The influences of increased CO2 and water supply on growth,biomass allocation and water use efficiency of Sinapis alba L. grown under different wind speeds

We examined how independent and interactive effects of CO₂ concentrations, water supply and wind speed affect growth rates, biomass partitioning, water use efficiency, diffusive conductance and stomatal density of plants. To test the prediction that wind stress will be ameliorated by increased CO₂ and/or by unrestricted water supply we grew Sinapis alba L. plants in controlled chambers under combinations of two levels of CO₂ (350 ppmv, 700 ppmv), two water regimes and two wind speeds (0.3 ms^–1, 3.7 ms^–1). We harvested at ten different dates over a period of 60 days. A growth analysis was carried out to evaluate treatment effects on plant responses. Plants grown both in increased CO₂ and in low wind conditions had significantly greater stem length, leaf area and dry weights of plant parts. Water supply significantly affected stem diameter, root weight and leaf area. CO₂ enrichment significantly increased the rate of biomass accumulation and the relative ratio of biomass increase to leaf area expansion. High wind speed significantly reduced plant growth rates and the rate of leaf area expansion was reduced more than the rate of biomass accumulation. Regression analysis showed significant CO₂ effects on the proportion of leaf and stem dry weight to total dry weight. A marked plant-age effect was dependent on water supply, wind speed and CO₂ concentration. A reduced water supply significantly decreased the stomatal conductance, and water use efficiency significantly increased with a limited water supply, low wind and increased CO₂. We found significant CO₂ x wind effects for water diffusion resistance, adaxial number of stomata and water use efficiencies and significant wind x water effect for water use efficiency. In conclusion, wind stress was ameliorated by growing in unrestricted water but not by growing in increased CO₂.     

Influence of elevated CO2 on canopy development and red:far-red ratios in two-storied stands ofRicinus communis

Vertical structure of plant stands and canopies may change under conditions of elevated CO₂ due to differential responses of overstory and understory plants or plant parts. In the long term, seedling recruitment, competition, and thus population or community structure may be affected. Aside from the possible differential direct effects of elevated CO₂ on photosynthesis and growth, both the quantity and quality of the light below the overstory canopy could be indirectly affected by CO₂-induced changes in overstory leaf area index (LAI) and/or changes in overstory leaf quality. In order to explore such possible interactions, we compared canopy leaf area development, canopy light extinction and the quality of light beneath overstory leaves of two-storied monospecific stands ofRicinus communis exposed to ambient (340 μl l⁻¹) and elevated (610 μl l⁻¹) CO₂. Plants in each stand were grown in a common soil as closed “artificial ecosystems” with a ground area of 6.7 m². LAI of overstory plants in all ecosystems more than doubled during the experiment but was not different between CO₂ treatments at the end. As a consequence, extinction of photosynthetically active radiation (PAR) was also not altered. However, under elevated CO₂ the red to far-red ratio (R:FR) measured beneath overstory leaves was 10% lower than in ecosystems treated with ambient CO₂. This reduction was associated with increased thickness of palisade layers of overstory leaves and appears to be a plausible explanation for the specific enhancement of stem elongation of understory plants (without a corresponding biomass response) under elevated CO₂. CO₂ enrichment led to increased biomass of overstory plants (mainly stem biomass) but had no effect on understory biomass. The results of this study raise the possibility of an important indirect effect of elevated CO₂ at the stand-level. We suggest that, under elevated CO₂, reductions in the R:FR ratio beneath overstory canopies may affect understory plant development independently of the effects of PAR extinction.     

Above- and below-ground responses of C3–C4 species mixtures to elevated CO2 and soil water availability

We evaluated the influences of CO₂[Control, ～ 370 µ mol mol ^{? 1}; 200 µ mol mol ^{? 1} above ambient applied by free‐air CO₂ enrichment (FACE)] and soil water (Wet, Dry) on above‐ and below‐ground responses of C₃ (cotton, Gossypium hirsutum) and C₄ (sorghum, Sorghum bicolor) plants in monocultures and two density mixtures. In monocultures, CO₂ enrichment increased height, leaf area, above‐ground biomass and reproductive output of cotton, but not sorghum, and was independent of soil water treatment. In mixtures, cotton, but not sorghum, above‐ground biomass and height were generally reduced compared to monocultures, across both CO₂ and soil water treatments. Density did not affect individual plant responses of either cotton or sorghum across the other treatments. Total (cotton + sorghum) leaf area and above‐ground biomass in low‐density mixtures were similar between CO₂ treatments, but increased by 17–21% with FACE in high‐density mixtures, due to a 121% enhancement of cotton leaf area and a 276% increase in biomass under the FACE treatment. Total root biomass in the upper 1.2 m of the soil was not influenced by CO₂ or by soil water in monoculture or mixtures; however, under dry conditions we observed significantly more roots at lower soil depths ( > 45 cm). Sorghum roots comprised 81–85% of the total roots in the low‐density mixture and 58–73% in the high‐density mixture. CO₂‐enrichment partly offset negative effects of interspecific competition on cotton in both low‐ and high‐density mixtures by increasing above‐ground biomass, with a greater relative increase in the high‐density mixture. As a consequence, CO₂‐enrichment increased total above‐ground yield of the mixture at high density. Individual plant responses to CO₂ enrichment in global change models that evaluate mixed plant communities should be adjusted to incorporate feedbacks for interspecific competition. Future field studies in natural ecosystems should address the role that a CO₂‐mediated increase in C₃ growth may have on subsequent vegetation change.     

The relationships between relative growth rate,meristematic potential and compensatory growth of semiarid-land shrubs

The submersed macrophyte Vallisneria americana was grown for seven weeks in a greenhouse to test for differences in the ability of three different sediments to support growth stimulation in response to CO₂ enrichment at low pH. Plants accumulated 21- to 24-fold greater biomass at 10 × ambient CO₂ concentrations than at ambient CO₂ on all sediments. At both CO₂ levels, plants grown on sediment from an acidified lake accumulated ca. 81%, and those grown on oligotrophic lake sediment ca. 47% as much biomass as plants grown on alkaline lake sediment. Despite striking CO₂ and sediment effects on biomass accumulation, there was no significant interaction (using log-transformed data) between CO₂ and sediment effects, indicating that all sediments allowed similar proportionate growth responses to CO₂ enrichment. Plants grown on the less fertile sediments showed greater relative allocation to horizontal versus vertical growth by producing more rosette-bearing stolons in relation to plant height (leaf length) than plants grown on relatively fertile, alkaline lake sediment. Tissue analysis suggested that sediment effects on Vallisneria growth could be attributed neither to mineral putrient (nitrogen and phosphorus) limitation nor to aluminum toxicity in these low pH treatments. In any case, CO₂ availability can be an important regulator of submersed macrophyte growth at low pH on a variety of sediment types, including those from oligotrophic and acidic lakes.     

Short photoperiod attenuates CO2 fertilization effect on shoot biomass in Arabidopsis thaliana

The level of carbon dioxide (CO₂) in the air can affect several traits in plants. Elevated atmospheric CO₂ (eCO₂) can enhance photosynthesis and increase plant productivity, including biomass, although there are inconsistencies regarding the effects of eCO₂ on the plant growth response. The compounding effects of ambient environmental conditions such as light intensity, photoperiod, water availability, and soil nutrient composition can affect the extent to which eCO₂ enhances plant productivity. This study aimed to investigate the growth response of Arabidopsis thaliana to eCO₂ (800 ppm) under short photoperiod (8/16 h, light/dark cycle). Here, we report an attenuated fertilization effect of eCO₂ on the shoot biomass of Arabidopsis plants grown under short photoperiod. The biomass of two-, three-, and four-week-old Arabidopsis plants was increased by 10%, 15%, and 28%, respectively, under eCO₂ compared to the ambient CO₂ (aCO₂, 400 ppm) i.e. control. However, the number of rosette leaves, rosette area, and shoot biomass were similar in mature plants under both CO₂ conditions, despite 40% higher photosynthesis in eCO₂ exposed plants. The levels of chlorophylls and carotenoids were similar in the fully expanded rosette leaves regardless of the level of CO₂. In conclusion, CO₂ enrichment moderately increased Arabidopsis shoot biomass at the juvenile stage, whereas the eCO₂-induced increment in shoot biomass was not apparent in mature plants. A shorter day-length can limit the source-to-sink resource allocation in a plant in age-dependent manner, hence diminishing the eCO₂ fertilization effect on the shoot biomass in Arabidopsis plants grown under short photoperiod.     

Carbon dioxide increase: the implications at the ecosystem level*

Abstract Possible effects of increased atmospheric concentrations of CO₂ on forest ecosystems are discussed and as an example a simulation case study using a set of mixed-age and mixed-species forest stand models is presented. The responses of the models to a simple scenario (uniform growth increase of all trees as a response to CO₂ enrichment) include increases in biomass that are considerably less than the increases in growth rate of the trees. These simulations and more general discussion of the possible effects of increased photosynthetic production identify the problem of scaling-up small time-scale and space-scale measurements of plant responses to CO₂ enrichment to the ecosystem level.     

Legume presence increases photosynthesis and N concentrations of co-occurring non-fixers but does not modulate their responsiveness to carbon dioxide enrichment

Legumes, with the ability to fix atmospheric nitrogen (N), may help alleviate the N limitations thought to constrain plant community response to elevated concentrations of atmospheric carbon dioxide (CO₂). To address this issue we assessed: (1) the effects of the presence of the perennial grassland N₂ fixer, Lupinus perennis, on biomass accumulation and plant N concentrations of nine-species plots of differing plant composition; (2) leaf-level physiology of co-occurring non-fixing species (Achillea millefolium, Agropyron repens, Koeleria cristata) in these assemblages with and without Lupinus; (3) the effects of elevated CO₂ on Lupinus growth and symbiotic N₂ fixation in both monoculture and the nine-species assemblages; and (4) whether assemblages containing Lupinus exhibit larger physiological and growth responses to elevated CO₂ than those without. This study was part of a long-term grassland field experiment (BioCON) that controls atmospheric CO₂ at current ambient and elevated (560 µmol mol^–1) concentrations using free-air CO₂ enrichment. Nine-species plots with Lupinus had 32% higher whole plot plant N concentrations and 26% higher total plant N pools than those without Lupinus, based on both above and belowground measurements. Co-occurring non-fixer leaf N concentrations increased 22% and mass-based net photosynthetic rates increased 41% in plots containing Lupinus compared to those without. With CO₂ enrichment, Lupinus monocultures accumulated 32% more biomass and increased the proportion of N derived from fixation from 44% to 57%. In nine-species assemblages, Lupinus N derived from fixation increased similarly from 43% to 54%. Although Lupinus presence enhanced photosynthetic rates and leaf N concentrations of co-occurring non-fixers, and increased overall plant N pools, Lupinus presence did not facilitate stronger photosynthetic responses of non-fixing species or larger growth responses of overall plant communities to elevated CO₂. Non-fixer leaf N concentrations declined similarly in response to elevated CO₂ with and without Lupinus present and the relationship between net photosynthesis and leaf N was not affected by Lupinus presence. Regardless of the presence or absence of Lupinus, CO₂ enrichment resulted in reduced leaf N concentrations and rates of net photosynthesis.     

[CO2]- and density-dependent competition between grassland species

The predicted ongoing increase of atmospheric carbon dioxide levels is considered to be one of the main threats to biodiversity due to potential changes in biotic interactions. We tested whether effects of intra‐ and interspecific planting density of the calcareous grassland perennials Bromus erectus and Carex flacca change in response to elevated [CO₂] (600 ppm) by using factorial combinations of seven densities (0, 1, 2, 4, 8, 16, 24 tillers per 8 × 8 cm² cell) of both species in plots with and without CO₂ enrichment. Although aboveground biomass of C. flacca was increased by 54% under elevated [CO₂], the combined aboveground biomass of the whole stand was not significantly increased. C. flacca tended to produce more tillers under elevated [CO₂] while B. erectus produced less tillers. The positive effect of [CO₂] on the number of tillers of C. flacca was strongest at high intraspecific densities. On the other hand, the negative effect of [CO₂] on the number of tillers of B. erectus was not present at intermediate intraspecific planting densities. Seed production of C. flacca was more than doubled under elevated [CO₂], while seed production of B. erectus was not affected. Moreover, the mass per seed of C. flacca was increased by elevated [CO₂] at intermediate interspecific planting densities while the mass per seed of B. erectus was decreased by elevated [CO₂] at high interspecific planting densities. Our results show that the responses of C. flacca and B. erectus to elevated [CO₂] depend in a complex way on initial planting densities of both species. In other words, competition between these two model species is both [CO₂]‐ and density dependent. On average, however, the effects of [CO₂] on the individual species indicate that the composition of calcareous grasslands is likely to change under elevated [CO₂] in favor of C. flacca.     

Effects of elevated CO2 and vascular plants on evapotranspiration in bog vegetation

We determined evapotranspiration in three experiments designed to study the effects of elevated CO₂ and increased N deposition on ombrotrophic bog vegetation. Two experiments used peat monoliths with intact bog vegetation in containers, with one experiment outdoors and the other in a greenhouse. A third experiment involved monocultures and mixtures of Sphagnum magellanicum and Eriophorum angustifolium in containers in the same greenhouse. To determine water use of the bog vegetation in July–August for each experiment and each year we measured water inputs and outputs from the containers. We studied the effects of elevated CO₂ and N supply on evapotranspiration in relation to vascular plant biomass and exposure of the moss surface (measured as height of the moss surface relative to the container edge). Elevated CO₂ reduced water use of the bog vegetation in all three experiments, but the CO₂ effect on evapotranspiration interacted with vascular plant biomass and exposure of the moss surface. Evapotranspiration in the outdoor experiment was largely determined by evaporation from the Sphagnum moss surface (as affected by exposure to wind) and less so by vascular plant transpiration. Nevertheless, elevated CO₂ significantly reduced evapotranspiration by 9–10% in the outdoor experiment. Vascular plants reduced evapotranspiration in the outdoor experiment, but increased water use in the greenhouse experiments. The relation between vascular plant abundance and evapotranspiration appears to depend on wind conditions; suggesting that vascular plants reduce water losses mainly by reducing wind speed at the moss surface. Sphagnum growth is very sensitive to changes in water level; low water availability can have deleterious effects. As a consequence, reduced evapotranspiration in summer, whether caused by elevated CO₂ or by small increases in vascular plant cover, is expected to favour Sphagnum growth in ombrotrophic bog vegetation.     

CO2 enrichment reduces reproductive dominance in competing stands of Ambrosia artemisiifolia (common ragweed)

Plants growing in dense stands may not equally acquire or utilize extra carbon gained in elevated CO₂. As a result, reproductive differences between dominant and subordinate plants may be altered under rising CO₂ conditions. We hypothesized that elevated CO₂ would enhance the reproductive allocation of shaded, subordinate Ambrosia artemisiifolia L. (Asteraceae) individuals more than that of light-saturated dominants. We grew stands of A. artemisiifolia at either 360 or 720 μL L⁻¹ CO₂ levels and measured the growth and reproductive responses of competing individuals. To test whether elevated CO₂ altered size and reproductive inequalities within stands, we compared stand-level coefficients of variation (CV) in height growth and final shoot, root, and reproductive organ biomasses. Elevated CO₂ enhanced biomass and reduced the CV for all aspects of plant growth, especially reproductive biomass. Allocation to reproduction was higher in the elevated CO₂ than in the ambient treatment, and this difference was more pronounced in small, rather than large plant positive relationships between the CV and total stand productivity declined under elevated CO₂, indicating that growth enhancements to smaller plants diminished the relative biomass advantages of larger plants in increasingly crowded conditions. We conclude that elevated CO₂ stimulates stand-level reproduction while CO₂-induced growth gains of subordinate A. artemisiifolia plants minimize differences in the reproductive output of small and large plants. Thus, more individuals are likely to produce greater amounts of seeds and pollen in future populations of this allergenic weed.     

Long-term effects of elevated atmospheric CO<Subscript>2</Subscript> on species composition and productivity of a southern African C<Subscript>4</Subscript> dominated grassland in the vicinity of a CO<Subscript>2</Subscript> exhalation

We describe the long-term effects of a CO₂ exhalation, created more than 70 years ago, on a natural C₄ dominated sub-tropical grassland in terms of ecosystem structure and functioning. We tested whether long-term CO₂ enrichment changes the competitive balance between plants with C₃ and C₄ photosynthetic pathways and how CO₂ enrichment has affected species composition, plant growth responses, leaf properties and soil nutrient, carbon and water dynamics. Long-term effects of elevated CO₂ on plant community composition and system processes in this sub-tropical grassland indicate very subtle changes in ecosystem functioning and no changes in species composition and dominance which could be ascribed to elevated CO₂ alone. Species compositional data and soil δ¹³C isotopic evidence suggest no detectable effect of CO₂ enrichment on C₃:C₄ plant mixtures and individual species dominance. Contrary to many general predictions C₃ grasses did not become more abundant and C₃ shrubs and trees did not invade the site. No season length stimulation of plant growth was found even after 5 years of exposure to CO₂ concentrations averaging 610 μmol mol⁻¹. Leaf properties such as total N decreased in the C₃ but not C₄ grass under elevated CO₂ while total non-structural carbohydrate accumulation was not affected. Elevated CO₂ possibly lead to increased end-of-season soil water contents and this result agrees with earlier studies despite the topographic water gradient being a confounding problem at our research site. Long-term CO₂ enrichment also had little effect on soil carbon storage with no detectable changes in soil organic matter found. There were indications that potential soil respiration and N mineralization rates could be higher in soils close to the CO₂ source. The conservative response of this grassland suggests that many of the reported effects of elevated CO₂ on similar ecosystems could be short duration experimental artefacts that disappear under long-term elevated CO₂ conditions.     

Transpiration of a 64-year old maritime pine stand in Portugal

Alpine plant species have been shown to exhibit a more pronounced increase in leaf photosynthesis under elevated CO₂ than lowland plants. In order to test whether this higher carbon fixation efficiency will translate into increased biomass production under CO₂ enrichment we exposed plots of narrow alpine grassland (Swiss Central Alps, 2470 m) to ambient (355 μl l^-1) and elevated (680 μl l^-1) CO₂ concentration using open top chambers. Part of the plost received moderate mineral nutrient additions (40 kg ha^-1 year^-1 of nitrogen in a complete fertilizer mix). Under natural nutrient supply CO₂ enrichment had no effect on biomass production per unit land area during any of the three seasons studied so far. Correspondingly, the dominant species Carex curvula and Leontodon helveticus as well as Trifolium alpinum did not show a growth response either at the population level or at the shoot level. However, the subdominant generalistic species Poa alpina strongly increased shoot growth (+47%). Annual root production (in ingrowth cores) was significantly enhanced in C. curvula in the 2nd and 3rd year of investigation (+43%) but was not altered in the bulk samples for all species. Fertilizer addition generally stimulated above-ground (+48%) and below-ground (+26%) biomass production right from the beginning. Annual variations in weather conditions during summer also strongly influenced above-ground biomass production (19–27% more biomass in warm seasons compared to cool seasons). However, neither nutrient availability nor climate had a significant effect on the CO₂ response of the plants. Our results do not support the hypothesis that alpine plants, due to their higher carbon uptake efficiency, will increase biomass production under future atmospheric CO₂ enrichment, at least not in such late successional communities. However, as indicated by the response of P. alpina, species-specific responses occur which may lead to altered community structure and perhaps ecosystem functioning in the long-term. Our findings further suggest that possible climatic changes are likely to have a greater impact on plant growth in alpine environments than the direct stimulation of photosynthesis by CO₂. Counter-intuitively, our results suggest that even under moderate climate warming or enhanced atmospheric nitrogen deposition positive biomass responses to CO₂ enrichment of the currently dominating species are unlikely.     

Effects of UV-B radiation on terrestrial plants and ecosystems: interaction with CO2 enrichment

UV-B radiation is just one of the environmental factors, that affect plant growth. It is now widely accepted that realistic assessment of plant responses to enhanced UV-B should be performed at sufficiently high Photosynthetically Active Radiation (PAR), preferably under field conditions. This will often imply, that responses of plants to enhanced UV-B in the field will be assessed under simultaneous water shortage, nutrient deficiency and variation of temperature. Since atmospheric CO₂ enrichment, global warming and increasing UV-B radiation represent components of global climatic change, interactions of UV-B with CO₂ enrichment and temperature are particularly relevant. Only few relevant UV-B× CO₂ interaction studies have been published. Most of these studies refer to greenhouse experiments. We report a significant CO₂ × UV-B interaction for the total plant dry weight and root dry weight of the C₃-grass Elymus athericus. At elevated CO₂ (720 mol mol^-1, plant growth was much less reduced by enhanced UV-B than at ambient atmospheric CO₂ although there were significant (positive) CO₂ effects and (negative) UV-B effects on plant growth. Most other CO₂ × UV-B studies do not report significant interactions on total plant biomass. This lack of CO₂ × UV-B interactions may result from the fact that primary metabolic targets for CO₂ and UVB are different. UV-B and CO₂ may differentially affect plant morphogenetic parameters: biomass allocation, branching, flowering, leaf thickness, emergence and senescence. Such more subtle interactions between CO₂ and UV-B need careful and long term experimentation to be detected. In the case of no significant CO₂× UV-B interactions, combined CO₂ and UV-B effects will be additive. Plants differ in their response to CO₂ and UV-B, they respond in general positively to elevated CO₂ and negatively to enhanced UV-B. Moreover, plant species differ in their responsiveness to CO₂ and UV-B. Therefore, even in case of additive CO₂ and UV-B effects, plant competitive relationships may change markedly under current climatic change with simultaneous enhanced atmospheric CO₂ and solar UV-B radiation.     

Enhanced root system C-sink activity,water relations and aspects of nutrient acquisition in mycotrophic Bouteloua gracilis subjected to CO2 enrichment

In order to better elucidate fixed-C partitioning, nutrient acquisition and water relations of prairie grasses under elevated [CO₂], we grew the C₄ grass Bouteloua gracilis (H.B.K.) lag ex Steud. from seed in soil-packed, column-lysimeters in two growth chambers maintained at current ambient [CO₂] (350 μL L⁻¹) and twice enriched [CO₂] (700 μL L⁻¹). Once established, plants were deficit irrigated; growth chamber conditions were maintained at day/night temperatures of 25/16°C, relative humidities of 35%/90% and a 14-hour photoperiod to simulate summer conditions on the shortgrass steppe in eastern Colorado. After 11 weeks of growth, plants grown under CO₂ enrichment had produced 35% and 65% greater total and root biomass, respectively, and had twice the level of vesicular-arbuscular mycorrhizal (VAM) infection (19.8% versus 10.8%) as plants grown under current ambient [CO₂]. The CO₂-enriched plants also exhibited greater leaf water potentials and higher plant water use efficiencies. Plant N uptake was reduced by CO₂ enrichment, while P uptake appeared little influenced by CO₂ regime. Under the conditions of the experiment, CO₂ enrichment increased root biomass and VAM infection via stimulated growth and adjustments in C partitioning below-ground. The U.S. Government right to retain a non-exclusive, royalty free licence in and to any copyright is acknowledged. The U.S. Government right to retain a non-exclusive, royalty free licence in and to any copyright is acknowledged.     

Responses of C4 grasses to atmospheric CO2 enrichment

Summary The growth and photosynethetic responses to atmospheric CO₂ enrichment of 4 species of C₄ grasses grown at two levels of irradiance were studied. We sought to determine whether CO₂ enrichment would yield proportionally greater growth enhancement in the C₄ grasses when they were grown at low irradiance than when grown at high irradiance. The species studied were Echinochloa crusgalli, Digitaria sanguinalis, Eleusine indica, and Setaria faberi. Plants were grown in controlled environment chambers at 350, 675 and 1,000 l 1^-1 CO₂ and 1,000 or 150 mol m^-2 s^-1 photosynthetic photon flux density (PPFD). An increase in CO₂ concentration and PPFD significantly affected net photosynthesis and total biomass production of all plants. Plants grown at low PPFD had significantly lower rates of photosynthesis, produced less biomass, and had reduced responses to increases in CO₂. Plants grown in CO₂-enriched atmosphere had lower photosynthetic capacity relative to the low CO₂ grown plants when exposed to lower CO₂ concentration at the time of measurement, but had greater rate of photosynthesis when exposed to increasing PPFD. The light level under which the plants were growing did not influence the CO₂ compensation point for photosynthesis.     

Effects of elevated [CO2] at the community level mediated by root symbionts

This review examines the effects of elevated [CO₂] on plant symbioses with mycorrhizal fungi and root nodule bacteria, with emphasis on community and ecosystem processes. The effects of elevated [CO₂] on the relationships between single plant species and root symbionts are considered first. There is some evidence that plant infection by and/or biomass of root symbionts are stimulated by elevated [CO₂], but growth enhancement of the host seemingly depends on its degree of dependence on symbiosis and on soil nutrient availability. Second, the effects of elevated [CO₂] on the relationships between plant multispecies assemblages and soil, and likely impacts on above-ground and belowground diversity, are analysed. Experimental and modelling work have suggested the existence of complex feedbacks in the responses of plants and the rhizosphere to CO₂ enrichment. By modifying C inputs from plants to soil, elevated [CO₂] may affect the biomass, the infectivity, and the species/isolate composition of root symbionts. This has the potential to alter community structure and ecosystem functioning. Finally, the incorporation of type and degree of symbiotic dependence into the definition of plant functional types, and into experimental work within the context of global change research, are discussed. More experimental work on the effects of elevated [CO₂] at the community/ecosystem level, explicitly considering the role of root symbioses, is urgently needed.     

Plant growth and reproduction along CO2 gradients: non-linear responses and implications for community change

The effects of rising atmospheric CO₂ concentrations on natural plant communities will depend upon the cumulative responses of plant growth and reproduction to gradual, incremental changes in climatic conditions. We analysed published studies of plant responses to elevated CO₂ to address whether reproductive and total biomass exhibit similar enhancement to elevated vs. ambient CO₂ concentrations, and to assess the patterns of plant response along gradients of CO₂ concentrations. In six annual plant species, mean enhancement at double ambient vs. ambient CO₂ was 1.13 for total biomass and 1.30 for reproductive biomass. The two measures were significantly correlated, but there was considerable scatter in the relationship, indicating that reproductive responses cannot be consistently predicted from enhancement of total biomass. Along experimental CO₂ gradients utilizing three concentrations, there was a great diversity of response patterns, including positive, negative, non-monotonic and non-significant (flat) responses. The distribution of response patterns differed for plants grown in stands compared to those grown individually. Positive responses were less frequent in competitive environments, and non-monotonic responses were more frequent. These results emphasize that interpolation of plant response based on enhancement ratios measured at elevated vs. ambient CO₂ concentrations is not sufficient to predict community responses to incremental changes in atmospheric conditions. The consequences of differential response patterns were assessed in a simulation of community dynamics for four species of annual plants. The model illustrates that the final community composition at a future point in time depends critically on both the magnitude and the rate of increase of atmospheric CO₂.     

The combined effects of elevated CO2 levels and UV-B radiation on growth characteristics ofElymus athericus (= E. pycnanathus)

Elymus athericus (Link) Kerguélen, a C₃ grass, was grown in a greenhouse experiment to determine the effect of enhanced atmospheric CO₂ and elevated UV-B radiation levels on plant growth. Plants were subjected to the following treatments; a) ambient CO₂-control UV-B, b) ambient CO₂-elevated UV-B, c) double CO₂-control UV-B, d) double CO₂-elevated UV-B. Elevated CO₂ concentrations stimulated plant growth, biomass production was 67% higher than at ambient CO₂. Elevated UV-B radiation had a negative effect on growth, biomass production was depressed by 31%. Enhanced CO₂ combined with elevated UV-B levels caused a biomass depression of 8% when compared with the control plants. UV-B induced growth depression can be modified by a growth stimulus caused by high CO₂ concentrations. Growth analysis has been performed and possible physiological mechanisms behind changing growth parameters are discussed.     

Interactive effects of atmospheric CO2 enrichment,salinity and flooding on growth of C3 (Elymus athericus) and C4 (Spartina anglica) salt marsh species

The growth response of Dutch salt marsh species (C₃ and C₄) to atmospheric CO₂ enrichment was investigated. Tillers of the C₃ speciesElymus athericus were grown in combinations of 380 and 720 11^-1 CO₂ and low (O) and high (300 mM NaCl) soil salinity. CO₂ enrichment increased dry matter production and leaf area development while both parameters were reduced at high salinity. The relative growth response to CO₂ enrichment was higher under saline conditions. Growth increase at elevated CO₂ was higher after 34 than 71 days. A lower response to CO₂ enrichment after 71 days was associated with a decreased specific leaf area (SLA). In two other experiments the effect of CO₂ (380 and 720 11^-1) on growth of the C₄ speciesSpartina anglica was studied. In the first experiment total plant dry weight was reduced by 20% at elevated CO₂. SLA also decreased at high CO₂. The effect of elevated CO₂ was also studied in combination with soil salinity (50 and 400 mM NaCl) and flooding. Again plant weight was reduced (10%) at elevated CO₂, except under the combined treatment high salinity/non-flooded. But these effects were not significant. High salinity reduced total plant weight while flooding had no effect. Causes of the salinity-dependent effect of CO₂ enrichment on growth and consequences of elevated CO₂ for competition between C₃ and C₄ species are discussed.     

Virus-induced differences in the response of oat plants to elevated carbon dioxide

Disease is an integral element of agricultural and natural systems, but the roles pathogens play in determining ecosystem response to elevated CO₂ have rarely been examined. To investigate whether disease can alter the response of plants to CO₂, we examined the effects of doubled CO₂ (～700 μmol mol^?1) on Avena sativa infected with barley yellow dwarf virus (BYDV), a common pathogen of cereals and grasses. Oats infected with BYDV showed a significantly greater biomass response to CO₂ enrichment than did healthy plants. Root mass of diseased plants increased by 37–60% with CO₂ enrichment, but was largely unaffected in healthy plants. CO₂ enrichment increased midday leaf-level photosynthesis and instantaneous water use efficiency by 34 and 93% in healthy plants and by 48 and 174% in infected plants. Foliar carbohydrates increased with both CO₂ enrichment and BYDV infection, but the two factors affected individual pools dissimilarly. CO₂ enrichment may alter the epidemiology of BYDV by increasing the persistence of infected plants.