor of 70%. Scaled parameters of two algorithms describing photosynthesis and respiration for each community type show strong similarity to those derived at the landscape level. Scaling photosynthesis, respiration, and net ecosystem CO₂ exchange from the community to landscape‐level over the season is within the maximum probable error of each methodological approach and helps substantiate the 1997 CO₂ budget. We explore the equilibrium response of net ecosystem CO₂ exchange of this fen to climatic change by examining the feedback of water table position on vegetation distribution and nitrogen availability. Based on the effective scaling factors computed for each community type, we hypothesize that a small decrease in mean water table position could nearly triple the net uptake of CO₂ at this wetland.     

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.     

Limited evidence for CO2‐related growth enhancement in northern Rocky Mountain lodgepole pine populations across climate gradients

Forests sequester large amounts of carbon annually and are integral in buffering against effects of global change. Increasing atmospheric CO₂ may enhance photosynthesis and/or decrease stomatal conductance (g_s) thereby enhancing intrinsic water‐use efficiency (iWUE), having potential indirect and direct benefits to tree growth. While increasing iWUE has been observed in most trees globally, enhanced growth is not ubiquitous, possibly due to concurrent climatic constraints on growth. To investigate our incomplete understanding of interactions between climate and CO₂ and their impacts on tree physiology and growth, we used an environmental gradient approach. We combined dendrochronology with carbon isotope analysis (δ¹³C) to assess the covariation of basal area increment (BAI) and iWUE over time in lodgepole pine. Trees were sampled at 18 sites spanning two climatically distinct elevation transects on the lee and windward sides of the Continental Divide, encompassing the majority of lodgepole pine's northern Rocky Mountain elevational range. We analyzed BAI and iWUE from 1950 to 2015, and explored correlations with monthly climate variables. As expected, iWUE increased at all sites. However, concurrent growth trends depended on site climatic water deficit (CWD). Significant growth increases occurred only at the driest sites, where increases in iWUE were strongest, while growth decreases were greatest at sites where CWD has been historically lowest. Late summer drought of the previous year negatively affected growth across sites. These results suggest that increasing iWUE, if strong enough, may indirectly benefit growth at drier sites by effectively extending the growing season via reductions in g_s. Strong growth decreases at high elevation windward sites may reflect increasing water stress as a result of decreasing snowpack, which was not offset by greater iWUE. Our results imply that increasing iWUE driven by decreasing g_s may benefit tree growth in limited scenarios, having implications for future carbon uptake potential of semiarid ecosystems.