Implications of CO2 fertilization for future climate change in a coupled climate–carbon model

The terrestrial carbon cycle plays a critical role in determining levels of atmospheric CO₂ that result from anthropogenic carbon emissions. Elevated atmospheric CO₂ is thought to stimulate terrestrial carbon uptake, through the process of CO₂ fertilization of vegetation productivity. This negative carbon cycle feedback results in reduced atmospheric CO₂ growth, and has likely accounted for a substantial portion of the historical terrestrial carbon sink. However, the future strength of CO₂ fertilization in response to continued carbon emissions and atmospheric CO₂ rise is highly uncertain. In this paper, the ramifications of CO₂ fertilization in simulations of future climate change are explored, using an intermediate complexity coupled climate–carbon model. It is shown that the absence of future CO₂ fertilization results in substantially higher future CO₂ levels in the atmosphere, as this removes the dominant contributor to future terrestrial carbon uptake in the model. As a result, climate changes are larger, though the radiative effect of higher CO₂ on surface temperatures in the model is offset by about 30% due to reduced positive dynamic vegetation feedbacks; that is, the removal of CO₂ fertilization results in less vegetation expansion in the model, which would otherwise constitute an important positive surface albedo‐temperature feedback. However, the effect of larger climate changes has other important implications for the carbon cycle – notably to further weaken remaining carbon sinks in the model. As a result, positive climate–carbon cycle feedbacks are larger when CO₂ fertilization is absent. This creates an interesting synergism of terrestrial carbon cycle feedbacks, whereby positive (climate–carbon cycle) feedbacks are amplified when a negative (CO₂ fertilization) feedback is removed.     

The response of soil CO2 flux to changes in atmospheric CO2, nitrogen supply and plant diversity

We measured soil CO₂ flux over 19 sampling periods that spanned two growing seasons in a grassland Free Air Carbon dioxide Enrichment (FACE) experiment that factorially manipulated three major anthropogenic global changes: atmospheric carbon dioxide (CO₂) concentration, nitrogen (N) supply, and plant species richness. On average, over two growing seasons, elevated atmospheric CO₂ and N fertilization increased soil CO₂ flux by 0.57 µmol m^?2 s^?1 (13% increase) and 0.37 µmol m^?2 s^?1 (8% increase) above average control soil CO₂ flux, respectively. Decreases in planted diversity from 16 to 9, 4 and 1 species decreased soil CO₂ flux by 0.23, 0.41 and 1.09 µmol m^?2 s^?1 (5%, 8% and 21% decreases), respectively. There were no statistically significant pairwise interactions among the three treatments. During 19 sampling periods that spanned two growing seasons, elevated atmospheric CO₂ increased soil CO₂ flux most when soil moisture was low and soils were warm. Effects on soil CO₂ flux due to fertilization with N and decreases in diversity were greatest at the times of the year when soils were warm, although there were no significant correlations between these effects and soil moisture. Of the treatments, only the N and diversity treatments were correlated over time; neither were correlated with the CO₂ effect. Models of soil CO₂ flux will need to incorporate ecosystem CO₂ and N availability, as well as ecosystem plant diversity, and incorporate different environmental factors when determining the magnitude of the CO₂, N and diversity effects on soil CO₂ flux.     

Natural CO2 springs in Italy: a resource for examining long-term response of vegetation to rising atmospheric CO2 concentrations

It is estimated that more than 100 geothermal CO₂ springs exist in central-western Italy. Eight springs were selected in which the atmospheric CO₂ concentrations were consistently observed to be above the current atmospheric average of 354μmol mol^-1. CO₂ concentration measurements at some of the springs are reported. The springs are described, and their major topographic and vegetational features are reported. Preliminary observations made on natural vegetation growing around the gas vents are then illustrated. An azonal pattern of vegetation distribution occurs around every CO₂ spring regardless of soil type and phytoclimatic areas. This is composed of pioneer populations of a Northern Eurasiatic species (Agrostis canina L.) which is often associated with Scirpus lacustris L. The potential of these sites for studying the long-term response of vegetation to rising atmospheric CO₂ concentrations is discussed.     

The density dependence of plant responses to elevated CO2

1 Stands of the annual Brassica kaber were grown at a range of six densities in both ambient and elevated CO₂ environments, and measurements of shoot growth were made from seedling emergence through to reproduction.
2 Early in stand development (21 days following emergence), CO₂ enhancement (β) for above-ground biomass was highly density-dependent, ranging from 1.41 at the lowest density (20 plants m−2) to 0.59 at the highest density (652 plants m−2).
3 As stands matured and total biomass exceeded a relatively low threshold level (<10.0 g m^{−2; c.}  20% of final yield), the density-dependence of β disappeared. Above this shoot biomass threshold, β-values remained remarkably stable (β = 0.34) across a broad range of stand biomass, independent of a stand's initial density or age.
4 Average stand-level reproductive β-values at a final harvest were very similar to biomass values (β = 0.38) and, as with biomass values at later stages, showed no apparent density-dependence.
5 These results highlight the importance of considering density and the time-course of stand development simultaneously when assessing the potential for CO₂-induced growth enhancements in plants.     

Sensitivity of a dynamic global vegetation model to climate and atmospheric CO2

The equilibrium carbon storage capacity of the terrestrial biosphere has been investigated by running the Lund–Potsdam–Jena Dynamic Global Vegetation Model to equilibrium for a range of CO₂ concentrations and idealized climate states. Local climate is defined by the combination of an observation-based climatology and perturbation patterns derived from a 4 × CO₂ warming simulations, which are linearly scaled to global mean temperature deviations, Δ T _glob. Global carbon storage remains close to its optimum for Δ T _glob in the range of ±3°C in simulations with constant atmospheric CO₂. The magnitude of the carbon loss to the atmosphere per unit change in global average surface temperature shows a pronounced nonlinear threshold behavior. About twice as much carbon is lost per degree warming for Δ T _glob above 3°C than for present climate. Tropical, temperate, and boreal trees spread poleward with global warming. Vegetation dynamics govern the distribution of soil carbon storage and turnover in the climate space. For cold climate conditions, the global average decomposition rate of litter and soil decreases with warming, despite local increases in turnover rates. This result is not compatible with the assumption, commonly made in global box models, that soil turnover increases exponentially with global average surface temperature, over a wide temperature range.     

CO2 uptake and fluorescence responses for a shade-tolerant cactus Hylocereus undatus under current and doubled CO2 concentrations

Hylocereus undatus (Haworth) Britton and Rose growing in controlled environment chambers at 370 and 740 μmol CO₂ mol^?1 air showed a Crassulacean acid metabolism (CAM) pattern of CO₂ uptake, with 34% more total daily CO₂ uptake under the doubled CO₂ concentration and most of the increase occurring in the late afternoon. For both CO₂ concentrations, 90% of the maximal daily CO₂ uptake occurred at a total daily photosynthetic photon flux density (PPFD) of only 10 mol m^?2 day^?1 and the best day/night air temperatures were 25/15°C. Enhancement of the daily net CO₂ uptake by doubling the CO₂ concentration was greater under the highest PPFD (30 mol m^?2 day^?1) and extreme day/night air temperatures (15/5 and 45/35°C). After 24 days of drought, daily CO₂ uptake under 370 μmol CO₂ mol^?1 was 25% of that under 740 μmol CO₂ mol^?1. The ratio of variable to maximal chlorophyll fluorescence (F_y/F_m) decreased as the PPFD was raised above 5 mol m^?2 day^?1, at extreme day/night temperatures and during drought, suggesting that stress occurred under these conditions. F_v/F_m was higher under the doubled CO₂ concentration, indicating that the current CO₂ concentration was apparently limiting for photosynthesis. Thus net CO₂ uptake by the shade-tolerant H. undatus, the photosynthetic efficiency of which was greatest at low PPFDs. showed a positive response to doubling the CO₂ concentration, especially under stressful environmental conditions.     

Simulated variations in atmospheric CO2 over a Wisconsin forest using a coupled ecosystem–atmosphere model

Ecosystem fluxes of energy, water, and CO₂ result in spatial and temporal variations in atmospheric properties. In principle, these variations can be used to quantify the fluxes through inverse modelling of atmospheric transport, and can improve the understanding of processes and falsifiability of models. We investigated the influence of ecosystem fluxes on atmospheric CO₂ in the vicinity of the WLEF‐TV tower in Wisconsin using an ecophysiological model (Simple Biosphere, SiB2) coupled to an atmospheric model (Regional Atmospheric Modelling System). Model parameters were specified from satellite imagery and soil texture data. In a companion paper, simulated fluxes in the immediate tower vicinity have been compared to eddy covariance fluxes measured at the tower, with meteorology specified from tower sensors. Results were encouraging with respect to the ability of the model to capture observed diurnal cycles of fluxes. Here, the effects of fluxes in the tower footprint were also investigated by coupling SiB2 to a high‐resolution atmospheric simulation, so that the model physiology could affect the meteorological environment. These experiments were successful in reproducing observed fluxes and concentration gradients during the day and at night, but revealed problems during transitions at sunrise and sunset that appear to be related to the canopy radiation parameterization in SiB2.     

Comparison of gas use efficiency and treatment uniformity in a forest ecosystem exposed to elevated [CO2] using pure and prediluted free-air CO2 enrichment technology

A direct comparison of treatment uniformity and CO₂ use of pure and prediluted free-air CO₂ enrichment (FACE) systems was conducted in a forest ecosystem. A vertical release pure CO₂ fumigation system was superimposed on an existing prediluted CO₂ fumigation system and operated on alternate days. The FACE system using prediluted CO₂ fumigation technology exhibited less temporal and spatial variability than the pure CO₂ fumigation system. The pure CO₂ fumigation system tended to over-fumigate the upwind portions of the plot and used 25% more CO₂ than the prediluted CO₂ fumigation system. The increased CO₂ use by the pure CO₂ system was exacerbated at low wind speeds. It is not clear if this phenomenon will also be observed in plots with smaller diameters and low-stature vegetation.     

Net ecosystem CO2 exchange in Mojave Desert shrublands during the eighth year of exposure to elevated CO2

Arid ecosystems, which occupy about 35% of the Earth's terrestrial surface area, are believed to be among the most responsive to elevated [CO₂]. Net ecosystem CO₂ exchange (NEE) was measured in the eighth year of CO₂ enrichment at the Nevada Desert Free‐Air CO₂ Enrichment (FACE) Facility between the months of December 2003–December 2004. On most dates mean daily NEE (24 h) (μmol CO₂ m^?2 s^?1) of ecosystems exposed to elevated atmospheric CO₂ were similar to those maintained at current ambient CO₂ levels. However, on sampling dates following rains, mean daily NEEs of ecosystems exposed to elevated [CO₂] averaged 23 to 56% lower than mean daily NEEs of ecosystems maintained at ambient [CO₂]. Mean daily NEE varied seasonally across both CO₂ treatments, increasing from about 0.1 μmol CO₂ m^?2 s^?1 in December to a maximum of 0.5–0.6 μmol CO₂ m^?2 s^?1 in early spring. Maximum NEE in ecosystems exposed to elevated CO₂ occurred 1 month earlier than it did in ecosystems exposed to ambient CO₂, with declines in both treatments to lowest seasonal levels by early October (0.09±0.03 μmol CO₂ m^?2 s^?1), but then increasing to near peak levels in late October (0.36±0.08 μmol CO₂ m^?2 s^?1), November (0.28±0.03 μmol CO₂ m^?2 s^?1), and December (0.54±0.06 μmol CO₂ m^?2 s^?1). Seasonal patterns of mean daily NEE primarily resulted from larger seasonal fluctuations in rates of daytime net ecosystem CO₂ uptake which were closely tied to plant community phenology and precipitation. Photosynthesis in the autotrophic crust community (lichens, mosses, and free‐living cyanobacteria) following rains were probably responsible for the high NEEs observed in January, February, and late October 2004 when vascular plant photosynthesis was low. Both CO₂ treatments were net CO₂ sinks in 2004, but exposure to elevated CO₂ reduced CO₂ sink strength by 30% (positive net ecosystem productivity=127±17 g C m^?2 yr^?1 ambient CO₂ and 90±11 g C m^?2 yr^?1 elevated CO₂, P=0.011). This level of net C uptake rivals or exceeds levels observed in some forested and grassland ecosystems. Thus, the decrease in C sequestration seen in our study under elevated CO₂– along with the extensive coverage of arid and semi‐arid ecosystems globally – points to a significant drop in global C sequestration potential in the next several decades because of responses of heretofore overlooked dryland ecosystems.     

Soil CO2 efflux in a boreal pine forest under atmospheric CO2 enrichment and air warming

The response of forest soil CO₂ efflux to the elevation of two climatic factors, the atmospheric concentration of CO₂ (↑CO₂ of 700 μmol mol⁻¹) and air temperature (↑ T with average annual increase of 5°C), and their combination (↑CO₂+↑ T ) was investigated in a 4-year, full-factorial field experiment consisting of closed chambers built around 20-year-old Scots pines ( Pinus sylvestris L.) in the boreal zone of Finland. Mean soil CO₂ efflux in May–October increased with elevated CO₂ by 23–37%, with elevated temperature by 27–43%, and with the combined treatment by 35–59%. Temperature elevation was a significant factor in the combined 4-year efflux data, whereas the effect of elevated CO₂ was not as evident. Elevated temperature had the most pronounced impact early and late in the season, while the influence of elevated CO₂ alone was especially notable late in the season. Needle area was found to be a significant predictor of soil CO₂ efflux, particularly in August, a month of high root growth, thus supporting the assumption of a close link between whole-tree physiology and soil CO₂ emissions. The decrease in the temperature sensitivity of soil CO₂ efflux observed in the elevated temperature treatments in the second year nevertheless suggests the existence of soil response mechanisms that may be independent of the assimilating component of the forest ecosystem. In conclusion, elevated atmospheric CO₂ and air temperature consistently increased forest soil CO₂ efflux over the 4-year period, their combined effect being additive, with no apparent interaction.     

Seventeen years of elevated CO2 exposure in a Chesapeake Bay Wetland: sustained but contrasting responses of plant growth and CO2 uptake

Increased atmospheric CO₂ concentration (Ca) produces a short‐term stimulation of photosynthesis and plant growth across terrestrial ecosystems. However, the long‐term response remains uncertain and is thought to depend on environmental constraints. In the longest experiment on natural ecosystem response to elevated Ca, we measured the shoot‐density, biomass and net CO₂ exchange (NEE) responses to elevated Ca from 1987 to 2003 in a Scirpus olneyi wetland sedge community of the Chesapeake Bay, MD, USA. Measurements were conducted in five replicated open‐top chambers per CO₂ treatment (ambient and elevated). In addition, unchambered control plots were monitored for shoot density. Responses of daytime NEE, Scirpus plant biomass and shoot density to elevated Ca were positive for any single year of the 17‐year period of study. Daytime NEE stimulation by elevated Ca rapidly dropped from 80% at the onset of the experiment to a long‐term stimulation average of about 35%. Shoot‐density stimulation by elevated Ca increased linearly with duration of exposure (r²=0.89), exceeding 120% after 17 years. Although of lesser magnitude, the shoot biomass response to elevated Ca was similar to that of the shoot density. Daytime NEE response to elevated Ca was not explained by the duration of exposure, but negatively correlated with salinity of the marsh, indicating that this elevated‐Ca response was decreased by water‐related stress. By contrast, circumstantial evidence suggested that salinity stress increased the stimulation of shoot density by elevated Ca, which highlights the complexity of the interaction between water‐related stresses and plant community responses to elevated Ca. Notwithstanding the effects of salinity stress, we believe that the most important finding of the present research is that a species response to elevated Ca can continually increase when this species is under stress and declining in its natural environment. This is particularly important because climate changes associated with elevated Ca are likely to increase environmental stresses on numerous species and modify their present distribution. Our results point to an increased resilience to change under elevated Ca when plants are exposed to adverse environmental conditions.     

Seasonal CO2 exchange patterns of developing peach (Prunus persica) fruits in response to temperature, light and CO2 concentration

CO₂ exchange rates per unit dry weight, measured in the field on attached fruits of the late-maturing Cal Red peach cultivar, at 1200 μmol photons m^?2S^?1 and in dark, and photosynthetic rates, calculated by the difference between the rates of CO₂ evolution in light and dark, declined over the growing season. Calculated photosynthetic rates per fruit increased over the season with increasing fruit dry matter, but declined in maturing fruits apparently coinciding with the loss of chlorophyll. Slight net fruit photosynthetic rates ranging from 0. 087 ± 0. 06 to 0. 003 ± 0. 05 nmol CO₂ (g dry weight)^?1 S^?1 were measured in midseason under optimal temperature (15 and 20°C) and light (1200 μmol photons m^?2 S^?1) conditions. Calculated fruit photosynthetic rates per unit dry weight increased with increasing temperatures and photon flux densities during fruit development. Dark respiration rates per unit dry weight doubled within a temperature interval of 10°C; the mean seasonal O₁₀ value was 2. 03 between 20 and 30°C. The highest photosynthetic rates were measured at 35°C throughout the growing season. Since dark respiration rates increased at high temperatures to a greater extent than CO₂ exchange rates in light, fruit photosynthesis was apparently stimulated by high internal CO₂ concentrations via CO₂ refixation. At 15°C, fruit photosynthetic rates tended to be saturated at about 600 μmol photons m^?2 S^?1. Young peach fruits responded to increasing ambient CO₂ concentrations with decreasing net CO₂ exchange rates in light, but more mature fruits did not respond to increases in ambient CO₂. Fruit CO₂ exchange rates in the dark remained fairly constant, apparently uninfluenced by ambient CO₂ concentrations during the entire growing season. Calculated fruit photosynthetic rates clearly revealed the difference in CO₂ response of young and mature peach fruits. Photosynthetic rates of younger peach fruits apparently approached saturation at 370 μl CO₂1^?2. In CO₂ free air, fruit photosynthesis was dependent on CO₂ refixation since CO₂ uptake by the fruits from the external atmosphere was not possible. The difference in photosynthetic rates between fruits in CO₂-free air and 370 μl CO₂ 1^?1 indicated that young peach fruits were apparently able to take up CO₂ from the external atmosphere. CO₂ uptake by peach fruits contributed between 28 and 16% to the fruit photosynthetic rate early in the season, whereas photosynthesis in maturing fruits was supplied entirely by CO₂ refixation.     

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

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

Elevated CO2 stimulates associative N2 fixation in a C3 plant of the Chesapeake Bay wetland

In this study, the response of N₂ fixation to elevated CO₂ was measured in Scirpus olneyi, a C₃ sedge, and Spartina patens, a C₄ grass, using acetylene reduction assay and ¹⁵N₂ gas feeding. Field plants grown in PVC tubes (25 cm long, 10 cm internal diameter) were used. Exposure to elevated CO₂ significantly (P < 0·05) caused a 35% increase in nitrogenase activity and 73% increase in ¹⁵N incorporated by Scirpus olneyi. In Spartina patens, elevated CO₂ (660 ± 1 μ mol mol ^{− 1}) increased nitrogenase activity and ¹⁵N incorporation by 13 and 23%, respectively. Estimates showed that the rate of N₂ fixation in Scirpus olneyi under elevated CO₂ was 611 ± 75 ng ¹⁵N fixed plant ^{− 1} h ^{− 1} compared with 367 ± 46 ng ¹⁵N fixed plant ^{− 1} h ^{− 1} in ambient CO₂ plants. In Spartina patens, however, the rate of N₂ fixation was 12·5 ± 1·1 versus 9·8 ± 1·3 ng ¹⁵N fixed plant ^{− 1} h ^{− 1} for elevated and ambient CO₂, respectively. Heterotrophic non-symbiotic N₂ fixation in plant-free marsh sediment also increased significantly (P < 0·05) with elevated CO₂. The proportional increase in ¹⁵N₂ fixation correlated with the relative stimulation of photosynthesis, in that N₂ fixation was high in the C₃ plant in which photosynthesis was also high, and lower in the C₄ plant in which photosynthesis was relatively less stimulated by growth in elevated CO₂. These results are consistent with the hypothesis that carbon fixation in C₃ species, stimulated by rising CO₂, is likely to provide additional carbon to endophytic and below-ground microbial processes.     

Large-scale plant light-use efficiency inferred from the seasonal cycle of atmospheric CO2

We combined atmospheric CO₂ measurements, satellite observations, and an atmospheric transport model in an inverse modeling framework to infer a key property of vegetation physiology, the light-use efficiency (LUE) of net primary production, for large geographic regions. We find the highest LUE in boreal regions and in the northern hemisphere tropics. Within boreal zones, Eurasian LUE is higher than North American LUE and has a distinctly different seasonal profile. This longitudinal asymmetry is consistent with ecological differences expected from the much greater cover of deciduous vegetation in boreal Eurasia caused by the vast Siberian forests of the deciduous conifer, Larch. Inferred LUE of the northern hemisphere tropics is also high and displays a seasonal profile consistent with variations of both cloud cover and C₄ vegetation activity.     

Elevated CO2 concentrations and stomatal density: observations from 17 plant species growing in a CO2 spring in central Italy

Stomatal density (SD) and stomatal conductance ( g _s) can be affected by an increase of atmospheric CO₂ concentration. This study was conducted on 17 species growing in a naturally enriched CO₂ spring and belonging to three plant communities. Stomatal conductance, stomatal density and stomatal index (SI) of plants from the spring, which were assumed to have been exposed for generations to elevated [CO₂], and of plants of the same species collected in a nearby control site, were compared. Stomatal conductance was significantly lower in most of the species collected in the CO₂ spring and this indicated that CO₂ effects on g _s are not of a transitory nature but persist in the long term and through plant generations. Such a decrease was, however, not associated with changes in the anatomy of leaves: SD was unaffected in the majority of species (the decrease was only significant in three out of the 17 species examined), and also SI values did not vary between the two sites with the exception of two species that showed increased SI in plants grown in the CO₂-enriched area. These results did not support the hypothesis that long-term exposure to elevated [CO₂] may cause adaptive modification in stomatal number and in their distribution.