Bibliography on CO2 Effects on Vegetation and Ecosystems: 1990-1999 Literature Michael H. Jones and Peter S. Curtis, editors ORNL/CDIAC-129 July 2000 (http://cdiac.esd.ornl.gov/epubs/cdiac/cdiac129/cdiac129.html) 1 Abarzua, S., R. Altenburger, R. Callies, L.H. Grimme, A. Mayer, D. Leibfritz, and U. Schiewer. 1993. Ammonium rhythm in cultures of the cyanobacterium microcystis- firma. Physiologia Plantarum 89(3):659-663. Over a period of several days, rhythmic changes in extracellular NH4+ concentration take place in cultures of the cyanobacterium Microcystis firma (Breb et Lenorm.) Schmidle, strain Gromov/St. Petersb. 398, under conditions of restricted CO2 supply and light/dark alternation. The changes are enhanced by nitrate supply. Among the various processes generating intracellular NH4+ (NH4+ uptake, NO3- reduction, protein and amino acid degradation, photorespiration), NO3- reduction appears as the one most important. This can be concluded from experiments with and without nitrate and/or ammonium in the medium. In the presence of saturating CO2, continuous light, or continuous darkness, rhythmic NH4+ oscillations are not induced. Studies of the incorporation of NH4+ nitrogen by in vivo N-15-NMR show that if CO2 is supplied, N-15 is accumulated in several components with the following time course: in the first hour in Gln (delta), in the second hour in the alpha- amino groups of most nonbranched amino acids, in the third hour in gamma-aminobutyric acid (GABA), Orn (delta) and Lys (epsilon), and in the sixth hour in Ala. Carbon limitation, however, results in accumulation of label in the amide nitrogen of glutamine only. KEYWORDS: METABOLISM, N-15, NMR-SPECTROSCOPY, NUCLEAR MAGNETIC- RESONANCE 2 Abdin, O.A., X.M. Zhou, B.E. Coulman, D. Cloutier, M.A. Faris, and D.L. Smith. 1998. Effect of sucrose supplementation by stem injection on the development of soybean plants. Journal of Experimental Botany 49(329):2013-2018. Over the past half decade several stem injection methods have been developed for cereal plants. These methods allow researchers to administer solutions to cereal plants to study their effects on plant physiology. However, little work has been done to extend this technique to non-cereals. An experiment was conducted to test an injection technique that could be suitable for soybean plants (Glycine max [L.] Merr.), and to study the effect of long-term injection of sucrose on the growth of soybean plants. An injection setup comprising a supporting stand and a fluid injection system was established. Pressure was applied to the plunger of a 5 ml syringe using ceramic bricks to force test solutions into the plants. Solutions of 0, 150, and 300 g sucrose I-1 were injected into soybean plants for 8 weeks starting at the seedling VC stage. Distilled water had the greatest uptake rate, followed by the 150, and then the 300 g sucrose I-1 solutions. The overall average uptake during the injection period was 77.3 ml. Average sucrose uptake values were 11.8 and 13.5 g per plant for the 150 and 300 g sucrose I-1 treatments. This represented approximately 65% of the total dry weight of the plants. Sucrose injection increased leaf area and pod number relative to the control plants. Nodule numbers were lower for sucrose injected treatments, but their dry weights were higher than the control. Above-soil dry matter was higher for the plants injected with 300 g sucrose I-1 than those injected with water. The injection system tested was able to administer concentrated solutions into soybean plants for most of their period of growth and development. The sucrose supplementation had positive effects on soybean growth but suppressed photosynthesis. KEYWORDS: CARBON-DIOXIDE ENRICHMENT, ELEVATED CO2, GROWTH, LEAVES, MAIZE, NITROGEN, PEDUNCLE PERFUSION, PHOTOSYNTHESIS, WHEAT, YIELD 3 Abdullaev, A.A., B.B. Dzhumaev, Z.N. Abdurakhmanova, V.L. Kaler, and I.M. Magmedov. 1992. Integral effect of environmental-factors on photosynthetic metabolism of carbon in cotton leaves. Soviet Plant Physiology 39(2):140-144. We used the method of mathematical experiment planning (a 2(3) scheme) to study the influence of environmental factors separately or in combination on the photosynthetic rate and distribution of C-14 among products of photosynthetic carbon metabolism in the cotton (Gossypium hirsutum L.) leaf Increase of light intensity during cultivation accelerated photosynthesis and stimulated incorporation of C-14 into phosphoglyceric acid (PGA), sugar diphosphate (SDP), fructose monophosphate (FMP), and malate, but suppressed incorporation of C-14 into sucrose, glucose monophosphate (GMP), and glycerate. Temperature increase by itself and in any combination with other factors at the upper level suppressed photosynthesis. Elevated temperature increased accumulation of the label in PGA, sucrose, and malate, but lowered it in GMP, alanine, glycine, and serine. Growing plants at enhanced CO2 concentration led to acceleration of photosynthesis and increase of the share of C-14 in SDP, GMP, and malate, but decrease of it in sucrose, alanine, glycine, and serine. Very perceptible effects of interaction are discernible in different combinations of factors. All three factors at the upper level appreciably induced activity of phosphoenolpyruvate carboxylase (PEPCase) in cotton leaves. KEYWORDS: PHYSIOLOGY 4 Aben, S.K., S.P. Seneweera, O. Ghannoum, and J.P. Conroy. 1999. Nitrogen requirements for maximum growth and photosynthesis of rice, Oryza sativa L-cv. Jarrah grown at 36 And 70 Pa CO2. Australian Journal of Plant Physiology 26(8):759-766. The hypothesis that growth of rice (Oryza sativa L. cv. Jarrah) at elevated atmospheric CO2 partial pressure alters leaf nitrogen (N) concentrations required to support maximum dry mass production and photosynthetic rates during the period of rapid tiller initiation was tested by growing plants for 30 days in unstirred sand/hydroponic culture with N concentrations of 5, 20, 40, 60 and 100 mg N L-1. Maximum growth and photosynthetic potential was greater at 70 than 36 Pa CO2 at all N concentrations in the solution. Elevated CO2 reduced leaf N concentrations required to support 90% of maximum growth and photosynthetic rates (critical concentration) from 40 to 27 g kg(-1) for growth and from 45 to 30 g kg(-1) for photosynthesis. Morphological changes at elevated CO2 included increased tiller numbers and reduced leaf area ratio. The latter could be explained by lower plant N concentrations which occurred at high CO2 at each N concentration in the solution, primarily due to lower leaf blade and root N concentrations. Changes in tiller numbers at high CO2 were unrelated to leaf or plant N but were strongly correlated with leaf soluble carbohydrate concentrations. We conclude that elevated CO2 alters the nutritional physiology of rice during the rapid tillering phase in a way that increases the efficiency of N utilisation for growth and photosynthesis. KEYWORDS: ACCLIMATION, CAPACITY, COTTON, ELEVATED CO2, LEAVES, NUTRITION, PLANTS, RIBULOSE-1;5-BISPHOSPHATE CARBOXYLASE-OXYGENASE, WHEAT 5 Aber, J.D., S.V. Ollinger, C.A. Federer, P.B. Reich, M.L. Goulden, D.W. Kicklighter, J.M. Melillo, and R.G. Lathrop. 1995. Predicting the effects of climate change on water yield and forest production in the northeastern United States. Climate Research 5(3):207-222. Rapid and simultaneous changes in temperature, precipitation and the atmospheric concentration of CO2 are predicted to occur over the next century. Simple, well-validated models of ecosystem function are required to predict the effects of these changes. This paper describes an improved version of a forest carbon and water balance model (PnET-II) and the application of the model to predict stand- and regional-level effects of changes in temperature, precipitation and atmospheric CO2 concentration. PnET-II is a simple, generalized, monthly time- step model of water and carbon balances (gross and net) driven by nitrogen availability as expressed through foliar N concentration. Improvements from the original model include a complete carbon balance and improvements in the prediction of canopy phenology, as well as in the computation of canopy structure and photosynthesis. The model was parameterized and run for 4 forest/site combinations and validated against available data for water yield, gross and net carbon exchange and biomass production. The validation exercise suggests that the determination of actual water availability to stands and the occurrence or non-occurrence of soil-based water stress are critical to accurate modeling of forest net primary production (NPP) and net ecosystem production (NEP). The model was then run for the entire NewEngland/New York (USA) region using a 1 km resolution geographic information system. predicted long- term NEP ranged from -85 to + 275 g Cm-2 yr(-1) for the 4 forest/site combinations, and from -150 to 350 g C m(-2) yr(-1) for the region, with a regional average of 76 g Cm-2 yr(-1). A combination of increased temperature (+6 degrees C), decreased precipitation (-15%) and increased water use efficiency (2x, due to doubling of CO2) resulted generally in increases in NPP and decreases in water yield over the region. KEYWORDS: DEPOSITION, ECOSYSTEMS, ELEVATED CO2, MODEL, REGIONAL-ANALYSIS, RESPONSES 6 Ackerly, D.D., and F.A. Bazzaz. 1995. Plant-growth and reproduction along co2 gradients - nonlinear responses and implications for community change. Global Change Biology 1(3):199-207. 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 (nat) 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. KEYWORDS: ANNUALS, ATMOSPHERIC CO2, CO2-INDUCED CLIMATE CHANGE, COMPETITION, ELEVATED CO2, ENRICHMENT, LIQUIDAMBAR- STYRACIFLUA, OLD- FIELD PERENNIALS, PINUS-TAEDA SEEDLINGS, RESOURCE USE 7 Ackerly, D.D., J.S. Coleman, S.R. Morse, and F.A. Bazzaz. 1992. Co2 and temperature effects on leaf-area production in 2 annual plant-species. Ecology 73(4):1260-1269. We studied leaf area production in two annual plant species, Abutilon theophrasti and Amaranthus retroflexus, under three day/night temperature regimes (18-degrees/14-degrees, 28- degrees/22- degrees, and 38-degrees/31-degrees-C) and two concentrations of carbon dioxide (400 and 700-mu- L/L). The production of whole-plant leaf area during the first 30 d of growth was analyzed in terms of the leaf initiation rate, leaf expansion, individual leaf area, and, in Amaranthus, production of branch leaves. Temperature and CO2 influenced leaf area production through effects on the rate of development, determined by the production of nodes on the main stem (the plastochron index), and through shifts in the relationship between whole-plant leaf area and the number of main stem nodes. In Abutilon, leaf initiation rate was highest at 38- degrees, but area of individual leaves was greatest at 28- degrees. Total leaf area was greatly reduced at 18-degrees due to slow leaf initiation rates. Elevated CO2 concentration increased leaf initiation rate at 28-degrees, resulting in an increase in whole-plant leaf area. In Amaranthus, leaf initiation rate increased with temperature, and was increased by elevated CO2 at 28-degrees. Individual leaf area was greatest at 28-degrees, and was increased by elevated CO2 at 28-degrees but decreased at 38-degrees. Branch leaf area displayed a similar response to CO2, but was greater at 38- degrees. Overall, whole-plant leaf area was slightly increased at 38-degrees relative to 28-degrees, and elevated CO2 levels resulted in increased leaf area at 28-degrees but decreased leaf area at 38-degrees. The effects on leaf area closely parallel rates of biomass accumulation in the same experiment, suggesting that responses of developmental processes to elevated CO2 and interacting factors may play an important role in mediating effects on plant growth. KEYWORDS: C-3, CANOPY, CARBON-DIOXIDE ENRICHMENT, ELEVATED CO2, GROWTH, LEAVES, LIGHT, PHOTOSYNTHESIS, RESPONSES, SUNFLOWER 8 Acock, B., M.C. Acock, and D. Pasternak. 1990. Interactions of CO2 enrichment and temperature on carbohydrate production and accumulation in muskmelon leaves. Journal of the American Society for Horticultural Science 115(4):525-529. 9 Acock, B., and G.W. Wall. 1995. A simple conductimetric co2 analyzer with automatic recalibration .1. Design, implementation, and functionality. Agronomy Journal 87(1):70-75. Controlled-environment plant growth cabinets may be used to investigate the long-term effect of elevated carbon dioxide concentration ([CO2]) on plant growth. Infrared gas analyzers (IRGAs) are normally used to monitor and control [CO2] in plant cabinets. With many cabinets in use, however, it soon becomes impractical to purchase an individual IRGA for each cabinet, A more economical method of monitoring and controlling [CO2] relies on the change in electrical conductivity when CO2 is dissolved in demineralized water, This work describes the design, implementation, and functionality of an inexpensive conductimetric system for controlling [CO2] in plant growth cabinets, Regressing electrical conductivity against [CO2] over the range 0 to 1000 mu L L(-1) yields a quadratic response. Calibration drift inherent in the conductimetric CO2 analyzer requires that each analyzer be recalibrated periodically. Automatically recalibrating with an IRGA every 900 s gave control of the [CO2] within the plant enclosures to within 10 to 15 mu L L(-1) of the set point, The [CO2] control system is robust enough to maintain this accuracy regardless of the desired [CO2] set point or the mass of plant material within the plant growth cabinet, In this approach, only one IRGA is required to control [CO2] in many plant growth cabinets if each cabinet has a dedicated conductimetric CO2 analyzer. 10 Adams, R.M., R.A. Fleming, C.C. Chang, B.A. McCarl, and C. Rosenzweig. 1995. A reassessment of the economic-effects of global climate-change on US agriculture. Climatic Change 30(2):147-167. This study uses recent GCM forecasts, improved plant science and water supply data and refined economic modeling capabilities to reassess the economic consequences of long-term climate change on U.S. agriculture. Changes in crop yields, crop water demand and irrigation water arising from climate change result in changes in economic welfare. Economic consequences of the three GCM scenarios are mixed; GISS and GFDL-QFlux result in aggregate economic gains, UKMO implies losses. As in previous studies, the yield enhancing effects of atmospheric CO2 are an important determinant of potential economic consequences. Inclusion of changes in world food production and associated export changes generally have a positive affect on U.S. agriculture. As with previous studies, the magnitude of economic effects estimated here are a small percentage of U. S. agricultural value. 11 Adamse, P., and S.J. Britz. 1992. Amelioration of uv-b damage under high irradiance .1. Role of photosynthesis. Photochemistry and photobiology 56(5):645-650. Sensitivity to ultraviolet-B radiation (UV-B, 280-315 nm) is generally reduced when background irradiance is high. We tested the involvement of photosynthesis in the amelioration of UV-B damage by treating plants at high PAR (photosynthetically- active radiation. 400-700 nm; 1000 mumol m-2 s-1) with supplemental UV-B at double ambient levels of biologically- effective radiation (18 kJ m-2 d-1) and either ''ambient'' (450 mumol mol-1) or short term elevated (750 mumol mol-1) CO2 levels. Responses to UV-B were assessed by photosynthetic gas exchange, leaf expansion and production of UV-absorbing compounds (presumptive flavonoids) in cultivars of cucumber (Cucumis sativus L.) previously demonstrated to be relatively sensitive (cv. Poinsett) and insensitive (cv. Ashley) to UV-B. Except for marginal leaf interveinal chlorosis observed in Poinsett, both cultivars responded similarly. UV-B had little direct effect on leaf photosynthesis, but it did cause reductions in leaf area and corresponding increases in leaf dry matter per area. Increased CO2 stimulated plant growth, counteracting the effect of UV-B on leaf growth and indicating an important role for photosynthesis. In contrast, the accumulation of UV-absorbing flavonoid compounds was enhanced by UV-B exposure but was not affected by CO2 enrichment. KEYWORDS: ACCLIMATION, CARBON DIOXIDE, CELL-SUSPENSION CULTURES, LIGHT, PETROSELINUM- HORTENSE, PHOTON FLUX- DENSITY, PHYTOCHROME, PLANTS, RADIATION, SENSITIVITY 12 Adamsen, F.J., P.J. Pinter, E.M. Barnes, R.L. LaMorte, G.W. Wall, S.W. Leavitt, and B.A. Kimball. 1999. Measuring wheat senescence with a digital camera. Crop Science 39(3):719-724. Documenting crop senescence rates is often difficult because of the need for frequent sampling during periods of rapid change and the subjective nature of human visual observations. The purpose of this study was to determine the feasibility of using images produced by a digital camera to measure the senescence rate of wheat and to compare the results with changes in greenness determined by two established methods. Measurements were made as part of an experiment to determine the effects of elevated CO2 and limited soil nitrogen on spring wheat (Triticum aestivum L.) at the University of Arizona's Maricopa Agricultural Center, near Phoenix, AZ. "Greenness" measurements were made during senescence of the crop with a color digital camera, a hand-held radiometer, and a SPAD chlorophyll meter. The green to red (GIR) for each pixel in an image was calculated and the average GIR computed for cropped images from a digital camera representing 1 m(2) for each treatment and sample date. The normalized difference vegetation index (NDVI) was calculated from the red and near-infrared canopy reflectances measured with a hand held radiometer. A SPAD reading was obtained from randomly selected flag leaves. All three methods of measuring plant greenness showed similar temporal trends. The relationships between GIR with NDVI and SPAD were linear over most of the range of GIR. However, NDVI was more sensitive at low values than GIR. GIR was more sensitive above G/R values of 1.2 than SPAD because the upper limits of SPAD measurements were constrained by the amount of chlorophyll in the leaf, while GIR responded to both chlorophyll concentration in the leaves as well as the number of leaves present. Color digital imaging appears useful for quantifying the senescence of crop canopies. The cost of color digital cameras is expected to decrease and the quality and convenience of use to improve. KEYWORDS: CHLOROPHYLL METER, CROP, EFFICIENCY, RED, VEGETATION INDEXES, WINTER-WHEAT, YIELD 13 Agar, I.T., J. Streif, and F. Bangerth. 1997. Effect of high CO2 and controlled atmosphere (CA) on the ascorbic and dehydroascorbic acid content of some berry fruits. Postharvest Biology and Technology 11(1):47-55. High CO2 concentrations as well as controlled atmosphere storage are widely used to extend the storage and shelf-life of many fruits. To investigate the effect of these storage procedures on several berry fruits, strawberries, raspberries, currants and blackberries were stored at three different elevated CO2 concentrations, with or without a parallel reduction in O-2. Vitamin C content (ascorbic acid plus dehydroascorbic acid) was reduced by high CO2 concentrations (10-30% CO2), particularly in strawberries. This reduction in vitamin C was moderate in black currants and blackberries and almost absent in raspberries and red currants when compared with strawberries. Reducing the O-2 concentration in the storage atmosphere in the presence of high CO2 had little effect on the vitamin C content. Ascorbic acid was more diminished al high CO2 than dehydroascorbic acid. This suggests a stimulating effect of high CO2 concentrations on the oxidation of ascorbic acid and/or an inhibition of mono- or dehydroascorbic acid reduction to ascorbic acid. (C) 1997 Elsevier Science B.V. KEYWORDS: HYDROGEN- PEROXIDE, O2, PLANTS 14 Aggangan, R.T., A.M. O'Connell, J.F. McGrath, and B. Dell. 1999. The effects of Eucalyptus globulus Labill. leaf letter on C and N mineralization in soils from pasture and native forest. Soil Biology and Biochemistry 31(11):1481-1487. The effects of addition of Eucalyptus globulus leaf litter on carbon and nitrogen mineralization in soils from a pasture and a native forest were evaluated using a long-term laboratory aerobic incubation assay (29 weeks at 20 degrees C) in leaching microlysimeters, The amount of added leaf litter significantly influenced microbial respiration, microbial biomass and N turnover in both the native forest and pasture soils. Cumulative CO2-C respired increased with increasing rate of leaf litter addition when leaf litter was mixed through the soil or placed on the soil surface. These increases were associated with increases in microbial biomass C content. Cumulative net N mineralization declined in ail treatments when litter was added and was lowest when leaf litter was mixed with soil. When leaf litter was added in increasing amounts to the soil surface, there was a concomitant increase in microbial biomass N content (r(2) = 0.79, n = 8), indicating that the reduction in net N mineralization was primarily due to immobilization of N in microbial tissues. In contrast, when litter was mixed with soil in increasing amounts, there was a decrease in microbial biomass N in forest soil and an increase in pasture soil. Consequently, changes in the rate of net N mineralization were not well related to changes in microbial biomass N content. It is suggested that this may be due to the greater activity and more rapid turnover of microorganisms where litter was incorporated resulting in more of the immobilized N being partitioned into metabolic products or dead microbial cells. Incorporation of litter may also have enhanced loss N through denitrification, (C) 1999 Published by Elsevier Science Ltd. All rights reserved. KEYWORDS: DECOMPOSITION, DENITRIFICATION, EXTRACTION METHOD, IMMOBILIZATION, LITTER, MICROBIAL BIOMASS CARBON, NITROGEN MINERALIZATION, PLANT RESIDUES, RESPIRATION, WESTERN-AUSTRALIA 15 Agren, G.I. 1996. Nitrogen productivity or photosynthesis minus respiration to calculate plant growth? Oikos 76(3):529-535. One approach to calculate plant growth rate is from models of photosynthesis, respiration and allocation. This requires that processes with characteristic time constants of seconds to minutes be scaled to hours or days. Another approach is to use aggregate models defined at the time scale of growth, hours and days. I use such an aggregate model, the nutrient productivity, to compare the performance of the two approaches on growth experiments with small, nitrogen-limited birch plants. The problems of error aggregation when using the large number of parameters required to scale from the detailed level of photosynthesis and respiration to the aggregate level of growth are in this case such that whole plant growth rate is more accurately predicted with the nutrient productivity model. KEYWORDS: ALLOCATION, BETULA-PENDULA ROTH, BIOMASS, BIRCH SEEDLINGS, CARBON, CLIMATE, ECOSYSTEMS, ELEVATED CO2, NUTRITION, STRESS 16 Agren, G.I., R.E. McMurtrie, W.J. Parton, J. Pastor, and H.H. Shugart. 1991. State-of-the-art of models of production decomposition linkages in conifer and grassland ecosystems. Ecological Applications 1(2):118-138. We review the state-of-the-art of models of forests and grasslands that could be used to predict the impact of a future climate change arising from increased atmospheric carbon dioxide concentration. Four levels of resolution are recognized: physiologically based models, population models, ecosystem models, and regional or global models. At the physiological level a number of important processes can be described in great detail, but these models often treat inadequately interactions with nutrient cycles, which operate on longer time scales. Population and ecosystem models can, on the other hand, encapsulate relationships between the plants and the soil system, but at the expense of requiring more ad hoc formulations of processes. At the regional and global scale we have so far only steady-state models, which cannot be used to predict transients caused by climate change. However, our conclusion is that, in spite of the gaps in knowledge, there are several models based on dominant processes that are well enough understood for the predictions of those models to be taken seriously. KEYWORDS: ATMOSPHERIC CO2 ENRICHMENT, BOUTELOUA-GRACILIS, CARBON DIOXIDE, CO2-INDUCED CLIMATE CHANGE, EVEN- AGED STANDS, LOLIUM-PERENNE L, NITROGEN PRODUCTIVITY, PLANT GROWTH, SIMULATION-MODEL, THEORETICAL- ANALYSIS 17 Ahmadi, H., W.V. Biasi, and E.J. Mitcham. 1999. Control of brown rot decay of nectarines with 15% carbon dioxide atmospheres. Journal of the American Society for Horticultural Science 124(6):708-712. Effects of short-term exposure to a 15% CO2 atmosphere on nectarines [Prunus persica (L.) Batsch (Nectarine Group) 'Summer Red'] inoculated with Monilinia fructicola (Wint.) Honey (causal agent of brown rot) were investigated, Nectarines were inoculated with spores of M.fructicola and incubated at 20 degrees C for 24, 48 or 72 hours and then transferred to storage in either air or air enriched with 15% CO2 at 5 degrees C. Fruit were removed from storage after 5 and 16 days and were examined for brown rot decay immediately and after ripening in air for 3 days at 20 degrees C. Noninoculated nectarines were stored and treated likewise for evaluation of postharvest fruit attributes to determine their tolerance to 15% CO2. Incubation period after inoculation, storage duration, and storage atmosphere had highly significant effects on fruit decay, 'Summer Red' nectarines tolerated a 15% CO2 atmosphere for 16 days at 5 degrees C. Development of brown rot decay in fruit inoculated 24 hours before 5 or 16 days storage in 15% CO2 at 5 OC was arrested. After 3 days ripening in air at 20 degrees C, the progression of brown rot disease was rapid in all inoculated nectarines, demonstrating the fungistatic effect of 15% CO2. The quantity of fungal cell wall materials (estimated by glucosamine concentration) was compared to visual estimation of decayed area and visual rating of fungal sporulation. The glucosamine assay defined the onset and progress of brown rot infection more precisely than either of the two visual tests. KEYWORDS: FRUIT, IPRODIONE, MOLD, SUPPRESSION, SWEET CHERRIES, TISSUE, TOMATO PRODUCTS 18 Ahmed, F.E., A.E. Hall, and M.A. Madore. 1993. Interactive effects of high-temperature and elevated carbon- dioxide concentration on cowpea [vigna-unguiculata (L) walp]. Plant, Cell and Environment 16(7):835-842. Limitations in carbohydrate supplies have been implicated as a factor responsible for reproductive failure under heat stress. Heat stress affects two stages of reproductive development in cowpea [Vigna unguiculata (L.) Walp.], and genotypes are available with tolerance and sensitivity to heat during these different stages. The objectives of this study were to determine the responses of these cowpea lines to ambient and elevated [CO2], under heat stress and optimal temperature, and test whether differences in carbohydrate supplies due to genotypes, CO2 enrichment and heat stress are associated with differences in sensitivity to heat during reproductive development. Plants were grown in reach-in growth chambers and subjected to day/night temperatures of either 33/20 or 33/30- degrees-C, and [CO2] levels of either 350 or 700 mumol mol-1. Under intermediate night temperature (33/20-degrees-C), all lines set substantial numbers of pods. Under high night temperature (33/30-degrees-C) with either ambient or elevated [CO2], one heat-sensitive line produced no flowers and the other set no pods, whereas the heat-tolerant line abundantly set pods. High night temperature reduced the overall carbohydrate content of the plants, especially peduncle sugars, and caused decreases in photosynthetic rates. The high pod set of the heat-tolerant line, under high night temperature, was associated with higher levels of sugars in peduncles compared with the heat-sensitive lines. The heat-tolerant line accumulated substantial shoot biomass, exhibited less accumulation of starch in leaves, and possibly had less down- regulation of photosynthesis in response to CO2 enrichment and heat stress than the heat-sensitive lines. Elevated [CO2] resulted in higher overall carbohydrate levels in heat- sensitive lines (starch in leaves, stems and peduncles), but it did not increase their heat tolerance with respect to flower production or pod set. Heat-induced damage to floral buds and anthers in the sensitive lines was associated with low sugars levels in peduncles, indicating that heat had greater effects on assimilate demand than on leaf assimilate supply. The heat- tolerant line was the most responsive genotype to elevated [CO2] with respect to pod production under either high or intermediate temperatures. KEYWORDS: ABSCISSION, ACCLIMATION, AIR- TEMPERATURE, CO2, COTTON, HEAT- STRESS, LEAVES, LONG-TERM EXPOSURE, PHOTOSYNTHETIC INHIBITION, REPRODUCTIVE RESPONSES 19 Aikman, D.P. 1996. A procedure for optimizing carbon dioxide enrichment of a glasshouse tomato crop. Journal of Agricultural Engineering Research 63(2):171-183. The procedure consists of two parts. A Gompertz model for the kinetics of fruit growth is used predict the time distribution of photosynthate subsequent harvests. This is combined with predictions of future market prices to compute estimates, one for each day from first anthesis, of a factor to convert CO2 assimilate to expected financial value, based on the worth anticipated from partitioning to fruit. A model of the climate and the crop regime is used to predict temperatures and hence allow for the temperature dependence of fruit growth. The conversion estimates are revised to include the deferred benefit given by additional photosynthesis through increasing early vegetative growth, and hence subsequent photosynthesis and yield. This revision also extends the set of conversion factors to include any period before first anthesis. Given the current environmental variables and conversion factor for that day, a real-time system can use a crop photosynthesis model to predict the cash benefit for any CO2 concentration. The cost of maintaining a concentration can be obtained from a prediction of the ventilation air exchange rate and the unit price of CO2. The CO2 set-point is evaluated as the concentration that maximizes the net profit rate. (C) 1996 Silsoe Research Institute KEYWORDS: CO2- ENRICHMENT, CUCUMBER, FRUIT- GROWTH, GREENHOUSES, LYCOPERSICON-ESCULENTUM MILL, MODEL, PLANTS, TEMPERATURE, VENTILATION 20 Akimoto, M., A. Shirai, K. Ohtaguchi, and K. Koide. 1998. Carbon dioxide fixation and polyunsaturated fatty acid production by the red alga Porphyridium cruentum. Applied Biochemistry and Biotechnology 73(2-3):269-278. Focusing on CO2 fixation, photoautotrophic cultivation of the red alga Porphyridium cruentum was investigated by means of a batch culture under a 5% CO2-enriched atmosphere. The alg-al growth kinetics was successfully described with a logistic model, and simulation of a continuous culture under the optimum growth conditions (30 degrees C, 12 klux and 1.18 g-cells/L) showed that the algal CO2-fixation activity could reach 0.66 g- CO2/(L X d). Under the same growth conditions, eicosapentaenoic acid (20:5 n-3, EPA) and arachidonic acid (20:4 n-6, ARA) yields were similarly calculated to be 3.6 mg-EPA/(L X d) and 6.5 mg-ARA/(L X d), respectively. KEYWORDS: CULTIVATION, GROWTH, LIGHT-INTENSITY, TEMPERATURE 21 Akin, D.E., B.A. Kimball, J.R. Mauney, R.L. Lamorte, G.R. Hendrey, K. Lewin, J. Nagy, and R.N. Gates. 1994. Influence of enhanced co2 concentration and irrigation on sudangrass digestibility. Agricultural and Forest Meteorology 70(1-4):279-287. An experimental line of sudangrass (Sorghum bicolor L. Moench) was included in the free-air CO2 enrichment (FACE) project in 1991 at the University of Arizona Maricopa Agricultural Center to evaluate the effect of ambient (approximately 370 mumol mol- 1) and enriched (550 mumol mol-1) CO2 in well-watered or water- stressed plots. Our specific objective was to determine modifications caused by these environmental effects on the percentages of morphological parts and the fiber components, and on the in vitro digestibility in vegetative and mature harvests. Enrichment with CO2 did not (P > 0.05) change the percentages of morphological parts or fiber components, or the digestibility of any of the morphological components. Protein levels tended to be lower in CO2- enriched plants. However, water-stressed plants tended to have a higher proportion of leaves (blades and sheaths) and a lower proportion of stems, were more digestible, and had lower amounts of anti- quality, aromatic compounds within the plant cell. Stems had the highest digestibility of all morphological components (about 75% in vegetative plants) despite the lowest levels of protein. Stems also showed the greatest changes caused by all treatments, including a 20% decline in digestibility from vegetative to mature samples. The results indicate that enriching CO2 to 550 mumol mol-1 did not reduce digestibility of sudangrass. KEYWORDS: CARBON DIOXIDE, ENRICHMENT 22 Akin, D.E., B.A. Kimball, W.R. Windham, P.J. Pinter, G.W. Wall, R.L. Garcia, R.L. Lamorte, and W.H. Morrison. 1995. Effect of free-air co2 enrichment (face) on forage quality of wheat. Animal Feed Science and Technology 53(1):29-43. Wheat (Triticum aestivum L., cultivar 'Yecora rojo') was grown in ambient (370 mu mol mol(-1)) or enriched (550 mu mol mol(- 1)) concentrations of CO2 in the free-air CO2 enrichment (FACE) project, and components were analyzed for in vitro digestibility, fiber constituents, and crude protein. Four replicated plots of each CO2 treatment were split for irrigation: 'wet' regions received 60 cm of water and 'dry' regions received 30 cm of water through underground tubes. Enriched CO2 concentrations had no effect on in vitro digestion of intact sections of young (26-32-day-old plants) leaf blades except at 24-27 h incubation, at which time enriched leaves were lower in digestibility than control ones. Enriched CO2 concentrations increased the content of acid detergent fiber (ADF) and cellulose of young wet leaves, Sections of main shoots at 26 days tended to have increased digestibility with elevated CO2 levels. Enriched CO2 concentrations did not alter the digestibility of flag leaves from 105-day-old plants or of flag leaves, uppermost stems, and sheaths from plants at full grain maturity, Enriched CO2 levels reduced the acid detergent lignin (ADL) and tended to reduce the protein of leaves from 105-day-old plants. For mature leaf blades, neutral detergent fiber, ADF, and cellulose were, or tended to be, higher while protein content tended to be lower in elevated CO2- grown plants; for both CO2 treatments, 'dry' leaves were higher in digestibility and lower in ADL than 'wet' samples. Mature stems plus sheaths had lower protein contents in plants grown in elevated CO2. Results indicated that enriched CO2 concentrations to 550 mu mol mol(-1) did not substantially alter wheat in vitro digestibility, regardless of irrigation treatment. Elevated CO2 altered fiber components and protein, but these were not consistent among parts and harvests. KEYWORDS: CELL-WALLS, ECOSYSTEMS, MICROSPECTROPHOTOMETRY, PHENOLIC CONSTITUENTS 23 Akin, D.E., L.L. Rigsby, G.R. Gamble, W.H. Morrison, B.A. Kimball, P.J. Pinter, G.W. Wall, R.L. Garcia, and R.L. Lamorte. 1995. Biodegradation of plant-cell walls, wall carbohydrates, and wall aromatics in wheat grown in ambient or enriched co2 concentrations. Journal of the Science of Food and Agriculture 67(3):399-406. Mature internodes from wheat (Triticum aestivum L) grown in control (ambient at c 370 mu mol mol(-1)) or enriched (to 550 mu mol mol(-1)) concentrations of atmospheric CO2 in the free- air CO2 enrichment (FACE) system were analyzed for potential changes in biodegradation of constituents due to predicted increases in atmospheric levels of CO2. The first internodes below the grain were incubated with the lignocellulose- degrading white rot fungus, Phanerochaete chrysosporium K-3, or incubated without microorganisms. Plant samples were then analyzed for dry weight loss, disposition of specific cell types to biodegradation using electron microscopy, carbohydrates and lignin using solid state NMR spectroscopy, and ester- and ether-linked aromatics using gas chromatography. Phanerochaete chrysosporium extensively degraded stems cells (c 75%) and both carbohydrate and aromatic portions of the wheat stems proportionately more carbohydrates were removed by the fungus from the stems. Enriched CO2 did not affect the chemical composition of wheat stems or the biodegradation by P chrysosporium of plant cell walls or wall components for the most part. Data from various methods all indicated that enriched CO2 did not substantially alter the biodegradation of wheat cell wall internodes or wall components. Evidence was not found for an influence on C cycling due to CO2 concentrations in this study. KEYWORDS: ECOSYSTEMS, LIGNINS, PHENOLIC CONSTITUENTS 24 Alagusundaram, K., D.S. Jayas, N.D.G. White, W.E. Muir, and R.N. Sinha. 1995. Controlling cryptolestes-ferrugineus (stephens) adults in wheat stored in bolted-metal bins using elevated carbon- dioxide. Canadian Agricultural Engineering 37(3):217-223. Experiments were conducted in two 5.56 m-diameter farm bins to determine the mortality of caged adult rusty grain beetles, Cryptolestes ferrugineus (Stephens) (Coleoptera: cucujidae), under elevated carbon dioxide (CO2) concentrations. The bins were filled with wheat to a depth of 2.5 m. Dry ice was used to create high CO2 concentrations in the wheat bulks. Two different modes of application of dry ice were used: (i) pellets on the grain surface and in the aeration duct and (ii) pellets on the grain surface and blocks in insulated boxes on the grain surface. The pellets exposed to the ambient conditions on the grain surface and in the aeration duct sublimated quickly and had to be replenished at frequent intervals. Dry ice blocks in insulated boxes, however, maintained high CO2 concentrations without replenishment for over 15 d. In both modes of application, the observed CO2 concentrations in the intergranular gas were about 15% and 30% (all the CO2 concentrations given in this article are on a volume basis) at 2.05 m and 0.55 m above the floor, respectively. At 0.55 m above the floor, the mortality of rusty grain beetle adults was more than 90% while in the top portions of the bulk (2.05 m above the floor) the mortality was only 30%. On an average about two thirds of the insects were killed. The use of controlled atmosphere treatment within an integrated pest management context is outlined. KEYWORDS: INSECTS 25 Alberto, A.M.P., L.H. Ziska, C.R. Cervancia, and P.A. Manalo. 1996. The influence of increasing carbon dioxide and temperature on competitive interactions between a C3 crop, rice (Oryza sativa) and a C-4 weed (Echinochloa glabrescens). Australian Journal of Plant Physiology 23(6):795-802. Many of the most troublesome weeds in agricultural systems are C-4 plants. As atmospheric CO2 increases it is conceivable that competitive ability of these weeds could be reduced relative to C-3 crops such as rice. At the International Rice Research Institute (IRRI) in the Philippines, rice (IR72) and one of its associated C-4 weeds, Echinochloa glabrescens, were grown from seeding to maturity using replacement series mixtures (100:0, 75:25, 50:50, 25:75, and 0:100, % rice:%weed) at two different CO2 concentrations (393 and 594 mu L L(-1)) in naturally sunlit glasshouses. Since increasing CO2 may also result in elevated growth temperatures, the response of rice to each CO2 concentration was also examined at day/night temperatures of 27/21 and 37/29 degrees C. At 27/21 degrees C, increasing the CO2 concentration resulted in a significant increase in above ground biomass (+47%) and seed yield (+55%) of rice when averaged over all mixtures. For E. glabrescens, the C-4 species, no significant effect of CO2 concentration on biomass or yield was observed. When grown in mixture, the proportion of rice biomass increased significantly relative to that of the C- 4 weed at all mixtures at elevated CO2. Evaluation of changes in competitiveness (by calculation of plant relative yield (PRY) and replacement series diagrams) of the two species demonstrated that, at elevated CO2, the competitiveness of rice was increased relative to that of E. glabrescens. However, at the higher growth temperature (37/29 degrees C), growth and reproductive stimulation of rice by elevated CO2 was reduced compared to the lower growth temperature. This resulted in a reduction in the proportion of rice:weed biomass present in all mixtures relative to 27/21 degrees C and a greater reduction in PRY in rice relative to E. glabrescens. Data from this experiment suggest that competitiveness could be enhanced in a C-3 crop (rice) relative to a C-4 weed (E. glabrescens) with elevated CO2 alone, but that simultaneous increases in CO2 and temperature could still favour a C-4 species. KEYWORDS: DRY-MATTER, ENRICHMENT, GROWTH, NITROGEN, PLANTS, WHEAT 26 Alcamo, J., G.J.J. Kreileman, M.S. Krol, and G. Zuidema. 1994. Modeling the global society- biosphere-climate system .1. Model description and testing. Water, Air, and Soil Pollution 76(1-2):1- 35. This paper describes the IMAGE 2.0 model, a multi-disciplinary, integrated model designed to simulate the dynamics of the global society-biosphere-climate system. The objectives of the model are to investigate linkages and feedbacks in the system, and to evaluate consequences of climate policies. Dynamic calculations are performed to year 2100, with a spatial scale ranging from grid (0.5- degrees x 0.5-degrees latitude- longitude) to world regional level, depending on the sub-model. The model consists of three fully linked sub-systems: Energy- Industry, Terrestrial Environment, and Atmosphere-Ocean. The Energy-Industry models compute the emissions of greenhouse gases in 13 world regions as a function of energy consumption and industrial production. End use energy consumption is computed from var-ious economic/demographic driving forces. The Terrestrial Environment models simulate the changes in global land cover on a grid-scale based on climatic and economic factors, and the flux of CO2 and other greenhouse gases from the biosphere to the atmosphere. The Atmosphere-Ocean models compute the buildup of greenhouse gases in the atmosphere and the resulting zonal-average temperature and precipitation patterns. The fully linked model has been tested against data from 1970 to 1990, and after calibration can reproduce the following observed trends: regional energy consumption and energy-related emissions, terrestrial flux of CO2 and emissions of greenhouse gases, concentrations of greenhouse gases in the atmosphere, and transformation of land cover. The model can also simulate long term zonal average surface and vertical temperatures. KEYWORDS: CARBON-CYCLE, CO2, SENSITIVITY 27 Alcamo, J., G.J. Vandenborn, A.F. Bouwman, B.J. Dehaan, K.K. Goldewijk, O. Klepper, J. Krabec, R. Leemans, J.G.J. Olivier, A.M.C. Toet, H.J.M. Devries, and H.J. Vanderwoerd. 1994. Modeling the global society-biosphere-climate system .2. Computed scenarios. Water, Air, and Soil Pollution 76(1-2):37-78. This paper presents scenarios computed with IMAGE 2.0, an integrated model of the global environment and climate change. Results are presented for selected aspects of the society- biosphere- climate system including primary energy consumption, emissions of various greenhouse gases, atmospheric concentrations of gases, temperature, precipitation, land cover and other indicators. Included are a ''Conventional Wisdom'' scenario, and three variations of this scenario: (i) the Conventional Wisdom scenario is a reference case which is partly based on the input assumptions of the IPCC's IS92a scenario; (ii) the ''Biofuel Crops'' scenario assumes that most biofuels will be derived from new cropland; (iii) the ''No Biofuels'' scenario examines the sensitivity of the system to the use of biofuels; and (iv) the ''Ocean Realignment'' scenario investigates the effect of a large-scale change in ocean circulation on the biosphere and climate. Results of the biofuel scenarios illustrate the importance of examining the impact of biofuels on the full range of greenhouse gases, rather than only CO2. These scenarios also indicate possible side effects of the land requirements for energy crops. The Ocean Realignment scenario shows that an unexpected, low probability event can both enhance the build-up of greenhouse gases, and at the same time cause a temporary cooling of surface air temperatures in the Northern Hemisphere. However, warming of the atmosphere is only delayed, not avoided. 28 Allen, D.J., I.F. McKee, P.K. Farage, and N.R. Baker. 1997. Analysis of limitations to CO2 assimilation on exposure of leaves of two Brassica napus cultivars to UV-B. Plant, Cell and Environment 20(5):633-640. Apex and Bristol cultivars of oilseed rape (Brassica napus) were irradiated with 0.63 W m(-2) of UV- B over 5 d. Analyses of the response of net leaf carbon assimilation to intercellular CO2 concentration were used to examine the potential limitations imposed by stomata, carboxylation velocity and capacity for regeneration of ribulose 1,5-bisphosphate on leaf photosynthesis. Simultaneous measurements of chlorophyll fluorescence were used to estimate the maximum quantum efficiency of photosystem II (PSII) photochemistry, the quantum efficiency of linear electron transport at steady-state photosynthesis, and the light and CO2-saturated rate of linear electron transport. Ribulose 1,5-bisphosphate carboxylase/oxygenase (Rubisco) content and activities were assayed in vitro. In both cultivars the UV-B treatment resulted in decreases in the light-saturated rate of CO2 assimilation, which were accompanied by decreases in carboxylation velocity and Rubisco content and activity. No major effects of UV-B were observed on end-product inhibition and stomatal limitation of photosynthesis or the rate of photorespiration relative to CO2 assimilation. In the Bristol cultivar, photoinhibition of PSII and loss of linear electron transport activity were observed when CO2 assimilation was severely inhibited, However, the Apex cultivar exhibited no major inhibition of PSII photochemistry or linear electron transport as the rate of CO2 assimilation decreased. It is concluded that loss of Rubisco is a primary factor in UV-B inhibition of CO2 assimilation. KEYWORDS: ENHANCED RADIATION, HIGHER-PLANTS, ORYZA-SATIVA, PHOTOSYNTHETIC ELECTRON-TRANSPORT, PHOTOSYSTEM, PISUM-SATIVUM, RIBULOSE-1;5-BISPHOSPHATE CARBOXYLASE, ULTRAVIOLET-RADIATION, VIGNA- SINENSIS L 29 Allen, L.H. 1992. Free-air co-2 enrichment field experiments - an historical overview. Critical Reviews in Plant Sciences 11(2-3):121-134. KEYWORDS: CARBON DIOXIDE, CO2, COTTON, CROPS, FUMIGATION, GROWN SOYBEANS, PHOTOSYNTHETIC ACCLIMATION, PLANTS, SULFUR-DIOXIDE, SYSTEM 30 Allen, L.H., E.C. Bisbal, and K.J. Boote. 1998. Nonstructural carbohydrates of soybean plants grown in subambient and superambient levels of CO2. Photosynthesis Research 56(2):143-155. Elevated carbon dioxide (CO2) concentration increases plant photosynthesis, biomass and carbohydrate accumulation. Since plants have grown in low CO2 (200 to 300 mu mol mol(-1)) for the last several million years, how will they use extra photoassimilate as the atmospheric CO(2 )continues to rise? The objectives were to determine the effects of past, present and projected future levels of CO2 on diurnal and seasonal patterns of total nonstructural carbohydrate (TNC) concentration of soybean [Glycine max (L.) Merr.] tissues. Plants were grown at 160, 220, 280, 330, 660 and 990 mu mol mol(-1) CO2 in outdoor, sunlit chambers wherein CO2 uptake rates were measured continuously. Early morning and fate afternoon plant samples were taken at eight dates. The TNC concentration of leaves, petioles and stems increased as CO:! increased. Canopy photosynthetic rates also increased with increasing CO2, apparently without any negative impact of increased leaf TNC. Concentrations of TNC in all vegetative tissues were lower in the morning than the afternoon, which indicates overnight mobilization and utilization of carbohydrates for growth processes. The concentration of TNC was lowest in ail plant components during rapid vegetative growth at Vg to R2 developmental stages. Leaves of all plants, especially those grown in superambient CO2, contained large pools of TNC at plant maturity, which indicated that not all of the reserves were utilized for seed yield. Soybean cultivars for the future should be designed to utilize carbohydrates more readily for seed production so that greater benefit can be realized from rising atmospheric CO2. KEYWORDS: CARBON-DIOXIDE ENRICHMENT, EXPORT, L MERR PLANTS, LEAF, LEAVES, LIGHT, PHOTOSYNTHETIC RESPONSE, REPRODUCTIVE GROWTH, STARCH CONCENTRATION, TEMPERATURE 31 Allen, L.H., E.C. Bisbal, K.J. Boote, and P.H. Jones. 1991. Soybean dry-matter allocation under subambient and superambient levels of carbon-dioxide. Agronomy Journal 83(5):875-883. Rising atmospheric carbon dioxide concentration [CO2] is expected to cause increases in crop growth and yield. The objective of this study was to investigate effects of subambient, as well as superambient, [CO2] on soybean [Glycine max (L.) Merr.] dry matter production and allocation for two reasons: to assess response of plants to prehistoric as well as future expected CO2 levels and to increase confidence in [CO2] response curves by imposing a wide range of [CO2] treatments. Soybean was grown in outdoor, sunlit, controlled- environment chambers at CO2 levels of 160, 220, 280, 330, 660, and 990-mu-mol (CO2) mol-1 (air). Total dry matter growth rates during the linear phase of vegetative growth were 5.0, 8.4, 10.9, 12.5, 18.2, and 20.7 g m-2 d-1 for the above respective [CO2]. Samples taken from 24 to 94 d after planting showed that the percentage of total plant mass in leaf trifoliolates decreased with increasing [CO2] whereas the percentage in structural components (petioles and stems) increased. At final harvest the respective [CO2] treatments resulted in 38, 53, 62, 100, 120, and 92% seed yield with respect to the 330-mu-mol mol-1 treatment. Total dry weight responses were similar. Late season spider mite damage of the 990 and 280-mu-mol mol-1 treatments reduced yields. These data confirm not only that rising CO2 should increase plant growth, but also that plant growth was probably seriously limited by atmospheric [CO2] in preindustrial revolution times back to the previous global glaciation. KEYWORDS: ATMOSPHERIC CO2 ENRICHMENT, CHAMBERS, DEVELOPMENTAL STAGES, PHOTOSYNTHESIS, PLANT GROWTH, TRANSPIRATION RESPONSES, WATER-USE, WEIGHT, YIELD 32 Allen, L.H., B.G. Drake, H.H. Rogers, and J.H. Shinn. 1992. Field techniques for exposure of plants and ecosystems to elevated co-2 and other trace gases. Critical Reviews in Plant Sciences 11(2-3):85-119. KEYWORDS: ATMOSPHERIC CO2 ENRICHMENT, CARBON-DIOXIDE CONCENTRATIONS, ESTUARINE MARSH, OPEN-AIR FUMIGATION, OPEN-TOP CHAMBERS, PORTABLE CHAMBER, SOYBEAN CANOPIES, TRANSPIRATION RESPONSES, VENTILATED CHAMBER, WATER-VAPOR EXCHANGE 33 Allen, L.H., R.R. Valle, J.W. Jones, and P.H. Jones. 1998. Soybean leaf water potential responses to carbon dioxide and drought. Agronomy Journal 90(3):375-383. Rising CO2 can have direct effects on crop water relations and indirect effects on water available for growth. We studied the effects of elevated CO2 and drought on leaf water relations of soybean [Glycine max (L.) Merr. cv. Bragg] and considered the hypothesis of osmotic adjustment mediated by increased photosynthesis (Hypothesis 1) vs. the hypothesis of water conservation mediated by decreased stomatal conductance (Hypothesis 2) to explain improved water relations of plants growing under elevated CO2. In Exp. 1, soybean was grown at 330, 450, 660, and 800 mu mol mol(-1) CO2 in sunlit, closed- circulation, controlled-environment chambers under well-watered conditions. Leaf total water potential (WP), osmotic potential (OP), and turgor potential (TP) were measured at midday during V4 to R6 stages of development. In Exp. 2 (well-watered, R1-R3) and Exp. 3 (13-d drying cycle, R6 seed filling), soybean was grown at 330 and 660 mu mol mol(-1) CO2 and WP, OP, and TP were measured five times per day on sunlit and shaded leaves. In Exp. 3, stomatal conductance (g(s)) and transpiration rate (TR) of leaves were also measured. Experiments 1 and 2 showed that elevated CO2 increased TP and decreased OP, but did not affect leaf WP, thus favoring Hypothesis 1. In Exp. 3, leaf WP was higher in elevated than ambient CO2. Diurnal TP was higher in elevated than ambient CO2 at the beginning of drought, and was maintained longer each day as drought progressed. At the end of drought, TP and WP was higher in elevated than ambient CO2. Elevated CO2 leaves had lower TR because of lower g(s) than ambient CO2 counterparts. Thus, Exp. 3 supported Hypothesis 2, that both stressed and nonstressed plants in elevated CO2 have a better water status (e.g., higher TP) than plants in ambient CO2 due to water conservation mediated by decreased g(s). Remobilization of leaf nutrients during seed filling may limit the capability for osmotic adjustment. Regardless of the mechanisms, growth of plants in elevated CO2 should be less affected by drought than plants in ambient CO2. KEYWORDS: DIFFERENT CO2 ENVIRONMENTS, FIELD, LEAVES, MAIZE, NITROGEN, OSMOTIC ADJUSTMENT, PLANT GROWTH, STRESS, USE EFFICIENCY, YIELD 34 Allen, L.H., R.R. Valle, J.W. Mishoe, and J.W. Jones. 1994. Soybean leaf gas-exchange responses to carbon-dioxide and water-stress. Agronomy Journal 86(4):625-636. As global carbon dioxide concentrations rise, we need to understand the combination of direct effects of this gas and the anticipated effects of climate change, including drought, on physiology and growth of all crops. Effects Of CO2 on plants begin at the leaf level; our objectives, therefore, were to determine interrelationships among factors governing gas exchange responses of soybean [Glycine max (L.) Merr.] leaves to elevated CO2 and water stress. Photosynthetic CO2 assimilation and transpiration rates were measured in cuvettes on leaflets of soybean (cv. Bragg) grown in controlled- environment chambers at 330 and 660 mumol CO2 Mol-1 air. Leaflets at high CO2, either water- stressed or well-watered, had higher photosynthetic and lower transpiration rates, and therefore higher water-use efficiencies (WUE), than those at Control CO2 levels. As irrigation was withheld during an 11-d period, WUE decreased about 30 to 50% with respect to the well- watered treatments. Midday leaf temperature and leaf-to-air vapor pressure gradient levels increased as the water stress progressed. For water stress treatments, midday leaf conductance (G(lw)) was generally higher and residual internal conductance (G(r)) was generally lower in low than in high CO2. Ratios of midday G(r)/G(lc), were nearly constant throughout the period in both the stressed and the well- watered treatments. The ratios of intercellular C(i), to ambient C(a), CO2 concentration (i.e., C(i)/C(a)) during the water stress period remained similar to the respective nonstressed treatments within each CO2 level. These findings support the concept that leaf conductances are governed by CO2 assimilation rates under water-stressed as well as unstressed conditions. KEYWORDS: ABSCISIC- ACID, CARBOXYLASE, DIFFERENT CO2 ENVIRONMENTS, FIELD, GROWTH, LEAVES, PHOTOSYNTHESIS, STOMATAL CONDUCTANCE, TRANSPIRATION RATE, WHEAT 35 Allen, L.H., R.R. Valle, J.W. Mishoe, J.W. Jones, and P.H. Jones. 1990. Soybean leaf gas- exchange responses to CO2 enrichment. Soil and Crop Science Society of Florida Proceedings 49:192-198. 36 Almeida, J.P.F., A. Luscher, M. Frehner, A. Oberson, and J. Nosberger. 1999. Partitioning of P and the activity of root acid phosphatase in white clover (Trifolium repens L.) are modified by increased atmospheric CO2 and P fertilisation. Plant and Soil 210(2):159-166. The growth response of white clover (Trifolium repens L.) to the expected increase in atmospheric partial pressure of CO2 (p(CO2)) may depend on P availability. A decrease in the rate of transpiration due to increased p(CO2) may reduce the amount of P transported to the shoot, thereby causing a change in the partitioning of P between the root and shoot. To test these hypotheses, four concentrations of P in the nutrient solution, combined with two p(CO2) treatments, were applied to nodulated white clover plants. Compared to ambient p(CO2) (35 Pa), twice ambient p(CO2) (70 Pa) reduced the rate of transpiration but did not impair the total P uptake per plant. However, at twice ambient p(CO2) and a moderate to high supply of P, concentrations of structural P and soluble P (Pi) were lower in the leaves and higher in the roots. The activity of root acid phosphatase was lower at twice ambient p(CO2) than at ambient p(CO2); it depended on the Pi concentration in the roots. At the highest P concentration, twice ambient p(CO2) stimulated photosynthesis and the growth rate of the plant without affecting the concentration of nonstructural carbohydrates in the leaves. However, at the lower P concentrations, plants at twice ambient p(CO2) lost their stimulation of photosynthesis in the afternoon, they accumulated nonstructural carbohydrates in the leaves and their growth rate was not stimulated; indicating C-sink limitation of growth. P nutrition will be crucial to the growth of white clover under the expected future conditions of increased p(CO2). KEYWORDS: AIR, BEAN-PLANTS, CARBON DIOXIDE, DRY-MATTER, ENRICHMENT, GROWTH, MAGNESIUM-DEFICIENCY, PHOSPHORUS, SOURCE-SINK RELATIONS, SUBTERRANEUM L 37 Alvarez, R., M. Alconada, and R. Lavado. 1999. Sewage sludge effects on carbon dioxide-carbon production from a desurfaced soil. Communications in Soil Science and Plant Analysis 30(13- 14):1861-1866. Desurfaced soils are found near cities in the Pampean Region of Argentina because A horizons were used for brick production. These soils are not suitable for agriculture. Application of sewage sludge is a tool for improving soil productivity, but its effects on the environment are not thoroughly understood. Production of carbon dioxide (CO2)-carbon (C) in the field from a desurfaced soil in which 25 Mg dry matter ha(-1) of sewage sludge were applied the first year and 10 Mg dry matter ha(-1), the second year was evaluated during a corn (Zea mays L.) growing cycle. Microbial biomass and metabolic activity were also measured. Sludge applications produced an increase of the CO2-C efflux in the field of 30-50% during summer. Microbial biomass was not affected by sludge some months after the application, but metabolic activity and organic matter mineralization were enhanced. The increase of the CO2-C emission from the soil represented 21% of the sludge C applied the year of the experiment and 15% of the C applied the year before. Consequently, an important quantity of the sludge C was retained in the soil. KEYWORDS: CROPS, GLUCOSE, HEAVY-METALS, MAIZE, MANURE, MICROBIAL BIOMASS DYNAMICS, RESIDUE 38 Ambus, P., and G.P. Robertson. 1999. Fluxes of CH4 and N2O in aspen stands grown under ambient and twice-ambient CO2. Plant and Soil 209(1):1-8. Elevated atmospheric CO2 has the potential to change below- ground nutrient cycling and thereby alter the soil-atmosphere exchange of biogenic trace gases. We measured fluxes of CH4 and N2O in trembling aspen (Populus tremuloides Michx.) stands grown in open-top chambers under ambient and twice-ambient CO2 concentrations crossed with `high' and low soil-N conditions. Flux measurements with small static chambers indicated net CH4 oxidation in the open-top chambers. Across dates, CH4 oxidation activity was significantly (P < 0.05) greater with ambient CO2 (8.7 mu g CH4-C m(-2) h(- 1)) than with elevated CO2 (6.5 mu g CH4-C m(-2) h(-1)) in the low N soil. Likewise, across dates and soil N treatments CH4 was oxidized more rapidly (P < 0.05) in chambers with ambient CO2 (9.5 mu g CH4-C m(-2) h(-1)) than in chambers with elevated CO2 (8.8 mu g CH4-C m(-2) h(-1)). Methane oxidation in soils incubated in serum bottles did not show any response to the CO2 treatment. We suggest that the depressed CH4 oxidation under elevated CO2 in the field chambers is due to soil moisture which tended to be higher in the twice-ambient CO2 treatment than in the ambient CO2 treatment. Phase I denitrification (denitrification enzyme activity) was 12-26% greater under elevated CO2 than under ambient CO2 in the `high' N soil; one sampling, however, showed a 39% lower enzyme activity with elevated CO2. In both soil N treatments, denitrification potentials measured after 24 or 48 h were between 11% and 21% greater (P < 0.05) with twice- ambient CO2 than with ambient CO2. Fluxes of N2O in the open- top chambers and in separate 44 cm(2) cores +/- N fertilization were not affected by CO2 treatment and soil N status. Our data show that elevated atmospheric CO2 may have a negative effect on terrestrial CH4 oxidation. The data also indicated temporary greater denitrification with elevated CO2 than with ambient CO2. In contrast, we found no evidence for altered fluxes of N2O in response to increases in atmospheric CO2. KEYWORDS: ATMOSPHERIC METHANE CONSUMPTION, DENITRIFICATION, ELEVATED CARBON-DIOXIDE, ENRICHMENT, GAS FLUXES, GRASSLAND, NITROUS-OXIDE, RESPONSES, TALLGRASS PRAIRIE, TEMPERATE FOREST SOILS 39 Amiro, B.D., J.I. MacPherson, and R.L. Desjardins. 1999. BOREAS flight measurements of forest-fire effects on carbon dioxide and energy fluxes. Agricultural and Forest Meteorology 96(4):199-208. Fire is the dominant stand-replacing agent in the Canadian boreal forest, but few quantitative measurements are available on the micrometeorological effects of fire. Airborne flux measurements during the BOREAS experiment were referenced to age of burn along a 500-km transect through Saskatchewan and Manitoba, Canada. These data for 1-, 5-, and 7-year-old burns were supplemented with 15- and 30-year-old-burn data from the BOREAS northern study site near Thompson, Manitoba. Data were available near midday only and included the June, July and September campaigns during 1994, and July of 1996. Surface radiometric temperature increased by up to 6 degrees C and remained elevated even 15 years after fire. Net radiation was largely unaffected whereas albedo decreased in the first year post-fire but recovered by the fifth year. Sensible heat flux increased by 10- 20% for the first few years after the fire and then decreased. Latent heat flux slightly decreased after the fire, causing the Bowen ratio to increase by ca. 50% for 7 years post-fire. The CO2 flux was reduced for the 15-year period after fire with the greatest reduction to ca. 25% of control areas during the year following fire. However, diurnal and annual data are needed to determine the total impact of fire on the boreal-forest carbon balance. (C) 1999 Elsevier Science B.V. All rights reserved. KEYWORDS: ASPEN FOREST, ATMOSPHERE, ECOSYSTEMS, EXCHANGES, MICROBIAL BIOMASS, NORTHERN, PINE FOREST, SOIL RESPIRATION, WATER-VAPOR, WILDFIRE 40 Amoroso, G., C. Weber, D. Sultemeyer, and H. Fock. 1996. Intracellular carbonic anhydrase activities in Dunaliella tertiolecta (Butcher) and Chlamydomonas reinhardtii (Dangeard) in relation to inorganic carbon concentration during growth: Further evidence for the existence of two distinct carbonic anhydrases associated with the chloroplasts. Planta 199(2):177-184. Using mass-spectrometric measurements of O-18 exchange from (CO2)-C-13-O-18 intracellular carbonic anhydrase (CA) activity was investigated in the unicellular green algae Dunaliella tertiolecta and Chlamydomonas reinhardtii which were either grown on air enriched with 5% CO2 (high-C-i cells) or on air (low-C-i cells). In D. tertiolecta high- and low-C-i cells had detectable levels of internal CA activity when measured under in-vivo conditions and this activity could be split up into three distinct forms. One CA was not associated with the chloroplasts, while two isozymes were found to be located within the plastids. The activities of all intracellular CAs were always about twofold higher in low than in high -C-i cells of D. tertiolecta and the chloroplastic enzymes were completely induced within 4 h of adaptation to air. One of the chloroplastic CAs was found to be soluble the other was insoluble. In addition to the physical differences, MgSO4 in vitro caused a more than twofold stimulation of the soluble activity while the insoluble form of CA remained rather unaffected. In C. reinhardtii, MgSO4 increased the soluble CA activity by 346% and the concentration of MgSO4 required for half-maximum stimulation was between 10 and 15 mM. Again, the insoluble CA activity was not affected by MgSO4. Furthermore, the soluble isoenzyme was considerably more sensitive to ethoxyzolamide, a potent inhibitor of CA, than the insoluble enzyme. The concentration of inhibitor causing 50% inhibition of soluble CA activity was 110 and 85 mu M ethoxyzolamide for D. tertiolecta and C. reinhardtii, respectively. From these data we conclude that the two chloroplast-associated CAs are distinct enzymes. KEYWORDS: CELL-SURFACE, CO2, CYANOBACTERIUM SYNECHOCOCCUS PCC7942, INCREASES, INTACT CHLOROPLASTS, MICROALGAE, O-18 EXCHANGE, PHOTOSYNTHESIS, SALINA, TRANSPORT 41 Amthor, J.S. 1991. Respiration in a future, higher-CO2 world. Plant, Cell and Environment 14(1):13-20. Apart from its impact on global warming, the annually increasing atmospheric [CO2] is of interest to plant scientists primarily because of its direct influence on photosynthesis and photorespiration in C3 species. But in addition, 'dark' respiration, another major component of the carbon budget of higher plants, may be affected by a change in [CO2] independent of an increase in temperature. Literature pertaining to an impact of [CO2] on respiration rate is reviewed. With an increase in [CO2], respiration rate is increased in some cases, but decreased in others. The effects of [CO2] on respiration rate may be direct or indirect. Mechanisms responsible for various observations are proposed. These proposed mechanisms relate to changes in: (1) levels of nonstructural carbohydrates, (2) growth rate and structural phytomass accumulation, (3) composition of phytomass, (4) direct chemical interactions between CO2 and respiratory enzymes, (5) direct chemical interactions between CO2 and other cellular components, (6) dark CO2 fixation rate, and (7) ethylene biosynthesis rate. Because a range of (possibly interactive) effects exist, and present knowledge is limited, the impact of future [CO2] on respiration rate cannot be predicted. Theoretical considerations and types of experiments that can lead to an increase in the understanding of this issue are outlined. KEYWORDS: CARBON DIOXIDE, CO2- ENRICHMENT, DARK RESPIRATION, ENERGY OVERFLOW, GAS-EXCHANGE, GROWTH, LEAF, PHOTOSYNTHESIS, PLANTS, TEMPERATURE 42 Amthor, J.S. 1994. Scaling co2-photosynthesis relationships from the leaf to the canopy. Photosynthesis Research 39(3):321-350. Responses of individual leaves to short-term changes in CO2 partial pressure have been relatively well studied. Whole-plant and plant community responses to elevated CO2 are less well understood and scaling up from leaves to canopies will be complicated if feedbacks at the small scale differ from feedbacks at the large scale. Mathematical models of leaf, canopy, and ecosystem processes are important tools in the study of effects on plants and ecosystems of global environmental change, and in particular increasing atmospheric CO2, and might be used to scale from leaves to canopies. Models are also important in assessing effects of the biosphere on the atmosphere. Presently, multilayer and big leaf models of canopy photosynthesis and energy exchange exist. Big leaf models - which are advocated here as being applicable to the evaluation of impacts of 'global change' on the biosphere - simplify much of the underlying leaf-level physics, physiology, and biochemistry, yet can retain the important features of plant- environment interactions with respect to leaf CO2 exchange processes and are able to make useful, quantitative predictions of canopy and community responses to environmental change. The basis of some big leaf models of photosynthesis, including a new model described herein, is that photosynthetic capacity and activity are scaled vertically within a canopy (by plants themselves) to match approximately the vertical profile of PPFD. The new big leaf model combines physically based models of leaf and canopy level transport processes with a biochemically based model of CO2 assimilation. Predictions made by the model are consistent with canopy CO2 exchange measurements, although a need exists for further testing of this and other canopy physiology models with independent measurements of canopy mass and energy exchange at the time scale of 1 h or less. KEYWORDS: C-3 PLANTS, CARBON DIOXIDE, DARK RESPIRATION, LIGHT-INTENSITY, PHOTOSYNTHETIC CO2 FIXATION, RIBULOSE-1;5-BISPHOSPHATE CARBOXYLASE, SOURCE-SINK RELATIONS, STOMATAL CONDUCTANCE, SUNFLOWER LEAVES, WATER- USE EFFICIENCY 43 Amthor, J.S. 1995. Predicting effects of atmospheric CO2 partial pressure on forest photosynthesis. Journal of Biogeography 22(2-3):269-280. A mechanistic (i.e. hierarchic or explanatory) model of forest canopy mass and energy exchange that has been previously tested with eddy-correlation measurements in the field - albeit only at present ambient CO2 partial pressure - was used to predict photosynthetic response of a deciduous Quercus- Acer forest in eastern North America to atmospheric CO2 partial pressure. Four partial pressures of CO2 were used in simulations: 28 (pre- industrial), 36 (present), 54 and 72 Pa. This is (one of) the first set(s) of predictions of forest photosynthetic response to CO2 partial pressure made by a mechanistic forest physiology model shown to accurately predict independent field measurements of whole-forest CO2 exchange at the hourly time scale. The model includes a biochemically based Farquhar-type model of leaf mesophyll CO2 assimilation, which is central to its ability to predict photosynthetic response to different CO2 partial pressures. Whole-forest photosynthesis was positively related to CO2 partial pressure, as expected. This was the case under both clear and cloudy skies, but the relative response to CO2 was greater under a clear sky compared to a cloudy sky (the clear sky day was also warmer). Instantaneous water use efficiency (mel CO2 assimilated per mol H2O transpired) was positively related to atmospheric CO2 partial pressure for all conditions included in the simulations. Model predictions indicate that (1) present forest photosynthesis and water use efficiency may be significantly greater than they were in pre- industrial times (per unit ground area of forest) and (2) future higher CO2 partial pressures could further stimulate forest photosynthesis and water use efficiency, unless future climatic changes have significant negative effects on photosynthesis or acclimation and adaptation processes markedly downregulate photosynthesis in response to greater CO2 partial pressure. KEYWORDS: C-3 PLANTS, CANOPY, DECIDUOUS FOREST, ELEVATED CO2, GAS- EXCHANGE, GROWTH, MODELS, RIBULOSE-1;5-BISPHOSPHATE CARBOXYLASE, RISING CO2, SOLAR RADIATION 44 Amthor, J.S. 1995. Terrestrial higher-plant response to increasing atmospheric [co2] in relation to the global carbon-cycle. Global Change Biology 1(4):243-274. Terrestrial higher plants exchange large amounts of CO2 with the atmosphere each year; c. 15% of the atmospheric pool of C is assimilated in terrestrial-plant photosynthesis each year, with an about equal amount returned to the atmosphere as CO2 in plant respiration and the decomposition of soil organic matter and plant litter. Any global change in plant C metabolism can potentially affect atmospheric CO2 content during the course of years to decades. In particular, plant responses to the presently increasing atmospheric CO2 concentration might influence the rate of atmospheric CO2 increase through various biotic feedbacks. Climatic changes caused by increasing atmospheric CO2 concentration may modulate plant and ecosystem responses to CO2 concentration. Climatic changes and increases in pollution associated with increasing atmospheric CO2 concentration may be as significant to plant and ecosystem C balance as CO2 concentration itself. Moreover, human activities such as deforestation and livestock grazing can have impacts on the C balance and structure of individual terrestrial ecosystems that far outweigh effects of increasing CO2 concentration and climatic change. In short-term experiments, which in this case means on the order of 10 years or less, elevated atmospheric CO2 concentration affects terrestrial higher plants in several ways. Elevated CO2 can stimulate photosynthesis, but plants may acclimate and (or) adapt to a change in atmospheric CO2 concentration. Acclimation and adaptation of photosynthesis to increasing CO2 concentration is unlikely to be complete, however. Plant water-use efficiency is positively related to CO2 concentration, implying the potential for more plant growth per unit of precipitation or soil moisture with increasing atmospheric CO2 concentration. Plant respiration may be inhibited by elevated CO2 concentration, and although a naive C balance perspective would count this as a benefit to a plant, because respiration is essential for plant growth and health, an inhibition of respiration can be detrimental. The net effect on terrestrial plants of elevated atmospheric CO2 concentration is generally an increase in growth and C accumulation in phytomass. Published estimations, and speculations about, the magnitude of global terrestrial- plant growth responses to increasing atmospheric CO2 concentration range from negligible to fantastic. Well-reasoned analyses point to moderate global plant responses to CO2 concentration. Transfer of C from plants to soils is likely to increase with elevated CO2 concentrations because of greater plant growth, but quantitative effects of those increased inputs to soils on soil C pool sizes are unknown. Whether increases in leaf-level photosynthesis and short-term plant growth stimulations caused by elevated atmospheric CO2 concentration will have, by themselves, significant long-term (tens to hundreds of years) effects on ecosystem C storage and atmospheric CO2 concentration is a matter for speculation, not firm conclusion. Longterm field studies of plant responses to elevated atmospheric CO2 are needed. These will be expensive, difficult, and by definition, results will not be forthcoming for at least decades. Analyses of plants and ecosystems surrounding natural geological CO2 degassing vents may provide the best surrogates for long-term controlled experiments, and therefore the most relevant information pertaining to long-term terrestrial-plant responses to elevated CO2 concentration, but pollutants associated with the vents are a concern in some cases, and quantitative knowledge of the history of atmospheric CO2 concentrations near vents is limited. On the whole, terrestrial higher-plant responses to increasing atmospheric CO2 concentration probably act as negative feedbacks on atmospheric CO2 concentration increases, but they cannot by themselves stop the fossil-fuel-oxidation-driven increase in atmospheric CO2 concentration. And, in the very long-term, atmospheric CO2 concentration is controlled by atmosphere-ocean C equilibrium rather than by terrestrial plant and ecosystem responses to atmospheric CO2 concentration. KEYWORDS: DIOXIDE CONCENTRATION, ELEVATED CO2, GAS-EXCHANGE, PARTIAL- PRESSURE, PAST 2 CENTURIES, PHOTOSYNTHETIC ACCLIMATION, RIBULOSE-1;5- BISPHOSPHATE CARBOXYLASE-OXYGENASE, STOMATAL DENSITY, VOSTOK ICE-CORE, WATER-USE EFFICIENCY 45 Amthor, J.S. 1998. Perspective on the relative insignificance of increasing atmospheric CO2 concentration to crop yield. Field Crops Research 58(2):109-127. Average yield of most crops in many countries increased significantly during the past 50 to 100 years. Although atmospheric CO2 concentration, [CO2](a), also increased during that time period, and although crop growth and yield can respond positively to [CO2](a) increase, yield increases were due mainly to factors other than increasing [CO2](a). Similarly, some yield increases prior to 1900 were also associated primarily with factors other than changes in [CO2](a). In particular, past national average yield increases were the result chiefly of technological advances such as nitrogen fertilization; selection of genotypes with increased harvest index and disease resistance; mechanization of planting, cultivation, and harvesting; and chemical weed and pest control. If technology continues to increase average yields at recent rates, near-future increases in [CO2](a) will have only small impacts on yield in comparison to technology in many countries. Conversely, if future increases in [CO2](a) are the main drivers of future yield increases, those yield increases will be small. These points are demonstrated through a comparison of (i) long-term records of yield, (ii) data from key controlled- [CO2] experiments, and (iii) records of past [CO2](a). Finally, it is noted that continued [CO2](a) increase may bring with it climatic changes that could have negative or positive impacts on future yield. (C) 1998 Elsevier Science B.V. All rights reserved. KEYWORDS: 18TH-CENTURY ENGLAND, AGRICULTURAL PRODUCTIVITY, CARBON DIOXIDE, CLIMATE, ENRICHMENT, PHOTOSYNTHESIS, RESPONSES, SPECULATIONS, TRENDS, WHEAT 46 Amthor, J.S., R.J. Mitchell, G.B. Runion, H.H. Rogers, S.A. Prior, and C.W. Wood. 1994. Energy content, construction cost and phytomass accumulation of glycine-max (L) merr and sorghum-bicolor (L) moench grown in elevated co2 in the field. New Phytologist 128(3):443-450. Grain sorghum [Sorghum] bicolor (L.) Moench, a C-4 crop] and soybean [Glycine max (L.) Merr. cv. Stonewall, a C-3 crop] plants were grown in ambient (c. 360 mu l l(-1)) and twice- ambient (c. 720 mu l l(-1)) CO2 levels in open-top chambers in soil without root constriction. Plant dry mass, energy content, composition and construction cost (i.e. amount of carbohydrate required to synthesize a unit of plant dry mass) were assessed at the end of the growing season. Elevated CO2 (a) increased phytomass accumulation (kg per plant) in both species, (b) had little affect on energy concentration (MJ kg(-1) plant) but caused large increases in the amount of plant energy per ground area (MJ m(-2) ground), and (c) did not alter specific growth cost (kg carbohydrate kg(-1) plant growth) but greatly increased growth cost per ground area (kg carbohydrate m(-2) ground) because growth was enhanced. For soybean, twice-ambient CO2 resulted in a 50 % increase in the amount of nitrogen and energy in grain (seed plus pod) per ground area. This response to elevated CO2 has important implications for agricultural productivity during the next century because the rate of human population growth is exceeding the rate of increase of land used for agriculture so that future food demands can only be met by greater production per ground area. KEYWORDS: CARBON DIOXIDE, ENRICHMENT, LEAVES, MAINTENANCE, NITROGEN, PHOTOSYNTHESIS, PLANTS, RESPIRATION, RESPONSES, YIELD 47 Andalo, C., B. Godelle, M. Lefranc, M. Mousseau, and I. TillBottraud. 1996. Elevated CO2 decreases seed germination in Arabidopsis thaliana. Global Change Biology 2(2):129-135. The impact of elevated [CO2] on seed germination was studied in different genotypes of Arabidopsis thaliana from natural populations. Two generations of seeds were studied: the maternal generation was produced in the greenhouse (present-day conditions), the offspring generation was produced in two chambers where the CO2 concentration was either the present atmospheric concentration (about 350 ppm) or elevated (700 ppm). The seeds were tested for proportion of germinated seeds and mean germination time in both chambers to study the impact of elevated [CO2] during seed production and germination. Elevated [CO2] during maturation of seeds on the mother-plants decreased the proportion of germinated seeds, while elevated [CO2] during germination had no effect on the proportion of germinated seeds. However, when seeds were both produced and germinated under elevated [CO2] (situation expected by the end of next century), germination was slow and low. Moreover, the effect of the [CO2] treatment differs among genotypes of Arabidopsis: there is a strong treatment x genotype interaction. This means that there is ample genetic variance for a selective response modiying the effects of high levels of [COP] in natural populations of Arabidopsis thaliana. The outcome at the community level will depend on what seeds are available, when they germinate and the resulting competition following germination. KEYWORDS: GROWTH, PLANTS 48 Andalo, C., C. Raquin, N. Machon, B. Godelle, and M. Mousseau. 1998. Direct and maternal effects of elevated CO2 on early root growth of germinating Arabidopsis thaliana seedlings. Annals of Botany 81(3):405-411. Individuals of Arabidopsis thaliana, collected in different natural populations, were grown in controlled and elevated CO2 in a glasshouse. Following germination, root growth of progeny of different lines of these populations was studied in control and elevated atmospheric CO2. No significant direct effect of atmospheric CO2 concentration could be demonstrated on root growth. An important parental effect was apparent, namely that root length and branching were decreased in seeds collected from a mother plant which had been grown in elevated CO2. This was correlated with smaller seeds, containing less nitrogen. These parental effects were genetically variable. We conclude that CO2 may affect plant fitness via parental effects on seed size and early root growth and that the genetic variability shown in our study demonstrates that Arabidopsis populations will evolve in the face of this new selective pressure. (C) 1998 Annals of Botany Company. KEYWORDS: ATMOSPHERIC CARBON-DIOXIDE, ELONGATION, ENRICHMENT, PLANT, QUALITY, SEED-GERMINATION, SENECIO-VULGARIS, TEMPERATURE, TRITICUM- AESTIVUM L, WHEAT 49 Anderson, J.M. 1991. The effects of climate change on decomposition processes in grassland and coniferous forests. Ecological Applications 1(3):326-347. Current models of climate change predict a reduction of area covered by northern coniferous forests and tundra, and an increase in grasslands. These scenarios also indicate a northerly shift in agricultural regions, bringing virgin soils under cultivation. The direct effects of man on tundra,boreal forest, and temperate grassland ecosystems are likely to result in less carbon mobilization from soils and vegetation than from tropical forests. However, as a consequence of climate change, carbon mineralization rates from arctic and sub-arctic soils could be very rapid under warmer and drier conditions because of low stabilization of soil organic matter (SOM) and enhanced microbial responses to small changes in soil moisture and temperature. Predicting the response of these systems to climate change is complicated where the edaphic environment regulating SOM dynamics is not a direct function of macroclimatic conditions. Grasslands contain a greater proportion of highly stabilized SOM than coniferous forests, distributed over greater depth in the soil profile, which is less susceptible to changes in mineralization rates. It is concluded that short-term responses of soil processes to climate change are more predictable in well-drained grassland and forest soils than in waterlogged soils of the tundra and boreal region. Over longer periods of time, however, plant species and soil types will alter in response to new temperature and moisture regimes above- and belowground interacting with the effects of carbon enrichment and changes in nutrient availability. The dynamics of these plant-soil interactions and the future status of soils in different life zones as sources or sinks of carbon is poorly understood. More data are also needed on the distribution of waterlogged forest soils in the boreal zone and responses to warming, which include the production of methane as well as CO2. The primary recommendation for future research is for integrated studies on plant and soil processes. KEYWORDS: ATMOSPHERIC CARBON-DIOXIDE, DOUGLAS-FIR, LEAF-LITTER DECOMPOSITION, LONG-TERM DECOMPOSITION, NITROGEN-AVAILABILITY, NORTHERN HARDWOODS, SCOTS PINE FOREST, SOIL ORGANIC MATTER, SPRUCE-LICHEN WOODLAND, TEMPERATE ECOSYSTEMS 50 Anderson, P.D., and P.T. Tomlinson. 1998. Ontogeny affects response of northern red oak seedlings to elevated CO2 and water stress - I. Carbon assimilation and biomass production. New Phytologist 140(3):477-491. The interactive influences of elevated carbon dioxide, water stress, and ontogeny on carbon assimilation and biomass production were investigated in northern red oak, a species having episodic shoot growth characteristics. Seedlings were grown from acorns through three shoot-growth hushes (8-11 wk) in controlled-environment chambers at 400, 530 or 700 mu mol mol(-1) CO2 and under well watered or water-stressed soil- moisture regimes. Increasing CO2 growth concentration from 400 to 700 mu mol mol(-1) resulted in a 34 % increase in net assimilation rate (A), a 31 % decrease in stomatal conductance to water vapour (g(s)) and a 141 % increase in water use efficiency (WUE) in well watered seedlings. In contrast, water- stressed seedlings grown at 700 mu mol mol(-1) CO2 demonstrated a 69 % increase in A, a 23 % decrease in g(s), and a 104 % increase in WUE. However, physiological responses to increased CO2 and water stress were strongly modified by ontogeny. During active third-flush shoot growth, A in first-flush and second- flush foliage of water- stressed seedlings increased relative to the quiescent phase following cessation of second-flush growth by an average of 115 %; g(s) increased by an average of 74 %. In contrast, neither A nor gs in comparable foliage of well watered seedlings changed in response to active third-flush growth. Whereas seedling growth was continuous through three flushes in well watered seedlings, growth of water-stressed seedlings was minimal following the leaf-expansion stage of the third flush. Through three growth flushes total seedling biomass and biomass allocation to root, shoot and foliage components were very similar in water-stressed seedlings grown at 700 mu mol mol(-1) CO2 and well watered seedlings grown at 400 mu mol mol(-1) CO2. Enhancement effects of elevated CO2 on seedling carbon (C) assimilation and biomass production may offset the negative impact of moderate water stress and are likely to be determined by ontogeny and stress impacts on carbon sink demand. KEYWORDS: ATMOSPHERIC CO2, DIOXIDE ENRICHMENT, DROUGHT, GAS-EXCHANGE, GROWTH-RESPONSE, LEAVES, PHOTOSYSTEM, QUERCUS-RUBRA L, RISING CO2, USE EFFICIENCY 51 Andersson, N.E. 1991. The influence of constant and diurnally changing CO2 concentrations on plant-growth and development. Journal of Horticultural Science 66(5):569-574. Plants of Ficus benjamina and miniature rose (Rosa hybrida cv. Red Minimo) were grown under four CO2 treatments. Two had constant CO2 levels (600 and 900 ppm) and the other two had diurnal changes in CO2 levels, one increasing from 600 to 1500 ppm and one decreasing from 1500 to 600 ppm, each in four steps of 300 ppm during the day-time. In all treatments 900 ppm CO2 was maintained during the night when supplementary light was used, except in the treatment with constant 600 ppm where 600 ppm was also continued throughout the night. Plant growth was monitored under both decreasing and increasing natural daylength and irradiance. The tallest plants and greatest increment in height for Ficus occurred with plants grown under constant CO2 concentration at 600 ppm and also with increasing CO2 concentration. In both experiments the dry weight per pot was lowest when plants were grown under a constant CO2 concentration at 900 ppm. In both experiments with miniature roses the number of flower buds was significantly increased under diurnally changing CO2 concentration or when the CO2 level was constant at 600 ppm compared with a constant 900 ppm. Time to flowering was decreased by constant CO2 at 900 ppm as compared with the other treatments. KEYWORDS: ATMOSPHERES, CARBON-DIOXIDE ENRICHMENT, DURATION, EXCHANGE, LIGHT-INTENSITY, ROSE, YIELD 52 Andrade, J.L., and P.S. Nobel. 1996. Habitat, CO2 uptake and growth for the CAM epiphytic cactus Epiphyllum phyllanthus in a Panamanian tropical forest. Journal of Tropical Ecology 12:291- 306. In the tropical forest of Barro Colorado Island, habitat characteristics, diel acidity changes, CO2 uptake and growth were investigated for the epiphytic cactus Epiphyllum phyllanthus (L.) Haw. It occurred most frequently in tree cavities with its roots in canopy soil and was especially abundant on two tree species: Platypodium elegans J. Vogel and Tabebuia guayacan (Seem.) Hemsl. Its maximum net CO2 uptake rates were low under natural conditions (1.4 mu mol m(-2) s(- 1)) but were comparable to those of other CAM and C-3 epiphytes under wet conditions in a screenhouse. Under both natural conditions and in the screenhouse, partial shade enhanced growth and CAM activity. When plants grew under a photosynthetic photon flux of c. 4 mol m(-2) d(-1), their nocturnal acidity increase and total net CO2 uptake were twice as much as for plants growing at lower (an average of 2.4 mol m(-2) d(-1)) and higher (7.7 mol m(-2) d(-1)) photosynthetic photon fluxes. Stem elongation was 27% greater at the intermediate photosynthetic photon flux. Seedlings of E. phyllanthus survived three months of drought and responded rapidly to rewetting, recovering fully within three days. Transpiration rates and nocturnal acidity increases also recovered to the values of well-watered plants a few days after rewetting, indicating that this species can take advantage of episodic rainfall during the dry season. KEYWORDS: ACCUMULATION, C-3 BROMELIADS, COMPARATIVE ECOPHYSIOLOGY, CRASSULACEAN ACID METABOLISM, LEAF, OPUNTIA FICUS INDICA, SHADE, VASCULAR EPIPHYTES 53 Andre, M., and H. Ducloux. 1993. Interaction of co2 enrichment and water limitations on photosynthesis and water efficiency in wheat. Plant Physiology and Biochemistry 31(1):103-112. Wheat plants (Triticum aestivum L. cv. Capitole) were grown in twin closed growth chambers with continuous monitoring of CO2 and water exchanges. During the vegetative stage the effect Of CO2 enrichment, from 330 to 660 mul-1, was studied under irradiance of 660 muE m-2 s-1 with an optimum watering. Comparisons were made with successive experiments in which daily water supply was fixed to a fraction (0.62-0.50-0.25) of the maximal transpiration of previous experiments. In a well- watered canopy, doubling CO2 decreased transpiration by only 8%. Water use efficiency was increased (factor 1.45) mainly by the stimulation of photosynthesis. Under restricted water supply, photosynthesis of plants was more limited than transpiration. The inhibition of photosynthesis and the increase of water use efficiency can be predicted by a simple diffusion model applied to the response curve of photosynthesis to CO2, measured on canopy in standard conditions of watering. The main hypothesis is that the equivalent stomatal conductance is reduced proportionally to the water availability, without closure by patching. Under enriched CO2, the same reduction of leaf surface by water limitation was observed. Photosynthesis was less affected. Therefore, water-use-efficiency was again increased. Doubling CO2 concentration can compensate for water stress inhibition on CO2 assimilation. That model also predicts interactions of CO2 and water stress observed on water-use- efficiency which was increased by a factor up to 5 in comparison with well-watered plants in standard atmosphere. The implications of this study for global change models are discussed. KEYWORDS: ASSIMILATION, CARBON-DIOXIDE ENRICHMENT, CONDUCTANCE, EXCHANGES, GROWTH, PHASEOLUS-VULGARIS L, PLANTS, SEEDLINGS, STRESS, YIELD 54 Andrews, J.A., K.G. Harrison, R. Matamala, and W.H. Schlesinger. 1999. Separation of root respiration from total soil respiration using carbon-13 labeling during Free-Air Carbon Dioxide Enrichment (FACE). Soil Science Society of America Journal 63(5):1429-1435. Soil respiration constitutes a major component of the global carbon cycle and is likely to be altered by climatic change. However, there is an incomplete understanding of the extent to which various processes contribute to total soil respiration, especially the contributions of root and rhizosphere respiration. Here, using a stable carbon isotope tracer, we separate thf relative contributions of root and soil heterotrophic respiration to total soil respiration in situ. The Free-Air Carbon dioxide Enrichment (FACE) facility in the Duke University Forest (NC) fumigates plots of an undisturbed loblolly pine (Pinus taeda L.) forest with CO2 that is strongly depleted in C-13. This labeled CO2 is found in the soil pore space through live root and mycorrhizal respiration and soil heterotroph respiration of labile root exudates. By measuring the depletion of (CO2)-C-13 in the soil system, we found that the rhizosphere contribution to soil CO2 reflected the distribution of fine roots in the soil and that late in the growing season roots contributed 55% of total soil respiration at the surface, This estimate may represent an upper limit on the contribution of roots to soil respiration because high atmospheric CO2 often increases in root density and/or root activity in the soil. KEYWORDS: CO2, DECIDUOUS FOREST, FATE, FLUXES, LITTER, ORGANIC-MATTER, PONDEROSA PINE, RHIZOSPHERE, SEEDLINGS 55 Andrews, T.J., G.S. Hudson, C.J. Mate, S. Voncaemmerer, J.R. Evans, and Y.B.C. Arvidsson. 1995. Rubisco - the consequences of altering its expression and activation in transgenic plants. Journal of Experimental Botany 46:1293-1300. Transgenic tobacco (Nicotiana tabacum W38) hemizygous for a single antisense gene directed against Rubisco's small subunit had 35% of the Rubisco content of control leaves (15% when homozygous). CO2 assimilation (at 1000 mu mol quanta m(-2) s(- 1) and 350 mu bar CO2) by the hemizygous leaves was reduced to 40% of that of the controls without material effect on stomatal conductance, chlorophyll content or other photosynthetic components. Leaf soluble protein was reduced commensurately with the reduction in Rubisco. CO2 assimilation rate in the hemizygous leaves remained limited by Rubisco activity at all, even very high, CO2 concentrations. This led to a simple, hyperbolic response of photosynthesis to intraplastid CO2 concentration from which the in vivo catalytic properties of Rubisco were inferred and compared with those of isolated Rubisco in vitro. Using a similar approach, the content of Rubisco activase was suppressed by incorporating a partial cDNA for activase into the tobacco genome in the antisense orientation with respect to a cauliflower mosaic virus 35S promoter. The progeny of a primary transformant with two anti- activase inserts had from <1% to 20% of the activase content of control plants. Quite severe suppression of activase, to less than 5% of the amount present in control leaves, was required before effects on photosynthesis and growth became apparent, indicating that one activase tetramer must be able to service, continuously, as many as 200 Rubisco octamers. Plants with lower activase contents could not grow unless the atmosphere was enriched with CO2. Their Rubisco was less carbamylated and they had lower CO2 assimilation rates than the controls. The rate of release of 2'-carboxyarabinitol-1- phosphate from Rubisco after illumination of the anti-activase leaves was also impaired. Older anti- activase plants accumulated increasing amounts of Rubisco in their younger leaves, but were unable to carbamylate it. The photosynthetic rate per carbamylated Rubisco active site in the strongly suppressed anti-activase leaves was only approximately 25% of that seen in control leaves, suggesting that activase may not only promote carbamylation of uncarbamylated Rubisco sites, but also accelerate turnover at carbamylated sites. KEYWORDS: 2-CARBOXYARABINITOL 1-PHOSPHATE, ACTIVITY INVIVO, ANTISENSE GENE, CATALYSIS, GROWTH, PHOTOSYNTHESIS, RIBULOSE BISPHOSPHATE CARBOXYLASE, RIBULOSE-1;5-BISPHOSPHATE CARBOXYLASE-OXYGENASE, SLOW INACTIVATION, TOBACCO NICOTIANA-TABACUM 56 Andriolo, J.L., J. LeBot, C. Gary, G. Sappe, P. Orlando, B. Brunel, and C. Sarrouy. 1996. An experimental set-up to study carbon, water and nitrate uptake rates by hydroponically grown plants. Journal of Plant Nutrition 19(10-11):1441-1462. The experimental system described allows concomitant hourly measurements of CO2, H2O, and NO3 uptake rates by plants grown hydroponically in a greenhouse. Plants are enclosed in an airtight chamber through which air flows at a controlled speed. Carbon dioxide exchange and transpiration rates are determined from respective differences of concentrations of CO2 and water vapor of the air at the system inlet and outlet. This set-up is based on the ''open-system'' principle with improvements made on existing systems. For instance, propeller anemometers are used to monitor air flow rates in the chamber. From their signal it is possible to continuously adjust air speed to changing environmental conditions and plant activity. The air temperature inside the system therefore never rises above that outside. Water and NO3 uptake rates are calculated at time intervals from changes in the volume and the NO3 concentration of the nutrient solution in contact with the roots. The precise measurement of the volume of solution is achieved using a balance which has a higher precision than any liquid level sensors. Nitrate concentration is determined in the laboratory from aliquots of solution sampled at time intervals. A number of test runs are reported which validate the measurements and confirm undisturbed conditions within the system. Results of typical diurnal changes in CO2, H2O, and NO3 uptake rates by fruiting tomato plants are also presented. KEYWORDS: CROP, DIOXIDE, ELEVATED CO2, GAS-EXCHANGE, LIFE-CYCLE, OPEN-TOP CHAMBERS, PHOTOSYNTHESIS, SYSTEM, TOMATO, TRANSPIRATION 57 Angell, R., and T. Svejcar. 1999. A chamber design for measuring net CO2 exchange on rangeland. Journal of Range Management 52(1):27-31. Net carbon exchange of terrestrial ecosystems will likely change as atmospheric CO2 concentration increases, Currently, little is known of the annual dynamics or magnitude of CO2 flux on many native and agricultural ecosystems. Remoteness of many ecosystems has limited our ability to measure CO2 flux on undisturbed vegetation. Today, many plant ecologists have portable photosynthesis systems with which they make single- leaf photosynthesis measurements. Utility of this equipment is enhanced when canopy-level CO2 flux is also measured. We designed a portable 1-m(3) closed chamber for use in measuring CO2 exchange in short statured vegetation with widely varied canopy structure. The design includes external ductwork equipped with doors which are used to open the chamber for ventilation with outside air between measurements. The chamber was tested on a Wyoming big sagebrush (Artemisia tridentata ssp. Wyomingensis Nutt.)/Thurber's needlegrass (Stipa thurberiana Piper) community using 10 plots equally divided between shrub and interspace, The ductwork and doors provided adequate ventilation to allow consecutive measurements of CO2 nux without removing the chamber from the plot. The chamber could differentiate CO2 flux between plots with sagebrush and those with grass only, even at relatively low fluxes, Net CO2 uptake per unit ground area was greater (P = 0.04) on sagebrush-grass plots (7.6 +/- 1.4 mu mol m(-2) s(-1)) than on interspace plots without sagebrush (3.1 +/- 1.0 pmol m(-2) s(- 1)). Chamber and leaf temperature increased by an average of 0.5 and 1.2 degrees C, respectively, during measurements. KEYWORDS: CARBON-CYCLE, FLUXES, SYSTEM, TUNDRA ECOSYSTEMS 58 Apel, P., and M. Peisker. 1995. Variability of photosynthetic gas exchange parameters, dark respiration, and stomatal numbers in species of Polygonum. Physiologia Plantarum 95(3):365-372. Within the genus Polygonum a large variation was found between species with regard to stomatal number, gas phase resistance, intracellular resistance and dark respiration. Interspecific variation in CO2 compensation concentration and intercellular CO2 concentration at constant external concentration were comparatively small. Correlations were found between stomatal number and gas phase resistance, stomatal number and Gamma, and Gamma and the product of dark respiration rate and intracellular resistance. The influence of dark respiration and stomatal number on photosynthetic gas exchange is discussed. It was concluded that dark respiration in light was enhanced by 22% as a mean value in 9 Polygonum species and by 62% in Polygonum lapathifolium. KEYWORDS: CONDUCTANCE, LEAVES, PLANTS 59 Apple, M.E., M.S. Lucash, D.M. Olszyk, and D.T. Tingey. 1998. Morphogenesis of Douglas-fir buds is altered at elevated temperature but not at elevated CO2. Environmental and Experimental Botany 40(2):159-172. Global climatic change as expressed by increased CO2 and temperature has the potential for dramatic effects on trees. To determine what its effects may be on Pacific Northwest forests, Douglas-fir (Pseudotsuga menziesii) seedlings were grown in sun-lit controlled environment chambers at ambient or elevated (+ 4 degrees C above ambient) temperature, and at ambient or elevated (+ 200 ppm above ambient) CO2. In 1995-1996 and 1996- 1997, elevated CO2 had no effect on vegetative bud morphology, while the following unusual morphological characteristics were found with greater frequency at elevated temperature than at ambient: rosetted buds with reflexed and loosened outer scales, convoluted inner scales, clusters of small buds, needles elongating between scales, needle primordia with white, hyaline apical extensions, and buds with hardened scales inside of unbroken buds. Buds became rosetted in elevated temperature chambers after temperatures exceeded 40 degrees C in July, 1996. Rosettes were induced within 48-h in buds placed in a 40 degrees C oven; fewer rosettes formed at 20 degrees C. Induction was reversible in buds transferred from 40 to 20 degrees C, implying that resetting is a physical rather than a growth phenomenon. It appears that rosettes form after long- term exposure to elevated temperature and after shorter periods of exposure to intense heat. Elevated temperature influences bud morphology and may therefore influence the overall branching structure of Douglas-fir seedlings. (C) 1998 Elsevier Science B.V. All rights reserved. KEYWORDS: ACCUMULATION, BUDBURST, CHILLING REQUIREMENT, DORMANCY, FROST DAMAGE, HEAT-SHOCK PROTEINS, INTERIOR, POPULATIONS, SEEDLINGS, SHOOT 60 Arakelyan, V.V., G.B. Ibragimova, and Y.S. Nasyrov. 1993. Effects of light, co2, and temperature on carbonic-anhydrase activity in C3-plants. Russian Journal of Plant Physiology 40(6):759-767. Carbonic anhydrase activity was studied in cotton (Gossypium hirsutum L.) and Triticale plants exposed to various light intensities, temperatures, and CO2 concentrations in the air. The activity was measured using an original method based on the HCO3- dehydration reaction, which is carried out in conditions resembling those occurring in the chloroplast stroma in vivo. Carbonic anhydrase activity in stromal fractions from cotton and triticale plant chloroplasts appears to respond to environmental changes. Plant exposure to increased light intensities and temperatures results in increased activity, whereas high ambient CO2 concentrations lower carbonic anhydrase activity. After examining in vitro the HCO3- dehydration reaction, which in vivo is catalyzed by carbonic anhydrase, we concluded that the physiological role of the stromal enzyme consists of preventing local CO2 depletion in the carboxylation sites. Thus, high temperatures and low ambient CO2 concentrations enhance carbonic anhydrase activity, while impeding CO2 transport from the air to the carboxylation sites in the leaf. This accelerates HCO3- dehydration and reduces its concentration in the stroma, thereby producing an additional driving force for HCO3- transport to the chloroplast. 61 Archer, S., D.S. Schimel, and E.A. Holland. 1995. Mechanisms of shrubland expansion - land-use, climate or co-2. Climatic Change 29(1):91-99. Encroachment of trees and shrubs into grasslands and the 'thicketization' of savannas has occurred worldwide over the past century. These changes in vegetation structure are potentially relevant to climatic change as they may be indicative of historical shifts in climate and as they may influence biophysical aspects of land surface-atmosphere interactions and alter carbon and nitro en cycles. Traditional explanations offered to account for the historic displacement of grasses by woody plants in many arid and semi-arid ecosystems have centered around changes in climatic, livestock grazing and fire regimes. More recently, it has been suggested that the increase in atmospheric CO2 since the industrial revolution has been the driving force. In this paper we evaluate the CO2 enrichment hypotheses and argue that historic, positive correlations between woody plant expansion and atmospheric CO2 are not cause and effect. KEYWORDS: AMERICAN SOUTHWEST, ATMOSPHERIC CO2, CARBON DIOXIDE, DESERTIFICATION, ECOSYSTEMS, ELEVATED CO2, GROWTH, INCREASING CO2, NATURAL VEGETATION, PAST 2 CENTURIES 62 Arienzo, M., G. Basile, R. Dandria, V. Magliulo, and A. Zena. 1995. Irrigation with carbonated water and nutrient availability - tests on strawberry plants. Agrochimica 39(1):61-72. A research was carried out to study the nutrient availability and yield performances of a strawberry crop cv. 'Chandler' in response to equivalent depths (100% of ETM) of CO2 enriched water and plain water applied with different irrigation frequencies. Plots were arranged in a complete randomized block design replicated four times, using mulch and a drip irrigation system adopting 4 1/h emitters. The crop was covered by a plastic tunnel following treatment differentiation. The statistical analysis revealed an increased availability of Cu, Zn, Ca, Mg, and Mn for the CO2 treatment, probably linked with the pH reduction (from 7,5 to 6,5). The increased nutrient uptake in the CO2 enriched water treatment may be the cause of the commercial yield enhancement (8,6 %) and reduction in the weight of deformed berries (-12,1 %). KEYWORDS: DIOXIDE 63 Arisi, A.C.M., G. Cornic, L. Jouanin, and C.H. Foyer. 1998. Overexpression of iron superoxide dismutase in transformed poplar modifies the regulation of photosynthesis at low CO2 partial pressures or following exposure to the prooxidant herbicide methyl viologen. Plant Physiology 117(2):565-574. Chloroplast-targeted overexpression of an Fe superoxide dismutase (SOD) from Arabidopsis thaliana resulted in substantially increased foliar SOD activities. Ascorbate peroxidase, glutathione reductase, and monodehydroascorbate reductase activities were similar in the leaves from all of the lines, but dehydroascorbate reductase activity was increased in the leaves of the FeSOD transformants relative to untransformed controls. Foliar H2O2, ascorbate, and glutathione contents were comparable in all lines of plants. Irradiance-dependent changes in net CO, assimilation and chlorophyll a fluorescence quenching parameters were similar in all lines both in air (21% O-2) and at low (1%) O-2. CO2- response curves for photosynthesis showed similar net CO2-exchange characteristics in all lines. In contrast, values of photochemical quenching declined in leaves from untransformed controls at intercellular CO2 (Ci) values below 200 mu L L-1 but remained constant with decreasing Ci in leaves of FeSOD transformants. When the O-2 concentration was decreased from 21 to 1%, the effect of FeSOD overexpression on photochemical quenching at limiting Ci was abolished. At high light (1000 mu mol m(-2) s(-1)) a progressive decrease in the ratio of variable (F-v) to maximal (F-m) fluorescence was observed with decreasing temperature. At 6 degrees C the high-light-induced decrease in the F-v/F-m ratio was partially prevented by low O-2 but values were comparable in all lines. Methyl viologen caused decreased F- v/F-m ratios, but this was less marked in the FeSOD transformants than in the untransformed controls. These observations suggest that the rate of superoxide dismutation limits flux through the Mehler-peroxidase cycle in certain conditions. KEYWORDS: ASCORBATE PEROXIDASE, CHLOROPHYLL FLUORESCENCE, ELEVATED LEVELS, HYDROGEN- PEROXIDE, OXIDATIVE STRESS TOLERANCE, PHOTOINHIBITION, QUANTUM YIELD, SPINACH-CHLOROPLASTS, TEMPERATURE, TRANSGENIC PLANTS 64 Arnone, J.A. 1997. Indices of plant N availability in an alpine grassland under elevated atmospheric CO2. Plant and Soil 190(1):61-66. The objective of this study was to estimate whether elevated atmospheric [CO2] alters plant N availability in a native high- elevation grassland in the Swiss Alps using two integrative, relatively non-disruptive methods. Estimates based on seasonal net plant N uptake, and those based on the amounts of NH4+-N plus NO3--N captured by ion exchange resin (IER) bags, did not differ in plots treated with ambient (355 mu L L-1) and elevated (680 mu L L-1) [CO2] in either the second (1993) or third (1994) growing season under treatment with elevated [CO2]. The results of this study suggest that the effects of rising atmospheric [CO2] on plant N availability may be negligible in this grassland. The results also contrast the relatively large effects of elevated atmospheric [CO2] (increases and decreases) reported for highly disturbed artificial systems. KEYWORDS: CARBON DIOXIDE, COMMUNITIES, EXCHANGE, FEEDBACK, GROWTH FORMS, NITROGEN-AVAILABILITY, NUTRIENT AVAILABILITY, RESPONSES, SOIL- NITROGEN, TUNDRA 65 Arnone, J.A. 1997. Temporal responses of community fine root populations to long- term elevated atmospheric CO2 and soil nutrient patches in model tropical ecosystems. Acta Oecologica- International Journal of Ecology 18(3):367-376. Biomass and length density of fine roots, as well as overall allocation of dry matter to root growth, of C-3 plants has been shown to increase under elevated CO2. However, it is uncertain whether the stimulatory effect of elevated CO2 on fine root population size in plant communities will persist, or whether fine root populations at high CO2 simply reach their maximum sooner (or possibly later) than those produced under ambient CO2. It is also unclear whether increased nutrient demand at the stand- level under elevated CO2 will lead to more intense nutrient foraging via enhanced fine root proliferation into relatively nutrient-rich soil microsites. I addressed these questions in a 530 day experiment with model tropical plant communities established in four equivalent ecosystem (17 m(3)) in which plants shared a common low fertility soil. Fine root (less than or equal to 2 mm empty set) populations (biomass and length density) in ecosystems maintained at elevated CO2 (610 mu l l(-1)) increased more rapidly than those in ecosystems maintained at ambient CO2 (340 mu l l(-1)) during the first half of the experiment and also remained greater over the entire experiment. The data also indicate that: (1) fine root populations at both CO2 levels eventually stabilize, (2) stabilization occurs sooner under elevated CO2 (occupation of the soil volume), and (3) steady-state populations under elevated CO2 may be slightly larger than those maintained under ambient CO2. Fine root proliferation into artifically nutrient- enriched microsites was dramatic in all ecosystems (22% to 75% greater than into non-enriched soil). However, proliferation into enriched microsites was not enhanced by elevated CO2. Thus, elevated CO2 may not enhance exploitation of nutrient- rich microsites even in low fertility soils, suggesting that increased plant nutrient capture under elevated CO2 also may be unlikely. KEYWORDS: AMAZONIAN FORESTS, BIOMASS, CARBON DIOXIDE, ENRICHMENT, GROWTH, MICROSITES, PLANT- COMMUNITIES, PROLIFERATION, RAIN-FOREST, UPTAKE KINETICS 66 Arnone, J.A. 1999. Symbiotic N-2 fixation in a high Alpine grassland: effects of four growing seasons of elevated CO2. Functional Ecology 13(3):383-387. 1. Increasing carbon dioxide concentration (E: 680 mu l CO2 litre(-1) vs ambient, A: 355 mu l CO2 litre(-1)) around late- successional Alpine sedge communities of the Swiss Central Alps (2450 m) for four growing seasons (1992-1995) had no detectable effect on symbiotic N-2 fixation in Trifolium alpinum-the sole N-2-fixing plant species in these communities (74 +/- 30 mg N m(-2) year(-1), A and E plots pooled). 2. This result is based on data collected in the fourth growing season showing that elevated CO2 had no effect on Trifolium above-ground biomass (4.4 +/- 1.7 g m(-2), A and E plots pooled, n = 24) or N content per unit land area (124 +/- 51 mg N m(-2), A and E pooled), or on the percentage of N Trifolium derived from the atmosphere through symbiotic N-2 fixation (%Ndfa: 61.0 +/- 4.1 across A and E plots) estimated using the N-15 dilution method. 3. Thus, it appears that N inputs to this ecosystem via symbiotic N-2 fixation will not be dramatically affected in the foreseeable future even as atmospheric CO2 continues to rise. KEYWORDS: ATMOSPHERIC CO2, ECOSYSTEM, ENRICHMENT, GAS-EXCHANGE, NITROGENASE ACTIVITY, NODULATION, REDUCTION, RESPONSES, TRIFOLIUM-REPENS L, TUNDRA 67 Arnone, J.A., and P.J. Bohlen. 1998. Stimulated N2O flux from intact grassland monoliths after two growing seasons under elevated atmospheric CO2. Oecologia 116(3):331-335. Long-term exposure of native vegetation to elevated atmospheric CO2 concentrations is expected to increase C inputs to the soil and, in ecosystems with seasonally dry periods, to increase soil moisture. We tested the hypothesis that these indirect effects of elevated CO2 (600 mu l l(-1) vs 350 mu l l(-1)) would improve conditions for microbial activity and stimulate emissions of nitrous oxide (N2O), a very potent and long-lived greenhouse gas. After two growing seasons, the mean N2O efflux from monoliths of calcareous grassland maintained at elevated CO2 was twice as high as that measured from monoliths maintained at current ambient CO2 (70 +/- 9 vs 37 +/- 4 mu g N2O m(-2) h(-1) in October, 27 +/- 5 vs 13 +/- 3 mu g N2O m(-2) h(-1) in November after aboveground harvest). The higher N2O emission rates at elevated CO2 were associated with increases in soil moisture, soil heterotrophic respiration, and plant biomass production, but appear to be mainly attributable to higher soil moisture. Our results suggest that rising atmospheric CO2 may contribute more to the total greenhouse effect than is currently estimated because of its plant- mediated effects on soil processes which may ultimately lead to increased N2O emissions from native grasslands. KEYWORDS: CALCAREOUS GRASSLAND, CARBON-DIOXIDE ENRICHMENT, DENITRIFICATION, ECOSYSTEMS, INCREASE, METHANE, NITROUS-OXIDE PRODUCTION, SHORTGRASS STEPPE, SOIL-NITROGEN, STOMATAL RESPONSES 68 Arnone, J.A., and J.C. Gordon. 1990. Effect of nodulation, nitrogen-fixation and CO2 enrichment on the physiology, growth and dry mass allocation of seedlings of alnus-rubra bong. New Phytologist 116(1):55-66. 69 Arnone, J.A., and G. Hirschel. 1997. Does fertilizer application alter the effects of elevated CO2 on Carex leaf litter quality and in situ decomposition in an alpine grassland? Acta Oecologica- International Journal of Ecology 18(3):201-206. The purpose of our investigation was to determine: (1) whether fertilization with NPK would result in an improvement in leaf litter quality of the dominant species (Carer curvula) in a high alpine grassland in Switzerland; and especially (2) if fertilization improves the quality of leaf litter produced under elevated atmospheric CO2 and compensates for the suppressive effects of high CO2 on the in situ decomposition rates of C. curvula litter, observed at this site in an earlier study. Fertilizer application (40 k(g) N ha(-1) yr(-1)) resulted in 34% higher leaf litter [N] but did not change C:N or lignin N ratios, when viewed across both CO2 treatments. Improvement in the mean N quality of litter produced under elevated CO2 resulting from fertilization appeared to lead to a significantly faster mean decomposition rate (+ 60%), but fertilization had no significant effect on decomposition of litter produced under ambient CO2. We conclude that the potential stimulatory effect of an increase in atmospheric N deposition on litter quality and decomposition rates may partially compensate for the inhibitory effects of rising atmospheric CO2 in these high alpine grassland ecosystems. KEYWORDS: ECOSYSTEMS, NITROGEN, RESPONSES 70 Arnone, J.A., and C. Kestenholz. 1997. Root competition and elevated CO2: Effects on seedling growth in Linum usitatissimum populations and Linum Silene cretica mixtures. Functional Ecology 11(2):209-214. 1. Root competition can be an important determinant of the performance of neighbours within plant populations and communities. Because plants often maintain larger root systems and allocate more of their carbon to root systems under elevated atmospheric CO2 than they do at lower CO2 concentrations, root-root interactions could play an increasingly important role in determining competitive outcomes among individuals and plant species as global CO2 concentration continues to rise. 2. We established 12 pure stands of Linum usitatissimum (flax) and 12 mixed stands of Linum and its naturally co-occurring weed species Silene cretica in opaque plastic trays each filled with the same amount of nutrient-rich soil mix. In half of the trays from each of these stand types, vertical waterproof partitions separated the root systems of individual plants from each other to prevent root competition, while in the other half no partitions were present. Half of the trays from all treatments were allowed to grow under low atmospheric CO2 concentration (320 mu ll(-1)) and the other half under elevated CO2 (600 mu ll(-1)), in daylight growth: chambers for 30 days from seedling emergence until harvest in mid-June. All trays received equal amounts of water so that soils in the low CO2 treatment were maintained at field capacity. 3. Our results indicate that under high soil fertilities: (1) intra-specific root-root interactions alone play a relatively insignificant role in determining plant biomass production within pure Linum populations and (2) the impact of an aggressive species (Silene) on co-occurring less aggressive species (Linum) becomes more severe under elevated CO2 as a result of amplified interspecific root competition. KEYWORDS: AMBIENT, ANNUALS, ATMOSPHERIC CO2, C-3, CARBON DIOXIDE, COMMUNITIES, ECOSYSTEMS, ENRICHMENT, PLANTS, RESPONSES 71 Arnone, J.A., and C. Korner. 1993. Influence of elevated co2 on canopy development and red - far- red ratios in 2-storied stands of ricinus-communis. Oecologia 94(4):510-515. 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 of Ricinus communis exposed to ambient (340 mul-1) and elevated (610 mul-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. Col 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. KEYWORDS: ATMOSPHERIC CO2, CARBON-DIOXIDE ENRICHMENT, CHLOROPHYLL CONTENT, CLOVER TRIFOLIUM-REPENS, GROWTH, LEAF ANATOMY, LIGHT-QUALITY, PLANTS, RAIN-FOREST, RESPONSES 72 Arnone, J.A., and C. Korner. 1995. Soil and biomass carbon pools in model communities of tropical plants under elevated co2. Oecologia 104(1):61-71. The experimental data presented here relate to the question of whether terrestrial ecosystems will sequester more C in their soils, litter and biomass as atmospheric CO2 concentrations rise. Similar to our previous study with relatively fertile growth conditions (Korner and Arnone 1992), we constructed four rather nutrient-limited model communities of moist tropical plant species in greenhouses (approximately 7 m(2) each). Plant communities were composed of seven species (77 individuals per community) representing major taxonomic groups and various life forms found in the moist tropics. Two ecosystems were exposed to 340 mu l CO2 l(-1) and two to 610 mu l l(-1) for 530 days of humid tropical growth conditions. In order to permit precise determination of C deposition in the soil, plant communities were initially established in C-free unwashed quartz sand. Soils were then amended with known amounts of organic matter (containing C and nutrients). Mineral nutrients were also supplied over the course of the experiment as timed-release full-balance fertilizer pellets. Soils represented by far the largest repositories for fixed C in all ecosystems. Almost 5 times more C (ca. 80% of net C fixation) was sequestered in the soil than in the biomass, but this did not differ between CO2 treatments. In addition, at the whole-ecosystem level we found a remarkably small and statistically non-significant increase in C sequestration (+4%; the sum of C accretion in the soil, biomass, litter and necromass). Total community biomass more than quadrupled during the experiment, but at harvest was, on average, only 8% greater (i.e. 6% per year; n.s.) under elevated CO2, mainly due to increased root biomass (+15%, P = 0.12). Time courses of leaf area index of all ecosystems suggested that canopy expansion was approaching steady state by the time systems were harvested. Net primary productivity (NPP) of all ecosystems - i.e. annual accumulation of biomass, necromass, and leaf litter (but not plant-derived soil organic matter) - averaged 815 and 910 g m(-2) year(-1) at ambient and elevated CO2, respectively. These NPPs are remarkably similar to those of many natural moist tropical forested ecosystems. At the same time net productivity of soil organic matter reached 7000 g dry matter equivalent per m(2) and year (i.e. 3500 g C m(-2) year(-1)). Very slight yet statistically significant CO2- induced shifts in the abundance of groups of species occurred by the end of the experiment, with one group of species (Elettaria cardamomum, Ficus benjamina, F: pumila, Epipremnum pinnatum) gaining slightly, and another group (Ctenanthe lubbersiana, Heliconia humilis, Cecropia peltata) losing. Our results show that: (1) enormous amounts of C can be deposited in the ground which are normally not accounted for in estimates of NPP and net ecosystem productivity; (2) any enhancement of C sequestration under elevated atmospheric CO2 may be substantially smaller than is believed will occur (yet still very important), especially under growth conditions which permit close to natural NPP; and (3) species dominance in plant communities is likely to change under elevated CO2, but that changes may occur rather slowly. KEYWORDS: AMBIENT, ATMOSPHERIC CO2, C-3, COMPETITION, DIOXIDE, ENRICHMENT, ESTUARINE MARSH, GROWTH, NITROGEN, TUSSOCK TUNDRA 73 Arnone, J.A., J.G. Zaller, C. Ziegler, H. Zandt, and C. Korner. 1995. Leaf quality and insect herbivory in model tropical plant- communities after long-term exposure to elevated atmospheric co2. Oecologia 104(1):72-78. Results from laboratory feeding experiments have shown that elevated atmospheric carbon dioxide can affect interactions between plants and insect herbivores, primarily through changes in leaf nutritional quality occurring at elevated CO2. Very few data are available on insect herbivory in plant communities where insects can choose among species and positions in the canopy in which to feed. Our objectives were to determine the extent to which CO2-induced changes in plant communities and leaf nutritional quality may affect herbivory at the level of the entire canopy. We introduced equivalent populations of fourth instar Spodoptera eridania, a lepidopteran generalist, to complex model ecosystems containing seven species of moist tropical plants maintained under low mineral nutrient supply. Larvae were allowed to feed freely for 14 days, by which time they had reached the seventh instar. Prior to larval introductions, plant communities had been continuously exposed to either 340 mu l CO2 l(-1) or to 610 mu l CO2 l(-1) for 1.5 years. No major shifts in leaf nutritional quality [concentrations of N, total non-structural carbohydrates (TNC), sugar, and starch; ratios of: C/N, TNC/N, sugar/N, starch/N; leaf toughness] were observed between CO2 treatments for any of the species. Furthermore, no correlations were observed between these measures of leaf quality and leaf biomass consumption. Total leaf area and biomass of all plant communities were similar when caterpillars were introduced. However, leaf biomass of some species was slightly greater - and for other species slightly less (e.g. Cecropia peltata) - in communities exposed to elevated CO2. Larvae showed the strongest preferenc