Gypsy moth feeding in the canopy of a CO2-enriched mature forest

Rising atmospheric carbon dioxide (CO₂) concentration is expected to change plant tissue quality with important implications for plant–insect interactions. Taking advantage of canopy access by a crane and long‐term CO₂ enrichment (530 μ mol mol^?1) of a natural old‐growth forest (web‐free air carbon dioxide enrichment), we studied the responses of a generalist insect herbivore feeding in the canopy of tall trees. We found that relative growth rates (RGR) of gypsy moth (Lymantria dispar) were reduced by 30% in larvae fed on high CO₂‐exposed Quercus petraea, but increased by 29% when fed on high CO₂‐grown Carpinus betulus compared with control trees at ambient CO₂ (370 μ mol mol^?1). In Fagus sylvatica, there was a nonsignificant trend for reduced RGR under elevated CO₂. Tree species‐specific changes in starch to nitrogen ratio, water, and the concentrations of proteins, condensed and hydrolyzable tannins in response to elevated CO₂ were identified to correlate with altered RGR of gypsy moth larvae. Our data suggest that rising atmospheric CO₂ will have strong species‐specific effects on leaf chemical composition of canopy trees in natural forests leading to contrasting responses of herbivores such as those reported here. A future change in host tree preference seems likely with far‐ranging consequences for forest community dynamics.     

Stable carbon isotopes in tree rings indicate improved water use efficiency and drought responses of a tropical dry forest tree species

Understanding the responses of tropical trees to increasing [CO₂] and climate change is important as tropical forests play an important role in carbon and hydrological cycles. We used stable carbon isotopes (δ¹³C) in tree rings to study the physiological responses of a tropical dry forest tree species in southern Mexico, Mimosa acantholoba to changes in atmospheric [CO₂] and variation in climate. Based on annual records of tree ring δ¹³C, we calculated intrinsic water use efficiency (W _i) and intercellular [CO₂] (c _i). Our results showed that trees responded strongly to the increase in atmospheric [CO₂] over the last four decades; W _i increased dramatically by 40%, while c _i remained largely constant. The maintenance of a constant c _i indicates that photosynthetic rates are unlikely to have increased in response to higher [CO₂], and that improvements in W _i are probably due to a reduction in stomatal conductance. This may have large consequences for the hydrological cycle. Inter-annual variation in c _i was strongly correlated with total annual rainfall (r = 0.70), and not influenced by temperature, solar radiation or cloud cover. Our results show that δ¹³C in tree rings of tropical dry forest trees may be a powerful tool to evaluate long-term responses of trees to increasing [CO₂] and to variation in climate.     

Increased leaf reflectance in tropical trees under elevated CO2

Globally increasing atmospheric CO₂ concentrations are known to affect many aspects of plant physiology and development; however, little attention has been given to leaf and canopy optical properties. Three tropical trees in the Leguminosae, an important canopy tree family in many tropical forests, responded similarly to an experimental doubling of CO₂ partial pressure with a 9–23% increase in spectral leaf reflectance to light in the visible (400–700 nm) waveband. Decreased leaf chlorophyll content under elevated CO₂ may explain part of the observed increase in reflectance. However, analyses that statistically corrected for chlorophyll content effects on reflectance still indicated a significant CO₂ effect. This results, in conjunction with the spectral pattern of the response, suggests that the primary mechanism is increased optical masking of chlorophyll under elevated CO₂. The magnitude of the increase in leaf reflectance is sufficient to suggest that increased canopy reflectance of tropical forests (and possibly other terrestrial ecosystems) may be an important negative feedback in the response of global net radiative climate forcing to increasing atmospheric CO₂.     

Global response of terrestrial ecosystem structure and function to CO2 and climate change: results from six dynamic global vegetation models

The possible responses of ecosystem processes to rising atmospheric CO₂ concentration and climate change are illustrated using six dynamic global vegetation models that explicitly represent the interactions of ecosystem carbon and water exchanges with vegetation dynamics. The models are driven by the IPCC IS92a scenario of rising CO₂ ( Wigley et al. 1991 ), and by climate changes resulting from effective CO₂ concentrations corresponding to IS92a, simulated by the coupled ocean atmosphere model HadCM2‐SUL. Simulations with changing CO₂ alone show a widely distributed terrestrial carbon sink of 1.4–3.8 Pg C y^?1 during the 1990s, rising to 3.7–8.6 Pg C y^?1 a century later. Simulations including climate change show a reduced sink both today (0.6–3.0 Pg C y^?1) and a century later (0.3–6.6 Pg C y^?1) as a result of the impacts of climate change on NEP of tropical and southern hemisphere ecosystems. In all models, the rate of increase of NEP begins to level off around 2030 as a consequence of the ‘diminishing return’ of physiological CO₂ effects at high CO₂ concentrations. Four out of the six models show a further, climate‐induced decline in NEP resulting from increased heterotrophic respiration and declining tropical NPP after 2050. Changes in vegetation structure influence the magnitude and spatial pattern of the carbon sink and, in combination with changing climate, also freshwater availability (runoff). It is shown that these changes, once set in motion, would continue to evolve for at least a century even if atmospheric CO₂ concentration and climate could be instantaneously stabilized. The results should be considered illustrative in the sense that the choice of CO₂ concentration scenario was arbitrary and only one climate model scenario was used. However, the results serve to indicate a range of possible biospheric responses to CO₂ and climate change. They reveal major uncertainties about the response of NEP to climate change resulting, primarily, from differences in the way that modelled global NPP responds to a changing climate. The simulations illustrate, however, that the magnitude of possible biospheric influences on the carbon balance requires that this factor is taken into account for future scenarios of atmospheric CO₂ and climate change.     

Endemic species and ecosystem sensitivity to climate change in Namibia

We present a first assessment of the potential impacts of anthropogenic climate change on the endemic flora of Namibia, and on its vegetation structure and function, for a projected climate in ～2050 and ～2080. We used both niche‐based models (NBM) to evaluate the sensitivity of 159 endemic species to climate change (of an original 1020 plant species modeled) and a dynamic global vegetation model (DGVM) to assess the impacts of climate change on vegetation structure and ecosystem functioning. Endemic species modeled by NBM are moderately sensitive to projected climate change. Fewer than 5% are predicted to experience complete range loss by 2080, although more than 47% of the species are expected to be vulnerable (range reduction >30%) by 2080 if they are assumed unable to migrate. Disaggregation of results by life‐form showed distinct patterns. Endemic species of perennial herb, geophyte and tree life‐formsare predicted to be negatively impacted in Namibia, whereas annual herb and succulent endemic species remain relatively stable by 2050 and 2080. Endemic annual herb species are even predicted to extend their range north‐eastward into the tree and shrub savanna with migration, and tolerance of novel substrates. The current protected area network is predicted to meet its mandate by protecting most of the current endemicity in Namibia into the future. Vegetation simulated by DGVM is projected to experience a reduction in cover, net primary productivity and leaf area index throughout much of the country by 2050, with important implications for the faunal component of Namibia's ecosystems, and the agricultural sector. The plant functional type (PFT) composition of the major biomes may be substantially affected by climate change and rising atmospheric CO₂– currently widespread deciduous broad leaved trees and C₄ PFTs decline, with the C₄ PFT particularly negatively affected by rising atmospheric CO₂ impacts by ～2080 and deciduous broad leaved trees more likely directly impacted by drying and warming. The C₃ PFT may increase in prominence in the northwestern quadrant of the country by ～2080 as CO₂ concentrations increase. These results suggest that substantial changes in species diversity, vegetation structure and ecosystem functioning can be expected in Namibia with anticipated climate change, although endemic plant richness may persist in the topographically diverse central escarpment region.     

Tree growth variation in the tropical forest: understanding effects of temperature,rainfall and CO2

Tropical forest responses to climatic variability have important consequences for global carbon cycling, but are poorly understood. As empirical, correlative studies cannot disentangle the interactive effects of climatic variables on tree growth, we used a tree growth model (IBTREE) to unravel the climate effects on different physiological pathways and in turn on stem growth variation. We parameterized the model for canopy trees of Toona ciliata (Meliaceae) from a Thai monsoon forest and compared predicted and measured variation from a tree‐ring study over a 30‐year period. We used historical climatic variation of minimum and maximum day temperature, precipitation and carbon dioxide (CO₂) in different combinations to estimate the contribution of each climate factor in explaining the inter‐annual variation in stem growth. Running the model with only variation in maximum temperature and rainfall yielded stem growth patterns that explained almost 70% of the observed inter‐annual variation in stem growth. Our results show that maximum temperature had a strong negative effect on the stem growth by increasing respiration, reducing stomatal conductance and thus mitigating a higher transpiration demand, and – to a lesser extent – by directly reducing photosynthesis. Although stem growth was rather weakly sensitive to rain, stem growth variation responded strongly and positively to rainfall variation owing to the strong inter‐annual fluctuations in rainfall. Minimum temperature and atmospheric CO₂ concentration did not significantly contribute to explaining the inter‐annual variation in stem growth. Our innovative approach – combining a simulation model with historical data on tree‐ring growth and climate – allowed disentangling the effects of strongly correlated climate variables on growth through different physiological pathways. Similar studies on different species and in different forest types are needed to further improve our understanding of the sensitivity of tropical tree growth to climatic variability and change.     

Solar radiation and ENSO predict fruiting phenology patterns in a 15‐year record from Kibale National Park,Uganda

Fruiting, flowering, and leaf set patterns influence many aspects of tropical forest communities, but there are few long‐term studies examining potential drivers of these patterns, particularly in Africa. We evaluated a 15‐year dataset of tree phenology in Kibale National Park, Uganda, to identify abiotic predictors of fruit phenological patterns and discuss our findings in light of climate change. We quantified fruiting for 326 trees from 43 species and evaluated these patterns in relation to solar radiance, rainfall, and monthly temperature. We used time‐lagged variables based on seasonality in linear regression models to assess the effect of abiotic variables on the proportion of fruiting trees. Annual fruiting varied over 3.8‐fold, and inter‐annual variation in fruiting is associated with the extent of fruiting in the peak period, not variation in time of fruit set. While temperature and rainfall showed positive effects on fruiting, solar radiance in the two‐year period encompassing a given year and the previous year was the strongest predictor of fruiting. As solar irradiance was the strongest predictor of fruiting, the projected increase in rainfall associated with climate change, and coincident increase in cloud cover suggest that climate change will lead to a decrease in fruiting. ENSO in the prior 24‐month period was also significantly associated with annual ripe fruit production, and ENSO is also affected by climate change. Predicting changes in phenology demands understanding inter‐annual variation in fruit dynamics in light of potential abiotic drivers, patterns that will only emerge with long‐term data.     

Water savings in mature deciduous forest trees under elevated CO2

Stomatal conductance of plants exposed to elevated CO₂ is often reduced. Whether this leads to water savings in tall forest‐trees under future CO₂ concentrations is largely unknown but could have significant implications for climate and hydrology. We used three different sets of measurements (sap flow, soil moisture and canopy temperature) to quantify potential water savings under elevated CO₂ in a ca. 35 m tall, ca. 100 years old mixed deciduous forest. Part of the forest canopy was exposed to 540 ppm CO₂ during daylight hours using free air CO₂ enrichment (FACE) and the Swiss Canopy Crane (SCC). Across species and a wide range of weather conditions, sap flow was reduced by 14% in trees subjected to elevated CO₂, yielding ca. 10% reduction in evapotranspiration. This signal is likely to diminish as atmospheric feedback through reduced moistening of the air comes into play at landscape scale. Vapour pressure deficit (VPD)‐sap flow response curves show that the CO₂ effect is greatest at low VPD, and that sap flow saturation tends to occur at lower VPD in CO₂‐treated trees. Matching stomatal response data, the CO₂ effect was largely produced by Carpinus and Fagus, with Quercus contributing little. In line with these findings, soil moisture at 10 cm depth decreased at a slower rate under high‐CO₂ trees than under control trees during rainless periods, with a reversal of this trend during prolonged drought when CO₂‐treated trees take advantage from initial water savings. High‐resolution thermal images taken at different heights above the forest canopy did detect reduced water loss through altered energy balance only at <5 m distance (0.44 K leaf warming of CO₂‐treated Fagus trees). Short discontinuations of CO₂ supply during morning hours had no measurable canopy temperature effects, most likely because the stomatal effects were small compared with the aerodynamic constraints in these dense, broad‐leaved canopies. Hence, on a seasonal basis, these data suggest a <10% reduction in water consumption in this type of forest when the atmosphere reaches 540% ppm CO₂.     

Flowering in grassland predicted by CO2 and resource effects on species aboveground biomass

Continuing enrichment of atmospheric CO₂ may change plant community composition, in part by altering the availability of other limiting resources including soil water, nutrients, or light. The combined effects of CO₂ enrichment and altered resource availability on species flowering remain poorly understood. We quantified flowering culm and ramet production and biomass allocation to flowering culms/ramets for 10 years in C₄‐dominated grassland communities on contrasting soils along a CO₂ concentration gradient spanning pre‐industrial to expected mid‐21st century levels (250–500 μl/L). CO₂ enrichment explained up to 77% of the variation in flowering culm count across soils for three of the five species, and was correlated with flowering culm count on at least one soil for four of five species. In contrast, allocation to flowering culms was only weakly correlated with CO₂ enrichment for two species. Flowering culm counts were strongly correlated with species aboveground biomass (AGB; R² = .34–.74), a measure of species abundance. CO₂ enrichment also increased soil moisture and decreased light levels within the canopy but did not affect soil inorganic nitrogen availability. Structural equation models fit across the soils suggested species‐specific controls on flowering in two general forms: (1) CO₂ effects on flowering culm count mediated by canopy light level and relative species AGB (species AGB/total AGB) or by soil moisture effects on flowering culm count; (2) effects of canopy light level or soil inorganic nitrogen on flowering and/or relative species AGB, but with no significant CO₂ effect. Understanding the heterogeneity in species responses to CO₂ enrichment in plant communities across soils in edaphically variable landscapes is critical to predict CO₂ effects on flowering and other plant fitness components, and species potential to adapt to future environmental changes.     

The space‐time continuum: the effects of elevated CO2 and temperature on trees and the importance of scaling

To predict how forests will respond to rising temperatures and atmospheric CO₂ concentrations, we need to understand how trees respond to both of these environmental factors. In this review, we discuss the importance of scaling, moving from leaf‐level responses to those of the canopy, and from short‐term to long‐term responses of vegetation to climate change. While our knowledge of leaf‐level, instantaneous responses of photosynthesis, respiration, stomatal conductance, transpiration and water‐use efficiency to elevated CO₂ and temperature is quite good, our ability to scale these responses up to larger spatial and temporal scales is less developed. We highlight which physiological processes are least understood at various levels of study, and discuss how ignoring differences in the spatial or temporal scale of a physiological process impedes our ability to predict how forest carbon and water fluxes forests will be altered in the future. We also synthesize data from the literature to show that light respiration follows a generalized temperature response across studies, and that the light compensation point of photosynthesis is reduced by elevated growth CO₂. Lastly, we emphasize the need to move beyond single factorial experiments whenever possible, and to combine both CO₂ and temperature treatments in studies of tree performance.     

Modelling the recent historical impacts of atmospheric CO2 and climate change on Mediterranean vegetation

During the past century, annual mean temperature has increased by 0.75°C and precipitation has shown marked variation throughout the Mediterranean basin. These historical climate changes may have had significant, but presently undefined, impacts on the productivity and structure of sclerophyllous shrubland, an important vegetation type in the region. We used a vegetation model for this functional type to examine climate change impacts, and their interaction with the concurrent historical rise in atmospheric CO₂. Using only climate and soil texture as data inputs, model predictions showed good agreement with observations of seasonal and regional variation in leaf and canopy physiology, net primary productivity (NPP), leaf area index (LAI) and soil water. Model simulations for shrubland sites indicated that potential NPP has risen by 25% and LAI by 7% during the past century, although the absolute increase in LAI was small. Sensitivity analysis suggested that the increase in atmospheric CO₂ since 1900 was the primary cause of these changes, and that simulated climate change alone had negative impacts on both NPP and LAI. Effects of rising CO₂ were mediated by significant increases in the efficiency of water‐use in NPP throughout the region, as a consequence of the direct effect of CO₂ on leaf gas exchange. This increase in efficiency compensated for limitation of NPP by drought, except in areas where drought was most severe. However, while water was used more efficiently, total canopy water loss rose slightly or remained unaffected in model simulations, because increases in LAI with CO₂ counteracted the effects of reduced stomatal conductance on transpiration. Model simulations for the Mediterranean region indicate that the recent rise in atmospheric CO₂ may already have had significant impacts on productivity, structure and water relations of sclerophyllous shrub vegetation, which tended to offset the detrimental effects of climate change in the region.     

Increased water‐use efficiency during the 20th century did not translate into enhanced tree growth

Aim The goals of this study are: (1) to determine whether increasing atmospheric CO₂ concentrations and changing climate increased intrinsic water use efficiency (iWUE, as detected by changes in Δ¹³C) over the last four decades; and if it did increase iWUE, whether it led to increased tree growth (as measured by tree‐ring growth); (2) to assess whether CO₂ responses are biome dependent due to different environmental conditions, including availability of nutrients and water; and (3) to discuss how the findings of this study can better inform assumptions of CO₂ fertilization and climate change effects in biospheric and climate models. Location A global range of sites covering all major forest biome types. Methods The analysis encompassed 47 study sites including boreal, wet temperate, mediterranean, semi‐arid and tropical biomes for which measurements of tree ring Δ¹³C and growth are available over multiple decades. Results The iWUE inferred from the Δ¹³C analyses of comparable mature trees increased 20.5% over the last 40 years with no significant differences between biomes. This increase in iWUE did not translate into a significant overall increase in tree growth. Half of the sites showed a positive trend in growth while the other half had a negative or no trend. There were no significant trends within biomes or among biomes. Main conclusions These results show that despite an increase in atmospheric CO₂ concentrations of over 50 p.p.m. and a 20.5% increase in iWUE during the last 40 years, tree growth has not increased as expected, suggesting that other factors have overridden the potential growth benefits of a CO₂‐rich world in many sites. Such factors could include climate change (particularly drought), nutrient limitation and/or physiological long‐term acclimation to elevated CO₂. Hence, the rate of biomass carbon sequestration in tropical, arid, mediterranean, wet temperate and boreal ecosystems may not increase with increasing atmospheric CO₂ concentrations as is often implied by biospheric models and short‐term elevated CO₂ experiments.     

Woody biomass production during the second rotation of a bio-energy Populus plantation increases in a future high CO2 world

The quickly rising atmospheric carbon dioxide (CO₂)‐levels, justify the need to explore all carbon (C) sequestration possibilities that might mitigate the current CO₂ increase. Here, we report the likely impact of future increases in atmospheric CO₂ on woody biomass production of three poplar species (Populus alba L. clone 2AS‐11, Populus nigra L. clone Jean Pourtet and Populus×euramericana clone I‐214). Trees were growing in a high‐density coppice plantation during the second rotation (i.e., regrowth after coppice; 2002–2004; POPFACE/EUROFACE). Six plots were studied, half of which were continuously fumigated with CO₂ (FACE; free air carbon dioxide enrichment of 550 ppm). Half of each plot was fertilized to study the interaction between CO₂ and nutrient fertilization. At the end of the second rotation, selective above‐ and belowground harvests were performed to estimate the productivity of this bio‐energy plantation. Fertilization did not affect growth of the poplar trees, which was likely because of the high rates of fertilization during the previous agricultural land use. In contrast, elevated CO₂ enhanced biomass production by up to 29%, and this stimulation did not differ between above‐ and belowground parts. The increased initial stump size resulting from elevated CO₂ during the first rotation (1999–2001) could not solely explain the observed final biomass increase. The larger leaf area index after canopy closure and the absence of any major photosynthetic acclimation after 6 years of fumigation caused the sustained CO₂‐induced biomass increase after coppice. These results suggest that, under future CO₂ concentrations, managed poplar coppice systems may exhibit higher potential for C sequestration and, thus, help mitigate climate change when used as a source of C‐neutral energy.     

Effects of Warming, Summer Drought, and CO2 Enrichment on Aboveground Biomass Production, Flowering Phenology, and Community Structure in an Upland Grassland Ecosystem

Future climate scenarios predict simultaneous changes in environmental conditions, but the impacts of multiple climate change drivers on ecosystem structure and function remain unclear. We used a novel experimental approach to examine the responses of an upland grassland ecosystem to the 2080 climate scenario predicted for the study area (3.5°C temperature increase, 20% reduction in summer precipitation, atmospheric CO₂ levels of 600 ppm) over three growing seasons. We also assessed whether patterns of grassland response to a combination of climate change treatments could be forecast by ecosystem responses to single climate change drivers. Effects of climate change on aboveground production showed considerable seasonal and interannual variation; April biomass increased in response to both warming and the simultaneous application of warming, summer drought, and CO₂ enrichment, whereas October biomass responses were either non-significant or negative depending on the year. Negative impacts of summer drought on production were only observed in combination with a below-average rainfall regime, and showed lagged effects on spring biomass. Elevated CO₂ had no significant effect on aboveground biomass during this study. Both warming and the 2080 climate change scenario were associated with a significant advance in flowering time for the dominant grass species studied. However, flowering phenology showed no significant response to either summer drought or elevated CO₂. Species diversity and equitability showed no response to climate change treatments throughout this study. Overall, our data suggest that single-factor warming experiments may provide valuable information for projections of future ecosystem changes in cool temperate grasslands.     

Gas exchange and water‐use efficiency in plant canopies

In this review, I first address the basics of gas exchange, water‐use efficiency and carbon isotope discrimination in C₃ plant canopies. I then present a case study of water‐use efficiency in northern Australian tree species. In general, C₃ plants face a trade‐off whereby increasing stomatal conductance for a given set of conditions will result in a higher CO₂ assimilation rate, but a lower photosynthetic water‐use efficiency. A common garden experiment suggested that tree species which are able to establish and grow in drier parts of northern Australia have a capacity to use water rapidly when it is available through high stomatal conductance, but that they do so at the expense of low water‐use efficiency. This may explain why community‐level carbon isotope discrimination does not decrease as steeply with decreasing rainfall on the North Australian Tropical Transect as has been observed on some other precipitation gradients. Next, I discuss changes in water‐use efficiency that take place during leaf expansion in C₃ plant leaves. Leaf phenology has recently been recognised as a significant driver of canopy gas exchange in evergreen forest canopies, and leaf expansion involves changes in both photosynthetic capacity and water‐use efficiency. Following this, I discuss the role of woody tissue respiration in canopy gas exchange and how photosynthetic refixation of respired CO₂ can increase whole‐plant water‐use efficiency. Finally, I discuss the role of water‐use efficiency in driving terrestrial plant responses to global change, especially the rising concentration of atmospheric CO₂. In coming decades, increases in plant water‐use efficiency caused by rising CO₂ are likely to partially mitigate impacts on plants of drought stress caused by global warming.     

Net biome production of the Amazon Basin in the 21st century

Global change includes multiple stressors to natural ecosystems ranging from direct climate and land‐use impacts to indirect degradation processes resulting from fire. Humid tropical forests are vulnerable to projected climate change and possible synergistic interactions with deforestation and fire, which may initiate a positive feedback to rising atmospheric CO₂. Here, we present results from a multifactorial impact analysis that combined an ensemble of climate change models with feedbacks from deforestation and accidental fires to quantify changes in Amazon Basin carbon cycling. Using the LPJmL Dynamic Global Vegetation Model, we modelled spatio‐temporal changes in net biome production (NBP); the difference between carbon fluxes from fire, deforestation, soil respiration and net primary production. By 2050, deforestation and fire (with no CO₂ increase or climate change) resulted in carbon losses of 7.4–20.3 Pg C with the range of uncertainty depending on socio‐economic storyline. During the same time period, interactions between climate and land use either compensated for carbon losses due to wetter climate and CO₂ fertilization or exacerbated carbon losses from drought‐induced forest mortality (?20.1 to +4.3 Pg C). By the end of the 21st century, depending on climate projection and the rate of deforestation (including its interaction with fire), carbon stocks either increased (+12.6 Pg C) or decreased (?40.6 Pg C). The synergistic effect of deforestation and fire with climate change contributed up to 26–36 Pg C of the overall decrease in carbon stocks. Agreement between climate projections (n=9), not accounting for deforestation and fire, in 2050 and 2098 was relatively low for the directional change in basin‐wide NBP (19–37%) and aboveground live biomass (13–24%). The largest uncertainty resulted from climate projections, followed by implementation of ecosystem dynamics and deforestation. Our analysis partitions the drivers of tropical ecosystem change and is relevant for guiding mitigation and adaptation policy related to global change.     

Canopy leaf area of a mature evergreen Eucalyptus woodland does not respond to elevated atmospheric [CO2] but tracks water availability

Canopy leaf area, quantified by the leaf area index (L), is a crucial driver of forest productivity, water use and energy balance. Because L responds to environmental drivers, it can represent an important feedback to climate change, but its responses to rising atmospheric [CO₂] and water availability of forests have been poorly quantified. We studied canopy leaf area dynamics for 28 months in a native evergreen Eucalyptus woodland exposed to free‐air CO₂ enrichment (the EucFACE experiment), in a subtropical climate where water limitation is common. We hypothesized that, because of expected stimulation of productivity and water‐use efficiency, L should increase with elevated [CO₂]. We estimated L from diffuse canopy transmittance, and measured monthly leaf litter production. Contrary to expectation, L did not respond to elevated [CO₂]. We found that L varied between 1.10 and 2.20 across the study period. The dynamics of L showed a quick increase after heavy rainfall and a steady decrease during periods of low rainfall. Leaf litter production was correlated to changes in L, both during periods of decreasing L (when no leaf growth occurred) and during periods of increasing L (active shedding of old foliage when new leaf growth occurred). Leaf lifespan, estimated from mean L and total annual litter production, was up to 2 months longer under elevated [CO₂] (1.18 vs. 1.01 years; P = 0.05). Our main finding that L was not responsive to elevated CO₂ is consistent with other forest FACE studies, but contrasts with the positive response of L commonly predicted by many ecosystem models.     

Conifer stem growth at the altitudinal treeline in response to four years of CO2 enrichment

Northern latitude and upper altitude climatic treelines have received increasing attention given their potential sensitivity to atmospheric and climate change. While greater radial stem growth at treeline sites in recent decades has been attributed largely to increasing temperature, rising atmospheric CO₂ concentration may also be contributing to this growth stimulation. Tree ring increments of mature Larix decidua and Pinus uncinata were measured over 4 years in a free air CO₂ enrichment experiment at treeline in the Swiss Central Alps (2180 m a.s.l.). In addition, a one‐time defoliation treatment in the second year (2002) of the experiment was used to simulate one of the common natural insect outbreak events. In response to elevated atmospheric CO₂, Larix showed a cumulative 4‐year growth response of+41%, with particularly strong responses in the third and fourth year. This increase in radial stem wood growth was the result of more latewood production, in particular, the formation of larger tracheids, rather than a greater number of cells. In contrast, Pinus showed no change in ring width to elevated [CO₂], neither in each of the treatment years, nor in the cumulative response over 4 years, although an increase in tracheid size was observed in the third year. Defoliation led to a pronounced decrease in annual ring width of both species, marked in particular by less latewood production, in the treatment, as well as subsequent years. There was no significant interaction between defoliation and CO₂ enrichment. Although Pinus showed no growth response to CO₂, the positive growth response observed in Larix after 4 years of CO₂ enrichment implies that the sensitivity of treeline trees to global change may not be purely temperature driven. We conclude that the open sparse canopy in the treeline ecotone favours the indeterminate growth strategy of the early successional Larix when neither weather nor carbon are limiting, whereas the later successional Pinus does not show any indication of more vigorous growth under future higher atmospheric CO₂ concentrations.     

Remote sensing of heterogeneity in photosynthetic efficiency, electron transport and dissipation of excess light in Populus deltoides stands under ambient and elevated CO2 concentrations, and in a tropical forest canopy, using a new laser-induced fluorescence transient device

Determining the spatial and temporal diversity of photosynthetic processes in forest canopies presents a challenge to the evaluation of biological feedbacks needed for improvement of carbon and climate models. Limited access with portable instrumentation, especially in the outer canopy, makes remote sensing of these processes a priority in experimental ecosystem and climate change research. Here, we describe the application of a new, active, chlorophyll fluorescence measurement system for remote sensing of light use efficiency, based on analysis of laser‐induced fluorescence transients (LIFT). We used mature stands of Populus grown at ambient (380 ppm) and elevated CO₂ (1220 ppm) in the enclosed agriforests of the Biosphere 2 Laboratory (B2L) to compare parameters of photosynthetic efficiency, photosynthetic electron transport, and dissipation of excess light measured by LIFT and by standard on‐the‐leaf saturating flash methods using a commercially available pulse‐modulated chlorophyll fluorescence instrument (Mini‐PAM). We also used LIFT to observe the diel courses of these parameters in leaves of two tropical forest dominants, Inga and Pterocarpus, growing in the enclosed model tropical forest of B2L. Midcanopy leaves of both trees showed the expected relationships among chlorophyll fluorescence‐derived photosynthetic parameters in response to sun exposure, but, unusually, both displayed an afternoon increase in nonphotochemical quenching in the shade, which was ascribed to reversible inhibition of photosynthesis at high leaf temperatures in the enclosed canopy. Inga generally showed higher rates of photosynthetic electron transport, but greater afternoon reduction in photosynthetic efficiency. The potential for estimation of the contribution of outer canopy photosynthesis to forest CO₂ assimilation, and assessment of its response to environmental stress using remote sensing devices such as LIFT, is briefly discussed.     

Long‐term increases in intrinsic water‐use efficiency do not lead to increased stem growth in a tropical monsoon forest in western Thailand

Rising atmospheric carbon dioxide [CO₂] can accelerate tree growth by stimulating photosynthesis and increasing intrinsic water‐use efficiency (iWUE). Little evidence exists, however, for the long‐term growth and gas‐exchange responses of mature trees in tropical forests to the combined effects of rising [CO₂] and other global changes such as warming. Using tree rings and stable isotopes of carbon and oxygen, we investigated long‐term trends in the iWUE and stem growth (basal area increment, BAI) of three canopy tree species in a tropical monsoon forest in western Thailand (Chukrasia tabularis, Melia azedarach, and Toona ciliata). To do this, we modelled the contribution of ontogenetic effects (tree diameter or age) and calendar year to variation in iWUE, oxygen isotopes, and BAI using mixed‐effects models. Although iWUE increased significantly with both tree diameter and calendar year in all species, BAI at a given tree diameter was lower in more recent years. For one species, C. tabularis, differences in crown dominance significantly influence stable isotopes and growth. Tree ring Δ¹⁸O increased with calendar year in all species, suggesting that increasing iWUE may have been driven by relatively greater reductions in stomatal conductance – leading to enrichment in Δ¹⁸O – than increases in photosynthetic capacity. Plausible explanations for the observed declines in growth include water stress resulting from rising temperatures and El Niño events, increased respiration, changes in allocation, or more likely, a combination of these factors.