The Effect of Atmospheric Carbon Dioxide Elevation on Plant Growth in Freshwater Ecosystems

We developed a dynamic model to investigate the effect of atmospheric carbon dioxide (CO₂) increase on plant growth in freshwater ecosystems. Steady-state simulations were performed to analyze the response of phytoplankton and submerged macrophytes to atmospheric CO₂ elevation from 350 to 700 ppm. We studied various conditions that may affect this response, such as alkalinity, the air–water exchange rate of CO₂, the community respiration rate, and the phosphorus (P) supply rate. The increase in atmospheric CO₂ could affect submerged plant growth only under relatively eutrophic conditions and at a low community respiration rate. Alkalinity had little effect on the response of the different species. When the air–water exchange was low, the proportional effect of the CO₂ increase on plant growth was higher. Under eutrophic conditions, algae and macrophytes using CO₂ and HCO₃^– may double their growth rate due to atmospheric CO₂ elevation, while the growth of macrophytes restricted to CO₂ assimilation may be threefold. The differences in response of the species under various conditions indicate that the elevation of atmospheric CO₂ may induce drastic changes in the productivity and species dominance in freshwater systems.     

Carbon fixation and carbon availability in marine phytoplankton

It is widely believed that inorganic C does not limit the rate of short-term photosynthesis, the net productivity, or the maximum biomass, of marine phytoplankton. This lack of inorganic C restriction is less widely believed to hold for phytoplankton in many low alkalinity freshwaters or for seaweed in nutrient-enriched rock pools. These views are examined in the context of the physical chemistry of the inorganic C system in natural waters and of the ways in which various taxa of phytoplankton deal with inorganic C and discriminate between ¹²C and ¹³C. Using this information to interpret data obtained in the ocean or in freshwater suggests that short-term photosynthesis, production rate, and achieved biomass, of phytoplankton are rarely limited by inorganic C supply but, rather, that the widely suggested factors of limited light, nitrogen or phosphorus supply are the resource inputs which restrict productivity. Global change, by increasing atmospheric CO₂ partial pressure and global mean temperatures, is likely to increase the mean CO₂ concentration in the atmosphere, but the corresponding change in the oceans will be much less. There are, however, genotypic differences in the handling of inorganic C among the diversity of marine phytoplankton, and in impact on use of limiting nutrients, so increases in the mean CO₂ and HCO₃ ^- concentrations in surface ocean waters could cause changes in species composition. However, the rarity of inorganic C limitation of marine phytoplankton short-term photosynthesis, net productivity, or the maximum biomass, in today's ocean means that global change is unlikely to increase these three values in the ocean.     

Rising CO2 Levels Will Intensify Phytoplankton Blooms in Eutrophic and Hypertrophic Lakes

Harmful algal blooms threaten the water quality of many eutrophic and hypertrophic lakes and cause severe ecological and economic damage worldwide. Dense blooms often deplete the dissolved CO₂ concentration and raise pH. Yet, quantitative prediction of the feedbacks between phytoplankton growth, CO₂ drawdown and the inorganic carbon chemistry of aquatic ecosystems has received surprisingly little attention. Here, we develop a mathematical model to predict dynamic changes in dissolved inorganic carbon (DIC), pH and alkalinity during phytoplankton bloom development. We tested the model in chemostat experiments with the freshwater cyanobacterium Microcystis aeruginosa at different CO₂ levels. The experiments showed that dense blooms sequestered large amounts of atmospheric CO₂, not only by their own biomass production but also by inducing a high pH and alkalinity that enhanced the capacity for DIC storage in the system. We used the model to explore how phytoplankton blooms of eutrophic waters will respond to rising CO₂ levels. The model predicts that (1) dense phytoplankton blooms in low- and moderately alkaline waters can deplete the dissolved CO₂ concentration to limiting levels and raise the pH over a relatively wide range of atmospheric CO₂ conditions, (2) rising atmospheric CO₂ levels will enhance phytoplankton blooms in low- and moderately alkaline waters with high nutrient loads, and (3) above some threshold, rising atmospheric CO₂ will alleviate phytoplankton blooms from carbon limitation, resulting in less intense CO₂ depletion and a lesser increase in pH. Sensitivity analysis indicated that the model predictions were qualitatively robust. Quantitatively, the predictions were sensitive to variation in lake depth, DIC input and CO₂ gas transfer across the air-water interface, but relatively robust to variation in the carbon uptake mechanisms of phytoplankton. In total, these findings warn that rising CO₂ levels may result in a marked intensification of phytoplankton blooms in eutrophic and hypertrophic waters.     

Physiological and ecological controls on carbon sequestering in terrestrial ecosystems

Carbon cycling processes in ecosystems are generally believed to be well understood. Carbon, hydrogen, oxygen and other essential elements are chemically converted from inorganic to organic compounds primarily in the process of photosynthesis. Secondary metabolic processes cycle carbon in and among organisms and carbon is ultimately released back to the environment as CO₂ by respiratory processes. Unfortunately, our understanding of this cycle was determined under the assumption that the primary inorganic form of C (CO₂ in the atmosphere) was relatively constant. With the emerging concensus that atmospheric carbon concentration is increasing, we must now reassess our understanding of the carbon cycle. How will plants, animals and decomposers respond to a doubling of carbon supply? Will biological productivity be accelerated? If plant productivity increases will a predictable percentage of the increase be accumulated as increased standing crop? Or, is it possible that doubling the availability of CO₂ will increase metabolic activity at all trophic levels resulting in no net increase in system standing crop? The purpose of this paper is to review evidence for physiological and growth responses of plants to carbon dioxide enhancement. Essentially no research has been completed on the ecological aspects of these questions. From this review, I conclude that accurate predictions of future ecosystem responses to increasing atmospheric carbon dioxide concentration are not possible without additional understanding of physiological and ecological mechanisms.     

Scaling up evolutionary responses to elevated CO2: lessons from Arabidopsis

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

Physiology of inorganic C acquisition and implications for resource use efficiency by marine phytoplankton: relation to increased CO2 and temperature

Abstract. Photosynthesis by many marine phytoplankton algae is saturated by the inorganic C concentration in air-equilibrated sea water. These organisms appear to use an active inorganic C transport process (CO₂-concentrating mechanism) which increases the CO₂ concentration around rubisco and saturates this enzyme with CO₂ and suppresses its oxygenase activity. A minority of marine phytoplankton algae have photosynthetic characteristics more suggestive of diffusive CO₂ entry; the inorganic C concentration present in sea water does not saturate photosynthesis by these organisms. Theoretical considerations, tested when possible against observation, suggest that the organisms with a CO₂-concentrating mechanism could have a lower cost of photons, nitrogen, iron, manganese and molybdenum to achieve a given rate of carbon accumulation by the cells than is the case for the organisms with diffusive CO₂ entry. Zinc and selenium costs may show the reverse effect. The increased sea-surface inorganic C, and CO₂ concentrations which will result from anthropogenic increases in atmospheric CO₂ content are predicted to increase the rate of photosynthesis, and of growth when other resources are abundant, and to reduce, or reverse, the higher resource (photons, nitrogen, iron, manganese and molybdenum) cost of a given rate of CO₂ assimilation in organisms with CO₂ diffusion relative to those which have CO₂ concentrating mechanisms and do not repress them at higher inorganic C concentrations. These effects may well alter species composition, and overall resource cost of growth, of phytoplankton; any influence that these effects may have on CO₂ removal from the atmosphere are severely constrained by other trophic levels and, especially, oceanic circulation patterns. Changed sea-surface temperatures are unlikely to qualitatively alter these conclusions.     

Net primary and ecosystem production and carbon stocks of terrestrial ecosystems and their responses to climate change

Evaluating the role of terrestrial ecosystems in the global carbon cycle requires a detailed understanding of carbon exchange between vegetation, soil, and the atmosphere. Global climatic change may modify the net carbon balance of terrestrial ecosystems, causing feedbacks on atmospheric CO₂ and climate. We describe a model for investigating terrestrial carbon exchange and its response to climatic variation based on the processes of plant photosynthesis, carbon allocation, litter production, and soil organic carbon decomposition. The model is used to produce geographical patterns of net primary production (NPP), carbon stocks in vegetation and soils, and the seasonal variations in net ecosystem production (NEP) under both contemporary and future climates. For contemporary climate, the estimated global NPP is 57.0 Gt C y^–1, carbon stocks in vegetation and soils are 640 Gt C and 1358 Gt C, respectively, and NEP varies from –0.5 Gt C in October to 1.6 Gt C in July. For a doubled atmospheric CO₂ concentration and the corresponding climate, we predict that global NPP will rise to 69.6 Gt C y^–1, carbon stocks in vegetation and soils will increase by, respectively, 133 Gt C and 160 Gt C, and the seasonal amplitude of NEP will increase by 76%. A doubling of atmospheric CO₂ without climate change may enhance NPP by 25% and result in a substantial increase in carbon stocks in vegetation and soils. Climate change without CO₂ elevation will reduce the global NPP and soil carbon stocks, but leads to an increase in vegetation carbon because of a forest extension and NPP enhancement in the north. By combining the effects of CO₂ doubling, climate change, and the consequent redistribution of vegetation, we predict a strong enhancement in NPP and carbon stocks of terrestrial ecosystems. This study simulates the possible variation in the carbon exchange at equilibrium state. We anticipate to investigate the dynamic responses in the carbon exchange to atmospheric CO₂ elevation and climate change in the past and future.     

Terrestrial higher-plant response to increasing atmospheric [CO2] in relation to the global carbon cycle

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. Long-term 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.     

Oceanic sinks for atmospheric CO2

There is approximately 50 times more inorganic carbon in the global ocean than in the atmosphere. On time scales of decades to millions of years, the interaction between these two geophysical fluids determines atmospheric CO₂ levels. During glacial periods, for example, the ocean serves as the major sink for atmospheric CO₂, while during glacial–interglacial transitions, it is a source of CO₂ to the atmosphere. The mechanisms responsible for determining the sign of the net exchange of CO₂ between the ocean and the atmosphere remain unresolved. There is evidence that during glacial periods, phytoplankton primary productivity increased, leading to an enhanced sedimentation of particulate organic carbon into the ocean interior. The stimulation of primary production in glacial episodes can be correlated with increased inputs of nutrients limiting productivity, especially aeolian iron. Iron directly enhances primary production in high nutrient (nitrate and phosphate) regions of the ocean, of which the Southern Ocean is the most important. This trace element can also enhance nitrogen fixation, and thereby indirectly stimulate primary production throughout the low nutrient regions of the central ocean basins. While the export flux of organic carbon to the ocean interior was enhanced during glacial periods, this process does not fully account for the sequestration of atmospheric CO₂. Heterotrophic oxidation of the newly formed organic carbon, forming weak acids, would have hydrolyzed CaCO₃ in the sediments, increasing thereby oceanic alkalinity which, in turn, would have promoted the drawdown of atmospheric CO₂. This latter mechanism is consistent with the stable carbon isotope pattern derived from air trapped in ice cores. The oceans have also played a major role as a sink for up to 30% of the anthropogenic CO₂ produced during the industrial revolution. In large part this is due to CO₂ solution in the surface ocean; however, some, poorly quantified fraction is a result of increased new production due to anthropogenic inputs of combined N, P and Fe. Based on ‘circulation as usual’, models predict that future anthropogenic CO₂ inputs to the atmosphere will, in part, continue to be sequestered in the ocean. Human intervention (large-scale Fe fertilization; direct CO₂ burial in the deep ocean) could increase carbon sequestration in the oceans, but could also result in unpredicted environmental perturbations. Changes in the oceanic thermohaline circulation as a result of global climate change would greatly alter the predictions of C sequestration that are possible on a ‘circulation as usual’ basis.     

Increased leaf reflectance in tropical trees under elevated CO2

Globally increasing atmospheric CO₂ concentrations are known to affect many aspects of plant physiology and development; however, little attention has been given to leaf and canopy optical properties. Three tropical trees in the Leguminosae, an important canopy tree family in many tropical forests, responded similarly to an experimental doubling of CO₂ partial pressure with a 9–23% increase in spectral leaf reflectance to light in the visible (400–700 nm) waveband. Decreased leaf chlorophyll content under elevated CO₂ may explain part of the observed increase in reflectance. However, analyses that statistically corrected for chlorophyll content effects on reflectance still indicated a significant CO₂ effect. This results, in conjunction with the spectral pattern of the response, suggests that the primary mechanism is increased optical masking of chlorophyll under elevated CO₂. The magnitude of the increase in leaf reflectance is sufficient to suggest that increased canopy reflectance of tropical forests (and possibly other terrestrial ecosystems) may be an important negative feedback in the response of global net radiative climate forcing to increasing atmospheric CO₂.     

Ecophysiological responses of the larch species in northern Japan to environmental changes as a basis for afforestation

For sustainable use and suitable management of larch plantations, we must clarify the ecophysiological responses of larch species to environmental changes. The physical environment has been changing dramatically, e.g., increase in atmospheric CO₂ concentration ([CO₂]), nitrogen (N) deposition, and atmospheric ozone concentration ([O₃]), and these changes may negatively affect growth of larch species. This review summarizes the previous experimental studies on the ecophysiological responses of larch species to elevated [CO₂], soil acidification, elevated [O₃], and N load. Based on the advanced studies, although elevated [CO₂] will stimulate the productivity of larch, increase of [O₃] and severe soil acidification will reduce it. Increase of N deposition, at least, will not negatively affect larch productivity. Finally, we propose the future direction for investigation to understand the mechanism of the responses of larch species and to predict the associated risk.     

Three Phases of Plant Response to Atmospheric CO(2) Enrichment

Several years of research on seven different plants (five terrestrial and two aquatic species) suggest that the beneficial effects of atmospheric CO₂ enrichment may be divided into three distinct growth response phases. First is a well-watered optimum-growth-rate phase where a 300 parts per million increase in the CO₂ content of the air generally increases plant productivity by approximately 30%. Next comes a nonlethal water-stressed phase where the same increase in atmospheric CO₂ is more than half again as effective in increasing plant productivity. Finally, there is a water-stressed phase normally indicative of impending death, where atmospheric CO₂ enrichment may actually prevent plants from succumbing to the rigors of the environment and enable them to maintain essential life processes, as life ebbs from corresponding ambient-treatment plants.     

The impact of elevated CO2 on yield loss from a C3 and C4 weed in field-grown soybean

Soybean (Glycine max) was grown at ambient and enhanced carbon dioxide (CO₂, + 250 μL L^?1 above ambient) with and without the presence of a C3 weed (lambsquarters, Chenopodium album L.) and a C4 weed (redroot pigweed, Amaranthus retroflexus L.), in order to evaluate the impact of rising atmospheric carbon dioxide concentration [CO₂] on crop production losses due to weeds. Weeds of a given species were sown at a density of two per metre of row. A significant reduction in soybean seed yield was observed with either weed species relative to the weed‐free control at either [CO₂]. However, for lambsquarters the reduction in soybean seed yield relative to the weed‐free condition increased from 28 to 39% as CO₂ increased, with a 65% increase in the average dry weight of lambsquarters at enhanced [CO₂]. Conversely, for pigweed, soybean seed yield losses diminished with increasing [CO₂] from 45 to 30%, with no change in the average dry weight of pigweed. In a weed‐free environment, elevated [CO₂] resulted in a significant increase in vegetative dry weight and seed yield at maturity for soybean (33 and 24%, respectively) compared to the ambient CO₂ condition. Interestingly, the presence of either weed negated the ability of soybean to respond either vegetatively or reproductively to enhanced [CO₂]. Results from this experiment suggest: (i) that rising [CO₂] could alter current yield losses associated with competition from weeds; and (ii) that weed control will be crucial in realizing any potential increase in economic yield of agronomic crops such as soybean as atmospheric [CO₂] increases.     

Long-term culture at elevated atmospheric CO2 fails to evoke specific adaptation in seven freshwater phytoplankton species

The concentration of CO₂ in the atmosphere is expected to double by the end of the century. Experiments have shown that this will have important effects on the physiology and ecology of photosynthetic organisms, but it is still unclear if elevated CO₂ will elicit an evolutionary response in primary producers that causes changes in physiological and ecological attributes. In this study, we cultured lines of seven species of freshwater phytoplankton from three major groups at current (approx. 380 ppm CO₂) and predicted future conditions (1000 ppm CO₂) for over 750 generations. We grew the phytoplankton under three culture regimes: nutrient-replete liquid medium, nutrient-poor liquid medium and solid agar medium. We then performed reciprocal transplant assays to test for specific adaptation to elevated CO₂ in these lines. We found no evidence for evolutionary change. We conclude that the physiology of carbon utilization may be conserved in natural freshwater phytoplankton communities experiencing rising atmospheric CO₂ levels, without substantial evolutionary change.     

Responses of CAM species to increasing atmospheric CO2 concentrations

Crassulacean acid metabolism (CAM) species show an average increase in biomass productivity of 35% in response to a doubled atmospheric CO₂ concentration. Daily net CO₂ uptake is similarly enhanced, reflecting in part an increase in chlorenchyma thickness and accompanied by an even greater increase in water‐use efficiency. The responses of net CO₂ uptake in CAM species to increasing atmospheric CO₂ concentrations are similar to those for C₃ species and much greater than those for C₄ species. Increases in net daily CO₂ uptake by CAM plants under elevated atmospheric CO₂ concentrations reflect increases in both Rubisco‐mediated daytime CO₂ uptake and phosphoenolpyruvate carboxylase (PEPCase)‐mediated night‐time CO₂ uptake, the latter resulting in increased nocturnal malate accumulation. Chlorophyll contents and the activities of Rubisco and PEPCase decrease under elevated atmospheric CO₂, but the activated percentage for Rubisco increases and the K_M(HCO₃ ^? ) for PEPCase decreases, resulting in more efficient photosynthesis. Increases in root:shoot ratios and the formation of additional photosynthetic organs, together with increases in sucrose‐Pi synthase and starch synthase activity in these organs under elevated atmospheric CO₂ concentrations, decrease the potential feedback inhibition of photosynthesis. Longer‐term studies for several CAM species show no downward acclimatization of photosynthesis in response to elevated atmospheric CO₂ concentrations. With increasing temperature and drought duration, the percentage enhancement of daily net CO₂ uptake caused by elevated atmospheric CO₂ concentrations increases. Thus net CO₂ uptake, productivity, and the potential area for cultivation of CAM species will be enhanced by the increasing atmospheric CO₂ concentrations and the increasing temperatures associated with global climate change.     

Laboratory incubations reveal potential responses of soil nitrogen cycling to changes in soil C and N availability in Mojave Desert soils exposed to elevated atmospheric CO2

Elevated atmospheric carbon dioxide (CO₂) has the potential to alter soil carbon (C) and nitrogen (N) cycling in arid ecosystems through changes in net primary productivity. However, an associated feedback exists because any sustained increases in plant productivity will depend upon the continued availability of soil N. We took soils from under the canopies of major shrubs, grasses, and plant interspaces in a Mojave Desert ecosystem exposed to elevated atmospheric CO₂ and incubated them in the laboratory with amendments of labile C and N to determine if elevated CO₂ altered the mechanistic controls of soil C and N on microbial N cycling. Net ammonification increased under shrubs exposed to elevated CO₂, while net nitrification decreased. Elevated CO₂ treatments exhibited greater fluxes of N₂O–N under Lycium spp., but not other microsites. The proportion of microbial/extractable organic N increased under shrubs exposed to elevated CO₂. Heterotrophic N₂‐fixation and C mineralization increased with C addition, while denitrification enzyme activity and N₂O–N fluxes increased when C and N were added in combination. Laboratory results demonstrated the potential for elevated CO₂ to affect soil N cycling under shrubs and supports the hypothesis that energy limited microbes may increase net inorganic N cycling rates as the amount of soil‐available C increases under elevated CO₂. The effect of CO₂ enrichment on N‐cycling processes is mediated by its effect on the plants, particularly shrubs. The potential for elevated atmospheric CO₂ to lead to accumulation of NH₄⁺ under shrubs and the subsequent volatilization of NH₃ may result in greater losses of N from this system, leading to changes in the form and amount of plant‐available inorganic N. This introduces the potential for a negative feedback mechanism that could act to constrain the degree to which plants can increase productivity in the face of elevated atmospheric CO₂.     

Genetic variation in germination, growth, and survivorship of red maple in response to subambient through elevated atmospheric CO2

Genetic variation in plant response to atmospheric carbon dioxide (CO₂) may have influenced paleo‐vegetation dynamics and could determine how future elevated CO₂ drives plant evolution and ecosystem productivity. We established how levels of relatedness – the maternal family, population, and provenance – affect variation in the CO₂ response of a species. This 2‐year growth chamber experiment focused on the germination, growth, biomass allocation, and survivorship responses of Acer rubrum to four concentrations of CO₂: 180, 270, 360, and 600 μL L^?1– representing Pleistocene through potential future conditions. We found that all levels of relatedness interacted with CO₂ to contribute to variation in response. Germination responses to CO₂ varied among families and populations, growth responses depended on families and regions of origin, and survivorship responses to CO₂ were particularly affected by regional identities. Differences among geographic regions accounted for 23% of the variation in biomass response to CO₂. If seeds produced under subambient CO₂ conditions behave similarly, our results suggest that A. rubrum may have experienced strong selection on seedling survivorship at Pleistocene CO₂ levels. Further, this species may evolve in response to globally rising CO₂ so as to increase productivity above that experimentally observed today. Species responses to future atmospheric CO₂ and the accompanying biotic effects on the global carbon cycle will vary among families, populations, and provenances.     

Growth and nitrogen uptake in an experimental community of annuals exposed to elevated atmospheric CO2

Rising levels of atmospheric CO₂ may alter patterns of plant biomass production. These changes will be dependent on the ability of plants to acquire sufficient nutrients to maintain enhanced growth. Species-specific differences in responsiveness to CO₂ may lead to changes in plant community composition and biodiversity. Differences in species-level growth responses to CO₂ may be, in a large part, driven by differences in the ability to acquire nutrients. To understand the mechanisms of how elevated CO₂ leads to changes in community-level productivity, we need to study the growth responses and patterns of nutrient acquisition for each of the species that comprise the community. In this paper, we present a study of how elevated CO₂ affects community-level and species-level patterns of nitrogen uptake and biomass production. As an experimental system we use experimental communities of 11 co-occurring annuals common to disturbed seasonal grasslands in south-western U.S.A. We established experimental communities with approximately even numbers of each species in three different atmospheric CO₂ concentrations (375, 550, and 700 ppm). We maintained these communities for 1, 1.5, and 2 months at which times we applied a ¹⁵N tracer (¹⁵NH₄¹⁵NO₃) to quantify the nitrogen uptake and then measured plant biomass, nitrogen content, and nitrogen uptake rates for the entire communities as well as for each species. Overall, community-level responses to elevated CO₂ were consistent with the majority of other studies of individual- and multispecies assemblages, where elevated CO₂ leads to enhanced biomass production early on, but this enhancement declines through time. In contrast, the responses of the individual species within the communities was highly variable, showing the full range of responses from positive to negative. Due to the large variation in size between the different species, community-level responses were generally determined by the responses of only one or a few species. Thus, while several of the smaller species showed trends of increased biomass and nitrogen uptake in elevated CO₂ at the end of the experiment, community-level patterns showed a decrease in these parameters due to the significant reduction in biomass and nitrogen content in the single largest species. The relationship between enhancement of nitrogen uptake and biomass production in elevated CO₂ was highly significant for both 550 ppm and 700 ppm CO₂. This relationship strongly suggests that the ability of plants to increase nitrogen uptake (through changes in physiology, morphology, architecture, or mycorrhizal symbionts) may be an important determinant of which species in a community will be able to respond to increased CO₂ levels with increased biomass production. The fact that the most dominant species within the community showed reduced enhancement and the smaller species showed increased enhancement suggest that through time, elevated CO₂ may lead to significant changes in community composition. At the community level, nitrogen uptake rates relative to plant nitrogen content were invariable between the three different CO₂ levels at each harvest. This was in contrast to significant reductions in total plant nitrogen uptake and nitrogen uptake relative to total plant biomass. These patterns support the hypothesis that plant nitrogen uptake is largely regulated by physiological activity, assuming that physiological activity is controlled by nitrogen content and thus protein and enzyme content.     

Stable carbon isotope variations in surface bloom scum and subsurface seston among shallow eutrophic lakes

Carbon stable isotope analysis of surface bloom scum and subsurface seston samples was conducted in shallow eutrophic lakes in China during warm seasons from 2003 to 2004. δ¹³C values of bloom scum were always higher (averaged 5‰) than those of seston in this study, and the possible reasons were attributed to (i) direct use of atmospheric CO₂ at the air–water interface, (ii) decrease in ¹³C fractionation due to higher carbon fixation, (iii) active CO₂ transport, and/or (iv) HCO₃⁻ accumulation. Negative correlation between δ¹³C_scum − δ¹³C_seston and pH in the test lakes indicated that phytoplankton at the subsurface water column increased isotopic enrichment under the carbon limitation along with the increase of pH, which might in turn decreased the differences in δ¹³C between the subsurface seston and the surface scums. Significant positive correlations of seston δ¹³C with total concentrations of nitrogen and phosphorus in water column suggested that the increase in δ¹³C of seston with trophic state was depending on nutrient (N or P, or both) supply. Our study showed that δ¹³C of phytoplankton was indicative of carbon utilization, primary productivity, and nutrient supply among the eutrophic lakes.     

Environmental productivity indices and productivity for Opuntia ficus-indica under current and elevated atmospheric CO2 levels*

Abstract. The productivity of the prickly-pear cactus Opuntia ficus-indica, which is cultivated worldwide for its fruits and stem segments, was predicted based on the responses of its net CO₂ uptake to soil water status, air temperature and photosynthetic photon flux density (PPFD). Each of these environmental factors was represented by an index with a maximum value of unity when that factor was not limiting net CO₂ uptake over a 24-h period. The water index, the temperature index, and the PPFD index were determined for 87 sites in the contiguous United States using data from 189 weather stations and for 148 sites worldwide using data from 1464 weather stations. The product of these three indices, the environmental productivity index (EPI), was used to predict the productivity of O. ficus-indica under current climatic conditions and under those accompanying a possible increase in the atmospheric CO₂ level to 650μumol mol^?1. Sites with temperatures always above -10°C and hence suitable for prickly-pear cultivation numbered 37 in the United States and 110 worldwide; such sites increased by 43 and 5%, respectively, for the global warming accompanying the elevated CO₂. Productivity of O. ficus-indica was at least 15 tonnes dry weight hectare^?1 year^?1, comparable to that of many agronomic crops, for 20 sites with temperatures always above -10°C in the contiguous United States and for 12 such sites worldwide under current climatic conditions; such sites increased by 85 and 117%, respectively, under the elevated CO₂ condition, mainly because of direct effects of the atmospheric CO₂ level on net CO₂ uptake. In summary, simulations based on EPI indicate that O. ficus-indica may presently be advantageously cultivated over a substantial fraction of the earth's surface, such regions increasing markedly with a future doubling in atmospheric CO₂ levels.