Responses of CAM species to increasing atmospheric CO2 concentrations

Crassulacean acid metabolism (CAM) species show an average increase in biomass productivity of 35% in response to a doubled atmospheric CO₂ concentration. Daily net CO₂ uptake is similarly enhanced, reflecting in part an increase in chlorenchyma thickness and accompanied by an even greater increase in water‐use efficiency. The responses of net CO₂ uptake in CAM species to increasing atmospheric CO₂ concentrations are similar to those for C₃ species and much greater than those for C₄ species. Increases in net daily CO₂ uptake by CAM plants under elevated atmospheric CO₂ concentrations reflect increases in both Rubisco‐mediated daytime CO₂ uptake and phosphoenolpyruvate carboxylase (PEPCase)‐mediated night‐time CO₂ uptake, the latter resulting in increased nocturnal malate accumulation. Chlorophyll contents and the activities of Rubisco and PEPCase decrease under elevated atmospheric CO₂, but the activated percentage for Rubisco increases and the K_M(HCO₃ ^? ) for PEPCase decreases, resulting in more efficient photosynthesis. Increases in root:shoot ratios and the formation of additional photosynthetic organs, together with increases in sucrose‐Pi synthase and starch synthase activity in these organs under elevated atmospheric CO₂ concentrations, decrease the potential feedback inhibition of photosynthesis. Longer‐term studies for several CAM species show no downward acclimatization of photosynthesis in response to elevated atmospheric CO₂ concentrations. With increasing temperature and drought duration, the percentage enhancement of daily net CO₂ uptake caused by elevated atmospheric CO₂ concentrations increases. Thus net CO₂ uptake, productivity, and the potential area for cultivation of CAM species will be enhanced by the increasing atmospheric CO₂ concentrations and the increasing temperatures associated with global climate change.     

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.     

N2 fixation by Acacia species increases under elevated atmospheric CO2

In the present study the effect of elevated CO₂ on growth and nitrogen fixation of seven Australian Acacia species was investigated. Two species from semi‐arid environments in central Australia (Acacia aneura and A. tetragonophylla) and five species from temperate south‐eastern Australia (Acacia irrorata, A. mearnsii, A. dealbata, A. implexa and A. melanoxylon) were grown for up to 148 d in controlled greenhouse conditions at either ambient (350 µmol mol^?1) or elevated (700 µmol mol^?1) CO₂ concentrations. After establishment of nodules, the plants were completely dependent on symbiotic nitrogen fixation. Six out of seven species had greater relative growth rates and lower whole plant nitrogen concentrations under elevated versus normal CO₂. Enhanced growth resulted in an increase in the amount of nitrogen fixed symbiotically for five of the species. In general, this was the consequence of lower whole‐plant nitrogen concentrations, which equate to a larger plant and greater nodule mass for a given amount of nitrogen. Since the average amount of nitrogen fixed per unit nodule mass was unaltered by atmospheric CO₂, more nitrogen could be fixed for a given amount of plant nitrogen. For three of the species, elevated CO₂ increased the rate of nitrogen fixation per unit nodule mass and time, but this was completely offset by a reduction in nodule mass per unit plant mass.     

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.     

Oceanic sinks for atmospheric CO2

There is approximately 50 times more inorganic carbon in the global ocean than in the atmosphere. On time scales of decades to millions of years, the interaction between these two geophysical fluids determines atmospheric CO₂ levels. During glacial periods, for example, the ocean serves as the major sink for atmospheric CO₂, while during glacial–interglacial transitions, it is a source of CO₂ to the atmosphere. The mechanisms responsible for determining the sign of the net exchange of CO₂ between the ocean and the atmosphere remain unresolved. There is evidence that during glacial periods, phytoplankton primary productivity increased, leading to an enhanced sedimentation of particulate organic carbon into the ocean interior. The stimulation of primary production in glacial episodes can be correlated with increased inputs of nutrients limiting productivity, especially aeolian iron. Iron directly enhances primary production in high nutrient (nitrate and phosphate) regions of the ocean, of which the Southern Ocean is the most important. This trace element can also enhance nitrogen fixation, and thereby indirectly stimulate primary production throughout the low nutrient regions of the central ocean basins. While the export flux of organic carbon to the ocean interior was enhanced during glacial periods, this process does not fully account for the sequestration of atmospheric CO₂. Heterotrophic oxidation of the newly formed organic carbon, forming weak acids, would have hydrolyzed CaCO₃ in the sediments, increasing thereby oceanic alkalinity which, in turn, would have promoted the drawdown of atmospheric CO₂. This latter mechanism is consistent with the stable carbon isotope pattern derived from air trapped in ice cores. The oceans have also played a major role as a sink for up to 30% of the anthropogenic CO₂ produced during the industrial revolution. In large part this is due to CO₂ solution in the surface ocean; however, some, poorly quantified fraction is a result of increased new production due to anthropogenic inputs of combined N, P and Fe. Based on ‘circulation as usual’, models predict that future anthropogenic CO₂ inputs to the atmosphere will, in part, continue to be sequestered in the ocean. Human intervention (large-scale Fe fertilization; direct CO₂ burial in the deep ocean) could increase carbon sequestration in the oceans, but could also result in unpredicted environmental perturbations. Changes in the oceanic thermohaline circulation as a result of global climate change would greatly alter the predictions of C sequestration that are possible on a ‘circulation as usual’ basis.     

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.     

Role of land cover changes for atmospheric CO2 increase and climate change during the last 150 years

We assess the role of changing natural (volcanic, aerosol, insolation) and anthropogenic (CO₂ emissions, land cover) forcings on the global climate system over the last 150 years using an earth system model of intermediate complexity, CLIMBER‐2. We apply several datasets of historical land‐use reconstructions: the cropland dataset by Ramankutty & Foley (1999) (R&F), the HYDE land cover dataset of Klein Goldewijk (2001) , and the land‐use emissions data from Houghton & Hackler (2002) . Comparison between the simulated and observed temporal evolution of atmospheric CO₂ and δ¹³CO₂ are used to evaluate these datasets. To check model uncertainty, CLIMBER‐2 was coupled to the more complex Lund–Potsdam–Jena (LPJ) dynamic global vegetation model. In simulation with R&F dataset, biogeophysical mechanisms due to land cover changes tend to decrease global air temperature by 0.26°C, while biogeochemical mechanisms act to warm the climate by 0.18°C. The net effect on climate is negligible on a global scale, but pronounced over the land in the temperate and high northern latitudes where a cooling due to an increase in land surface albedo offsets the warming due to land‐use CO₂ emissions. Land cover changes led to estimated increases in atmospheric CO₂ of between 22 and 43 ppmv. Over the entire period 1800–2000, simulated δ¹³CO₂ with HYDE compares most favourably with ice core during 1850–1950 and Cape Grim data, indicating preference of earlier land clearance in HYDE over R&F. In relative terms, land cover forcing corresponds to 25–49% of the observed growth in atmospheric CO₂. This contribution declined from 36–60% during 1850–1960 to 4–35% during 1960–2000. CLIMBER‐2‐LPJ simulates the land cover contribution to atmospheric CO₂ growth to decrease from 68% during 1900–1960 to 12% in the 1980s. Overall, our simulations show a decline in the relative role of land cover changes for atmospheric CO₂ increase during the last 150 years.     

Future atmospheric CO2 leads to delayed autumnal senescence

Growing seasons are getting longer, a phenomenon partially explained by increasing global temperatures. Recent reports suggest that a strong correlation exists between warming and advances in spring phenology but that a weaker correlation is evident between warming and autumnal events implying that other factors may be influencing the timing of autumnal phenology. Using freely rooted, field‐grown Populus in two Free Air CO₂ Enrichment Experiments (AspenFACE and PopFACE), we present evidence from two continents and over 2 years that increasing atmospheric CO₂ acts directly to delay autumnal leaf coloration and leaf fall. In an atmosphere enriched in CO₂ (by ～45% of the current atmospheric concentration to 550 ppm) the end of season decline in canopy normalized difference vegetation index (NDVI) – a commonly used global index for vegetation greenness – was significantly delayed, indicating a greener autumnal canopy, relative to that in ambient CO₂. This was supported by a significant delay in the decline of autumnal canopy leaf area index in elevated as compared with ambient CO₂, and a significantly smaller decline in end of season leaf chlorophyll content. Leaf level photosynthetic activity and carbon uptake in elevated CO₂ during the senescence period was also enhanced compared with ambient CO₂. The findings reveal a direct effect of rising atmospheric CO₂, independent of temperature in delaying autumnal senescence for Populus, an important deciduous forest tree with implications for forest productivity and adaptation to a future high CO₂ world.     

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.     

Increase of atmospheric CO2 promotes phytoplankton productivity

It is usually thought that unlike terrestrial plants, phytoplankton will not show a significant response to an increase of atmospheric CO₂. Here we suggest that this view may be biased by a neglect of the effects of carbon (C) assimilation on the pH and the dissociation of the C species. We show that under eutrophic conditions, productivity may double as a result of doubling of the atmospheric CO₂ concentration. Although in practice productivity increase will usually be less, we still predict a productivity increase of up to 40% in marine species with a low affinity for bicarbonate. In eutrophic freshwater systems doubling of atmospheric CO₂ may result in an increase of the productivity of more than 50%. Freshwaters with low alkalinity appeared to be very sensitive to atmospheric CO₂ elevation. Our results suggest that the aquatic C sink may increase more than expected, and that nuisance phytoplankton blooms may be aggravated at elevated atmospheric CO₂ concentrations.     

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.     

Soil microbial responses to increased concentrations of atmospheric CO2

Terrestrial ecosystems respond to an increased concentration of atmospheric CO₂. While elevated atmospheric CO₂ has been shown to alter plant growth and productivity, it also affects ecosystem structure and function by changing below-ground processes. Knowledge of how soil microbiota respond to elevated atmospheric CO₂ is of paramount importance for understanding global carbon and nutrient cycling and for predicting changes at the ecosystem-level. An increase in the atmospheric CO₂ concentration not only alters the weight, length, and architecture of plant roots, but also affects the biotic and abiotic environment of the root system. Since the concentration of CO₂ in soil is already 10–50 times higher than that in the atmosphere, it is unlikely that increasing atmospheric CO₂ will directly influence the rhizosphere. Rather, it is more likely that elevated atmospheric CO₂ will affect the microbe–soil–plant root system indirectly by increasing root growth and rhizodeposition rates, and decreasing soil water deficit. Consequently, the increased amounts and altered composition of rhizosphere-released materials will have the potential to alter both population and community structure, and activity of soil- and rhizosphere-associated microorganisms. This occurrence could in turn affect plant health and productivity and plant community structure. This review covers current knowledge about the response of soil microbes to elevated concentrations of atmospheric CO₂.     

Respiratory responses of higher plants to atmospheric CO2 enrichment

Although the respiratory response of native and agricultural plants to atmospheric CO₂ enrichment has been reported over the past 75 years, only recently have these effects emerged as prominent measures of plant and ecosystem response to the earth's changing climate. In this review we discuss this rapidly expanding field of study and propose that both increasing and decreasing rates of leaf and whole-plant respiration are likely to occur in response to rising CO₂ concentrations. While the stimulatory effects of CO₂ on respiration are consistent with our knowledge of leaf carbohydrate status and plant metabolism, we wish to emphasize the rather surprising short-term inhibition of leaf respiration by elevated CO₂ and the reported effects of long-term CO₂ exposure on growth and maintenance respiration. As is being found in many studies, it is easier to document the respiratory response of higher plants to elevated CO₂ than it is to assign a mechanistic basis for the observed effects. Despite this gap in our understanding of how respiration is affected by CO₂ enrichment, data are sufficient to suggest that changes in leaf and whole-plant respiration may be important considerations in the carbon dynamics of terrestrial ecosystems as global CO₂ continues to rise. Suggestions for future research that would enable these and other effects of CO₂ on respiration to be unravelled are presented.     

Biomass production and species composition change in a tallgrass prairie ecosystem after long-term exposure to elevated atmospheric CO2

To determine the long-term impact of elevated CO₂ on primary production of native tallgrass prairie, we compared the responses of tallgrass prairie at ambient and twice-ambient atmospheric CO₂ levels over an 8-year period. Plots in open-top chambers (4.5 m diameter) were exposed continuously (24 h) to ambient and elevated CO₂ from early April to late October each year. Unchambered plots were monitored also. Above-ground peak biomass was determined by clipping each year in early August, and root growth was estimated by harvesting roots from root ingrowth bags. Plant community composition was censused each year in early June. In the last 2 years of the study, subplots were clipped on 1 June or 1 July, and regrowth was harvested on 1 October. Volumetric soil water content of the 0–100 cm soil layer was determined using neutron scattering, and was generally higher in elevated CO₂ plots than ambient. Peak above-ground biomass was greater on elevated CO₂ plots than ambient CO₂ plots with or without chambers during years with significant plant water stress. Above-ground regrowth biomass was greater under elevated CO₂ than under ambient CO₂ in a year with late-season water stress, but did not differ in a wetter year. Root ingrowth biomass was also greater in elevated CO₂ plots than ambient CO₂ plots when water stress occurred during the growing season. The basal cover and relative amount of warm-season perennial grasses (C4) in the stand changed little during the 8-year period, but basal cover and relative amount of cool-season perennial grasses (C3) in the stand declined in the elevated CO₂ plots and in ambient CO₂ plots with chambers. Forbs (C3) and members of the Cyperaceae (C3) increased in basal cover and relative amount in the stand at elevated compared to ambient CO₂. Greater biomass production under elevated CO₂ in C4-dominated grasslands may lead to a greater carbon sequestration by those ecosystems and reduce peak atmospheric CO₂ concentrations in the future.     

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.     

Soil acidification induced by elevated atmospheric CO2

Soil acidification is a very important process in the functioning of earth's ecosystems. A major source of soil acidity is CO₂, derived from the respiration of plant roots and microbes, which forms carbonic acid in soil waters. Because elevated atmospheric CO₂ often stimulates respiration of soil biota in experiments that test ecosystem effects of elevated atmospheric CO₂, we hypothesize that rising atmospheric CO₂ (which has increased from ～200 ppm since the interglacial and may exceed 550 ppm by the end of the 21st century) is significantly increasing acid inputs to soils. Here, using column‐leaching experiments with contrasting soils, we demonstrate that soil CO₂ is a much more potent agent of soil acidification than is generally appreciated, capable of displacing almost all exchangeable base cations in soils, and even elevating Al(III) concentrations in H₂CO₃‐acidified soil waters. The potent soil acidifying potential of soil H₂CO₃ is attributed to the low pK_a,1 of molecular H₂CO₃ (3.76 at 25°C), which contrasts greatly with that of (a convention that combines CO₂ (aq) and molecular H₂CO₃, the pK_a,1 of which is 6.36 at 25°C). This distinction is significant for soil systems because of soil's greatly elevated CO₂, their variety of sinks for H⁺, and the wide range of contact times between soil solids, water, and gas. Modelling suggests that a doubling of atmospheric CO₂ may increase acid inputs from carbonic acid leaching by up to 50%. Combined with the results of CO₂ studies in whole ecosystems, this implies that increases in atmospheric CO₂ since the interglacial have gradually acidified soils, especially poorly buffered soils, throughout the world.