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.     

The effects of increased atmospheric carbon dioxide and temperature on carbon partitioning, source-sink relations and respiration

Abstract. Herbaceous C₃ plants grown in elevated CO₂ show increases in carbon assimilation and carbohydrate accumulation (particularly starch) within source leaves. Although changes in the partitioning of biomass between root and shoot occur, the proportion of this extra assimilate made available for sink growth is not known. Root:shoot ratios tend to increase for CO₂-enriched herbaceous plants and decrease for CO₂-enriched trees. Root:shoot ratios for cereals tend to remain constant. In contrast, elevated temperatures decrease carbohydrate accumulation within source and sink regions of a plant and decrease root:shoot ratios. Allometric analysis of at least two species showing changes in root: shoot ratios due to elevated CO₂ show no alteration in the whole-plant partitioning of biomass. Little information is available for interactions between temperature and CO₂. Cold-adapted plants show little response to elevated levels of CO₂, with some species showing a decline in biomass accumulation. In general though, increasing temperature will increase sucrose synthesis, transport and utilization for CO₂-enriched plants and decrease carbohydrate accumulation within the leaf. Literature reports are discussed in relation to the hypothesis that sucrose is a major factor in the control of plant carbon partitioning. A model is presented in support.     

Root competition and elevated CO2: effects on seedling growth in Linum usitatissimum populations and Linum–Silene cretica mixtures

1. Root competition can be an important determinant of the performance of neighbours within plant populations and communities. Because plants often maintain larger root systems and allocate more of their carbon to root systems under elevated atmospheric CO₂ than they do at lower CO₂ concentrations, root–root interactions could play an increasingly important role in determining competitive outcomes among individuals and plant species as global CO₂ concentration continues to rise.
2. We established 12 pure stands of Linum usitatissimum (flax) and 12 mixed stands of Linum and its naturally co-occurring weed species Silene cretica in opaque plastic trays each filled with the same amount of nutrient-rich soil mix. In half of the trays from each of these stand types, vertical waterproof partitions separated the root systems of individual plants from each other to prevent root competition, while in the other half no partitions were present. Half of the trays from all treatments were allowed to grow under low atmospheric CO₂ concentration (320μll^–1) and the other half under elevated CO₂ (600μll^–1), in daylight growth chambers for 30 days from seedling emergence until harvest in mid-June. All trays received equal amounts of water so that soils in the low CO₂ treatment were maintained at field capacity.
3. Our results indicate that under high soil fertilities: (1) intra-specific root–root interactions alone play a relatively insignificant role in determining plant biomass production within pure Linum populations and (2) the impact of an aggressive species ( Silene ) on co-occurring less aggressive species ( Linum ) becomes more severe under elevated CO₂ as a result of amplified interspecific root competition.     

Elevated atmospheric CO2 influences the interaction between the parasitic angiosperm Orobanche minor and its host Trifolium repens

The influence of the root holoparasitic angiosperm Orobanche minor Sm. on the biomass, photosynthesis, carbohydrate and nitrogen content of Trifolium repens L. was determined for plants grown at two CO₂ concentrations (350 and 550 μmol mol⁻¹). Infected plants accumulated less biomass than their uninfected counterparts, although early in the association there was a transient stimulation of growth. Infection also influenced biomass allocation both between tissues (infected plants had lower root:shoot ratios) and within tissues:infected roots were considerably thicker before the point of parasite attachment and thinner below. Higher concentrations of starch were also found in roots above the point of attachment, particularly for plants grown in elevated CO₂. Elevated CO₂ stimulated the growth of T. repens only during the early stages of development. There was a significant interaction between infection and CO₂ on growth, with infected plants showing a greater response, such that elevated CO₂ partly alleviated the effects of the parasite on host growth. Elevated CO₂ did not affect total O. minor biomass per host, the number of individual parasites supported by each host, or their time of attachment to the host root system. Photosynthesis was stimulated by elevated CO₂ but was unaffected by O. minor . There was no evidence of down-regulation of photosynthesis in T. repens grown at elevated CO₂ in either infected or uninfected plants. The data are discussed with regard to the influence of elevated CO₂ on other parasitic angiosperm-host associations and factors which control plant responses to elevated CO₂.     

Interspecific differences in the response of arbuscular mycorrhizal fungi to Artemisia tridentata grown under elevated atmospheric CO2

Arbuscular mycorrhizal (AM) fungi form mutualistic symbioses with the root systems of most plant species. These mutualisms regulate nutrient exchange in the plant–soil interface and might influence the way in which plants respond to increasing atmospheric CO₂. In other experiments, mycorrhizal responses to elevated CO₂ have been variable, so in this study we test the hypothesis that different genera of AM fungi differ in their response, and in turn alter the plant's response, to elevated CO₂. Four species from three genera of AM fungi were tested. Artemisia tridentata Nutt. seedlings were inoculated with either Glomus intraradices Schenck & Smith, Glomus etunicatum Becker & Gerdemann, Acaulospora sp. or Scutellospora calospora (Nicol. & Gerd.) Walker & Sanders and grown at either ambient CO₂ (350 ppm) or elevated CO₂ (700 ppm). Several significant inter-specific responses were detected. Elevated CO₂ caused percent arbuscular and hyphal colonization to increase for the two Glomus species, but not for Acaulospora sp. or S. calospora . Vesicular colonization was not affected by elevated CO₂ for any fungal species. In the extra-radical phase, the two Glomus species produced a significantly higher number of spores in response to elevated CO₂, whereas Acaulospora sp. and S. calospora developed significantly higher hyphal lengths. These data show that AM fungal taxa differ in their growth allocation strategies and in their responses to elevated CO₂, and that mycorrhizal diversity should not be overlooked in global change research.     

Elevated CO2 and conifer roots: effects on growth, life span and turnover

Elevated CO₂ increases root growth and fine (diam. 2 mm) root growth across a range of species and experimental conditions. However, there is no clear evidence that elevated CO₂ changes the proportion of C allocated to root biomass, measured as either the root:shoot ratio or the fine root:needle ratio. Elevated CO₂ tends to increase mycorrhizal infection, colonization and the amount of extramatrical hyphae, supporting their key role in aiding the plant to more intensively exploit soil resources, providing a route for increased C sequestration. Only two studies have determined the effects of elevated CO₂ on conifer fine-root life span, and there is no clear trend. Elevated CO₂ increases the absolute fine-root turnover rates; however, the standing crop root biomass is also greater, and the effect of elevated CO₂ on relative turnover rates (turnover:biomass) ranges from an increase to a decrease. At the ecosystem level these changes could lead to increased C storage in roots. Increased fine-root production coupled with increased absolute turnover rates could also lead to increases in soil organic C as greater amounts of fine roots die and decompose. Although CO₂ can stimulate fine-root growth, it is not known if this stimulation persists over time. Modeling studies suggest that a doubling of the atmospheric CO₂ concentration initially increases biomass, but this stimulation declines with the response to elevated CO₂ because increases in assimilation are not matched by increases in nutrient supply.     

Coppicing shifts CO2 stimulation of poplar productivity to above-ground pools: a synthesis of leaf to stand level results from the POP/EUROFACE experiment

A poplar short rotation coppice (SRC) grown for the production of bioenergy can combine carbon (C) storage with fossil fuel substitution. Here, we summarize the responses of a poplar ( Populus ) plantation to 6 yr of free air CO₂ enrichment (POP/EUROFACE consisting of two rotation cycles). We show that a poplar plantation growing in nonlimiting light, nutrient and water conditions will significantly increase its productivity in elevated CO₂ concentrations ([CO₂]). Increased biomass yield resulted from an early growth enhancement and photosynthesis did not acclimate to elevated [CO₂]. Sufficient nutrient availability, increased nitrogen use efficiency (NUE) and the large sink capacity of poplars contributed to the sustained increase in C uptake over 6 yr. Additional C taken up in high [CO₂] was mainly invested into woody biomass pools. Coppicing increased yield by 66% and partly shifted the extra C uptake in elevated [CO₂] to above-ground pools, as fine root biomass declined and its [CO₂] stimulation disappeared. Mineral soil C increased equally in ambient and elevated [CO₂] during the 6 yr experiment. However, elevated [CO₂] increased the stabilization of C in the mineral soil. Increased productivity of a poplar SRC in elevated [CO₂] may allow shorter rotation cycles, enhancing the viability of SRC for biofuel production.     

The effect of different growth conditions on dark and light carbon assimilation in Littorella uniflora

The effect of long-term exposure to different inorganic carbon, nutrient and light regimes on CAM activity and photosynthetic performance in the submerged aquatic plant, Littorella uniflora (L.) Aschers was investigated. The potential CAM activity of Littorella was highly plastic and was reduced upon exposure to low light intensities (43 μmol m⁻² s⁻¹), high CO₂ concentrations (5.5 mM, pH 6.0) or low levels of inorganic nutrients, which caused a 25–80% decline in the potential maximum CAM activity relative to the activity in the control experiments (light: 450 μmol m⁻² s⁻¹; free CO₂: 1.5 mM). The CAM activity was regulated more by light than by CO₂, while nutrient levels only affected the activity to a minor extent. The minor effect of low nutrient regimes may be due to a general adaptation of isoetid species to low nutrient levels.
The photosynthetic capacity and CO₂ affinity was unaffected or increased by exposure to low CO₂, irrespective of nutrient levels. High CO₂, low nutrient and low light, however, reduced the capacity by 22–40% and the CO₂ affinity by 35-45%, relative to control.
The parallel effect of growth conditions on CAM activity and photosynthetic performance of Littorella suggest that light and dark carbon assimilation are interrelated and constitute an integrated part of the carbon assimilation physiology of the plant. The results are consistent with the hypothesis that CAM is a carbon-conserving mechanism in certain aquatic plants. The investment in the CAM enzyme system is beneficial to the plants during growth at high light and low CO₂ conditions.     

Nodulation and nitrogenase activity in nitrogen-fixing woody plants stimulated by CO2 enrichment of the atmosphere

The responses of three species of nitrogen-fixing trees to CO₂ enrichment of the atmosphere were investigated under nutrient-poor conditions. Seedlings of the legume, Robinia pseudoacacia L. and the actinorhizal species, Alnus glutinosa (L.) Gaertn. and Elaeagnus angustifolia L. were grown in an infertile forest soil in controlled-environment chambers with atmospheric CO₂ concentrations of 350 μl ⁻¹ (ambient) or 700 μl ⁻¹. In R. pseudoacacia and A. glutinosa , total nitrogenase (N₂ reduction) activity per plant, assayed by the acetylene reduction method, was significantly higher in elevated CO₂, because the plants were larger and had more nodule mass than did plants in ambient CO₂. The specific nitrogenase activity of the nodules, however, was not consistently or significantly affected by CO₂ enrichment. Substantial increases in plant growth occurred with CO₂ enrichment despite probable nitrogen and phosphorus deficiencies. These results support the premises that nutrient limitations will not preclude growth responses of woody plants to elevated CO₂ and that stimulation of symbiotic activity by CO₂ enrichment of the atmosphere could increase nutrient availability in infertile habitats.     

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.     

Photosynthesis, growth and nutrient changes in non-nodulated Phaseolus vulgaris grown under atmospheric and elevated carbon dioxide conditions

The response of Phaseolus vulgaris L. cv. Contender grown under controlled environment at either ambient or elevated (360 and 700 μmol mol^-1, respectively) CO₂ concentrations ([CO₂]), was monitored from 10 days after germination (DAG) until the onset of senescence. Elevated CO₂ had a pronounced effect on total plant height (TPH), leaf area (LA), leaf dry weight (LD), total plant biomass (TB) accumulation and specific leaf area (SLA). All of these were significantly increased under elevated carbon dioxide with the exception of SLA which was significantly reduced. Other than high initial growth rates in CO₂-enriched plants, relative growth rates remained relatively unchanged throughout the growth period. While the trends in growth parameters were clearly different between [CO₂], some physiological processes were largely transient, in particular, net assimilation rate (NAR) and foliar nutrient concentrations of N, Mg and Cu. CO₂ enrichment significantly increased NAR, but from 20 DAG, a steady decline to almost similar levels to those measured in plants grown under ambient CO₂ occurred. A similar trend was observed for leaf N content where the loss of leaf nitrogen in CO₂-enriched plants after 20 DAG, was significantly greater than that observed for ambient-CO₂ plants. Under enhanced CO₂, the foliar concentrations of K and Mn were increased significantly whilst P, Ca, Fe and Zn were reduced significantly. Changes in Mg and Cu concentrations were insignificant. In addition. high CO₂ grown plants exhibited a pronounced leaf discoloration or chlorosis, coupled with a significant reduction in leaf longevity.     

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.     

Growth and physiological responses of canola (Brassica napus) to UV-B and CO2 under controlled environment conditions

Few studies have investigated the interaction of ultraviolet (UV)-B radiation and CO₂ concentration on plants. We studied the combined effects of UV-B radiation and CO₂ concentration on canola ( Brassica napus cv. 46A65) under four growth conditions – ambient CO₂ with UV-B (control), elevated CO₂ with UV-B, ambient CO₂ without UV-B, and elevated CO₂ without UV-B – to determine whether the adverse effects of UV-B are mitigated by elevated CO₂. Elevated CO₂ significantly increased plant height and seed yield, whereas UV-B decreased them. Elevated CO₂ ameliorated the adverse effects of UV-B in plant height. UV-B did not affect the physical characteristics of leaf but CO₂ did. Certain flower and fruit characteristics were affected negatively by UV-B and positively by CO₂. UV-B did not affect net photosynthesis, transpiration and stomatal conductance but decreased water use efficiency (WUE). Elevated CO₂ significantly increased net photosynthesis and WUE. Neither UV-B nor CO₂ affected chlorophyll (Chl) fluorescence. UV-B significantly decreased Chl b and increased the ratio of Chl a / b . Elevated CO₂ decreased only the ratio of Chl a / b . UV-B significantly increased UV-absorbing compounds while CO₂ had no effect on them. Both UV-B and CO₂ significantly increased epicuticular wax content. Many significant relationships were found between morphological, physiological, and chemical parameters. This study showed that elevated CO₂ can partially ameliorate some of the adverse effects of UV-B radiation in B . napus .     

Biomass, reproductive output, and physiological responses of rapid-cycling Brassica (Brassica rapa) to ozone and modified root temperature

Brassica rapa L. (rapid-cycling Brassica), was grown in environmentally controlled chambers to determine the interactive effects of ozone (O₃) and increased root temperature (RT) on biomass, reproductive output, and photosynthesis. Plants were grown with or without an average treatment of 63 ppb O₃. RT treatments were 13°C (LRT) and 18°C (HRT). Air temperatures were 25°C/15°C day/night for all RT treatments.
Ozone affected plant biomass more than did root temperature. Plants in O₃ had significantly smaller total plant d. wt, shoot weight, leaf weight, leaf area and leaf number than plants grown without O₃. LRT plants tended to have slightly smaller total plant d. wt, shoot weight, root weight, leaf weight, leaf area, and leaf number than HRT plants. For all variables, LRT plants grown in O₃ had the smallest biomass, and plants grown in HRT without O₃ had the largest biomass.
Ozone reduced both fruit weight and fruit number; LRT also reduced fruit weight but had no effect on fruit number. Ozone reduced photosynthesis but RT had no effect. Conductance and internal CO₂ were unaffected by O₃ or RT.
These studies indicate that plant growth with LRT might be more reduced in the presence of O₃ than growth in plants with HRT, which might be able to compensate for O₃-caused reductions in photosynthesis to avoid decreased biomass and reproductive output.     

The effect of an elevated atmospheric CO2 concentration on growth, photosynthesis and respiration of Plantago major

The effect of an elevated atmospheric CO₂ concentration on growth, photosynthesis and root respiration of Plantago major L. ssp. major L. was investigated. Plants were grown in a nutrient solution in growth chambers at 350 and 700 μl I⁻¹ CO₂ during 7 weeks. The total dry weight of the Co₂-enriched plants at the end of this period was 50% higher than that of control plants. However, the relative growth rate (RGR) was stimulated only during the first half of the growing period. The transient nature of the stimulation of the RGR was not likely to be due to end-product inhibition of photosynthesis. It is suggested that in P. major , a rosette plant, self-shading causes a decline in photosynthesis and results in an increase in the shoot: root ratio and a decrease in RGR. CO₂-enriched plants grow faster and cosequently suffer more from self-shading. Corrected for this ontogenetic drift, high CO₂ concentrations stimulated the RGR of P. major throughout the entire experiment.     

Effect of root zone carbon dioxide enrichment on ethylene inhibition of carbon assimilation in potato plants

Potato plants ( Solanum tuberosum L. var. Russet Burbank) treated with 1 μl ethylene 1⁻¹ of air showed an inhibition of CO₂ assimilation by 18%. The inhibition occurred after 3 h of exposure to ethylene and was not mediated through closure of the stomata. The enrichment of the root zone with CO₂ almost completely abolished the ethylene inhibition of CO₂ assimilation which was apparently due to an increase in the intercellular concentration of CO₂ in leaves following enrichment. The effect of application of CO₂ to the root zone on ethylene inhibition of CO₂ assimilation seemed to last for a few days. Potato plants treated with aminoethoxyvinlglycine (AVG) showed an increase in fresh and dry weight as compared to non-treated plants. Our results indicate that both CO₂ and AVG alter the effect of ethylene and promote growth in plants by inhibiting ethylene action and biosynthesis, respectively.     

Why do fast- and slow-growing grass species differ so little in their rate of root respiration,considering the large differences in rate of growth and ion uptake?

Herbaceous plants grown with free access to nutrients exhibit inherent differences in maximum relative growth rate (RGR) and rate of nutrient uptake. Measured rates of root respiration are higher in fast-growing species than in slow-growing ones. Fast-growing herbaceous species, however, exhibit lower rates of respiration than would be expected from their high rates of growth and nitrate uptake. We investigated why the difference in root O₂ uptake between fast- and slow-growing species is relatively small. Inhibition of respiration by the build-up of CO₂ in closed cuvettes, diurnal variation in respiration rates or an increasing ratio of respiratory CO₂ release to O₂ uptake (RQ) with increasing RGR failed to explain the relatively low root respiration rates in fast-growing grasses. Furthermore, differences in alternative pathway activity can at most only partly explain why the difference in root respiration between fast- and slow-growing grasses is relatively small. Although specific respiratory costs for maintenance of biomass are slightly higher in the fast-growing Dactylis glomerata L. than those in the slow-growing Festuca ovina L., they account for 50% of total root respiration in both species. The specific respiratory costs for ion uptake in the fast-growing grass are one-third of those in the slow-growing grass [0·41 versus 1·22 mol O₂ mol (NO₃^–)^–1]. We conclude that this is the major cause of the relatively low rates of root respiration in fast-growing grasses.     

Increasing growth temperature reduces the stimulatory effect of elevated CO2 on photosynthesis or biomass in two perennial species

We examined how anticipated changes in CO₂ concentration and temperature interacted to alter plant growth, harvest characteristics and photosynthesis in two cold-adapted herbaceous perennials, alfalfa ( Medicago sativa L. cv. Arc) and orchard grass ( Dactylis glomerata L. cv. Potomac). Plants were grown at two CO₂ concentrations (362 [ambient] and 717 [elevated] μmol mol⁻¹ CO₂) and four constant day/night temperatures of 15, 20, 25 and 30°C in controlled environmental chambers. Elevated CO₂ significantly increased total plant biomass and protein over a wide range of temperatures in both species. Stimulation of photosynthetic rate, however, was eliminated at the highest growth temperature in M. sativa and relative stimulation of plant biomass and protein at high CO₂ declined as temperature increased in both species. Lack of a synergistic effect between temperature and CO₂ was unexpected since elevated CO₂ reduces the amount of carbon lost via photorespiration and photorespiration increases with temperature. Differences between anticipated stimulatory effects of CO₂ and temperature and whole plant single and leaf measurements are discussed. Data from this study suggest that stimulatory effects of atmospheric CO₂ on growth and photosynthesis may decline with anticipated increases in global temperature, limiting the degree of carbon storage in these two perennial species.     

Responses to elevated temperature and CO2 in the perennial grass Agrostis curtisii in relation to population origin

1. Evolutionary responses to climate change will depend on the presence of heritable variation within species populations for traits that increase fitness under the changing conditions. Patterns of ecotypic differentiation in relation to latitude in some species suggest that such variation exists in relation to temperature responses. Response to elevated CO₂, whether heritable or not, is not expected to be related to latitudinal or climatic differences within temperate regions.
2. To test these ideas, seeds were collected from 10 populations of the outbreeding perennial grass Agrostis curtisii across its range in Europe from south Wales to Portugal. Plants were grown under ambient and elevated temperature and CO₂ conditions, in a factorial design, in solardomes; two half sibs from each population were planted in separate pots in each of the two replicate domes with each combination of treatments. One half sib was harvested at the end of the first summer, the second at the end of the second summer.
3. Survival was uniformly high and flowering uniformly low across treatments and populations.
4. Responses to temperature and CO₂ treatments varied over time for almost all populations. Treatment effects were not significant on plants harvested in year 1, although there was a trend towards higher shoot biomass under the elevated temperature and CO₂ treatment. In year 2 shoot biomass was significantly higher under the elevated temperature treatment across all populations and there was a strong trend towards decreased biomass under elevated CO₂.
5. There were no significant correlations of plant response to either CO₂ or temperature with climate at origin.
6. These results warn of the dangers of extrapolating evolutionary plant responses to CO₂ from short-term experiments.     

The cellular basis of guard cell sensing of rising CO2

Numerous studies conducted on both whole plants and isolated epidermes have documented stomatal sensitivity to CO₂. In general, CO₂ concentrations below ambient stimulate stomatal opening, or an inhibition of stomatal closure, while CO₂ concentrations above ambient have the opposite effect. The rise in atmospheric CO₂ concentrations which has occurred since the industrial revolution, and which is predicted to continue, will therefore alter rates of transpirational water loss and CO₂ uptake in terrestrial plants. An understanding of the cellular basis for guard cell CO₂ sensing could allow us to better predict, and perhaps ultimately to manipulate, such vegetation responses to climate change. However, the mechanisms by which guard cells sense and respond to the CO₂ signal remain unknown. It has been hypothesized that cytosolic pH and malate levels, cytosolic Ca²⁺ levels, chloroplastic zeaxanthin levels, or plasma-membrane anion channel regulation by apoplastic malate are involved in guard cell perception and response to CO₂. In this review, these hypotheses are discussed, and the evidence for guard cell acclimation to prevailing CO₂ concentrations is also considered.