Enhanced isoprene emission capacity and altered light responsiveness in aspen grown under elevated atmospheric CO2 concentration

Controversial evidence of CO₂‐responsiveness of isoprene emission has been reported in the literature with the response ranging from inhibition to enhancement, but the reasons for such differences are not understood. We studied isoprene emission characteristics of hybrid aspen (Populus tremula x P. tremuloides) grown under ambient (380 μmol mol^?1) and elevated (780 μmol mol^?1) [CO₂] to test the hypothesis that growth [CO₂] effects on isoprene emission are driven by modifications in substrate pool size, reflecting altered light use efficiency for isoprene synthesis. A novel in vivo method for estimation of the pool size of the immediate isoprene precursor, dimethylallyldiphosphate (DMADP) and the activity of isoprene synthase was used. Growth at elevated [CO₂] resulted in greater leaf thickness, more advanced development of mesophyll and moderately increased photosynthetic capacity due to morphological “upregulation”, but isoprene emission rate under growth light and temperature was not significantly different among ambient‐ and elevated‐[CO₂]‐grown plants independent of whether measured at 380 μmol mol^?1 or 780 μmol mol^?1 CO₂. However, DMADP pool size was significantly less in elevated‐[CO₂]‐grown plants, but this was compensated by increased isoprene synthase activity. Analysis of CO₂ and light response curves of isoprene emission demonstrated that the [CO₂] for maximum isoprene emission was shifted to lower [CO₂] in elevated‐[CO₂]‐grown plants. The light‐saturated isoprene emission rate (I_max,Q) was greater, but the quantum efficiency at given I_max,Q was less in elevated‐[CO₂]‐grown plants, especially at higher CO₂ measurement concentration, reflecting stronger DMADP limitation at lower light and higher [CO₂]. These results collectively demonstrate important shifts in light and CO₂‐responsiveness of isoprene emission in elevated‐[CO₂]‐acclimated plants that need consideration in modeling isoprene emissions in future climates.     

Profiles of isoprene emission and photosynthetic parameters in hybrid poplars exposed to free‐air CO2 enrichment†

Poplar (Populus × euroamericana) saplings were grown in the field to study the changes of photosynthesis and isoprene emission with leaf ontogeny in response to free air carbon dioxide enrichment (FACE) and soil nutrient availability. Plants growing in elevated [CO₂] produced more leaves than those in ambient [CO₂]. The rate of leaf expansion was measured by comparing leaves along the plant profile. Leaf expansion and nitrogen concentration per unit of leaf area was similar between nutrient treatment, and this led to similar source–sink functional balance. Consequently, soil nutrient availability did not cause downward acclimation of photosynthetic capacity in elevated [CO₂] and did not affect isoprene synthesis. Photosynthesis assessed in growth [CO₂] was higher in plants growing in elevated than in ambient [CO₂]. After normalizing for the different number of leaves over the profile, maximal photosynthesis was reached and started to decline earlier in elevated than in ambient [CO₂]. This may indicate a [CO₂]‐driven acceleration of leaf maturity and senescence. Isoprene emission was adversely affected by elevated [CO₂]. When measured on the different leaves of the profile, isoprene peak emission was higher and was reached earlier in ambient than in elevated [CO₂]. However, a larger number of leaves was emitting isoprene in plant growing in elevated [CO₂]. When integrating over the plant profile, emissions in the two [CO₂] levels were not different. Normalization as for photosynthesis showed that profiles of isoprene emission were remarkably similar in the two [CO₂] levels, with peak emissions at the centre of the profile. Only the rate of increase of the emission of young leaves may have been faster in elevated than in ambient [CO₂]. Our results indicate that elevated [CO₂] may overall have a limited effect on isoprene emission from young seedlings and that plants generally regulate the emission to reach the maximum at the centre of the leaf profile, irrespective of the total leaf number. In comparison with leaf expansion and photosynthesis, isoprene showed marked and repeatable differences among leaves of the profile and may therefore be a useful trait to accurately monitor changes of leaf ontogeny as a consequence of elevated [CO₂].     

Leaf isoprene emission rate as a function of atmospheric CO2 concentration

There is considerable interest in modeling isoprene emissions from terrestrial vegetation, because these emissions exert a principal control over the oxidative capacity of the troposphere. We used a unique field experiment that employs a continuous gradient in CO₂ concentration from 240 to 520 ppmv to demonstrate that isoprene emissions in Eucalyptus globulus were enhanced at the lowest CO₂ concentration, which was similar to the estimated CO₂ concentrations during the last Glacial Maximum, compared with 380 ppmv, the current CO₂ concentration. Leaves of Liquidambar styraciflua did not show an increase in isoprene emission at the lowest CO₂ concentration. However, isoprene emission rates from both species were lower for trees grown at 520 ppmv CO₂ compared with trees grown at 380 ppmv CO₂. When grown in environmentally controlled chambers, trees of Populus deltoides and Populus tremuloides exhibited a 30–40% reduction in isoprene emission rate when grown at 800 ppmv CO₂, compared with 400 ppmv CO₂. P. tremuloides exhibited a 33% reduction when grown at 1200 ppmv CO₂, compared with 600 ppmv CO₂. We used current models of leaf isoprene emission to demonstrate that significant errors occur if the CO₂ inhibition of isoprene is not taken into account. In order to alleviate these errors, we present a new model of isoprene emission that describes its response to changes in atmospheric CO₂ concentration. The model logic is based on assumed competition between cytosolic and chloroplastic processes for pyruvate, one of the principal substrates of isoprene biosynthesis.     

The effects of glacial atmospheric CO2 concentrations and climate on isoprene emissions by vascular plants

Isoprene (C₅H₈) emissions by terrestrial vegetation vary with temperature and light intensity, and play an important role in biosphere–chemistry–climate interactions. Such interactions were probably substantially modified by Pleistocene climate and CO₂ cycles. Central to understanding the nature of these modifications is assessment and analysis of how emissions changed under glacial environmental conditions. Currently, even the net direction of change is difficult to predict because a CO₂‐depleted atmosphere may have stimulated emissions compensating for the negative impacts of a cooler climate. Here, we address this issue and attempt to determine the direction of change from an experimental standpoint by investigating the interaction between isoprene emissions and plant growth of two known isoprene‐emitting herbaceous species (Mucuna pruriens and Arundo donax) grown at glacial (180 ppm) to present (366 ppm) CO₂ levels. We found a significant enhancement of isoprene emissions per unit leaf area in M. pruriens under subambient CO₂ concentrations relative to ambient controls but not for A. donax. In contrast, canopy emissions remained unaltered for both plant species because enhanced leaf emissions were offset by reductions in biomass and leaf area. Separate growth experiments with M. pruriens revealed that lowering day/night temperatures by 5°C decreased canopy isoprene emissions irrespective of the CO₂ level. Incorporation of these results into a simple canopy emissions model highlights their potential to attenuate reductions in the total isoprene flux from forests under glacial conditions predicted by standard models.     

Interactive effects of elevated CO2 and soil fertility on isoprene emissions from Quercus robur

The effects of global change on the emission rates of isoprene from plants are not clear. A factor that can influence the response of isoprene emission to elevated CO₂ concentrations is the availability of nutrients. Isoprene emission rate under standard conditions (leaf temperature: 30°C, photosynthetically active radiation (PAR): 1000 μmol photons m^?2 s^?1), photosynthesis, photosynthetic capacity, and leaf nitrogen (N) content were measured in Quercus robur grown in well‐ventilated greenhouses at ambient and elevated CO₂ (ambient plus 300 ppm) and two different soil fertilities. The results show that elevated CO₂ enhanced photosynthesis but leaf respiration rates were not affected by either the CO₂ or nutrient treatments. Isoprene emission rates and photosynthetic capacity were found to decrease with elevated CO₂, but an increase in nutrient availability had the converse effect. Leaf N content was significantly greater with increased nutrient availability, but unaffected by CO₂. Isoprene emission rates measured under these conditions were strongly correlated with photosynthetic capacity across the range of different treatments. This suggests that the effects of CO₂ and nutrient levels on allocation of carbon to isoprene production and emission under near‐saturating light largely depend on the effects on photosynthetic electron transport capacity.     

Isoprenoid emission in trees of Quercus pubescens and Quercus ilex with lifetime exposure to naturally high CO2 environment†

The long‐term effect of elevated atmospheric CO₂ on isoprenoid emissions from adult trees of two Mediterranean oak species (the monoterpene‐emitting Quercus ilex L. and the isoprene‐emitting Quercus pubescens Willd.) native to a high‐CO₂ environment was investigated. During two consecutive years, isoprenoid emission was monitored both at branch level, measuring the actual emissions under natural conditions, and at leaf level, measuring the basal emissions under the standard conditions of 30 °C and at light intensity of 1000 µmol m^?2 s^?1. Long‐term exposure to high atmospheric levels of CO₂ did not significantly affect the actual isoprenoid emissions. However, when leaves of plants grown in the control site were exposed for a short period to an elevated CO₂ level by rapidly switching the CO₂ concentration in the gas‐exchange cuvette, both isoprene and monoterpene basal emissions were clearly inhibited. These results generally confirm the inhibitory effect of elevated CO₂ on isoprenoid emission. The absence of a CO₂ effect on actual emissions might indicate higher leaf temperature at elevated CO₂, or an interaction with multiple stresses some of which (e.g. recurrent droughts) may compensate for the CO₂ effect in Mediterranean ecosystems. Under elevated CO₂, isoprene emission by Q. pubescens was also uncoupled from the previous day's air temperature. In addition, pronounced daily and seasonal variations of basal emission were observed under elevated CO₂ underlining that correction factors may be necessary to improve the realistic estimation of isoprene emissions with empirical algorithms in the future. A positive linear correlation of isoprenoid emission with the photosynthetic electron transport and in particular with its calculated fraction used for isoprenoid synthesis was found. The slope of this relationship was different for isoprene and monoterpenes, but did not change when plants were grown in either ambient or elevated CO₂. This suggests that physiological algorithms may usefully predict isoprenoid emission also under rising CO₂ levels.     

Competition between isoprene emission and pigment synthesis during leaf development in aspen

In growing leaves, lack of isoprene synthase (IspS) is considered responsible for delayed isoprene emission, but competition for dimethylallyl diphosphate (DMADP), the substrate for both isoprene synthesis and prenyltransferase reactions in photosynthetic pigment and phytohormone synthesis, can also play a role. We used a kinetic approach based on post‐illumination isoprene decay and modelling DMADP consumption to estimate in vivo kinetic characteristics of IspS and prenyltransferase reactions, and to determine the share of DMADP use by different processes through leaf development in Populus tremula. Pigment synthesis rate was also estimated from pigment accumulation data and distribution of DMADP use from isoprene emission changes due to alendronate, a selective inhibitor of prenyltransferases. Development of photosynthetic activity and pigment synthesis occurred with the greatest rate in 1‐ to 5‐day‐old leaves when isoprene emission was absent. Isoprene emission commenced on days 5 and 6 and increased simultaneously with slowing down of pigment synthesis. In vivo Michaelis–Menten constant (K_m) values obtained were 265 nmol m^?2 (20 μm ) for DMADP‐consuming prenyltransferase reactions and 2560 nmol m^?2 (190 μm ) for IspS. Thus, despite decelerating pigment synthesis reactions in maturing leaves, isoprene emission in young leaves was limited by both IspS activity and competition for DMADP by prenyltransferase reactions.     

Response of isoprene emission to ambient CO2 changes and implications for global budgets

We explore the potential role of atmospheric carbon dioxide (CO₂) on isoprene emissions using a global coupled land–atmosphere model [Community Atmospheric Model–Community Land Model (CAM–CLM)] for recent (year 2000, 365 ppm CO₂) and future (year 2100, 717 ppm CO₂) conditions. We incorporate an empirical model of observed isoprene emissions response to both ambient CO₂ concentrations in the long‐term growth environment and short‐term changes in intercellular CO₂ concentrations into the MEGAN biogenic emission model embedded within the CLM. Accounting for CO₂ inhibition has little impact on predictions of present‐day global isoprene emission (increase from 508 to 523 Tg C yr^?1). However, the large increases in future isoprene emissions typically predicted in models, which are due to a projected warmer climate, are entirely offset by including the CO₂ effects. Projected global isoprene emissions in 2100 drop from 696 to 479 Tg C yr^?1 when this effect is included, maintaining future isoprene sources at levels similar to present day. The isoprene emission response to CO₂ is dominated by the long‐term growth environment effect, with modulations of 10% or less due to the variability in intercellular CO₂ concentration. As a result, perturbations to isoprene emissions associated with changes in ambient CO₂ are largely aseasonal, with little diurnal variability. Future isoprene emissions increase by more than a factor of two in 2100 (to 1242 Tg C yr^?1) when projected changes in vegetation distribution and leaf area density are included. Changing land cover and the role of nutrient limitation on CO₂ fertilization therefore remain the largest source of uncertainty in isoprene emission prediction. Although future projections suggest a compensatory balance between the effects of temperature and CO₂ on isoprene emission, the enhancement of isoprene emission due to lower ambient CO₂ concentrations did not compensate for the effect of cooler temperatures over the last 400 thousand years of the geologic record (including the Last Glacial Maximum).     

Impact of rising CO2 on emissions of volatile organic compounds: isoprene emission from Phragmites australis growing at elevated CO2 in a natural carbon dioxide spring†

Isoprene basal emission (the emission of isoprene from leaves exposed to a light intensity of 1000 µmol m^?2 s^?1 and maintained at a temperature of 30 °C) was measured in Phragmites australis plants growing under elevated CO₂ in the Bossoleto CO₂ spring at Rapolano Terme, Italy, and under ambient CO₂ at a nearby control site. Gas exchange and biochemical measurements were concurrently taken. Isoprene emission was lower in the plants growing at elevated CO₂ than in those growing at ambient CO₂. Isoprene emission and isoprene synthase activity (IsoS) were very low in plants growing at the bottom of the spring under very rich CO₂ and increased at increasing distance from the spring (and decreasing CO₂ concentration). Distance from the spring did not significantly affect photosynthesis making it therefore unlikely that there is carbon limitation to isoprene formation. The isoprene emission rate was very quickly reduced after rapid switches from elevated to ambient CO₂ in the gas‐exchange cuvette, whereas it increased when switching from ambient to elevated CO₂. The rapidity of the response may be consistent with post‐translational modifications of enzymes in the biosynthetic pathway of isoprene formation. Reduction of IsoS activity is interpreted as a long‐term response. Basal emission of isoprene was not constant over the day but showed a diurnal course opposite to photosynthesis, with a peak during the hottest hours of the day, independent of stomatal conductance and probably dependent on external air temperature or temporary reduction of CO₂ concentration. The present experiments show that basal emission rate of isoprene is likely to be reduced under future elevated CO₂ levels and allow improvement in the modelling of future isoprene emission rates.     

The effect of elevated atmospheric CO2 and drought on sources and sinks of isoprene in a temperate and tropical rainforest mesocosm

Isoprene is the most abundant volatile hydrocarbon emitted by many tree species and has a major impact on tropospheric chemistry, leading to formation of pollutants and enhancing the lifetime of methane, a powerful greenhouse gas. Reliable estimates of global isoprene emission from different ecosystems demand a clear understanding of the processes of both production and consumption. Although the biochemistry of isoprene production has been studied extensively and environmental controls over its emission are relatively well known, the study of isoprene consumption in soil has been largely neglected. Here, we present results on the production and consumption of isoprene studied by measuring the following different components: (1) leaf and soil and (2) at the whole ecosystem level in two distinct enclosed ultraviolet light‐depleted mesocosms at the Biosphere 2 facility: a cottonwood plantation with trees grown at ambient and elevated atmospheric CO₂ concentrations and a tropical rainforest, under well watered and drought conditions. Consumption of isoprene by soil was observed in both systems. The isoprene sink capacity of litter‐free soil of the agriforest stands showed no significant response to different CO₂ treatments, while isoprene production was strongly depressed by elevated atmospheric CO₂ concentrations. In both mesocosms, drought suppressed the sink capacity, but the full sink capacity of dry soil was recovered within a few hours upon rewetting. We conclude that soil uptake of atmospheric isoprene is likely to be modest but significant and needs to be taken into account for a comprehensive estimate of the global isoprene budget. More studies investigating the capacity of soils to uptake isoprene in natural conditions are clearly needed.     

Acclimation of photosynthesis to elevated CO2 in onion (Allium cepa) grown at a range of temperatures

Onion (Allium cepa) was grown in the field within temperature gradient tunnels (providing about ‐2.5°C to +2.5°C from outside temperatures) maintained at either 374 or 532 μmol mol^?1 CO₂. Plant leaf area was determined non‐destructively at 7 day intervals until the time of bulbing in 12 combinations of temperature and CO₂ concentration. Gas exchange was measured in each plot at the time of bulbing, and the carbohydrate content of the leaf (source) and bulb (sink) was determined. Maximum rate of leaf area expansion increased with mean temperature. Leaf area duration and maximum rate of leaf area expansion were not significantly affected by CO₂. The light‐saturated rates of leaf photosynthesis (A_sat) were greater in plants grown at normal than at elevated CO₂ concentrations at the same measurement CO₂ concentration. Acclimation of photosynthesis decreased with an increase in growth temperature, and with an increase in leaf nitrogen content at elevated CO₂. The ratio of intercellular to atmospheric CO₂ (C₁/C₃ ratio) was 7.4% less for plants grown at elevated compared with normal CO₂. A_sat in plants grown at elevated CO₂ was less than in plants grown at normal CO₂ when compared at the same C₁. Hence, acclimation of photosynthesis was due both to stomatal acclimation and to limitations to biochemical CO₂ fixation. Carbohydrate content of the onion bulbs was greater at elevated than at normal CO₂. In contrast, carbohydrate content was less at elevated compared with normal CO₂ in the leaf sections in which CO₂ exchange was measured at the same developmental stage. Therefore, acclimation of photosynthesis in fully expanded onion leaves was detected despite the absence of localised carbohydrate accumulation in these field‐grown crops.     

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.     

Effects of doubled atmospheric CO2 concentration on the growth and photosynthesis of Chlamydomonas reinhardtii (Volvocales, Chlorophyceae)

The freshwater microalga, Chlamydomonas reinhardtii Dangeard, was cultured under 350 and 700 ppmv CO₂ to determine the impact of doubled atmospheric CO₂ concentration on its growth and photosynthesis. No significant difference was observed in the specific growth rate, photosynthetic efficiency, maximal net photo‐synthetic rate and light‐saturating point between the low and high CO₂ cultures. Both the low‐ and high‐CO₂‐grown cells showed reduced light‐dependent O₂ evolution rate and photochemical efficiency (F_v/F_m) owing to photoinhibition when exposed to high photon flux density. However, high‐CO₂‐grown cells were less photoinhibited, and showed better recovery in dim light or darkness during the initial period of the recovery process.     

Photosynthetic acclimation to elevated CO2 in Phaseolus vulgaris L. is altered by growth response to nitrogen supply

Abstract Long‐term exposure of plants to elevated CO₂ often leads to downward photosynthetic acclimation. Nitrogen (N) deficiency could potentially exacerbate this response by reducing growth rate and the sink for photosynthates, but this has not always been observed. Experimentally, the interpretation of N effects on CO₂ responses can be confounded by increasing severity of tissue N deficiency over time when N supply is not adjusted as demand increases. In this study, N supply ranged from sub‐ to supra‐optimal (20–540 kgN ha^–l equivalent), and relatively stable levels of tissue N concentration were obtained in all treatments by varying twice‐weekly application rates in proportion to plant growth. The effects of N on photosynthesis and growth of beans (Phaseolus vulgaris L.) raised at ambient (35 Pa) and three elevated (70, 105, 140 Pa) CO₂ partial pressures (pCO₂) were evaluated. Averaging across N treatments, leaf total non‐structural carbohydrates (TNC) were 2.5‐ to 3‐fold higher and leaf N concentrations were 31–35% lower at elevated compared to ambient pCO₂. Light‐saturated net CO₂ assimilation rates measured at growth pCO₂ (A_satg) were significantly higher (26–40% depending on N supply) in plants grown at elevated compared to ambient pCO₂. When measured at a common pCO₂ of 35 Pa, the A_sat of plants grown at elevated CO₂ was 15–29% less than that of plants grown at 35 Pa, indicative of downward photosynthetic acclimation. The magnitude of downward photosynthetic acclimation to elevated CO₂ was greater in plants grown at high (180 and 540 kgN ha^–l) compared to low (20 and 60 kgN ha^–l) N supply, and this was associated with a higher A_sat at growth pCO₂, higher leaf area ratio (leaf area/total biomass), and higher TNC in leaves of high‐N plants. Our results indicate that the effect of N on acclimation to CO₂ will depend on the balance between supply and demand for N during the growing period, and the effect this has on biomass allocation and source‐sink C balance at the whole‐plant level.     

Effects of elevated CO2 concentration and water deficit on fructan metabolism in Viguiera discolor Baker

Elevated [CO₂] is suggested to mitigate the negative effects of water stress in plants; however responses vary among species. Fructans are recognised as protective compounds against drought and other stresses, as well as having a role as reserve carbohydrates. We analysed the combined effects of elevated [CO₂] and water deficit on fructan metabolism in the Cerrado species Viguiera discolor Baker. Plants were cultivated for 18 days in open‐top chambers (OTC) under ambient (～380 ppm), and high (～760 ppm) [CO₂]. In each OTC, plants were submitted to three treatments: (i) daily watering (control), (ii) withholding water (WS) for 18 days and (iii) re‐watering (RW) on day 11. Analyses were performed at time 0 and days 5, 8, 11, 15 and 18. High [CO₂] increased photosynthesis in control plants and increased water use efficiency in WS plants. The decline in soil water content was more distinct in WS 760 (WS under 760 ppm), although the leaf and tuberous root water status was similar to WS 380 plants (WS under 380 ppm). Regarding fructan active enzymes, 1‐SST activity decreased in WS plants in both CO₂ concentrations, a result consistent with the decline in photosynthesis and, consequently, in substrate availability. Under WS and both [CO₂] treatments, 1‐FFT and 1‐FEH seemed to act in combination to generate osmotically active compounds and thus overcome water deficit. The proportion of hexoses to sucrose, 1‐kestose and nystose (SKN) was higher in WS plants. In WS 760, this increase was higher than in WS 380, and was not accompanied by decreases in SKN at the beginning of the treatment, as observed in WS 380 plants. These results suggest that the higher [CO₂] in the atmosphere contributed to maintain, for a longer period, the pool of hexoses and of low DP fructans, favouring the maintenance of the water status and plant survival under drought.     

Leaf temperature of soybean grown under elevated CO2 increases Aphis glycines (Hemiptera: Aphididae) population growth

Abstract Plants grown under elevated carbon dioxide (CO₂) experience physiological changes that influence their suitability as food for insects. To determine the effects of living on soybean (Glycine max Linnaeus) grown under elevated CO₂, population growth of the soybean aphid (Aphis glycines Matsumura) was determined at the SoyFACE research site at the University of Illinois, Urbana‐Champaign, Illinois, USA, grown under elevated (550 μL/L) and ambient (370 μL/L) levels of CO₂. Growth of aphid populations under elevated CO₂ was significantly greater after 1 week, with populations attaining twice the size of those on plants grown under ambient levels of CO₂. Soybean leaves grown under elevated levels of CO₂ were previously demonstrated at SoyFACE to have increased leaf temperature caused by reduced stomatal conductance. To separate the increased leaf temperature from other effects of elevated CO₂, air temperature was lowered while the CO₂ level was increased, which lowered overall leaf temperatures to those measured for leaves grown under ambient levels of CO₂. Aphid population growth on plants grown under elevated CO₂ and reduced air temperature was not significantly greater than on plants grown under ambient levels of CO₂. By increasing Glycine max leaf temperature, elevated CO₂ may increase populations of Aphis glycines and their impact on crop productivity.     

Combined effects of ocean acidification and solar UV radiation on photosynthesis,growth, pigmentation and calcification of the coralline alga Corallina sessilis (Rhodophyta)

Previous studies have shown that increasing atmospheric CO₂ concentrations affect calcification in some planktonic and macroalgal calcifiers due to the changed carbonate chemistry of seawater. However, little is known regarding how calcifying algae respond to solar UV radiation (UVR, UVA+UVB, 280–400 nm). UVR may act synergistically, antagonistically or independently with ocean acidification (high CO₂/low pH of seawater) to affect their calcification processes. We cultured the articulated coralline alga Corallina sessilis Yendo at 380 ppmv (low) and 1000 ppmv (high) CO₂ levels while exposing the alga to solar radiation treatments with or without UVR. The presence of UVR inhibited the growth, photosynthetic O₂ evolution and calcification rates by13%, 6% and 3% in the low and by 47%, 20% and 8% in the high CO₂ concentrations, respectively, reflecting a synergistic effect of CO₂ enrichment with UVR. UVR induced significant decline of pH in the CO₂‐enriched cultures. The contents of key photosynthetic pigments, chlorophyll a and phycobiliproteins decreased, while UV‐absorptivity increased under the high pCO₂/low pH condition. Nevertheless, UV‐induced inhibition of photosynthesis increased when the ratio of particulate inorganic carbon/particulate organic carbon decreased under the influence of CO₂‐acidified seawater, suggesting that the calcified layer played a UV‐protective role. Both UVA and UVB negatively impacted photosynthesis and calcification, but the inhibition caused by UVB was about 2.5–2.6 times that caused by UVA. The results imply that coralline algae suffer from more damage caused by UVB as they calcify less and less with progressing ocean acidification.     

Effects of atmospheric CO2 enrichment and root restriction on leaf gas exchange and growth of banana (Musa)

The effects of atmospheric CO₂ enrichment and root restriction on photosynthetic characteristics and growth of banana (Musa sp. AAA cv. Gros Michel) plants were investigated. Plants were grown aeroponically in root chambers in controlled environment glasshouse rooms at CO₂ concentrations of 350 or 1 000 μmol CO₂ mol^-1. At each CO₂ concentration, plants were grown in large (2001) root chambers that did not restrict root growth or in small (20 1) root chambers that restricted root growth. Plants grown at 350 μmol CO₂ mol^-1 generally had a higher carboxylation efficiency than plants grown at 1 000 μmol CO₂ mol^-1 although actual net CO₂ assimilation (A) was higher at the higher ambient CO₂ concentration due to increased intercellular CO₂ concentrations (C_i resulting from CO₂ enrichment. Thus, plants grown at 1 000 μmol CO₂ mol^-1 accumulated more leaf area and dry weight than plants grown at 350 μmol CO₂ mol^-1. Plants grown in the large root chambers were more photosynthetically efficient than plants grown in the small root chambers. At 350 μmol CO₂ mol^-1, leaf area and dry weights of plant organs were generally greater for plants in the large root chambers compared to those in the small root chambers. Atmospheric CO₂ enrichment may have compensated for the effects of root restriction on plant growth since at 1 000 μmol CO₂ mol^-1 there was generally no effect of root chamber size on plant dry weight.     

PHOTOSYNTHETIC UTILIZATION OF INORGANIC CARBON IN THE ECONOMIC BROWN ALGA,HIZIKIA FUSIFORME (SARGASSACEAE) FROM THE SOUTH CHINA SEA1

The mechanism of inorganic carbon (C_i) acquisition by the economic brown macroalga, Hizikia fusiforme (Harv.) Okamura (Sargassaceae), was investigated to characterize its photosynthetic physiology. Both intracellular and extracellular carbonic anhydrase (CA) were detected, with the external CA activity accounting for about 5% of the total. Hizikia fusiforme showed higher rates of photosynthetic oxygen evolution at alkaline pH than those theoretically derived from the rates of uncatalyzed CO₂ production from bicarbonate and exhibited a high pH compensation point (pH 9.66). The external CA inhibitor, acetazolamide, significantly depressed the photosynthetic oxygen evolution, whereas the anion‐exchanger inhibitor 4,4′‐diisothiocyano‐stilbene‐2,2′‐disulfonate had no inhibitory effect on it, implying the alga was capable of using HCO₃^? as a source of C_i for its photosynthesis via the mediation of the external CA. CO₂ concentrations in the culture media affected its photosynthetic properties. A high level of CO₂ (10,000 ppmv) resulted in a decrease in the external CA activity; however, a low CO₂ level (20 ppmv) led to no changes in the external CA activity but raised the intracellular CA activity. Parallel to the reduction in the external CA activity at the high CO₂ was a reduction in the photosynthetic CO₂ affinity. Decreased activity of the external CA in the high CO₂ grown samples led to reduced sensitiveness of photosynthesis to the addition of acetazolamide at alkaline pH. It was clearly indicated that H. fusiforme, which showed CO₂‐limited photosynthesis with the half‐saturating concentration of C_i exceeding that of seawater, did not operate active HCO₃^? uptake but used it via the extracellular CA for its photosynthetic carbon fixation.     

Monoterpene emission and monoterpene synthase activities in the Mediterranean evergreen oak Quercus ilex L. grown at elevated CO2 concentrations

Monoterpene emissions, monoterpene synthase activities, photosynthesis, fluorescence yield in the dark and drought stress indicators (stomatal conductance and mid‐day water potential) were concurrently measured under similar temperature and illumination in current‐year leaves of Quercus ilex L. of plants grown in open‐top chambers at ambient (350 ppm) and elevated (700 ppm) CO₂. The study was undertaken to understand the effect of CO₂ on monoterpene biosynthesis, and to predict and parameterize the biogenic emissions at growing CO₂ concentrations. The results of the 1998 and 1999 studies show that at elevated CO₂, and in the absence of persistent environmental stresses, photosynthesis was stimulated with respect to ambient CO₂, but that the emission of the three most abundantly emitted monoterpenes (α‐pinene, sabinene and β‐pinene) was inhibited by approximately 68%. The enzyme activities of the monoterpene synthases catalysing the formation of the three monoterpenes were also inhibited at elevated CO₂ and an excellent relationship was found between monoterpene emission and activity of the corresponding enzyme both at ambient and elevated CO₂. Interestingly, however, limonene emission was enhanced in conditions of elevated CO₂ as it was also the corresponding synthase. The ratio between enzyme activity and emission of the three main monoterpenes was high (above 20) at ambient CO₂ but it was below 10 at elevated CO₂ and, for limonene, on both treatments. Our results indicate that the overall emission of monoterpenes at elevated CO₂ will be inhibited because of a concurrent, strong down‐regulation of monoterpene synthase activities. When the enzyme activity does not change, as for limonene, the high photosynthetic carbon availability at elevated CO₂ conditions may even stimulate emission. The results of the 1997 study show that severe and persistent drought, as commonly occurs in the Mediterranean, may inhibit both photosynthesis and monoterpene (α‐pinene) emission, particularly at ambient CO₂. Thus, emission is probably limited by photosynthetic carbon availability; the effect of elevated CO₂per se is not apparent if drought, and perhaps other environmental stresses, are also present.