Flowering phenology in a species-rich temperate grassland is sensitive to warming but not elevated CO2

* Flowering is a critical stage in plant life cycles, and changes might alter processes at the species, community and ecosystem levels. Therefore, likely flowering-time responses to global change drivers are needed for predictions of global change impacts on natural and managed ecosystems. * Here, the impact of elevated atmospheric CO2 concentration ([CO2]) (550 micromol mol(-1)) and warming (+2 masculineC) is reported on flowering times in a native, species-rich, temperate grassland in Tasmania, Australia in both 2004 and 2005. * Elevated [CO2] did not affect average time of first flowering in either year, only affecting three out of 23 species. Warming reduced time to first flowering by an average of 19.1 d in 2004, acting on most species, but did not significantly alter flowering time in 2005, which might be related to the timing of rainfall. Elevated [CO2] and warming treatments did not interact on flowering time. * These results show elevated [CO2] did not alter average flowering time or duration in this grassland; neither did it alter the response to warming. Therefore, flowering phenology appears insensitive to increasing [CO2] in this ecosystem, although the response to warming varies between years but can be strong.     

The effects of elevated CO2 on symbiotic N2 fixation: a link between the carbon and nitrogen cycles in grassland ecosystems

The response of plants to elevated CO₂ is dependent on the availability of nutrients, especially nitrogen. It is generally accepted that an increase in the atmospheric CO₂ concentration increases the C:N ratio of plant residues and exudates. This promotes temporary N-immobilization which might, in turn, reduce the availability of soil nitrogen. In addition, both a CO₂ stimulated increase in plant growth (thus requiring more nitrogen) and an increased N demand for the decomposition of soil residues with a large C:N will result under elevated CO₂ in a larger N-sink of the whole grassland ecosystem. One way to maintain the balance between the C and N cycles in elevated CO₂ would be to increase N-import to the grassland ecosystem through symbiotic N₂ fixation. Whether this might happen in the context of temperate ecosystems is discussed, by assessing the following hypothesis: i) symbiotic N₂ fixation in legumes will be enhanced under elevated CO₂, ii) this enhancement of N₂ fixation will result in a larger N-input to the grassland ecosystem, and iii) a larger N-input will allow the sequestration of additional carbon, either above or below-ground, into the ecosystem. Data from long-term experiments with model grassland ecosystems, consisting of monocultures or mixtures of perennial ryegrass and white clover, grown under elevated CO₂ under free-air or field-like conditions, supports the first two hypothesis, since: i) both the percentage and the amount of fixed N increases in white clover grown under elevated CO₂, ii) the contribution of fixed N to the nitrogen nutrition of the mixed grass also increases in elevated CO₂. Concerning the third hypothesis, an increased nitrogen input to the grassland ecosystem from N₂ fixation usually promotes shoot growth (above-ground C storage) in elevated CO₂. However, the consequences of this larger N input under elevated CO₂ on the below-ground carbon fluxes are not fully understood. On one hand, the positive effect of elevated CO₂ on the quantity of plant residues might be overwhelming and lead to an increased long-term below-ground C storage; on the other hand, the enhancement of the decomposition process by the N-rich legume material might favour carbon turn-over and, hence, limit the storage of below-ground carbon.     

The carbon balance of tropical, temperate and boreal forests

Forest biomes are major reserves for terrestrial carbon, and major components of global primary productivity. The carbon balance of forests is determined by a number of component processes of carbon acquisition and carbon loss, and a small shift in the magnitude of these processes would have a large impact on the global carbon cycle. In this paper, we discuss the climatic influences on the carbon dynamics of boreal, temperate and tropical forests by presenting a new synthesis of micrometeorological, ecophysiological and forestry data, concentrating on three case-study sites. Historical changes in the carbon balance of each biome are also reviewed, and the evidence for a carbon sink in each forest biome and its likely behaviour under future global change are discussed. We conclude that there have been significant advances in determining the carbon balance of forests, but there are still critical uncertainties remaining, particularly in the behaviour of soil carbon stocks.     

Elevated [CO2], temperature increase and N supply effects on the accumulation of below-ground carbon in a temperate grassland ecosystem

The effects of elevated [CO₂] (700 μl l⁻¹ [CO₂]) and temperature increase (+3 °C) on carbon accumulation in a grassland soil were studied at two N-fertiliser supplies (160 and 530 kgN ha⁻¹ year⁻¹) in a long-term experiment (2.5 years) on well established ryegrass swards (Lolium perenne L.,) supplied with the same amounts of irrigation water. For all experimental treatments, the C:N ratio of the top soil organic matter fractions increased with their particle size. Elevated CO₂ concentration increased the C:N ratios of the below-ground phytomass and of the macro-organic matter. A supplemental fertiliser N or a 3 °C increase in elevated [CO₂] reduced it. At the last sampling date, elevated [CO₂] did not affect the C:N ratio of the soil organic matter fractions, but increased significantly the accumulation of roots and of macro-organic matter above 200 μm (MOM). An increased N-fertiliser supply stimulated the accumulation of the non harvested plant phytomass and of the OM between 2 and 50 μm, without positive effect on the macro-organic matter >200 μm. Elevated [CO₂2] increased C accumulation in the OM fractions above 50 μm by +2.1 tC ha⁻¹, on average, whereas increasing the fertiliser N supply led to an average supplemental accumulation of +0.8 tC ha⁻¹. There was no significant effect of a 3 °C temperature increase under elevated [CO₂] on C accumulation in the OM fractions above 50 μm. This revised version was published online in June 2006 with corrections to the Cover Date.     

Insights into mycorrhizal colonisation at elevated CO2: a simple carbon partitioning model

A simulation model was used to investigate the effect of an increased rate of plant photosynthesis at enhanced atmospheric CO2 concentration on a non-leguminous plant-mycorrhizal fungus association. The model allowed the user to modify carbon allocation patterns at three levels: (1) within the plant (shoot–root), (2) between the plant and the mycorrhizal fungus and (3) within the mycorrhizal fungus (intraradical–extraradical structures). Belowground (root and fungus) carbon losses via respiration (and turnover) could also be manipulated. The specific objectives were to investigate the dynamic nature of the potential effects of elevated CO2 on mycorrhizal colonisation and to elucidate some of the various mechanisms by which these effects may be negated. Many of the simulations showed that time (i.e. plant age) had a more significant effect on the observed stimulation of mycorrhizal colonisation by elevated CO2 than changes in carbon allocation patterns or belowground carbon losses. There were two main mechanisms which negated a stimulatory effect of elevated CO2 on internal mycorrhizal colonisation: an increased mycorrhizal carbon allocation to the external hyphal network and an increased rate of mycorrhizal respiration. The results are discussed in relation to real experiments. The need for studies consisting of multiple harvests is emphasised, as is the use of allometric analysis. Implications at the ecosystem level are discussed and key areas for future research are presented.     

Strong ecological but weak evolutionary effects of elevated CO2 on a recombinant inbred population of Arabidopsis thaliana

Increases in atmospheric CO2 concentration have an impact on plant communities by influencing plant growth and morphology, species interactions, and ecosystem processes. These ecological effects may be accompanied by evolutionary change if elevated CO2 (eCO2) alters patterns of natural selection or expression of genetic variation. Here, a statistically powerful quantitative genetic experiment and manipulations of CO2 concentrations in a field setting were used to investigate how eCO2 impacts patterns of selection on ecologically important traits in Arabidopsis thaliana; heritabilities, which influence the rate of response to selection; and genetic covariances between traits, which may constrain responses to selection. CO2 had strong phenotypic effects; plants grown in eCO2 were taller and produced more biomass and fruits. Also, significant directional selection was observed on many traits and significant genetic variation was observed for all traits. However, no evolutionary effect of eCO2 was detected; patterns of selection, heritabilities and genetic correlations corresponded closely in ambient and elevated CO2 environments. The data suggest that patterns of natural selection and the quantitative genetic parameters of this A. thaliana population are robust to increases in CO2 concentration and that responses to eCO2 will be primarily ecological.     

The fate of photosynthetically-fixed carbon in Lolium perenne grassland as modified by elevated CO2 and sward management

Prediction of the impact of climate change requires the response of carbon (C) flow in plant-soil systems to increased CO(2) to be understood. A mechanism by which grassland C sequestration might be altered was investigated by pulse-labelling Lolium perenne swards, which had been subject to CO(2) enrichment and two levels of nitrogen (N) fertilization for 10 yr, with (14)CO(2). Over a 6-d period 40-80% of the (14)C pulse was exported from mature leaves, 1-2% remained in roots, 2-7% was lost as below-ground respiration, 0.1% was recovered in soil solution, and 0.2-1.5% in soil. Swards under elevated CO(2) with the lower N supply fixed more (14)C than swards grown in ambient CO(2), exported more fixed (14)C below ground and respired less than their high-N counterparts. Sward cutting reduced root (14)C, but plants in elevated CO(2) still retained 80% more (14)C below ground than those in ambient CO(2). The potential for below-ground C sequestration in grasslands is enhanced under elevated CO(2), but any increase is likely to be small and dependent upon grassland management.     

A trade-off between nitrogen uptake and use increases responsiveness to elevated CO2 in infrequently cut mixed C3 grasses

The aim of this study was to evaluate whether the responsiveness of mixed C3 grass species to elevated CO2 is related more to nitrogen uptake or to N-use efficiency. Nitrogen uptake and whole-plant N-use efficiency were investigated with two binary mixtures: Lolium perenne was mixed either with Festuca arundinacea or with Holcus lanatus. The swards were grown on sand with or without CO2 doubling, and subjected to two cutting frequencies. A C20 alcohol was used as a marker to determine species proportion in the total root mass of the mixtures. The mean residence time of N was calculated from that of 15N-labelled fertilizer. Lolium perenne took up significantly more N per unit root mass than its grass competitors, but its N-use efficiency was lower. Elevated CO2 significantly reduced the N uptake of the three grass species. A trade-off between N capture and use was found, as N-use efficiency and N-uptake rate were negatively correlated. A high N-use efficiency, and conversely low N uptake appeared to favour the responsiveness to elevated CO2 of the infrequently cut grasses.     

Regenerating temperate forest mesocosms in elevated CO2: belowground growth and nitrogen cycling

The response of temperate forest ecosystems to elevated atmospheric CO₂ concentrations is important because these ecosystems represent a significant component of the global carbon cycle. Two important but not well understood processes which elevated CO₂ may substantially alter in these systems are regeneration and nitrogen cycling. If elevated CO₂ leads to changes in species composition in regenerating forest communities then the structure and function of these ecosystems may be affected. In most temperate forests, nitrogen appears to be a limiting nutrient. If elevated CO₂ leads to reductions in nitrogen cycling through increased sequestration of nitrogen in plant biomass or reductions in mineralization rates, long-term forest productivity may be constrained. To study these processes, we established mesocosms of regenerating forest communities in controlled environments maintained at either ambient (375 ppm) or elevated (700 ppm) CO₂ concentrations. Mesocosms were constructed from intact monoliths of organic forest soil. We maintained these mesocosms for 2 years without any external inputs of nitrogen and allowed the plants naturally present as seeds and rhizomes to regenerate. We used ¹⁵N pool dilution techniques to quantify nitrogen fluxes within the mesocosms at the end of the 2 years. Elevated atmospheric CO₂ concentration significantly affected a number of plant and soil processes in the experimental regenerating forest mesocosms. These changes included increases in total plant biomass production, plant C/N ratios, ectomycorrhizal colonization of tree fine roots, changes in tree fine root architecture, and decreases in plant NH₄ ⁺ uptake rates, gross NH₄ ⁺ mineralization rates, and gross NH₄ ⁺ consumption rates. In addition, there was a shift in the relative biomass contribution of the two dominant regenerating tree species; the proportion of total biomass contributed by white birch (Betula papyrifera) decreased and the proportion of total biomass contributed by yellow birch (B. alleghaniensis) increased. However, elevated CO₂ had no significant effect on the total amount of nitrogen in plant and soil microbial biomass. In this study we observed a suite of effects due to elevated CO₂, some of which could lead to increases in potential long term growth responses to elevated CO₂, other to decreases. The reduced plant NH₄ ⁺ uptake rates we observed are consistent with reduced NH₄ ⁺ availability due to reduced gross mineralization rates. Reduced NH₄ ⁺ mineralization rates are consistent with the increases in C/N ratios we observed for leaf and fine root material. Together, these data suggest the positive increases in plant root architectural parameters and mycorrhizal colonization may not be as important as the potential negative effects of reduced nitrogen availability through decreased decomposition rates in a future atmosphere with elevated CO₂. Received: 10 January 1997 / Accepted: 25 July 1997     

Seedling survival in a northern temperate forest understory is increased by elevated atmospheric carbon dioxide and atmospheric nitrogen deposition

We tested the main and interactive effects of elevated carbon dioxide concentration ([CO₂]), nitrogen (N), and light availability on leaf photosynthesis, and plant growth and survival in understory seedlings grown in an N‐limited northern hardwood forest. For two growing seasons, we exposed six species of tree seedlings (Betula papyrifera, Populus tremuloides, Acer saccharum, Fagus grandifolia, Pinus strobus, and Prunus serotina) to a factorial combination of atmospheric CO₂ (ambient, and elevated CO₂ at 658 μmol CO₂ mol⁻¹) and N deposition (ambient and ambient +30 kg N ha⁻¹ yr⁻¹) in open‐top chambers placed in an understory light gradient. Elevated CO₂ exposure significantly increased apparent quantum efficiency of electron transport by 41% (P<0.0001), light‐limited photosynthesis by 47% (P<0.0001), and light‐saturated photosynthesis by 60% (P<0.003) compared with seedlings grown in ambient [CO₂]. Experimental N deposition significantly increased light‐limited photosynthesis as light availability increased (P<0.037). Species differed in the magnitude of light‐saturated photosynthetic response to elevated N and light treatments (P<0.016). Elevated CO₂ exposure and high N availability did not affect seedling growth; however, growth increased slightly with light availability (R²=0.26, P<0.0001). Experimental N deposition significantly increased average survival of all species by 48% (P<0.012). However, seedling survival was greatest (85%) under conditions of both high [CO₂] and N deposition (P<0.009). Path analysis determined that the greatest predictor for seedling survival in the understory was total biomass (R²=0.39, P<0.001), and that carboxylation capacity (V_cmax) was a better predictor for seedling growth and survival than maximum photosynthetic rate (A_max). Our results suggest that increasing [CO₂] and N deposition from fossil fuel combustion could alter understory tree species recruitment dynamics through changes in seedling survival, and this has the potential to alter future forest species composition.     

Elevated [CO2], temperature increase and N supply effects on the turnover of below-ground carbon in a temperate grassland ecosystem

The effects of elevated [CO₂] (700 μl l^-1 CO₂) and temperature increase (+3 °C) on carbon turnover in grassland soils were studied during 2.5 years at two N fertiliser supplies (160 and 530 kg N ha^-1 y^-1) in an experiment with well-established ryegrass swards (Lolium perenne) supplied with the same amounts of irrigation water. During the growing season, swards from the control climate (350 μl l^-1 [CO₂] at outdoor air temperature) were pulse labelled by the addition of ¹³CO₂. The elevated [CO₂] treatments were continuously labelled by the addition of fossil-fuel derived CO₂ (^{13 C} of -40 to -50 ‰). Prior to the start of the experimental treatments, the carbon accumulated in the plant parts and in the soil macro-organic matter (‘old’ C) was at −32‰. During the experiment, the carbon fixed in the plant material (‘new’ C) was at −14 and −54‰ in the ambient and elevated [CO₂] treatments, respectively. During the experiment, the ¹³C isotopic mass balance method was used to calculate, for the top soil (0–15 cm), the carbon turnover in the stubble and roots and in the soil macro-organic matter above 200 μ (MOM). Elevated [CO₂] stimulated the turnover of organic carbon in the roots and stubble and in the MOM at N+, but not at N−. At the high N supply, the mean replacement time of ‘old’ C by ‘new’ C declined in elevated, compared to ambient [CO₂], from 18 to 7 months for the roots and stubble and from 25 to 17 months for the MOM. This resulted from increased rates of ‘new’ C accumulation and of ‘old’ C decay. By contrast, at the low N supply, despite an increase in the rate of accumulation of ‘new’ C, the soil C pools did not turnover faster in elevated [CO₂], as the rate of ‘old’ C decomposition was reduced. A 3 °C temperature increase in elevated [CO₂] decreased the input of fresh C to the roots and stubble and enhanced significantly the exponential rate for the ‘old’ C decomposition in the roots and stubble. An increased fertiliser N supply reduced the carbon turnover in the roots and stubble and in the MOM, in ambient but not in elevated [CO₂]. The respective roles for carbon turnover in the coarse soil OM fractions, of the C:N ratio of the litter, of the inorganic N availability and of a possible priming effect between C-substrates are discussed. This revised version was published online in June 2006 with corrections to the Cover Date.     

Mycorrhizal dynamics under elevated CO2 and nitrogen fertilization in a warm temperate forest

We examined the response of mycorrhizal fungi to free-air CO₂ enrichment (FACE) and nitrogen (N) fertilization in a warm temperate forest to better understand potential influences over plant nutrient uptake and soil carbon (C) storage. In particular, we hypothesized that mycorrhizal fungi and glomalin would become more prevalent under elevated CO₂ but decrease under N fertilization. In addition, we predicted that N fertilization would mitigate any positive effects of elevated CO₂ on mycorrhizal abundance. Overall, we observed a 14% increase in ectomycorrhizal (ECM) root colonization under CO₂ enrichment, which implies that elevated CO₂ results in greater C investments in these fungi. Arbuscular mycorrhizal (AM) hyphal length and glomalin stocks did not respond substantially to CO₂ enrichment, and effects of CO₂ on AM root colonization varied by date. Nitrogen effects on AM fungi were not consistent with our hypothesis, as we found an increase in AM colonization under N fertilization. Lastly, neither glomalin concentrations nor ECM colonization responded significantly to N fertilization or to an N-by-CO₂ interaction. A longer duration of N fertilization may be required to detect effects on these parameters.     

Temperate forest responses to carbon dioxide, temperature and nitrogen: a model analysis

The ITE Edinburgh Forest Model, which describes diurnal and seasonal changes in the pools and fluxes of C, N and water in a fully coupled forest–soil system, was parametrized to simulate a managed conifer plantation in upland Britain. The model was used to examine (i) the transient effects on forest growth of an IS92a scenario of increasing [CO₂] and temperature over two future rotations, and (ii) the equilibrium (sustainable) effects of all combinations of increases in [CO₂] from 350 to 550 and 750 μmol mol^?1, mean annual temperature from 7.5 to 8.5 and 9.5°C and annual inputs of 20 or 40 kg N ha^?1. Changes in underlying processes represented in the model were then used to explain the responses. Eight conclusions were supported by the model for this forest type and climate.

• 1 Increasing temperatures above 3°C alone may cause forest decline owing to water stress.
• 2 Elevated [CO₂] can protect trees from water stress that they may otherwise suffer in response to increased temperature.
• 3 In N-limiting conditions, elevated [CO₂] can increase allocation to roots with little increase in leaf area, whereas in N-rich conditions elevated [CO₂] may not increase allocation to roots and generally increases leaf area.
• 4 Elevated [CO₂] can decrease water use by forests in N-limited conditions and increase water use in N-rich conditions.
• 5 Elevated [CO₂] can increase forest productivity even in N-limiting conditions owing to increased N acquisition and use efficiency.
• 6 Rising temperatures (along with rising [CO₂]) may increase or decrease forest productivity depending on the supply of N and changes in water stress.
• 7 Gaseous losses of N from the soil can increase or decrease in response to elevated [CO₂] and temperature.
• 8 Projected increases in [CO₂] and temperature (IS92a) are likely to increase net ecosystem productivity and hence C sequestration in temperate forests.

     

Response of soil surface CO2 flux in a boreal forest to ecosystem warming

Soil surface carbon dioxide (CO₂) flux (R_S) was measured for 2 years at the Boreal Soil and Air Warming Experiment site near Thompson, MB, Canada. The experimental design was a complete random block design that consisted of four replicate blocks, with each block containing a 15 m × 15 m control and heated plot. Black spruce [Picea mariana (Mill.) BSP] was the overstory species and Epilobium angustifolium was the dominant understory. Soil temperature was maintained (～5 °C) above the control soil temperature using electric cables inside water filled polyethylene tubing for each heated plot. Air inside a 7.3‐m‐diameter chamber, centered in the soil warming plot, contained approximately nine black spruce trees was heated ～5 °C above control ambient air temperature allowing for the testing of soil‐only warming and soil+air warming. Soil surface CO₂ flux (R_S) was positively correlated (P < 0.0001) to soil temperature at 10 cm depth. Soil surface CO₂ flux (R_S) was 24% greater in the soil‐only warming than the control in 2004, but was only 11% greater in 2005, while R_S in the soil+air warming treatments was 31% less than the control in 2004 and 23% less in 2005. Live fine root mass (< 2 mm diameter) was less in the heated than control treatments in 2004 and statistically less (P < 0.01) in 2005. Similar root mass between the two heated treatments suggests that different heating methods (soil‐only vs. soil+air warming) can affect the rate of decomposition.     

The effects of water table manipulation and elevated temperature on the net CO2 flux of wet sedge tundra ecosystems

In situ manipulations were conducted in a naturally drained lake on the arctic coastal plain near Prudhoe Bay, Alaska (70 °21.98′ N, 148 °33.72′ W) to assess the potential short-term effects of decreased water table and elevated temperature on net ecosystem CO₂ flux. The experiments were conducted over a 2-year period, and during that time, water table depth of drained plots was maintained on average 7 cm lower than the ambient water table, and surface temperatures of plots exposed to elevated temperature were increased on average 0.5 °C. Water table drainage, and to a lesser extent elevated temperature, resulted in significant increases in ecosystem respiration (ER) rates, and only small and variable changes in gross ecosystem productivity (GEP). As a result, drained plots were net sources of ≈ 40 gC m^–2 season^–1 over both years of manipulation, while control plots were net sinks of atmospheric CO₂ of about 10 gC m^–2 season^–1 (growing season length was an estimated 125 days). Control plots exposed to elevated temperatures accumulated slightly more carbon than control plots exposed to ambient temperatures. The direct effects of elevated temperature on net CO₂ flux, ER, and GEP were small, however, elevated temperature appeared to interact with drainage to exacerbate the amount of net carbon loss. These data suggest that many currently saturated or nearly saturated wet sedge ecosystems of the north slope of Alaska may become significant sources of CO₂ to the atmosphere if climate change predictions of increased evapotranspiration and reduced soil water status are realized. There is ample evidence that this may be already occurring in arctic Alaska, as a change in net carbon balance has been observed for both tussock and wet-sedge tundra ecosystems over the last 2–3 decades, which coincides with a recent increase in surface temperature and an associated decrease in soil water content. In contrast, if precipitation increases relatively more than evapotranspiration, then increases in soil moisture content will likely result in greater carbon accumulation.     

CO2 stabilization,climate change and the terrestrial carbon sink

A nonequilibrium, dynamic, global vegetation model, Hybrid v4.1, with a subdaily timestep, was driven by increasing CO₂ and transient climate output from the UK Hadley Centre GCM (HadCM2) with simulated daily and interannual variability. Three IPCC emission scenarios were used: (i) IS92a, giving 790 ppm CO₂ by 2100, (ii) CO₂ stabilization at 750 ppm by 2225, and (iii) CO₂ stabilization at 550 ppm by 2150. Land use and future N deposition were not included. In the IS92a scenario, boreal and tropical lands warmed 4.5 °C by 2100 with rainfall decreased in parts of the tropics, where temperatures increased over 6 °C in some years and vapour pressure deficits (VPD) doubled. Stabilization at 750 ppm CO₂ delayed these changes by about 100 years while stabilization at 550 ppm limited the rise in global land surface temperature to 2.5 °C and lessened the appearance of relatively hot, dry areas in the tropics. Present‐day global predictions were 645 PgC in vegetation, 1190 PgC in soils, a mean carbon residence time of 40 years, NPP 47 PgC y^?1 and NEP (the terrestrial sink) about 1 PgC y^?1, distributed at both high and tropical latitudes. With IS92a emissions, the high latitude sink increased to the year 2100, as forest NPP accelerated and forest vegetation carbon stocks increased. The tropics became a source of CO₂ as forest dieback occurred in relatively hot, dry areas in 2060–2080. High VPDs and temperatures reduced NPP in tropical forests, primarily by reducing stomatal conductance and increasing maintenance respiration. Global NEP peaked at 3–4 PgC y^?1 in 2020–2050 and then decreased abruptly to near zero by 2100 as the tropical source offset the high‐latitude sink. The pattern of change in NEP was similar with CO₂ stabilization at 750 ppm, but was delayed by about 100 years and with a less abrupt collapse in global NEP. CO₂ stabilization at 550 ppm prevented sustained tropical forest dieback and enabled recovery to occur in favourable years, while maintaining a similar time course of global NEP as occurred with 750 ppm stabilization. By lessening dieback, stabilization increased the fraction of carbon emissions taken up by the land. Comparable studies and other evidence are discussed: climate‐induced tropical forest dieback is considered a plausible risk of following an unmitigated emissions scenario.     

Interactive effects of nitrogen deposition, tropospheric ozone, elevated CO2 and land use history on the carbon dynamics of northern hardwood forests

Temperate forests are affected by a wide variety of environmental factors that stem from human industrial and agricultural activities. In the north‐eastern US, important change agents include tropospheric ozone, atmospheric nitrogen deposition, elevated CO₂, and historical human land use. Although each of these has received attention for its effects on forest carbon dynamics, integrated analyses that examine their combined effects are rare. To examine the relative importance of all of these factors on current forest growth and carbon balances, we included them individually and in combination in a forest ecosystem model that was applied over the period of 1700–2000 under different scenarios of air pollution and land use history. Results suggest that historical increases in CO₂ and N deposition have stimulated forest growth and carbon uptake, but to different degrees following agriculture and timber harvesting. These differences resulted from the effects of each land use scenario on soil C and N pools and on the resulting degree of growth limitations by carbon vs. nitrogen. Including tropospheric ozone in the simulations offset a substantial portion of the increases caused by CO₂ and N deposition. This result is particularly relevant given that ozone pollution is widespread across much of the world and because broad‐scale spatial patterns of ozone are coupled with patterns of nitrogen oxide emissions. This was demonstrated across the study region by a significant correlation between ozone exposure and rates of N deposition and suggests that the reduction of N‐induced carbon sinks by ozone may be a common phenomenon in other regions. Collectively, the combined effects of all physical and chemical factors we addressed produced growth estimates that were surprisingly similar to estimates obtained in the absence of any form of disturbance. The implication of this result is that intact forests may show relatively little evidence of altered growth since preindustrial times despite substantial changes in their physical and chemical environment.