Elevated CO2 stimulates cells to divide in grass meristems: a differential effect in two natural populations of Dactylis glomerata

In this study, we tested the hypothesis that elevated [CO₂] shortens the cell cycle in meristems of Dactylis glomerata, more in a Portuguese population (38°53′N) than in a Swedish population (63°09′N). In the shoot meristem, the cell cycle shortened to about the same extent (～ 26%) in both populations exposed to the elevated [CO₂] treatment. In the root meristem, the cell cycle shortened by 17% in the Portuguese and by 8% in the Swedish population. However, the proportion of rapidly cycling cells increased in the Portuguese much more than in the Swedish population in both meristems. In the root meristem, there was a 1.86-fold increase in the Portuguese compared with a 1.31-fold increase in the Swedish. In the shoot meristem, the increases were 1.5–3-fold for the Portuguese and 1.2-fold for the Swedish. The data are consistent in showing that a major response to the elevated [CO₂] treatment was an increase in the proportion of cells that were cycling and that this was more marked for the Portuguese population. A more general response to the elevated [CO₂] treatment was a shortening of the cell cycle regardless of population.     

Effects of source-sink relations on photosynthetic acclimation to elevated CO2

Abstract. While photosynthesis of C₃ plants is stimulated by an increase in the atmospheric CO₂ concentration, photosynthetic capacity is often reduced after long-term exposure to elevated CO₂. This reduction appears to be brought about by end product inhibition, resulting from an imbalance in the supply and demand of carbohydrates. A review of the literature revealed that the reduction of photosynthetic capacity in elevated CO₂ was most pronounced when the increased supply of carbohydrates was combined with small sink size. The volume of pots in which plants were grown affected the sink size by restricting root growth. While plants grown in small pots had a reduced photosynthetic capacity, plants grown in the field showed no reduction or an increase in this capacity. Pot volume also determined the effect of elevated CO₂ on the root/shoot ratio: the root/shoot ratio increased when root growth was not restricted and decreased in plants grown in small pots. The data presented in this paper suggest that plants growing in the field will maintain a high photosynthetic capacity as the atmospheric CO₂ level continues to rise.     

A mechanistic evaluation of photosynthetic acclimation at elevated CO2

Plants grown at elevated pCO₂ often fail to sustain the initial stimulation of net CO₂ uptake rate (A). This reduced, acclimated, stimulation of A often occurs concomitantly with a reduction in the maximum carboxylation velocity (V_c,max) of Rubisco. To investigate this relationship we used the Farquhar model of C₃ photosynthesis to predict the minimum V_c,max capable of supporting the acclimated stimulation in A observed at elevated pCO₂. For a wide range of species grown at elevated pCO₂ under contrasting conditions we found a strong correlation between observed and predicted values of V_c,max. This exercise mechanistically and quantitatively demonstrated that the observed acclimated stimulation of A and the simultaneous decrease in V_c,max observed at elevated pCO₂ is mechanistically consistent. With the exception of plants grown at a high elevated pCO₂ (> 90 Pa), which show evidence of an excess investment in Rubisco, the failure to maintain the initial stimulation of A is almost entirely attributable to the decrease in V_c,max and investment in Rubisco is coupled to requirements.     

Inhibition of whole plant respiration by elevated CO2 as modified by growth temperature

Plants of alfalfa (Medicago sativa) and orchard grass (Dactylus glomerata) were grown in controlled environment chambers at two CO₂ concentrations (350 and 700 μmol mol^-1) and 4 constant day/night growth temperatures of 15, 20, 25 and 30°C for 50–90 days to determine changes in growth and whole plant CO₂ efflux (dark respiration). To facilitate comparisons with other studies, respiration data were expressed on the basis of leaf area, dry weight and protein. Growth at elevated CO₂ increased total plant biomass at all temperatures relative to ambient CO₂, but the relative enhancement declined (P≤0.05) as temperature increased. Whole plant respiration (R_d) at elevated CO₂ declined at 15 and 20°C in D. glomerata on an area, weight or protein basis and in M. sativa on a weight or protein basis when compared to ambient CO₂. Separation of R_d into respiration required for growth (R_g) and maintenance (R_m) showed a significant effect of elevated CO₂ on both components. R_m was reduced in both species but only at lower temperatures (15°C in M. sativa and 15 and 20°C in D. glomerata). The effect on R_m could not be accounted for by protein content in either species. R_g was also reduced with elevated CO₂; however no particular effect of temperature was observed, i. e. R_g was reduced at 20, 25 and 30°C in M. sativa and at 15 and 25°C in D. glomerata. For the two perennial species used in the present study, the data suggest that both R_g and R_m can be reduced by anticipated increases in atmospheric CO₂; however, CO₂ inhibition of total plant respiration may decline as a function of increasing temperature     

Direct and indirect inhibition of single leaf respiration by elevated CO2 concentrations: Interaction with temperature

Two herbaceous perennials, alfalfa (Medicago sativa L. cv. Arc) and orchard grass (Dactylus glomerata L. cv. Potomac), were grown at ambient (367 μmol mol⁻¹) and elevated (729 μmol mol⁻¹) CO₂ concentrations at constant temperatures of 15, 20, 25 and 30°C in order to examine direct and indirect changes in nighttime CO₂ efflux rate (respiration) of single leaves. Direct (biochemical) effects of CO₂ on nighttime respiration were determined for each growth condition by brief (<30 min) exposure to each CO₂ concentration. If no direct inhibition of respiration was observed, then long-term reductions in CO₂ efflux between CO₂ treatments were presumed to be due to indirect inhibition, probably related to long-term changes in leaf composition. By this criterion, indirect effects of CO₂ on leaf respiration were observed at 15 and 20°C for M. sativa on a weight basis, but not on a leaf area or protein basis. Direct effects however, were observed at 15, 20 and 25°C in D. glomerata; therefore the observed reductions in respiration for leaves grown and measured at elevated relative to ambient CO₂ concentrations could not be distinguished as indirect inhibition. No inhibition of respiration at elevated CO₂ was observed at the highest growth temperature (30°C) in either species. CO₂ efflux increased with measurement and growth temperature for M. sativa at both CO₂ concentrations; however, CO₂ efflux in D. glomerata showed complete acclimation to growth temperature. Stimulation of leaf area and weight by elevated CO₂ levels declined with growth temperature in both species. Data from the present study suggest that both direct and indirect inhibition of respiration are possible with future increases in atmospheric CO₂, and that the degree of each type of respiratory inhibition is a function of growth temperature.     

Seasonal response of photosynthesis to elevated CO2 in loblolly pine (Pinus taeda L.) over two growing seasons

Trees growing in natural systems undergo seasonal changes in environmental factors that generate seasonal differences in net photosynthetic rates. To examine how seasonal changes in the environment affect the response of net photosynthetic rates to elevated CO₂, we grew Pinus taeda L. seedlings for three growing seasons in open-top chambers continuously maintained at either ambient or ambient + 30 Pa CO₂. Seedlings were grown in the ground, under natural conditions of light, temperature nd nutrient and water availability. Photosynthetic capacity was measured bimonthly using net photosynthetic rate vs. intercellular CO₂ partial pressure (A-C_i) curves. Maximum Rubisco activity (Vc_max) and ribulose 1,5-bisphosphate regeneration capacity mediated by electron transport (J_max) and phosphate regeneration (PiRC) were calculated from A-C_i curves using a biochemically based model. Rubisco activity, activation state and content, and leaf carbohydrate, chlorophyll and nitrogen concentrations were measured concurrently with photosynthesis measurements. This paper presents results from the second and third years of treatment. Mean leaf nitrogen concentrations ranged from 13.7 to 23.8 mg g^?1, indicating that seedlings were not nitrogen deficient. Relative to ambient CO₂ seedlings, elevated CO₂ increased light-saturated net photosynthetic rates 60–110% during the summer, but < 30% during the winter. A relatively strong correlation between leaf temperature and the relative response of net photosynthetic rates to elevated CO₂ suggests a strong effect of leaf temperature. During the third growing season, elevated CO₂ reduced Rubisco activity 30% relative to ambient CO₂ seedlings, nearly completely balancing Rubisco and RuBP-regeneration regulation of photosynthesis. However, reductions in Rubisco activity did not eliminate the seasonal pattern in the relative response of net photosynthetic rates to elevated CO₂. These results indicate that seasonal differences in the relative response of net photosynthetic rates to elevated CO₂ are likely to occur in natural systems.     

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.     

Effect of CO2 enrichment and elevated temperature on methane emissions from rice, Oryza sativa

Methane emissions from rice grown within Temperature Gradient Greenhouse Tunnels under doubled CO₂ concentrations were 10–45 times less than emissions from control plants grown under ambient CO₂. For two cultivars of rice (cvs. Lemont and IR-72), methane emissions increased with a temperature increase of 2°, from outdoor ambient temperatures to the first cell of the ambient CO₂ tunnel (ambient temperature + 2 °C). Within both tunnels and for both cultivars methane emissions decreased with further temperature increases (from 2° to 5 °C above ambient). Carbon dioxide enrichment stimulated both above- and below-ground production. Our original hypothesis was that increased CO₂ would stimulate plant productivity and therefore stimulate methane emission, since direct linkages between these parameters have been observed. We hypothesize that CO₂ enrichment led to the attenuation of methane production due to increased delivery of oxygen to the rhizosphere because of increased root biomass and porosity. The increased root biomass due to elevated CO₂ may have more effectively aerated the soil, suppressing methane production. However, this study may be unique because the low organic content (< 1%) of the sandy soils in which the rice was grown created very little oxygen demand.     

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.     

Mineral nitrogen cycling through earthworm casts in a grazed pasture under elevated atmospheric CO2

We used the New Zealand grazed pasture free air CO₂ enrichment experiment to determine the effects of elevated CO₂ on earthworm (Aporrectodea caliginosa and Lumbricus rubellus) cast production and mineral nitrogen (N) concentration over a 5‐week period in the spring. Elevated CO₂ did not affect earthworm biomass or the amount of cast material produced. However, cast mineral N concentrations were 18% lower, resulting in 27% less mineral N being deposited on the soil surface under elevated CO₂. An analysis of the earthworms' potential diet showed that a reduction in the N content of sheep dung at elevated CO₂ was the most likely cause of the lower cast N concentrations. Earthworm casts made only a small contribution to mineral N cycling in our system; however, their quality may act as a sensitive indicator of reduced N availability under elevated CO₂ which is consistent with the hypothesised process of progressive N limitation.     

Scaling up evolutionary responses to elevated CO2: lessons from Arabidopsis

Results from norm of reaction studies and selection experiments indicate that elevated CO₂ will act as a selective agent on natural plant populations, especially for C₃ species that are most sensitive to changes in atmospheric CO₂ concentration. Evolutionary responses to CO₂ may alter plant physiology, development rate, growth, and reproduction in ways that cannot be predicted from single generation studies. Moreover, ecological and evolutionary changes in plant communities will have a range of consequences at higher spatial scales and may cause substantial deviations from ecosystem level predictions based on short‐term responses to elevated CO₂. Therefore, steps need to be taken to identify the plant traits that are most likely to evolve at elevated CO₂, and to understand how these changes may affect net primary productivity within ecosystems. These processes may range in scale from molecular and physiological changes that occur among genotypes at the individual and population levels, to changes in community‐ and ecosystem‐level productivity that result from the integrative effects of different plant species evolving simultaneously. In this review, we (1) synthesize recent studies investigating the role of atmospheric CO₂ as a selective agent on plants, (2) discuss possible control points during plant development that may change in response to selection at elevated CO₂ with an emphasis at the primary molecular level, and (3) provide a quantitative framework for scaling the evolutionary effects of CO₂ on plants in order to determine changes in community and ecosystem productivity. Furthermore, this review points out that studies integrating the effects of plant evolution in response to elevated CO₂ are lacking, and therefore more attention needs be devoted to this issue among the global change research community.     

Combined effects of elevated CO2 and air temperature on carbon assimilation of Pinus taeda trees

Branches of 22-year-old loblolly pine (Pinus taeda, L.) trees growing in a plantation were exposed to ambient CO₂, ambient + 165 μmol mol^?1 CO₂ or ambient + 330 μmol mol^?1 CO₂ concentrations in combination with ambient or ambient + 2°C air temperatures for 3 years. Field measurements in the third year indicated that net carbon assimilation was enhanced in the elevated CO₂ treatments in all seasons. On the basis of A/C_i, curves, there was no indication of photosynthetic down-regulation. Branch growth and leaf area also increased significantly in the elevated CO₂ treatments. The imposed 2°C increase in air temperature only had slight effects on net assimilation and growth. Compared with the ambient CO₂ treatment, rates of net assimilation were ～1·6 times greater in the ambient + 165 μmol mol^?1 CO₂ treatment and 2·2 times greater in the ambient + 330 μmol mol^?1 CO₂ treatment. These ratios did not change appreciably in measurements made in all four seasons even though mean ambient air temperatures during the measurement periods ranged from 12·6 to 28·2°C. This indicated that the effect of elevated CO₂ concentrations on net assimilation under field conditions was primarily additive. The results also indicated that the effect of elevated CO₂ (+ 165 or + 330 μmol mol^?1) was much greater than the effect of a 2°C increase in air temperature on net assimilation and growth in this species.     

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.     

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

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

Effects of elevated CO2 on development and larval food-plant preference in the butterfly Coenonympha pamphilus (Lepidoptera, Satyridae)

The objective of this study was to determine how increasing atmospheric CO₂ change plant tissue quality in four native grassland grass species (Agrostis stolonifera, Anthoxanthum odoratum, Festuca rubra, Poa pratensis) which are all larval food‐plants of Coenonympha pamphilus (Lepidoptera, Satyridae). We assessed the effect of these changes on the performance and larval food‐plant preference of C. pamphilus in a greenhouse experiment. Furthermore, we tested the interactive effects of elevated CO₂ and soil nutritional availability in F. rubra and its effect an larval development of C. pamphilus. In general, elevated CO₂ decreased leaf water concentration, nitrogen concentration and specific leaf area (SLA), while leaf starch concentration was increased in all grass species. A species‐specific reaction to elevated CO₂ was only found for foliar starch concentration. P. pratensis did not increase its starch concentration under elevated CO₂ conditions, whereas the other three species did. Fertilisation, investigated only for F. rubra, increased leaf nitrogen concentration and amplified the CO₂‐induced decrease in leaf nitrogen. Development time of C. pamphilus was on the average prolonged by two days under elevated CO₂ and the prolongation differed from 0.7 to 5.3 days among food‐plant species. Pupal fresh weight differed marginally between CO₂ treatments. Fertilisation of the larval food‐plant F. rubra shortened development time by one day and significantly increased pupal and adult fresh weights. C. pamphilus larvae showed a clear food‐plant preference among grass species at the age of 36 h or older. Additionally, a change of food‐plant preference under elevated CO₂ was found. Larvae at ambient CO₂ preferred Agrostis stolonifera and F. rubra, while under elevated CO₂Anthoxanthum odoratum and P. pratensis were preferred. The present study demonstrates that larval development of C. pamphilus is affected by food‐plant species and CO₂ induced changes in foliar chemistry. Although we found some species‐specific reactions to elevated CO₂ for foliar chemistry, no such CO₂ by species interaction was found for insect development. The change in food‐plant preference of larvae under elevated CO₂ implies potential changes in selection pressure for grass species and might therefore affect evolutionary processes.     

Responses of multiple generations of Gastrophysa viridula, feeding on Rumex obtusifolius, to elevated CO2

Rumex obtusifolius plants and three generations of the tri-voltine leaf beetle Gastrophysa viridula were simultaneously exposed to elevated CO₂ (600 ppm) to determine its effect on plant quality and insect performance. This exposure resulted in a reduction in leaf nitrogen, an increase in the C/N ratio and lower concentrations of oxalate in the leaves than in ambient air (350 ppm). Despite these changes in food quality, the effect of elevated CO₂ on larvae of Gastrophysa viridula over three generations was minimal. However, the effect of CO₂ did differ slightly between the generations of the insect. For the first generation, the results obtained were different from many of the published results in that elevated CO₂ had no measurable effects on performance, except that third instar larvae showed compensatory feeding. Food quality, including leaf nitrogen content, declined over time in material grown in both ambient and elevated CO₂. The results obtained for the second generation were similar to the first except that first instar larvae showed reduced relative growth rate in elevated CO₂. Development time from hatching to pupation decreased over each generation, probably as a result of increasing temperatures. Measurements of adult performance showed that fecundity at the end of the second generation was reduced relative to the first, in line with the reduction in food quality. In addition at the end of the second generation, but not at the end of the first generation, adult females in elevated CO₂ laid 30% fewer eggs per day and the eggs laid were 15% lighter than those in ambient conditions. These lighter eggs, coupled with no effect of elevated CO₂ on growth during the third generation, meant that the larvae were consistently smaller in elevated CO₂ during this generation. These results offer further insights into the effect that elevated CO₂ will have on insect herbivores and provide a more detailed basis for population predictions.