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.     

Potassium limits potential growth of bog vegetation under elevated atmospheric CO2 and N deposition

The free air carbon dioxide enrichment (FACE) and N deposition experiments on four ombrotrophic bogs in Finland, Sweden, the Netherlands and Switzerland, revealed that after three years of treatment: (1) elevated atmospheric CO₂ concentration had no significant effect on the biomass growth of Sphagnum and vascular species; and (2) increased N deposition reduced Sphagnum growth, because it increased the cover of vascular plants and the tall moss Polytrichum strictum, while vascular plant biomass growth was not affected. This paper focuses on water chemistry, plant nutrient content, and litter decomposition rates. Potassium limitation, or low supply of K and P, may have prevented a significant increase of Sphagnum growth under elevated CO₂ and N deposition. Vascular plant growth under elevated CO₂ and N deposition was also limited by K, or by K in combination with P or N (N in CO₂ experiment). Elevated CO₂ and N deposition had no effect on decomposition rates of Sphagnum and vascular plant litter. Aside from a possible effect of N deposition on light competition between species, we expect that elevated atmospheric CO₂ and N deposition concentrations will not affect Sphagnum and vascular plant growth in bogs of north‐west Europe due to K‐, or K in combination with N‐ or P‐, limited growth. For the same reason we expect no effect of elevated CO₂ and N deposition on litter decomposition. Net primary production of raised ombrotrophic bogs that are at or close to steady state, is regulated by input of nutrients through atmospheric deposition. Therefore, we hypothesize that the expected increase of plant growth under elevated CO₂ and N deposition is diminished by current levels of K (and to some extent P and N) in atmospheric deposition.     

The impact of nitrogen deposition on carbon sequestration in European forests and forest soils

An estimate of net carbon (C) pool changes and long‐term C sequestration in trees and soils was made at more than 100 intensively monitored forest plots (level II plots) and scaled up to Europe based on data for more than 6000 forested plots in a systematic 16 km × 16 km grid (level I plots). C pool changes in trees at the level II plots were based on repeated forest growth surveys At the level I plots, an estimate of the mean annual C pool changes was derived from stand age and available site quality characteristics. C sequestration, being equal to the long‐term C pool changes accounting for CO₂ emissions because of harvest and forest fires, was assumed 33% of the overall C pool changes by growth. C sequestration in the soil were based on calculated nitrogen (N) retention (N deposition minus net N uptake minus N leaching) rates in soils, multiplied by the C/N ratio of the forest soils, using measured data only (level II plots) or a combination of measurements and model calculations (level I plots). Net C sequestration by forests in Europe (both trees and soil) was estimated at 0.117 Gton yr^?1, with the C sequestration in stem wood being approximately four times as high (0.094 Gton yr^?1) as the C sequestration in the soil (0.023 Gton yr^?1). The European average impact of an additional N input on the net C sequestration was estimated at approximately 25 kg C kg^?1 N for both tree wood and soil. The contribution of an average additional N deposition on European forests of 2.8 kg ha^?1 yr^?1 in the period 1960–2000 was estimated at 0.0118 Gton yr^?1, being equal to 10% of the net C sequestration in both trees and soil in that period (0.117 Gton yr^?1). The C sequestration in trees increased from Northern to Central Europe, whereas the C sequestration in soil was high in Central Europe and low in Northern and Southern Europe. The result of this study implies that the impact of forest management on tree growth is most important in explaining the C pool changes in European forests.     

Tree ring responses to elevated CO2 and increased N deposition in Picea abies

Four- to seven-year-old spruce trees (Picea abies) were exposed to three CO₂ concentrations (280, 420 and 560 cm³ m^?3) and three rates of wet N deposition (0, 30 and 90 kg ha^?1 year^?1) for 3 years in a simulated montane forest climate. Six trees from each of six clones were grown in competition in each of nine 100 × 70 × 36 cm model ecosystems with nutrient-poor natural forest soil. Stem dises were analysed using X-ray densitometry. The radial stem increment was not affected by [CO₂] but increased with increasing rates of N deposition. Wood density was increased by [CO₂], but decreased by N deposition. Wood-starch concentration increased, and wood nitrogen concentration decreased with increasing [CO₂], but neither was affected by N deposition. The lignin concentration in wood was affected by neither [CO₂] nor N deposition. Our results suggest that, under natural growth conditions, rising atmospheric [CO₂] will not lead to enhanced radial stem growth of spruce, but atmospheric N deposition will, and in some regions is probably already doing so. Elevated [CO₂], however, will lead to denser wood unless this effect is compensated by massive atmospheric N deposition. If can be speculated that greater wood density under elevated [CO₂] may alter the mechanical properties of wood, and higher ratios of C/N and lignin/N in wood grown at elevated [CO₂] may affect nutrient cycles of forest ecosystems.     

Quantifying the response of photosynthesis to changes in leaf nitrogen content and leaf mass per area in plants grown under atmospheric CO2 enrichment

Previous modelling exercises and conceptual arguments have predicted that a reduction in biochemical capacity for photosynthesis (A_area) at elevated CO₂ may be compensated by an increase in mesophyll tissue growth if the total amount of photosynthetic machinery per unit leaf area is maintained (i.e. morphological upregulation). The model prediction was based on modelling photosynthesis as a function of leaf N per unit leaf area (N_area), where N_area = N_mass×LMA. Here, N_mass is percentage leaf N and is used to estimate biochemical capacity and LMA is leaf mass per unit leaf area and is an index of leaf morphology. To assess the relative importance of changes in biochemical capacity versus leaf morphology we need to control for multiple correlations that are known, or that are likely to exist between CO₂ concentration, N_area, N_mass, LMA and A_area. Although this is impractical experimentally, we can control for these correlations statistically using systems of linear multiple-regression equations. We developed a linear model to partition the response of A_area to elevated CO₂ into components representing the independent and interactive effects of changes in indexes of biochemical capacity, leaf morphology and CO₂ limitation of photosynthesis. The model was fitted to data from three pine and seven deciduous tree species grown in separate chamber-based field experiments. Photosynthetic enhancement at elevated CO₂ due to morphological upregulation was negligible for most species. The response of A_area in these species was dominated by the reduction in CO₂ limitation occurring at higher CO₂ concentration. However, some species displayed a significant reduction in potential photosynthesis at elevated CO₂ due to an increase in LMA that was independent of any changes in N_area. This morphologically based inhibition of A_area combined additively with a reduction in biochemical capacity to significantly offset the direct enhancement of A_area caused by reduced CO₂ limitation in two species. This offset was 100% for Acer rubrum, resulting in no net effect of elevated CO₂ on A_area for this species, and 44% for Betula pendula. This analysis shows that interactions between biochemical and morphological responses to elevated CO₂ can have important effects on photosynthesis.     

Elevated CO2 and temperature alter nitrogen allocation in Douglas-fir

The effects of elevated CO₂ and temperature on principal carbon constituents (PCC) and C and N allocation between needle, woody (stem and branches) and root tissue of Pseudotsuga menziesii Mirb. Franco seedlings were determined. The seedlings were grown in sun‐lit controlled‐environment chambers that contained a native soil. Chambers were controlled to reproduce ambient or ambient +180 ppm CO₂ and either ambient temperature or ambient +3.5 °C for 4 years. There were no significant CO₂ × temperature interactions; consequently the data are presented for the CO₂ and temperature effects. At the final harvest, elevated CO₂ decreased the nonpolar fraction of the PCC and increased the polar fraction and amount of sugars in the needles. In contrast, elevated temperature increased the nonpolar fraction of the PCC and decreased sugars in needles. There were no CO₂ or temperature effects on the PCC fractions in the woody tissue or root tissue. Elevated CO₂ and temperature had no significant effects on the C content of any of the plant tissues or fractions. In contrast, the foliar N content declined under elevated CO₂ and increased under elevated temperature; there were no significant effects in other tissues. The changes in the foliar N concentrations were in the cellulose and lignin fractions, the fractions, which contain protein, and are the consequences of changes in N allocation under the treatments. These results indicate reallocation of N among plant organs to optimize C assimilation, which is mediated via changes in the selectivity of Rubisco and carbohydrate modulation of gene expression.     

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.     

Belowground competition and the response of developing forest communities to atmospheric CO2 and O3

As human activity continues to increase CO₂ and O₃, broad expanses of north temperate forests will be simultaneously exposed to elevated concentrations of these trace gases. Although both CO₂ and O₃ are potent modifiers of plant growth, we do not understand the extent to which they alter competition for limiting soil nutrients, like nitrogen (N). We quantified the acquisition of soil N in two 8‐year‐old communities composed of trembling aspen genotypes (n= 5) and trembling aspen–paper birch which were exposed to factorial combinations of CO₂ (ambient and 560 μL L⁻¹) and O₃ (ambient = 30–40 vs. 50–60 nL L⁻¹). Tracer amount of ¹⁵NH₄⁺ were applied to soil to determine how these trace gases altered the competitive ability of genotypes and species to acquire soil N. One year after isotope addition, we assessed N acquisition by measuring the amount of ¹⁵N tracer contained in the plant canopy (i.e. recent N acquisition), as well as the total amount of canopy N (i.e. cumulative N acquisition). Exposure to elevated CO₂ differentially altered recent and cumulative N acquisition among aspen genotypes, changing the rank order in which they obtained soil N. Elevated O₃ also altered the rank order in which aspen genotypes obtained soil N by eliciting increases, decreases and no response among genotypes. If aspen genotypes respond similarly under field conditions, then rising concentrations of CO₂ and O₃ could alter the structure of aspen populations. In the aspen–birch community, elevated CO₂ increased recent N (i.e. ¹⁵N) acquisition in birch (68%) to a greater extent than aspen (19%), suggesting that, over the course of this experiment, birch had gained a competitive advantage over aspen. The response of genotypes and species to rising CO₂ and O₃ concentrations, and how these responses are modified by competitive interactions, has the potential to change the future composition and productivity of northern temperate forests.     

Elevated atmospheric CO2 and increased nitrogen deposition: effects on C and N metabolism and growth of the peat moss Sphagnum recurvum P. Beauv. var. mucronatum (Russ.) Warnst

Sphagnum bogs play an important role when considering the impacts of global change on global carbon and nitrogen cycles. Sphagnum recurvum P. Beauv. var. mucronatum (Russ.) was grown at 360 (ambient) and 700 μL L^?1 (elevated) atmospheric [CO₂] in combination with different nitrogen deposition rates (6, 15, 23 g N m^?2 y^?1), in a short‐ and long‐term growth chamber experiment. After 6 months, elevated atmospheric [CO₂] in combination with the lowest nitrogen deposition rate, increased plant dry mass by 17%. In combination with a high nitrogen deposition rate, biomass production was not significantly stimulated. At the start of the experiment, photosynthesis was stimulated by elevated atmospheric [CO₂], but it was downregulated to control levels after three days of exposure. Elevated [CO₂] substantially reduced dark respiration, which resulted in a continuous increase in soluble sugar content in capitula. Differences in growth response among different nitrogen and CO₂ treatments could not be related to measured carbon exchange rates, which was mainly due to interference of microbial respiration. Doubling atmospheric [CO₂] reduced total nitrogen content in capitula but not in stems at all nitrogen deposition rates. Reduction in total nitrogen content coincided with a decrease in amino acids, but soluble protein levels remained unaffected. Thus, elevated [CO₂] induced a substantial shift in the partitioning of nitrogen compounds in capitula. Soluble sugar concentration was negatively correlated with total nitrogen content, which implies that the reduction in amino acid content in capitula, exposed to elevated [CO₂], might be caused by the accumulation of soluble sugars. Growth was not stimulated by increased nitrogen deposition. High nitrogen deposition, resulting in a capitulum nitrogen content in excess of 15 mg g^?1 dw, was detrimental to photosynthesis, reduced water content and induced necrosis. We propose a capitulum nitrogen content of 15 mg g^?1 dw as a possible bioindicator for the detection of nitrogen pollution stress in oligo‐mesotrophic peat bog ecosystems. At the lowest nitrogen deposition level, nitrogen recovery was higher than 100%, which indicates substantial dry deposition and/or gaseous nitrogen fixation by bacteria, associated with Sphagnum. Increasing nitrogen deposition rates decreased nitrogen recovery percentages, which indicates reduced efficiency of nitrogen fixation.     

Raised atmospheric CO2 levels and increased N deposition cause shifts in plant species composition and production in Sphagnum bogs

Part of the missing sink in the global CO₂ budget has been attributed to the positive effects of CO₂ fertilization and N deposition on carbon sequestration in Northern Hemisphere terrestrial ecosystems. The genus Sphagnum is one of the most important groups of plant species sequestrating carbon in temperate and northern bog ecosystems, because of the low decomposability of the dead material it produces. The effects of raised CO₂ and increased atmospheric N deposition on growth of Sphagnum and other plants were studied in bogs at four sites across Western Europe. Contrary to expectations, elevated CO₂ did not significantly affect Sphagnum biomass growth. Increased N deposition reduced Sphagnum mass growth, because it increased the cover of vascular plants and the tall moss Polytrichum strictum. Such changes in plant species composition may decrease carbon sequestration in Sphagnum‐dominated bog ecosystems.     

Dry nitrogen deposition estimates over a forest experiencing free air CO2 enrichment

The quantification of atmospheric additions of nitrogen (N) to an ecosystem is often desirable, but difficult for many locations including many ecological manipulation experiments. Ideal methodologies for the complete and chemically speciated quantification of dry N deposition (e.g. tunable diode laser absorption spectrometer, thermal‐dissociation laser‐induced fluorescence) are expensive, technologically challenging to maintain, and rarely colocated with important global change manipulation experiments. Here we present an alternative method for obtaining an approximation of total N deposition using short‐term eddy flux and concentration measurements, annual regional concentration estimates, and modeling for the Duke Experimental Forest. The motivation for generating estimates for this location was to inform the long‐term elevated CO₂ experiment conducted at Duke Forest. We estimated the total annual atmospheric N deposition to the forest to be 13.7 kg N ha^?1. Of this total, ～58% was in dry‐deposited forms. Surprisingly, and contrary to some previous predictions, nitric acid (HNO₃) was not the dominant portion of the total dry‐deposited N, implying strongly that other forms of gaseous N like organic peroxy and alkyl nitrate compounds are a significant portion of the total flux. Furthermore, this study sheds light on the completeness of the estimates derived from Clean Air Status and Trends Network (CASTNet) and other dry deposition networks. CASTNet does not measure organic forms of dry deposition. In fact, CASTNet only quantifies HNO₃ in the gas phase and may significantly underestimate total N deposition in many environments.     

Mycorrhizal fungi have a potential role in soil carbon storage under elevated CO2 and nitrogen deposition

In this review, we discuss the potential for mycorrhizal fungi to act as a source or sink for carbon (C) under elevated CO₂ and nitrogen deposition. Mycorrhizal tissue has been estimated to comprise a significant fraction of soil organic matter and below-ground biomass in a range of systems. The current body of literature indicates that in many systems exposed to elevated CO₂, mycorrhizal fungi might sequester increased amounts of C in living, dead and residual hyphal biomass in the soil. Through this process, the fungi might serve as a negative feedback on the rise in atmospheric CO₂ levels caused by fossil fuel burning and deforestation. By contrast, a few preliminary studies suggest that N deposition might increase turnover rates of fungal tissue and negate CO₂ effects on hyphal biomass. If these latter responses are consistent among ecosystems, C storage in hyphae might decline in habitats surrounding agricultural and urban areas. When N additions occur without CO₂ enrichment, effects on mycorrhizal growth are inconsistent. We note that analyses of hyphal decomposition under elevated CO₂ and N additions are extremely sparse but are critical in our understanding of the impact of global change on the cycling of mycorrhizal C. Finally, shifts in the community composition of arbuscular and ectomycorrhizal fungi with increasing CO₂ or N availability are frequently documented. Since mycorrhizal groups vary in growth rate and tissue quality, these changes in species assemblages could produce unforeseeable impacts on the productivity, survivorship, or decomposition of mycorrhizal biomass.     

Effects of elevated CO2, nitrogen deposition, and decreased species diversity on foliar fungal plant disease

Three components of global change, elevated CO₂, nitrogen addition, and decreased plant species richness (‘diversity’), increased the percent leaf area infected by fungi (pathogen load) for much to all of the plant community in one year of a factorial grassland experiment. Decreased plant diversity had the broadest effect, increasing pathogen load across the plant community. Decreased diversity increased pathogen load primarily by allowing remaining plant species to increase in abundance, facilitating spread of foliar fungal pathogens specific to each plant species. Changes in plant species composition also strongly influenced community pathogen load, with communities that lost less disease prone plant species increasing more in pathogen load. Elevated CO₂ increased pathogen load of C₃ grasses, perhaps by decreasing water stress, increasing leaf longevity, and increasing photosynthetic rate, all of which can promote foliar fungal disease. Decreased plant diversity further magnified the increase in C₃ grass pathogen load under elevated CO₂. Nitrogen addition increased pathogen load of C₄ grasses by increasing foliar nitrogen concentration, which can enhance pathogen infection, growth, and reproduction. Because changes in foliar fungal pathogen load can strongly influence grassland ecosystem processes, our study suggests that increased pathogen load can be an important mechanism by which global change affects grassland ecosystems.     

Acquisition and allocation of carbon and nitrogen by Danthonia richardsonii in response to restricted nitrogen supply and CO2 enrichment

Dry weight (DW) and nitrogen (N) accumulation and allocation were measured in isolated plants of Danthonia richardsonii (Wallaby Grass) for 37 d following seed imbibition. Plants were grown at ≈ 365 or 735 μ L L^–1 CO₂ with N supply of 0·05, 0·2 or 0·5 mg N plant^–1 d^–1. Elevated CO₂ increased DW accumulation by 28% (low-N) to 103% (high-N), following an initial stimulation of relative growth rate. Net assimilation rate and leaf nitrogen productivity were increased by elevated CO₂, while N concentration was reduced. N uptake per unit root surface area was unaffected by CO₂ enrichment. The ratio of leaf area to root surface area was decreased by CO₂ enrichment. Allometric analysis revealed a decrease in the shoot-N to root-N ratio at elevated CO₂, while the shoot-DW to root-DW ratio was unchanged. Allometric analysis showed leaf area was reduced, while root surface area was unchanged by elevated CO₂, indicating a down-regulation of total plant capacity for carbon gain rather than a stimulation of mineral nutrient acquisition capacity. Overall, growth in elevated CO₂ resulted in changes in plant morphology and nitrogen use, other than those associated simply with changing plant size and non-structural carbohydrate content.     

Comparison of photosynthetic acclimation to elevated CO2 and limited nitrogen supply in soybean

Plants grown at elevated CO₂ often acclimate such that their photosynthetic capacities are reduced relative to ambient CO₂-grown plants. Reductions in synthesis of photosynthetic enzymes could result either from reduced photosynthetic gene expression or from reduced availability of nitrogen-containing substrates for enzyme synthesis. Increased carbohydrate concentrations resulting from increased photosynthetic carbon fixation at elevated CO₂ concentrations have been suggested to reduce the expression of photosynthetic genes. However, recent studies have also suggested that nitrogen uptake may be depressed by elevated CO₂, or at least that it is not increased enough to keep pace with increased carbohydrate production. This response could induce a nitrogen limitation in elevated-CO₂ plants that might account for the reduction in photosynthetic enzyme synthesis. If CO₂ acclimation were a response to limited nitrogen uptake, the effects of elevated CO₂ and limiting nitrogen supply on photosynthesis and nitrogen allocation should be similar. To test this hypothesis we grew non-nodulating soybeans at two levels each of nitrogen and CO₂ concentration and measured leaf nitrogen contents, photosynthetic capacities and Rubisco contents. Both low nitrogen and elevated CO₂ reduced nitrogen as a percentage of total leaf dry mass but only low nitrogen supply produced significant decreases in nitrogen as a percentage of leaf structural dry mass. The primary effect of elevated CO₂ was to increase non-structural carbohydrate storage rather than to decrease nitrogen content. Both low nitrogen supply and elevated CO₂ also decreased carboxylation capacity (V_cmax) and Rubisco content per unit leaf area. However, when V_cmax and Rubisco content were expressed per unit nitrogen, low nitrogen supply generally caused them to increase whereas elevated CO₂ generally caused them to decrease. Finally, elevated CO₂ significantly increased the ratio of RuBP regeneration capacity to V_cmax whereas neither nitrogen supply nor plant age had a significant effect on this parameter. We conclude that reductions in photosynthetic enzyme synthesis in elevated CO₂ appear not to result from limited nitrogen supply but instead may result from feedback inhibition by increased carbohydrate contents.     

Sensing of atmospheric CO2 by plants

Abstract. Despite recent interest in the effects of high CO₂ on plant growth and physiology, very little is known about the mechanisms by which plants sense changes in the concentration of this gas. Because atmospheric CO₂ concentration is relatively constant and because the conductance of the cuticle to CO₂ is low, sensory mechanisms are likely to exist only for intercellular CO₂ concentration. Therefore, responses of plants to changes in atmospheric CO₂ will depend on the effect of these changes on intercellular CO₂ concentration. Although a variety of plant responses to atmospheric CO₂ concentration have been reported, most of these can be attributed to the effects of intercellular CO₂ on photosynthesis or stomatal conductance. Short-term and long-term effects of CO₂ on photosynthesis and stomatal conductance are discussed as sensory mechanisms for responses of plants to atmospheric CO₂. Available data suggest that plants do not fully realize the potential increases in productivity associated with increased atmospheric CO₂. This may be because of genetic and environmental limitations to productivity or because plant responses to CO₂ have evolved to cope with variations in intercellular CO₂ caused by factors other than changes in atmospheric CO₂.