Canopy photosynthesis of crops and native plant communities exposed to long-term elevated CO2

Abstract. There have been seven studies of canopy photosynthesis of plants grown in elevated atmospheric CO₂: three of seed crops, two of forage crops and two of native plant ecosystems. Growth in elevated CO₂ increased canopy photosynthesis in all cases. The relative effect of CO₂ was correlated with increasing temperature: the least stimulation occurred in tundra vegetation grown at an average temperature near 10°C and the greatest in rice grown at 43°C. In soybean, effects of CO₂ were greater during leaf expansion and pod fill than at other stages of crop maturation. In the longest running experiment with elevated CO₂ treatment to date, monospecific stands of a C₃ sedge, Scirpus olneyi (Grey), and a C₄ grass, Spartina patens (Ait.) Muhl., have been exposed to twice normal ambient CO₂ concentrations for four growing seasons, in open top chambers on a Chesapeake Bay salt marsh. Net ecosystem CO₂ exchange per unit green biomass (NCE_b) increased by an average of 48% throughout the growing season of 1988, the second year of treatment. Elevated CO₂ increased net ecosystem carbon assimilation by 88% in the Scirpus olneyi community and 40% in the Spartina patens community.     

The effects of elevated atmospheric CO2 on the amount and depth distribution of plant water uptake in a California annual grassland

Soil moisture profiles can affect species composition and ecosystem processes, but the effects of increased concentrations of atmospheric carbon dioxide ([CO₂]) on the vertical distribution of plant water uptake have not been studied. Because plant species composition affects soil moisture profiles, and is likely to shift under elevated [CO₂], it is also important to test whether the indirect effects of [CO₂] on soil water content may depend on species composition. We examined the effects of elevated [CO₂] and species composition on soil moisture profiles in an annual grassland of California. We grew monocultures and a mixture of Avena barbata and Hemizonia congesta– the dominant species of two phenological groups – in microcosms exposed to ambient (～370 μmol mol^?1) and elevated (～700 μmol mol^?1) [CO₂]. Both species increased intrinsic and yield‐based water use efficiency under elevated [CO₂], but soil moisture increased only in communities with A. barbata, the dominant early‐season annual grass. In A. barbata monocultures, the [CO₂] treatment did not affect the depth distribution of soil water loss. In contrast to communities with A. barbata, monocultures of H. congesta, a late‐season annual forb, did not conserve water under elevated [CO₂], reflecting the increased growth of these plants. In late spring, elevated [CO₂] also increased the efficiency of deep roots in H. congesta monocultures. Under ambient [CO₂], roots below 60 cm accounted for 22% of total root biomass and were associated with 9% of total water loss, whereas in elevated [CO₂], 16% of total belowground biomass was associated with 34% of total water loss. Both soil moisture and isotope data showed that H. congesta monocultures grown under elevated [CO₂] began extracting water from deep soils 2 weeks earlier than plants in ambient [CO₂].     

Leaf carbon management in slow-growing plants exposed to elevated CO2

Two slow‐growing plant species (Chamaerops humilis, L. and Cycas revoluta Thunb.) were exposed to elevated CO₂ conditions over a 20‐month period in order to study the CO₂ effect on growth, photosynthetic capacity and leaf carbon (C) management. The ambient isotopic ¹³C/¹²C composition (δ¹³C) of the greenhouse module corresponding to elevated CO₂ (800 μmol mol^?1 CO₂) conditions was changed from δ¹³C ca. ?12.8±0.3‰ to ca. ?19.2±0.2‰. Exposure of these plants to elevated CO₂ enhanced dry mass (DM) by 82% and 152% in Chamerops and Cycas, respectively, mainly as a consequence of increases in plant level photosynthetic rates. However, analyses of A–C_i curve parameters revealed that elevated CO₂ diminished leaf photosynthetic rates of Chamaerops whereas in Cycas, no photosynthetic acclimation was detected. The fact that Chamaerops plants had a lower DM increase, together with a longer leaf C residence time and a diminished capacity to respire recently fixed C, suggests that this species was unable to increase C sink strength. Furthermore, the consequent C source/sink imbalance in Chamaerops might have induced the downregulation of Rubisco. Cycas plants were capable of avoiding photosynthetic downregulation due to a greater ability to increase C sink strength, as was confirmed by DM values, and ¹²C‐enriched CO₂ labeling data. Cycas developed the ability to respire a larger proportion of recently fixed C and to reallocate the recently fixed C away from leaves to other plant tissues. These findings suggest that leaf C management is a key factor in the responsiveness of slow‐growing plants to future CO₂ scenarios.     

The turnover of carbon pools contributing to soil CO2 and soil respiration in a temperate forest exposed to elevated CO2 concentration

Soil carbon is returned to the atmosphere through the process of soil respiration, which represents one of the largest fluxes in the terrestrial C cycle. The effects of climate change on the components of soil respiration can affect the sink or source capacity of ecosystems for atmospheric carbon, but no current techniques can unambiguously separate soil respiration into its components. Long‐term free air CO₂ enrichment (FACE) experiments provide a unique opportunity to study soil C dynamics because the CO₂ used for fumigation has a distinct isotopic signature and serves as a continuous label at the ecosystem level. We used the ¹³C tracer at the Duke Forest FACE site to follow the disappearance of C fixed before fumigation began in 1996 (pretreatment C) from soil CO₂ and soil‐respired CO₂, as an index of belowground C dynamics during the first 8 years of the experiment. The decay of pretreatment C as detected in the isotopic composition of soil‐respired CO₂ and soil CO₂ at 15, 30, 70, and 200 cm soil depth was best described by a model having one to three exponential pools within the soil system. The majority of soil‐respired CO₂ (71%) originated in soil C pools with a turnover time of about 35 days. About 55%, 50%, and 68% of soil CO₂ at 15, 30, and 70 cm, respectively, originated in soil pools with turnover times of less than 1 year. The rest of soil CO₂ and soil‐respired CO₂ originated in soil pools that turn over at decadal time scales. Our results suggest that a large fraction of the C returned to the atmosphere through soil respiration results from dynamic soil C pools that cannot be easily detected in traditionally defined soil organic matter standing stocks. Fast oxidation of labile C substrates may prevent increases in soil C accumulation in forests exposed to elevated [CO₂] and may consequently result in shorter ecosystem C residence times.     

Growth in elevated CO2 protects photosynthesis against high-temperature damage

We present evidence that plant growth at elevated atmospheric CO₂ increases the high‐temperature tolerance of photosynthesis in a wide variety of plant species under both greenhouse and field conditions. We grew plants at ambient CO₂ (~ 360 μ mol mol ^{? 1}) and elevated CO₂ (550–1000 μ mol mol ^{? 1}) in three separate growth facilities, including the Nevada Desert Free‐Air Carbon Dioxide Enrichment (FACE) facility. Excised leaves from both the ambient and elevated CO₂ treatments were exposed to temperatures ranging from 28 to 48 °C. In more than half the species examined (4 of 7, 3 of 5, and 3 of 5 species in the three facilities), leaves from elevated CO₂‐grown plants maintained PSII efficiency (F_v/F_m) to significantly higher temperatures than ambient‐grown leaves. This enhanced PSII thermotolerance was found in both woody and herbaceous species and in both monocots and dicots. Detailed experiments conducted with Cucumis sativus showed that the greater F_v/F_m in elevated versus ambient CO₂‐grown leaves following heat stress was due to both a higher F_m and a lower F_o, and that F_v/F_m differences between elevated and ambient CO₂‐grown leaves persisted for at least 20 h following heat shock. Cucumis sativus leaves from elevated CO₂‐grown plants had a critical temperature for the rapid rise in F_o that averaged 2·9 °C higher than leaves from ambient CO₂‐grown plants, and maintained a higher maximal rate of net CO₂ assimilation following heat shock. Given that photosynthesis is considered to be the physiological process most sensitive to high‐temperature damage and that rising atmospheric CO₂ content will drive temperature increases in many already stressful environments, this CO₂‐induced increase in plant high‐temperature tolerance may have a substantial impact on both the productivity and distribution of many plant species in the 21st century.     

Plant and microbial N acquisition under elevated atmospheric CO2 in two mesocosm experiments with annual grasses

The impact of elevated CO₂ on terrestrial ecosystem C balance, both in sign or magnitude, is not clear because the resulting alterations in C input, plant nutrient demand and water use efficiency often have contrasting impacts on microbial decomposition processes. One major source of uncertainty stems from the impact of elevated CO₂ on N availability to plants and microbes. We examined the effects of atmospheric CO₂ enrichment (ambient+370 μmol mol^?1) on plant and microbial N acquisition in two different mesocosm experiments, using model plant species of annual grasses of Avena barbata and A. fatua, respectively. The A. barbata experiment was conducted in a N‐poor sandy loam and the A. fatua experiment was on a N‐rich clayey loam. Plant–microbial N partitioning was examined through determining the distribution of a ¹⁵N tracer. In the A. barbata experiment, ¹⁵N tracer was introduced to a field labeling experiment in the previous year so that ¹⁵N predominantly existed in nonextractable soil pools. In the A. fatua experiment, ¹⁵N was introduced in a mineral solution [(¹⁵NH₄)₂SO₄ solution] during the growing season of A. fatua. Results of both N budget and ¹⁵N tracer analyses indicated that elevated CO₂ increased plant N acquisition from the soil. In the A. barbata experiment, elevated CO₂ increased plant biomass N by ca. 10% but there was no corresponding decrease in soil extractable N, suggesting that plants might have obtained N from the nonextractable organic N pool because of enhanced microbial activity. In the A. fatua experiment, however, the CO₂‐led increase in plant biomass N was statistically equal to the reduction in soil extractable N. Although atmospheric CO₂ enrichment enhanced microbial biomass C under A. barbata or microbial activity (respiration) under A. fatua, it had no significant effect on microbial biomass N in either experiment. Elevated CO₂ increased the colonization of A. fatua roots by arbuscular mycorrhizal fungi, which coincided with the enhancement of plant competitiveness for soluble soil N. Together, these results suggest that elevated CO₂ may tighten N cycling through facilitating plant N acquisition. However, it is unknown to what degree results from these short‐term microcosm experiments can be extrapolated to field conditions. Long‐term studies in less‐disturbed soils are needed to determine whether CO₂‐enhancement of plant N acquisition can significantly relieve N limitation over plant growth in an elevated CO₂ environment.     

Seed production and seed quality in a calcareous grassland in elevated CO2

In diverse plant communities the relative contribution of species to community biomass may change considerably in response to elevated CO₂. Along with species‐specific biomass responses, reproduction is likely to change as well with increasing CO₂ and might further accelerate shifts in species composition. Here, we ask if, after 5 years of CO₂ exposure, seed production and seed quality in natural nutrient‐poor calcareous grassland are affected by elevated CO₂ (650 μ L L^?1 vs 360 μ L L^?1) and how this might affect long‐term community dynamics. The effect of elevated CO₂ on the number of flowering shoots (+ 24%, P < 0.01) and seeds (+ 29%, P = 0.06) at the community level was similar to above ground biomass responses in this year, suggesting that the overall allocation to sexual reproduction remained unchanged. Compared among functional groups of species we found a 42% increase in seed number (P < 0.01) of graminoids, a 33% increase (P = 0.07) in forbs, and no significant change in legumes (? 38%, n.s.) under elevated CO₂. Large responses particularly of two graminoid species and smaller responses of many forb species summed up to the significant or marginally significant increase in seed number of graminoids and forbs, respectively. In several species the increase in seed number resulted both from an increase in flowering shoots and an increase in inflorescence size. In most species, seeds tended to be heavier (+ 12%, P < 0.01), and N‐concentration of seeds was significantly reduced in eight out of 13 species. The fraction of germinating seeds did not differ between seeds produced in ambient and elevated CO₂, but time to germination was significantly shortened in two species and prolonged in one species when seeds had been produced in elevated CO₂. Results suggest that species specific increases in seed number and changes in seed quality will exert substantial cumulative effects on community composition in the long run.     

Responses of net ecosystem CO2 exchange in managed grassland to long-term CO2 enrichment, N fertilization and plant species

The effects of elevated pCO₂ on net ecosystem CO₂ exchange were investigated in managed Lolium perenne (perennial ryegrass) and Trifolium repens (white clover) monocultures that had been exposed continuously to elevated pCO₂ (60 Pa) for nine growing seasons using Free Air CO₂ Enrichment (FACE) technology. Two levels of nitrogen (N) fertilization were applied. Midday net ecosystem CO₂ exchange (mNEE) and night-time ecosystem respiration (NER) were measured in three growing seasons using an open-flow chamber system. The annual net ecosystem carbon (C) input resulting from the net CO₂ fluxes was estimated for one growing season. In both monocultures and at both levels of N supply, elevated pCO₂ stimulated mNEE by up to 32%, the exact amount depending on intercepted PAR. The response of mNEE to elevated pCO₂ was larger than that of harvestable biomass. Elevated pCO₂ increased NER by up to 39% in both species at both levels of N supply. NER, which was affected by mNEE of the preceding day, was higher in T. repens than in L. perenne. High N increased NER compared to low N supply. According to treatment, the annual net ecosystem C input ranged between 210 and 631 g C m⁻² year⁻¹ and was not significantly affected by the level of pCO₂. Low N supply led to a higher net C input than high N supply. We demonstrated that at the ecosystem level, there was a long-term stimulation in the net C assimilation during daytime by elevated pCO₂. However, because NER was also stimulated, net ecosystem C input was not significantly increased at elevated pCO₂. The annual net ecosystem C input was primarily affected by the amount of N supplied.     

Increasing CO2 from subambient to elevated concentrations increases grassland respiration per unit of net carbon fixation

Respiration (carbon efflux) by terrestrial ecosystems is a major component of the global carbon (C) cycle, but the response of C efflux to atmospheric CO₂ enrichment remains uncertain. Respiration may respond directly to an increase in the availability of C substrates at high CO₂, but also may be affected indirectly by a CO₂‐mediated alteration in the amount by which respiration changes per unit of change in temperature or C uptake (sensitivity of respiration to temperature or C uptake). We measured CO₂ fluxes continuously during the final 2 years of a 4‐year experiment on C₃/C₄ grassland that was exposed to a 200–560 μmol mol^?1 CO₂ gradient. Flux measurements were used to determine whether CO₂ treatment affected nighttime respiration rates and the response of ecosystem respiration to seasonal changes in net C uptake and air temperature. Increasing CO₂ from subambient to elevated concentrations stimulated grassland respiration at night by increasing the net amount of C fixed during daylight and by increasing either the sensitivity of C efflux to daily changes in C fixation or the respiration rate in the absence of C uptake (basal ecosystem respiration rate). These latter two changes contributed to a 30–47% increase in the ratio of nighttime respiration to daytime net C influx as CO₂ increased from subamient to elevated concentrations. Daily changes in net C uptake were highly correlated with variation in temperature, meaning that the shared contribution of C uptake and temperature in explaining variance in respiration rates was large. Statistically controlling for collinearity between temperature and C uptake reduced the effect of a given change in C influx on respiration. Conversely, CO₂ treatment did not affect the response of grassland respiration to seasonal variation in temperature. Elevating CO₂ concentration increased grassland respiration rates by increasing both net C input and respiration per unit of C input. A better understanding of how C efflux varies with substrate supply thus may be required to accurately assess the C balance of terrestrial ecosystems.     

Responses of a C3 and a C4 perennial grass to elevated CO2 and temperature under different water regimes

An experiment was carried out to determine the effects of elevated CO₂, elevated temperatures, and altered water regimes in native shortgrass steppe. Intact soil cores dominated by Bouteloua gracilis, a C₄ perennial grass, or Pascopyrum smithii, a C₃ perennial grass, were placed in growth chambers with 350 or 700 μL L^?1 atmospheric CO₂, and under either normal or elevated temperatures. The normal regime mimicked field patterns of diurnal and seasonal temperatures, and the high-temperature regime was 4 °C warmer. Water was supplied at three different levels in a seasonal pattern similar to that observed in the field. Total biomass after two growing seasons was 19% greater under elevated CO₂, with no significant difference between the C₃ and C₄ grass. The effect of elevated CO₂ on biomass was greatest at the intermediate water level. The positive effect of elevated CO₂ on shoot biomass was greater at normal temperatures in B. gracilis, and greater at elevated temperatures in P. smithii. Neither root-to-shoot ratio nor production of seed heads was affected by elevated CO₂. Plant tissue N and soil inorganic N concentrations were lower under elevated Co₂, but no more so in the C₃ than the C₄ plant. Elevated CO₂ appeared to increase plant N limitation, but there was no strong evidence for an increase in N limitation or a decrease in the size of the CO₂ effect from the first to the second growing season. Autumn samples of large roots plus crowns, the perennial organs, had 11% greater total N under elevated CO₂, in spite of greater N limitation.     

Elevated CO2 and plant structure: a review

Consequences of increasing atmospheric CO₂ concentration on plant structure, an important determinant of physiological and competitive success, have not received sufficient attention in the literature. Understanding how increasing carbon input will influence plant developmental processes, and resultant form, will help bridge the gap between physiological response and ecosystem level phenomena. Growth in elevated CO₂ alters plant structure through its effects on both primary and secondary meristems of shoots and roots. Although not well established, a review of the literature suggests that cell division, cell expansion, and cell patterning may be affected, driven mainly by increased substrate (sucrose) availability and perhaps also by differential expression of genes involved in cell cycling (e.g. cyclins) or cell expansion (e.g. xyloglucan endotransglycosylase). Few studies, however, have attempted to elucidate the mechanistic basis for increased growth at the cellular level. Regardless of specific mechanisms involved, plant leaf size and anatomy are often altered by growth in elevated CO₂, but the magnitude of these changes, which often decreases as leaves mature, hinges upon plant genetic plasticity, nutrient availability, temperature, and phenology. Increased leaf growth results more often from increased cell expansion rather than increased division. Leaves of crop species exhibit greater increases in leaf thickness than do leaves of wild species. Increased mesophyll and vascular tissue cross-sectional areas, important determinates of photosynthetic rates and assimilate transport capacity, are often reported. Few studies, however, have quantified characteristics more reflective of leaf function such as spatial relationships among chlorenchyma cells (size, orientation, and surface area), intercellular spaces, and conductive tissue. Greater leaf size and/or more leaves per plant are often noted; plants grown in elevated CO₂ exhibited increased leaf area per plant in 66% of studies, compared to 28% of observations reporting no change, and 6% reported a decrease in whole plant leaf area. This resulted in an average net increase in leaf area per plant of 24%. Crop species showed the greatest average increase in whole plant leaf area (+ 37%) compared to tree species (+ 14%) and wild, nonwoody species (+ 15%). Conversely, tree species and wild, nontrees showed the greatest reduction in specific leaf area (– 14% and – 20%) compared to crop plants (– 6%). Alterations in developmental processes at the shoot apex and within the vascular cambium contributed to increased plant height, altered branching characteristics, and increased stem diameters. The ratio of internode length to node number often increased, but the length and sometimes the number of branches per node was greater, suggesting reduced apical dominance. Data concerning effects of elevated CO₂ on stem/branch anatomy, vital for understanding potential shifts in functional relationships of leaves with stems, roots with stems, and leaves with roots, are too few to make generalizations. Growth in elevated CO₂ typically leads to increased root length, diameter, and altered branching patterns. Altered branching characteristics in both shoots and roots may impact competitive relationships above and below the ground. Understanding how increased carbon assimilation affects growth processes (cell division, cell expansion, and cell patterning) will facilitate a better understanding of how plant form will change as atmospheric CO₂ increases. Knowing how basic growth processes respond to increased carbon inputs may also provide a mechanistic basis for the differential phenotypic plasticity exhibited by different plant species/functional types to elevated CO₂.     

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.     

The sensitivity of C3 photosynthesis to increasing CO2 concentration: a theoretical analysis of its dependence on temperature and background CO2 concentration

The atmospheric CO₂ concentration has increased from the pre-industrial concentration of about 280 μmol mol⁻¹ to its present concentration of over 350 μmol mol⁻¹, and continues to increase. As the rate of photosynthesis in C₃ plants is strongly dependent on CO₂ concentration, this should have a marked effect on photosynthesis, and hence on plant growth and productivity. The magnitude of photo-synthetic responses can be calculated based on the well-developed theory of photosynthetic response to intercellular CO₂ concentration. A simple biochemically based model of photosynthesis was coupled to a model of stomatal conductance to calculate photosynthetic responses to ambient CO₂ concentration. In the combined model, photosynthesis was much more responsive to CO₂ at high than at low temperatures. At 350 μmol mol⁻¹, photosynthesis at 35°C reached 51% of the rate that would have been possible with non-limiting CO₂, whereas at 5°C, 77% of the CO₂ non-limited rate was attained. Relative CO₂ sensitivity also became smaller at elevated CO₂, as CO₂ concentration increased towards saturation. As photosynthesis was far from being saturated at the current ambient CO₂ concentration, considerable further gains in photosynthesis were predicted through continuing increases in CO₂ concentration. The strong interaction with temperature also leads to photosynthesis in different global regions experiencing very different sensitivities to increasing CO₂ concentrations.     

The effect of growth at elevated CO2 concentrations on photosynthesis in wheat

Rising levels of atmospheric CO₂ will have profound, direct effects on plant carbon metabolism. In this study we used gas exchange measurements, models describing the instantaneous response of leaf net CO₂ assimilation rate (A) to intercellular CO₂ partial pressure (C_i), in vitro enzyme activity assay, and carbohydrate assay in order to investigate the photosynthetic responses of wheat (Triticum aestivum L., cv. Wembley) to growth under elevated partial pressures of atmospheric CO₂ (C_a). At flag leaf ligule emergence, the modelled, in vivo, maximum carboxylation velocity for RuBisCO was significantly lower in plants grown at elevated C_a than in plants grown at ambient C_a (70 Pa compared with 40 Pa). By 12 d after ligule emergence, no significant difference in this parameter was detectable. At ligule emergence, plants grown at elevated C_a exhibited reduced in vitro initial activities and activation states of RuBisCO. At their respective growth C_i values, the photosynthesis of 40-Pa-grown plants was sensitive to p(O₂) and to p(CO₂) whereas that of 70-Pa-grown plants was insensitive. Both sucrose and starch accumulated more rapidly in the leaves of plants grown at 70 Pa. At flag leaf ligule emergence, modelled non-photorespiratory respiration in the light (R_d) was significantly higher in 70-Pa-grown plants than in 40-Pa-grown plants. By 12 d after ligule emergence no significant differences in R_d were detectable.