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.     

Tree responses to rising CO2 in field experiments: implications for the future forest

The need to assess the role of forests in the global cycling of carbon and how that role will change as the atmospheric concentration of CO₂ increases has spawned many experiments over a range of scales. Experiments using open-top chambers have been established at many sites to test whether the short-term responses of tree seedlings described in controlled environments would be sustained over several growing seasons under field conditions. Here we review the results of those experiments, using the framework of the interacting cycles of carbon, water and nutrients, because that is the framework of the ecosystem models that are being used to address the decades-long response of forests. Our analysis suggests that most of what was learned in seedling studies was qualitatively correct. The evidence from field-grown trees suggests a continued and consistent stimulation of photosynthesis of about 60% for a 300 p.p.m. increase in [CO₂], and there is little evidence of the long-term loss of sensitivity to CO₂ that was suggested by earlier experiments with tree seedlings in pots. Despite the importance of respiration to a tree's carbon budget, no strong scientific consensus has yet emerged concerning the potential direct or acclimation response of woody plant respiration to CO₂ enrichment. The relative effect of CO₂ on above-ground dry mass was highly variable and greater than that indicated by most syntheses of seedling studies. Effects of CO₂ concentration on static measures of response are confounded with the acceleration of ontogeny observed in elevated CO₂. The trees in these open-top chamber experiments were in an exponential growth phase, and the large growth responses to elevated CO₂ resulted from the compound interest associated with an increasing leaf area. This effect cannot be expected to persist in a closed-canopy forest where growth potential is constrained by a steady-state leaf area index. A more robust and informative measure of tree growth in these experiments is the annual increment in wood mass per unit leaf area, which increased 27% in elevated CO₂. There is no support for the conclusion from many studies of seedlings that root-to-shoot ratio is increased by elevated CO₂; the production of fine roots may be enhanced, but it is not clear that this response would persist in a forest. Foliar nitrogen concentrations were lower in CO₂-enriched trees, but to a lesser extent than was indicated in seedling studies and only when expressed on a leaf mass basis. The prediction that leaf litter C/N ratio would increase was not supported in field experiments. Also contrasting with seedling studies, there is little evidence from the field studies that stomatal conductance is consistently affected by CO₂; however, this is a topic that demands more study. Experiments with trees in open-top chambers under field conditions have provided data on longer-term, larger-scale responses of trees to elevated CO₂ under field conditions, confirmed some of the conclusions from previous seedling studies, and challenged other conclusions. There remain important obstacles to using these experimental results to predict forest responses to rising CO₂, but the studies are valuable nonetheless for guiding ecosystem model development and revealing the critical questions that must be addressed in new, larger-scale CO₂ experiments.     

System-level adjustments to elevated CO2 in model spruce ecosystems

Atmospheric carbon dioxide enrichment and increasing nitrogen deposition are often predicted to increase forest productivity based on currently available data for isolated forest tree seedlings or their leaves. However, it is highly uncertain whether such seedling responses will scale to the stand level. Therefore, we studied the effects of increasing CO₂ (280, 420 and 560 μL L^-1) and increasing rates of wet N deposition (0, 30 and 90 kg ha^-1 y^-1) on whole stands of 4-year-old spruce trees (Picea abies). One tree from each of six clones, together with two herbaceous understory species, were established in each of nine 0.7 m² model ecosystems in nutrient poor forest soil and grown in a simulated montane climate for two years. Shoot level light-saturated net photosynthesis measured at growth CO₂ concentrations increased with increasing CO₂, as well as with increasing N deposition. However, predawn shoot respiration was unaffected by treatments. When measured at a common CO₂ concentration of 420 μL L^-1 37% down-regulation of photosynthesis was observed in plants grown at 560 μL CO₂ L^-1. Length growth of shoots and stem diameter were not affected by CO₂ or N deposition. Bud burst was delayed, leaf area index (LAI) was lower, needle litter fall increased and soil CO₂ efflux increased with increasing CO₂. N deposition had no effect on these traits. At the ecosystem level the rate of net CO₂ exchange was not significantly different between CO₂ and N treatments. Most of the responses to CO₂ studied here were nonlinear with the most significant differences between 280 and 420 μL CO₂ L^-1 and relatively small changes between 420 and 560 μL CO₂ L^-1. Our results suggest that the lack of above-ground growth responses to elevated CO₂ is due to the combined effects of physiological down-regulation of photosynthesis at the leaf level, allometric adjustment at the canopy level (reduced LAI), and increasing strength of below-ground carbon sinks. The non-linearity of treatment effects further suggests that major responses of coniferous forests to atmospheric CO₂ enrichment might already be under way and that future responses may be comparatively smaller.     

Foliar gas exchange responses of two deciduous hardwoods during 3 years of growth in elevated CO2: no loss of photosynthetic enhancement

Responses of photosynthesis and stomatal conductance were monitored throughout a 3-year field exposure of Liriodendron tulipifera (yellow-poplar) and Quercus alba (white oak) to elevated concentrations of atmospheric CO₂. Exposure to atmospheres enriched with +150 and +300 umol mol^-1 CO₂ increased net photosynthesis by 12–144% over the course of the study. Net photosynthesis was consistently higher at +300 than at +150 umol mol^-1 CO₂. The effect of CO₂ enrichment on stomatal conductance was limited, but instantaneous leaf-level water use efficiency increased significantly. No decrease in the responsiveness of photosynthesis to CO₂ enrichment over time was detected, and the responses were consistent throughout the canopy and across successive growth flushes and seasons. The relationships between internal CO₂ concentration and photosynthesis (e.g. photosynthetic capacity and carboxylation efficiency) were not altered by growth at elevated concentrations of CO₂. No alteration in the timing of leaf senescence or abscission was detected, suggesting that the seasonal duration of effective gas-exchange was unaffected by CO₂ treatment. These results are consistent with data previously reported for these species in controlled-environment studies, and suggest that leaf-level photosynthesis does not down-regulate in these species as a result of acclimation to CO₂ enrichment in the field. This sustained enhancement of photosynthesis provides the opportunity for increased growth and carbon storage by trees as the atmospheric concentration of CO₂ rises, but many additional factors interact in determining whole-plant and forest responses to global change.     

The Direct Effect of Increased CO2 on Gas Exchange and Growth of Forest Tree Species

CO₂ enrichment of the atmosphere is now well documented andits effect on the growth of world forests is being questionedby the scientific community. The direct effects of increasedCO₂ on tree species are reviewed: the different experimentalapproaches are described, as well as the principal results alreadyobtained. Short-term experiments have shown an increased photosyntheticrate, as predicted by leaf models. In longer experiments thisincrease is reduced after a few weeks or months by mechanismsthat remain to be found. Elevated CO₂ seems to decrease thedark respiration rate, but the results are still controversial.Biomass partitioning in elevated CO₂ is clearly related to themineral supply of the trees: An increase in root investmentin elevated CO₂ is related to a poor mineral status. The mineralcontent of trees grown in elevated CO2 is generally loweredcompared to controls. No general rule has yet been found forthe effect of increased CO₂ on leaf area development. The paper emphasizes large areas of ignorance: the reasons forthe different responses of different species, which may be relatedto their developmental strategies, are largely ignored. Muchexperimental effort is needed to parameterize all the physiologicalprocesses which are susceptible to change with an increase inatmospheric CO₂, leading to a change in forest tree growth. Key words: Elevated CO₂, tree, forest, photosynthesis, respiration, biomass, partitioning, mineral nutrition     

高浓度CO2对树木生理生态的影响研究进展

大气CO2浓度升高已成为世界范围内的重要环境问题。CO2浓度升高势必会对植物的生理生态变化产生重要影响。综述了国内外有关高浓度CO2对树木生理生态影响研究的最新进展,具体包括高浓度CO2对树木生长发育、光合和呼吸作用、抗氧化系统、树木代谢物质、挥发性有机化合物以及树木凋落物等方面的影响。高浓度CO2一般会促进树木地上植株的生长和发育,但也因树种差异而有所不同。最新研究表明,高浓度CO2促进了树木细根周转,树木根系生长在大气CO2浓度升高条件下表现为促进作用,这种作用加快了全球森林生态系统的C循环。高浓度CO2虽然在一定程度上促进树木光合速率的增加,但长期熏蒸也往往会发生光合驯化,这种现象产生的生理学机制目前仍无定论。高浓度CO2对树木呼吸作用尤其是根系呼吸的影响将是未来研究的重点和难点。高浓度CO2一般会提高树木抗氧化酶活性与抗氧化剂含量,但不同树种响应高浓度CO2的过程和机理也有所差异。研究表明,高浓度CO2一般对树木凋落物的分解产生不利影响,但也因树种而异。需要强调的是,目前关于树木地下部分、树木对高浓度CO2的适应机理和重要过程(碳氮水耦合及基因调控等)以及多个树种包括不同类型树种及不同品种之间比较研究较少;关于某一重要生理生态机制(如根系生理代谢)尤其是多个生态因子复合条件下缺乏长期深入的研究。在此基础上给出了大气CO2浓度升高下树木生理生态学研究的未来发展方向,包括高CO2浓度条件下树木根系生理代谢及机制、树木碳氮水耦合的生理过程及机制、不同生态因子复合作用对树木生理影响机制以及树木分子作用机理等方面的研究。这些研究不仅将丰富森林树木应对未来气候变化的有关科学理论,也为全球气候变化背景下实现森林树种生态功能的优化选择及森林生态系统的可持续发展与经营提供重要的生理生态学理论依据和参考。     

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.     

Differential anatomical responses to elevated CO2 in saplings of four hardwood species

To determine whether an elevated carbon dioxide concentration ([CO₂]) can induce changes in the wood structure and stem radial growth in forest trees, we investigated the anatomical features of conduit cells and cambial activity in 4‐year‐old saplings of four deciduous broadleaved tree species – two ring‐porous (Quercus mongolica and Kalopanax septemlobus) and two diffuse‐porous species (Betula maximowicziana and Acer mono) – grown for three growing seasons in a free‐air CO₂ enrichment system. Elevated [CO₂] had no effects on vessels, growth and physiological traits of Q. mongolica, whereas tree height, photosynthesis and vessel area tended to increase in K. septemlobus. No effects of [CO₂] on growth, physiological traits and vessels were seen in the two diffuse‐porous woods. Elevated [CO₂] increased larger vessels in all species, except B. maximowicziana and number of cambial cells in two ring‐porous species. Our results showed that the vessel anatomy and radial stem growth of Q. mongolica, B. maximowicziana and A. mono were not affected by elevated [CO₂], although vessel size frequency and cambial activity in Q. mongolica were altered. In contrast, changes in vessel anatomy and cambial activity were induced by elevated [CO₂] in K. septemlobus. The different responses to elevated [CO₂] suggest that the sensitivity of forest trees to CO₂ is species dependent.     

A simple method for testing leaf responses of tall tropical forest trees to elevated CO2

The effects of atmospheric CO₂ enrichment on mature trees in their natural environment are largely unknown. Here we present a new, and inexpensive technique which can be used in situ to address some key physiological questions related to the CO₂ problem. Small, light-weight cups mounted on the lower side of rigid leaves at the top of tall tropical forest trees were supplied with CO₂-enriched air derived from a low-technology air mixing device utilizing forest floor CO₂ evolution. We present the scientific rationale for such field experiments, technical details, an assessment of potential cup artifacts and first results illustrating effects of elevated CO₂ on stomata and carbohydrate accumulation in the canopies of mature trees.     

First-year growth response of trees in an intact forest exposed to elevated CO2

Although elevated atmospheric CO₂ has been shown to increase growth of tree seedlings and saplings, the response of intact forest ecosystems and established trees is unclear. We report results from the first large-scale experimental system designed to study the effects of elevated CO₂ on an intact forest with the full complement of species interactions and environmental stresses. During the first year of exposure to ^ 1.5 Ë ambient CO₂, canopy loblolly pine (Pinus taeda, L.) trees increased basal area growth rate by 24% but understorey trees of loblolly pine, sweetgum (Liquidambar styraciflua L.), and red maple (Acer rubrum L.) did not respond. Winged elm (Ulmus alata Michx.) had a marginally significant increase in growth rate (P = 0.069). These data suggest that this ecosystem has the capacity to respond immediately to a step increase in atmospheric CO₂; however, as exposure time increases, nutrient limitations may reduce this initial growth stimulation.     

Responses of N fluxes and pools to elevated atmospheric CO2 in model forest ecosystems with acidic and calcareous soils

The objectives of this study were to estimate how soil type, elevated N deposition (0.7 vs. 7 g N m^–2y^–1) and tree species influence the potential effects of elevated CO₂ (370 vs. 570 mol CO₂ mol^–1) on N pools and fluxes in forest soils. Model spruce-beech forest ecosystems were established on a nutrient-rich calcareous sand and on a nutrient-poor acidic loam in large open-top chambers. In the fourth year of treatment, we measured N concentrations in the soil solution at different depths, estimated N accumulation by ion exchange resin (IER) bags, and quantified N export in drainage water, denitrification, and net N uptake by trees. Under elevated CO₂, concentrations of N in the soil solution were significantly reduced. In the nutrient-rich calcareous sand, CO₂ enrichment decreased N concentrations in the soil solution at all depths (–45 to –100%). In the nutrient-poor acidic loam, the negative CO₂ effect was restricted to the uppermost 5 cm of the soil. Increasing the N deposition stimulated the negative impact of CO₂ enrichment on soil solution N in the acidic loam at 5 cm depth from –20% at low N inputs to –70% at high N inputs. In the nutrient-rich calcareous sand, N additions did not influence the CO₂ effect on soil solution N. Accumulation of N by IER bags, which were installed under individual trees, was decreased at high CO₂ levels under spruce in both soil types. Under beech, this decrease occurred only in the calcareous sand. N accumulation by IER bags was negatively correlated with current-years foliage biomass, suggesting that the reduction of soil N availability indices was related to a CO₂-induced growth enhancement. However, the net N uptake by trees was not significantly increased by elevated CO₂. Thus, we suppose that the reduced N concentrations in the soil solution at elevated CO₂ concentrations were rather caused by an increased N immobilisation in the soil. Denitrification was not influenced by atmospheric CO₂ concentrations. CO₂ enrichment decreased nitrate leaching in drainage by 65%, which suggests that rising atmospheric CO₂ potentially increases the N retention capacity of forest ecosystems.     

Growth and phenology of mature temperate forest trees in elevated CO2

Are mature forest trees carbon limited at current CO₂ concentrations? Will ‘mid‐life’, 35 m tall deciduous trees grow faster in a CO₂‐enriched atmosphere? To answer these questions we exposed ca. 100‐year‐old temperate forest trees at the Swiss Canopy Crane site near Basel, Switzerland to a ca. 540 ppm CO₂ atmosphere using web‐FACE technology. Here, we report growth responses to elevated CO₂ for 11 tall trees (compared with 32 controls) of five species during the initial four treatment years. Tested across all trees, there was no CO₂ effect on stem basal area (BA) increment (neither when tested per year nor cumulatively for 4 years). In fact, the 4th year means were almost identical for the two groups. Stem growth data were standardized by pretreatment growth (5 years) in order to account for a priori individual differences in vigor. Although this experiment was not designed to test species specific effects, one species, the common European beech, Fagus sylvatica, showed a significant growth enhancement in the first year, which reoccurred during a centennial drought in the third year. None of the other dominant species (Quercus petraea, Carpinus betulus) showed a growth response to CO₂ in any of the 4 years or for all years together. The inclusion or exclusion of single individuals of Prunus avium and Tilia platyphyllos did not change the picture. In elevated CO₂, lateral branching in terminal shoots was higher in Fagus in 2002, when shoots developed from buds that were formed during the first season of CO₂ enrichment (2001), but there was no effect in later years and no change in lateral branching in any of the other species. In Quercus, there was a steady stimulation of leading shoot length in high‐CO₂ trees. Phenological variables (bud break, leaf fall, leaf duration) were highly species specific and were not affected by elevated CO₂ in any consistent way. Our 4‐year data set reflects a very dynamic and species‐specific response of tree growth to a step change in CO₂ supply. Stem growth after 4 years of exposure does not support the notion that mature forest trees will accrete wood biomass at faster rates in a future CO₂‐enriched atmosphere.     

Elevated CO<Subscript>2</Subscript> reduces sap flux in mature deciduous forest trees

We enriched in CO₂ the canopy of 14 broad-leaved trees in a species-rich, ca. 30-m-tall forest in NW Switzerland to test whether elevated CO₂ reduces water use in mature forest trees. Measurements of sap flux density (J_S) were made prior to CO₂ enrichment (summer 2000) and throughout the first whole growing season of CO₂ exposure (2001) using the constant heat-flow technique. The short-term responses of sap flux to brief (1.5–3 h) interruptions of CO₂ enrichment were also examined. There were no significant a priori differences in morphological and physiological traits between trees which were later exposed to elevated CO₂ (n=14) and trees later used as controls (n=19). Over the entire growing season, CO₂ enrichment resulted in an average 10.7% reduction in mean daily J_S across all species compared to control trees. Responses were most pronounced in Carpinus, Acer, Prunus and Tilia, smaller in Quercus and close to zero in Fagus trees. The J_S of treated trees significantly increased by 7% upon transient exposure to ambient CO₂ concentrations at noon. Hence, responses of the different species were, in the short term, similar in magnitude to those observed over the whole season (though opposite because of the reversed treatment). The reductions in mean J_S of CO₂-enriched trees were high (22%) under conditions of low evaporative demand (vapour pressure deficit, VPD <5 hPa) and small (2%) when mean daily VPD was greater than 10 hPa. During a relatively dry period, the effect of elevated CO₂ on J_S even appeared to be reversed. These results suggest that daily water savings by CO₂-enriched trees may have accumulated to a significantly improved water status by the time when control trees were short of soil moisture. Our data indicate that the magnitude of CO₂ effects on stand transpiration will depend on rainfall regimes and the relative abundance of the different species, being more pronounced under humid conditions and in stands dominated by species such as Carpinus and negligible in mono-specific Fagus forests.     

Water relations in grassland and desert ecosystems exposed to elevated atmospheric CO2

Atmospheric CO₂ enrichment may stimulate plant growth directly through (1) enhanced photosynthesis or indirectly, through (2) reduced plant water consumption and hence slower soil moisture depletion, or the combination of both. Herein we describe gas exchange, plant biomass and species responses of five native or semi-native temperate and Mediterranean grasslands and three semi-arid systems to CO₂ enrichment, with an emphasis on water relations. Increasing CO₂ led to decreased leaf conductance for water vapor, improved plant water status, altered seasonal evapotranspiration dynamics, and in most cases, periodic increases in soil water content. The extent, timing and duration of these responses varied among ecosystems, species and years. Across the grasslands of the Kansas tallgrass prairie, Colorado shortgrass steppe and Swiss calcareous grassland, increases in aboveground biomass from CO₂ enrichment were relatively greater in dry years. In contrast, CO₂-induced aboveground biomass increases in the Texas C₃/C₄ grassland and the New Zealand pasture seemed little or only marginally influenced by yearly variation in soil water, while plant growth in the Mojave Desert was stimulated by CO₂ in a relatively wet year. Mediterranean grasslands sometimes failed to respond to CO₂-related increased late-season water, whereas semiarid Negev grassland assemblages profited. Vegetative and reproductive responses to CO₂ were highly varied among species and ecosystems, and did not generally follow any predictable pattern in regard to functional groups. Results suggest that the indirect effects of CO₂ on plant and soil water relations may contribute substantially to experimentally induced CO₂-effects, and also reflect local humidity conditions. For landscape scale predictions, this analysis calls for a clear distinction between biomass responses due to direct CO₂ effects on photosynthesis and those indirect CO₂ effects via soil moisture as documented here.     

Predicting tree biomass growth in the temperate–boreal ecotone: Is tree size,age, competition,or climate response most important?

As global temperatures rise, variation in annual climate is also changing, with unknown consequences for forest biomes. Growing forests have the ability to capture atmospheric CO₂ and thereby slow rising CO₂ concentrations. Forests’ ongoing ability to sequester C depends on how tree communities respond to changes in climate variation. Much of what we know about tree and forest response to climate variation comes from tree‐ring records. Yet typical tree‐ring datasets and models do not capture the diversity of climate responses that exist within and among trees and species. We address this issue using a model that estimates individual tree response to climate variables while accounting for variation in individuals’ size, age, competitive status, and spatially structured latent covariates. Our model allows for inference about variance within and among species. We quantify how variables influence aboveground biomass growth of individual trees from a representative sample of 15 northern or southern tree species growing in a transition zone between boreal and temperate biomes. Individual trees varied in their growth response to fluctuating mean annual temperature and summer moisture stress. The variation among individuals within a species was wider than mean differences among species. The effects of mean temperature and summer moisture stress interacted, such that warm years produced positive responses to summer moisture availability and cool years produced negative responses. As climate models project significant increases in annual temperatures, growth of species like Acer saccharum, Quercus rubra, and Picea glauca will vary more in response to summer moisture stress than in the past. The magnitude of biomass growth variation in response to annual climate was 92–95% smaller than responses to tree size and age. This means that measuring or predicting the physical structure of current and future forests could tell us more about future C dynamics than growth responses related to climate change alone.     

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.     

Mangrove growth in New Zealand estuaries: the role of nutrient enrichment at sites with contrasting rates of sedimentation

Mangrove forest coverage is increasing in the estuaries of the North Island of New Zealand, causing changes in estuarine ecosystem structure and function. Sedimentation and associated nutrient enrichment have been proposed to be factors leading to increases in mangrove cover, but the relative importance of each of these factors is unknown. We conducted a fertilization study in estuaries with different sedimentation histories in order to determine the role of nutrient enrichment in stimulating mangrove growth and forest development. We expected that if mangroves were nutrient-limited, nutrient enrichment would lead to increases in mangrove growth and forest structure and that nutrient enrichment of trees in our site with low sedimentation would give rise to trees and sediments that converged in terms of functional characteristics on control sites in our high sedimentation site. The effects of fertilizing with nitrogen (N) varied among sites and across the intertidal zone, with enhancements in growth, photosynthetic carbon gain, N resorption prior to leaf senescence and the leaf area index of canopies being significantly greater at the high sedimentation sites than at the low sedimentation sites, and in landward dwarf trees compared to seaward fringing trees. Sediment respiration (CO₂ efflux) was higher at the high sedimentation site than at the low one sedimentation site, but it was not significantly affected by fertilization, suggesting that the high sedimentation site supported greater bacterial mineralization of sediment carbon. Nutrient enrichment of the coastal zone has a role in facilitating the expansion of mangroves in estuaries of the North Island of New Zealand, but this effect is secondary to that of sedimentation, which increases habitat area and stimulates growth. In estuaries with high sediment loads, enrichment with N will cause greater mangrove growth and further changes in ecosystem function.     

Towards a better experimental basis for upscaling plant responses to elevated CO2 and climate warming

Few of the most common assumptions used in models of responses of plants and ecosystems to elevated CO₂ and climate warming have been tested under realistic life con-ditions. It is shown that some unexpected discrepancies between predictions and experimental findings exist, suggesting that a better empirical basis is required for predictions. The following ten suggestions may improve our potential to scale up from experimental scales to the real world. (1) Experiments should be timed to account for non-linearity in system responsiveness, asynchrony of responses and developmental differences. (2) By altering mineral nutrient supply, a wide range of CO₂ responses can be ‘produced’, thus requiring realistic soil conditions. (3) Distinctions should be made between ‘doubling CO₂ sup-ply’ and biologically effective degrees of CO₂ enrichment. (4) Because of the non-linearity of plant responses to CO₂, studies of at least three instead of two CO2 concentrations are necessary to describe future trends adequately. (5) Edge effects, in particular unscreened side light, may lead to allometric anomalies, strongly constraining up-scaling to stand-scale CO₂ responses. (6) Variables such as growth, yield, net primary production and C turnover are often confused with carbon pools, carbon sequestration or net ecosystem production. (7) Mono- and interspecific interactions between individuals may lead to completely unpredictable CO₂ responses. (8) Experiments with seedlings benefit from the absence of prehistory effects but are likely to be irrelevant for the responses of larger trees which, on the other hand, may be constrained by carry-over effects. Tree ring research indicates immediate sensitivity of large trees to environmental changes, supporting their usefulness in short-term CO₂-enrichment experiments. (9) In predicting temperature responses, acclimation deserves more attention. (10) The significance of developmental responses is largely under-represented in experimental research, although these responses may overrule many of the other effects of atmospheric change. Results of more realistic experiments which account for these problems will provide a better basis for modelling the future of the biosphere.     

Disentangling the effects of acidic air pollution,atmospheric CO2, and climate change on recent growth of red spruce trees in the Central Appalachian Mountains

In the 45 years after legislation of the Clean Air Act, there has been tremendous progress in reducing acidic air pollutants in the eastern United States, yet limited evidence exists that cleaner air has improved forest health. Here, we investigate the influence of recent environmental changes on the growth and physiology of red spruce (Picea rubens Sarg.) trees, a key indicator species of forest health, spanning three locations along a 100 km transect in the Central Appalachian Mountains. We incorporated a multiproxy approach using 75‐year tree ring chronologies of basal tree growth, carbon isotope discrimination (?¹³C, a proxy for leaf gas exchange), and δ¹⁵N (a proxy for ecosystem N status) to examine tree and ecosystem level responses to environmental change. Results reveal the two most important factors driving increased tree growth since ca. 1989 are reductions in acidic sulfur pollution and increases in atmospheric CO₂, while reductions in pollutant emissions of NO_x and warmer springs played smaller, but significant roles. Tree ring ?¹³C signatures increased significantly since 1989, concurrently with significant declines in tree ring δ¹⁵N signatures. These isotope chronologies provide strong evidence that simultaneous changes in C and N cycling, including greater photosynthesis and stomatal conductance of trees and increases in ecosystem N retention, were related to recent increases in red spruce tree growth and are consequential to ecosystem recovery from acidic pollution. Intrinsic water use efficiency (iWUE) of the red spruce trees increased by ~51% across the 75‐year chronology, and was driven by changes in atmospheric CO₂ and acid pollution, but iWUE was not linked to recent increases in tree growth. This study documents the complex environmental interactions that have contributed to the recovery of red spruce forest ecosystems from pervasive acidic air pollution beginning in 1989, about 15 years after acidic pollutants started to decline in the United States.     

In deep shade, elevated CO2 increases the vigor of tropical climbing plants

Climbing plants have profound influences on tropical forest dynamics and may take particular advantage from atmospheric CO₂ enrichment, thus potentially enhancing tree turnover. Here we test the effect of a four‐step CO₂‐enrichment on growth of three typical Yucatan (Mexico) climbers, across two low photon flux densities, representing typical understory situations. In pairs of two, species of Gonolobus (Asclepiadaceae), Ceratophytum (Bignoniaceae) and Thinouia (Sapindaceae) were grown on Yucatan forest soil in growth cabinets, which simulated the diurnal climate variation. Biomass increased non‐linearly in response to CO₂ enrichment from 280 (preindustrial) to 420 ppm and 560 ppm, but then (700 ppm) leveled off. The relative effect of CO₂‐enrichment between the two lower (280–420 ppm) CO₂ concentrations was 63% at low light (LL == 42 µmol m^2?2 s^2?1), compared to 37% at high light (HL = 87 µmol m^2?2 s^2?1). This overall response of species pairs was the combined effect of linear and non‐linear responses of the individual species across CO₂ treatments. Plant biomass was 61% larger in HL compared to LL. The species‐specific response depended on the neighbor, a species grew with h, irrespective of plant size. Stem length increased, but not consistently across species and light conditions. Specific stem length (SSL, length per dry mass) declined non‐linearly in all three species as CO₂ concentration increased (more pronounced at LL than at HL). SLA (leaf area per unit leaf dry mass) became lower as CO₂ concentration increased (more pronounced in HL). Enhanced vigor of climbers under elevated CO₂ as documented here may accelerate tropical forest dynamics and lead to greater abundance of early succesional tree species. This could reduce forest carbon stocking in the long run.