Plant responses to elevated CO<Subscript>2</Subscript> concentration at different scales: leaf,whole plant,canopy, and population

Elevated CO₂ enhances photosynthesis and growth of plants, but the enhancement is strongly influenced by the availability of nitrogen. In this article, we summarise our studies on plant responses to elevated CO₂. The photosynthetic capacity of leaves depends not only on leaf nitrogen content but also on nitrogen partitioning within a leaf. In Polygonum cuspidatum, nitrogen partitioning among the photosynthetic components was not influenced by elevated CO₂ but changed between seasons. Since the alteration in nitrogen partitioning resulted in different CO₂-dependence of photosynthetic rates, enhancement of photosynthesis by elevated CO₂ was greater in autumn than in summer. Leaf mass per unit area (LMA) increases in plants grown at elevated CO₂. This increase was considered to have resulted from the accumulation of carbohydrates not used for plant growth. With a sensitive analysis of a growth model, however, we suggested that the increase in LMA is advantageous for growth at elevated CO₂ by compensating for the reduction in leaf nitrogen concentration per unit mass. Enhancement of reproductive yield by elevated CO₂ is often smaller than that expected from vegetative growth. In Xanthium canadense, elevated CO₂ did not increase seed production, though the vegetative growth increased by 53%. As nitrogen concentration of seeds remained constant at different CO₂ levels, we suggest that the availability of nitrogen limited seed production at elevated CO₂ levels. We found that leaf area development of plant canopy was strongly constrained by the availability of nitrogen rather than by CO₂. In a rice field cultivated at free-air CO₂ enrichment, the leaf area index (LAI) increased with an increase in nitrogen availability but did not change with CO₂ elevation. We determined optimal LAI to maximise canopy photosynthesis and demonstrated that enhancement of canopy photosynthesis by elevated CO₂ was larger at high than at low nitrogen availability. We also studied competitive asymmetry among individuals in an even-aged, monospecific stand at elevated CO₂. Light acquisition (acquired light per unit aboveground mass) and utilisation (photosynthesis per unit acquired light) were calculated for each individual in the stand. Elevated CO₂ enhanced photosynthesis and growth of tall dominants, which reduced the light availability for shorter subordinates and consequently increased size inequality in the stand.     

Involvement of nitrogen and cytokinins in photosynthetic acclimation to elevated CO2 of spring wheat

Acclimation of photosynthetic capacity to elevated CO₂ involves a decrease of the leaf Rubisco content. In the present study, it was hypothesized that nitrogen uptake and partitioning within the leaf and among different aboveground organs affects the down-regulation of Rubisco. Given the interdependence of nitrogen and cytokinin signals at the whole plant level, it is also proposed that cytokinins affect the nitrogen economy of plants under elevated CO₂, and therefore the acclimatory responses. Spring wheat received varying levels of nitrogen and cytokinin in field chambers with ambient (370 μmol mol⁻¹) or elevated (700 μmol mol⁻¹) atmospheric CO₂. Gas exchange, Rubisco, soluble protein and nitrogen contents were determined in the top three leaves in the canopy, together with total nitrogen contents per shoot. Growth in elevated CO₂ induced decreases in photosynthetic capacity only when nitrogen supply was low. However, the leaf contents of Rubisco, soluble protein and total nitrogen on an area basis declined in elevated CO₂ regardless of nitrogen supply. Total nitrogen in the shoot was no lower in elevated than ambient CO₂, but the fraction of this nitrogen located in flag and penultimate leaves was lower in elevated CO₂. Decreased Rubisco: chlorophyll ratios accompanied losses of leaf Rubisco with CO₂ enrichment. Cytokinin applications increased nitrogen content in all leaves and nitrogen allocation to senescing leaves, but decreased Rubisco contents in flag leaves at anthesis and in all leaves 20 days later, together with the amount of Rubisco relative to soluble protein in all leaves at both growth stages. The results suggest that down regulation of Rubisco in leaves at elevated CO₂ is linked with decreased allocation of nitrogen to the younger leaves and that cytokinins cause a fractional decrease of Rubisco and therefore do not alleviate acclimation to elevated CO₂.     

Climate change and cereal aphids: the relative effects of increasing CO2 and temperature on aphid population dynamics

Experimental evidence regarding the responses of cereal aphids to rising atmospheric CO₂ has been ambiguous. Some studies suggest increased population sizes under future CO₂ levels, others suggest decreased population sizes, and still others suggest little or no difference. Recently, Newman et al. (2003) constructed a general mathematical model of the aphid–grass interaction to investigate whether or not we should, in fact, expect a general aphid response to rising CO₂. They concluded that aphid populations are likely to be larger under future CO₂ concentrations if soil N levels are high, the aphid species' nitrogen requirement is low and the aphid species' density‐dependent response in winged morph production is weak. In that model, and in field experiments, CO₂ concentration influences aphid population dynamics through the effect it has on plant quality. However, future CO₂ concentrations are also likely to be accompanied by higher ambient temperatures, a combination that has received little focus to date. In the present paper, the Newman et al. model is used to consider the combined effects of increased CO₂ concentrations and temperature on aphid population sizes. It is concluded that, when both factors are elevated, aphid population dynamics will be more similar to current ambient conditions than expected from the results of experiments studying either factor alone. This result has important implications for future experimentation.     

Effect of elevated CO<Subscript>2</Subscript> levels on leaf starch,nitrogen and photosynthesis of plants growing at three natural CO<Subscript>2</Subscript> springs in Japan

Plant communities around natural CO₂ springs have been exposed to elevated CO₂ levels over many generations and give us a unique opportunity to investigate the effects of long-term elevated CO₂ levels on wild plants. We searched for natural CO₂ springs in cool temperate climate regions in Japan and found three springs that were suitable for studying long-term responses of plants to elevated levels of CO₂: Ryuzin-numa, Yuno-kawa and Nyuu. At these CO₂ springs, the surrounding air was at high CO₂ concentration with no toxic gas emissions throughout the growth season, and there was natural vegetation around the springs. At each site, high-CO₂ (HC) and low-CO₂ (LC) plots were established, and three dominant species at the shrub layers were used for physiological analyses. Although the microenvironments were different among the springs, dicotyledonous species growing at the HC plots tended to have more starch and less nitrogen per unit dry mass in the leaves than those growing at the LC plots. In contrast, monocotyledonous species growing in the HC and LC plots had similar starch and nitrogen concentrations. Photosynthetic rates at the mean growth CO₂ concentration were higher in HC plants than LC plants, but photosynthetic rates at a common CO₂ concentration were lower in HC plants. Efficiency of water and nitrogen use of leaves at growth CO₂ concentration was greatly increased in HC plants. These results suggest that natural plants growing in elevated CO₂ levels under cool temperate climate conditions have down-regulated their photosynthetic capacity but that they increased photosynthetic rates and resource use efficiencies due to the direct effect of elevated CO₂ concentration.     

Growth and nitrogen accretion of dinitrogen-fixing Alnus glutinosa (L.) Gaertn. under elevated carbon dioxide

Short-term studies of tree growth at elevated CO₂ suggest that forest productivity may increase as atmospheric CO₂ concentrations rise, although low soil N availability may limit the magnitude of this response. There have been few studies of growth and N₂ fixation by symbiotic N₂-fixing woody species under elevated CO₂ and the N inputs these plants could provide to forest ecosystems in the future. We investigated the effect of twice ambient CO₂ on growth, tissue N accretion, and N₂ fixation of nodulated Alnus glutinosa (L.) Gaertn. grown under low soil N conditions for 160 d. Root, nodule, stem, and leaf dry weight (DW) and N accretion increased significantly in response to elevated CO₂. Whole-plant biomass and N accretion increased 54% and 40%, respectively. Delta-¹⁵N analysis of leaf tissue indicated that plants from both treatments derived similar proportions of their total N from symbiotic fixation suggesting that elevated CO₂ grown plants fixed approximately 40% more N than did ambient CO₂ grown plants. Leaves from both CO₂ treatments showed similar relative declines in leaf N content prior to autumnal leaf abscission, but total N in leaf litter increased 24% in elevated compared to ambient CO₂ grown plants. These results suggest that with rising atmospheric CO₂ N₂-fixing woody species will accumulate greater amounts of biomass N through N₂ fixation and may enhance soil N levels by increased litter N inputs.     

Loblolly pine grown under elevated CO2 affects early instar pine sawfly performance

Seedlings of loblolly pine Pinus taeda (L.), were grown in open-topped field chambers under three CO₂ regimes: ambient, 150 l l^–1 CO₂ above ambient, and 300 l l^–1 CO₂ above ambient. A fourth, non-chambered ambient treatment was included to assess chamber effects. Needles were used in 96 h feeding trials to determine the performance of young, second instar larvae of loblolly pine's principal leaf herbivore, red-headed pine sawfly, Neodiprion lecontei (Fitch). The relative consumption rate of larvae significantly increased on plants grown under elevated CO₂, and needles grown in the highest CO₂ regime were consumed 21% more rapidly than needles grown in ambient CO₂. Both the significant decline in leaf nitrogen content and the substantial increase in leaf starch content contributed to a significant increase in the starch:nitrogen ratio in plants grown in elevated CO₂. Insect consumption rate was negatively related to leaf nitrogen content and positively related to the starch:nitrogen ratio. Of the four volatile leaf monoterpenes measured, only -pinene exhibited a significant CO₂ effect and declined in plants grown in elevated CO₂. Although consumption changed, the relative growth rates of larvae were not different among CO₂ treatments. Despite lower nitrogen consumption rates by larvae feeding on the plants grown in elevated CO₂, nitrogen accumulation rates were the same for all treatments due to a significant increase in nitrogen utilization efficiency. The ability of this insect to respond at an early, potentially susceptible larval stage to poorer food quality and declining levels of a leaf monoterpene suggest that changes in needle quality within pines in future elevated-CO₂ atmospheres may not especially affect young insects and that tree-feeding sawflies may respond in a manner similar to herb-feeding lepidopterans.     

Effects of combined nitrogen on anapleurotic carbon assimilation and bleeding sap composition in Phaseolus vulgaris L.

Dwarf french beans, Phaseolus vulgaris L., were grown with or without inoculation with rhizobia (strain 3644), and with or without a combined nitrogen source (nitrate or ammonium ions). The distribution of radioactivity into products of dark ¹⁴CO₂ assimilation was studied in roots or nodules from these plants. A detailed study was also made of the distribution and rates of excretion of nitrogen in xylem bleeding sap in 28 day old plants grown on the various sources of nitrogen. Whereas detached nodules accumulated radioactive glycine, serine and glutamate when incubated with ¹⁴CO₂, bleeding sap from plants root fed ¹⁴CO₂ contained low levels of radioactivity in these compounds but higher levels in allantoin. Chemical analysis showed allantoin to be the major compound transported in the xylem of nodulated plants, whether or not they were fed on combined nitrogen. In contrast uninoculated plants accumulated mainly amino acids in the bleeding sap, the amount and chemical composition of which depended on the combined nitrogen source.Abbreviations PEP phosphoenol pyruvate - OAA oxaloacetate     

Aphid suitability for the predatory hoverfly Episyrphus balteatus altered with elevating atmospheric CO2 and sinigrin

The fitness of natural enemies should be altered in response to changes in herbivore quality induced by the impact of increased atmospheric CO₂ levels on plants. We studied the effect of different CO₂ levels on the aphid predator Episyrphus balteatus DeGeer fed either specialist or generalist aphids reared on either of two host plants under laboratory conditions. In the host plant that contains sinigrin (black mustard), elevated CO₂ increased the sinigrin content of both host plant and the specialist aphid, but reduced the already very low levels in the generalist aphid. Predator development time increased with elevated CO₂, while fecundity decreased. Consequently, individual fitness decreased slightly with increasing atmospheric CO₂. Sinigrin significantly decreased fecundity and increased development time of the predator. As a result, fitness was significantly lower too. The consumption rate was influenced significantly by plant and prey solely and the interactions of host plant × prey type and CO₂ level × prey type. Further research on the effects of climate change parameters (e.g. greenhouse gases such as CO₂, ozone (O₃) and nitrogen dioxide (NO₂), etc.) separately and jointly under controlled environmental conditions will help to understand the nature and direction of their effects on natural enemies as part of the tritrophic system.     

Cisgenic overexpression of cytosolic glutamine synthetase improves nitrogen utilization efficiency in barley and prevents grain protein decline under elevated CO2

Cytosolic glutamine synthetase (GS1) plays a central role in nitrogen (N) metabolism. The importance of GS1 in N remobilization during reproductive growth has been reported in cereal species but attempts to improve N utilization efficiency (NUE) by overexpressing GS1 have yielded inconsistent results. Here, we demonstrate that transformation of barley (Hordeum vulgare L.) plants using a cisgenic strategy to express an extra copy of native HvGS1‐1 lead to increased HvGS1.1 expression and GS1 enzyme activity. GS1 overexpressing lines exhibited higher grain yields and NUE than wild‐type plants when grown under three different N supplies and two levels of atmospheric CO₂. In contrast with the wild‐type, the grain protein concentration in the GS1 overexpressing lines did not decline when plants were exposed to elevated (800–900 μL/L) atmospheric CO₂. We conclude that an increase in GS1 activity obtained through cisgenic overexpression of HvGS1‐1 can improve grain yield and NUE in barley. The extra capacity for N assimilation obtained by GS1 overexpression may also provide a means to prevent declining grain protein levels under elevated atmospheric CO₂.     

How do aphids respond to elevated CO2?

The performance of herbivore insects is determined directly by the quality of host plants. Elevated CO₂ induced a decline in foliar nitrogen, which reduced the growth of chewing insects. Phloem-sucking insects (i.e. aphid), however, had species-specific responses to elevated CO₂ and were the only feeding guild to respond positively to elevated CO₂. Although many studies attempt to illuminate the interaction between aphids and plants under elevated CO₂, few studies can explain why some aphids are more successful than other chewing insects in elevated CO₂. Elevated CO₂ leads to a re-allocation of the carbon and nitrogen resources in plant tissue, which increases the thickness of the microscopic structures of leaves, reduces amino acids content of leaf phloem sap and increases the secondary metabolites. Considering the complexity of aphid–plant interactions, it is difficult and unreasonable to predict the general response of aphids to elevated CO₂ using a single plant component. Instead, it is more likely that aphids are able to overcome the disadvantages of the indirect effects of elevated CO₂ by reducing developmental times and increasing fecundity under elevated CO₂ conditions. Our results provide several clues to why some aphids are successful in elevated CO₂ conditions. We review recent studies of the effects of elevated CO₂ on aphids and discuss the effects of elevated CO₂ on aphid performance on crops using cotton and cereal aphids as examples.     

Controls of biomass partitioning between roots and shoots: Atmospheric CO2 enrichment and the acquisition and allocation of carbon and nitrogen in wild radish

Summary The effects of CO₂ enrichment on plant growth, carbon and nitrogen acquisition and resource allocation were investigated in order to examine several hypotheses about the mechanisms that govern dry matter partitioning between shoots and roots. Wild radish plants (Raphanus sativus × raphanistrum) were grown for 25 d under three different atmospheric CO₂ concentrations (200 ppm, 330 ppm and 600 ppm) with a stable hydroponic 150 mol 1^–1 nitrate supply. Radish biomass accumulation, photosynthetic rate, water use efficiency, nitrogen per unit leaf area, and starch and soluble sugar levels in leaves increased with increasing atmospheric CO₂ concentration, whereas specific leaf area and nitrogen concentration of leaves significantly decreased. Despite substantial changes in radish growth, resource acquisition and resource partitioning, the rate at which leaves accumulated starch over the course of the light period and the partitioning of biomass between roots and shoots were not affected by CO₂ treatment. This phenomenon was consistent with the hypothesis that root/shoot partitioning is related to the daily rate of starch accumulation by leaves during the photoperiod, but is inconsistent with hypotheses suggesting that root/shoot partitioning is controlled by some aspect of plant C/N balance.     

Pea plant responsiveness under elevated [CO2] is conditioned by the N source (N2 fixation versus NO3− fertilization)

The main goal of this study was to test the effect of [CO₂] on C and N management in different plant organs (shoots, roots and nodules) and its implication in the responsiveness of exclusively N₂-fixing and NO₃⁻-fed plants. For this purpose, exclusively N₂-fixing and NO₃⁻-fed (10 mM) pea (Pisum sativum L.) plants were exposed to elevated [CO₂] (1000 μmol mol⁻¹ versus 360 μmol mol⁻¹ CO₂). Gas exchange analyses, together with carbohydrate, nitrogen, total soluble proteins and amino acids were determined in leaves, roots and nodules. The data obtained revealed that although exposure to elevated [CO₂] increased total dry mass (DM) in both N treatments, photosynthetic activity was down-regulated in NO₃⁻-fed plants, whereas N₂-fixing plants were capable of maintaining enhanced photosynthetic rates under elevated [CO₂]. In the case of N₂-fixing plants, the enhanced C sink strength of nodules enabled the avoidance of harmful leaf carbohydrate build up. On the other hand, in NO₃⁻-fed plants, elevated [CO₂] caused a large increase in sucrose and starch. The increase in root DM did not contribute to stimulation of C sinks in these plants. Although N₂ fixation matched plant N requirements with the consequent increase in photosynthetic rates, in NO₃⁻-fed plants, exposure to elevated [CO₂] negatively affected N assimilation with the consequent photosynthetic down-regulation.     

Elevated atmospheric partial pressure of CO2 and plant growth

Cotton plants were grown in late spring under full sunlight in glasshouses containing normal ambient partial pressure of CO₂ (32±2Pa) and enriched partial pressure of CO₂ (64±1.5Pa) and at four levels of nitrogen nutrition. Thirty-five days after planting, the total dry weights of high CO₂-grown plants were 2- to 3.5-fold greater than plants grown in normal ambient CO₂ partial pressure. Depending on nitrogen nutrition level, non-structural carbohydrate content (mainly starch) in the leaves of plants grown in normal CO₂ was between 4 and 37% of the total leaf dry weight compared to 39 to 52% in the leaves of high CO₂-grown plants. Specific leaf weight calculated using total dry weight was 1.6- to 2-fold greater than that based on structural dry weight. In high CO₂-grown plants the amount of non-structural carbohydrate translocated from the leaves at night was between 10 and 20% of the level at the end of the photoperiod. This suggests that the plant was unable to utilize all the carbohydrate it assimilated in elevated CO₂ atmosphere. While there was a 1.5-fold enhancement in the rate of CO₂ assimilation in plants grown in 64 Pa CO₂, there was, however, some evidence to suggest that the activities of other metabolic pathways in the plants were not stimulated to the same extent by the enriched CO₂ atmosphere. This resulted in massive accumulation of non-structural carbohydrate, particularly at low level of nitrogen nutrition.Abbreviations A rate of CO₂ assimilation - PPFD photosynthetic photo flux density - NAR net assimilation rate - pCO₂ partial pressure of CO₂ - RGR relative growth rate     

The response of fast- and slow-growing Acacia species to elevated atmospheric CO2: an analysis of the underlying components of relative growth rate

In this study we assessed the impact of elevated CO₂ with unlimited water and complete nutrient on the growth and nitrogen economy of ten woody Acacia species that differ in relative growth rate (RGR). Specifically, we asked whether fast- and slow-growing species systematically differ in their response to elevated CO₂. Four slow-growing species from semi-arid environments (Acacia aneura, A. colei, A. coriacea and A. tetragonophylla) and six fast-growing species from mesic environments (Acacia dealbata, A. implexa, A. mearnsii, A. melanoxylon, A. irrorata and A. saligna) were grown in glasshouses with either ambient (˜350 ppm) or elevated (˜700 ppm) atmospheric CO₂. All species reached greater final plant mass with the exception of A. aneura, and RGR, averaged across all species, increased by 10% over a 12-week period when plants were exposed to elevated CO₂. The stimulation of RGR was evident throughout the 12-week growth period. Elevated CO₂ resulted in less foliage area per unit foliage dry mass, which was mainly the result of an increase in foliage thickness with a smaller contribution from greater dry matter content per unit fresh mass. The net assimilation rate (NAR, increase in plant mass per unit foliage area and time) of the plants grown at elevated CO₂ was higher in all species (on average 30% higher than plants in ambient CO₂) and was responsible for the increase in RGR. The higher NAR was associated with a substantial increase in foliar nitrogen productivity in all ten Acacia species. Plant nitrogen concentration was unaltered by growth at elevated CO₂ for the slow-growing Acacia species, but declined by 10% for faster-growing species. The rate of nitrogen uptake per unit root mass was higher in seven of the species when grown under elevated CO₂, and leaf area per unit root mass was reduced by elevated CO₂ in seven of the species. The absolute increase in RGR due to growth under elevated CO₂ was greater for fast- than for slow-growing Acacia species. Received: 21 December 1998 / Accepted: 31 May 1999     

Elevated CO2 reduces the drought effect on nitrogen metabolism in barley plants during drought and subsequent recovery

The objective of this study was to determine the response of nitrogen metabolism to drought and recovery upon rewatering in barley (Hordeum vulgare L.) plants under ambient (350 μmol mol⁻¹) and elevated (700 μmol mol⁻¹) CO₂ conditions. Barley plants of the cv. Iranis were subjected to drought stress for 9, 13, or 16 days. The effects of drought under each CO₂ condition were analysed at the end of each drought period, and recovery was analysed 3 days after rewatering 13-day droughted plants. Soil and plant water status, protein content, maximum (NR_max) and actual (NR_act) nitrate reductase, glutamine synthetase (GS), and aminant (NADH-GDH) and deaminant (NAD-GDH) glutamate dehydrogenase activities were analysed. Elevated CO₂ concentration led to reduced water consumption, delayed onset of drought stress, and improved plant water status. Moreover, in irrigated plants, elevated CO₂ produced marked changes in plant nitrogen metabolism. Nitrate reduction and ammonia assimilation were higher at elevated than at ambient CO₂, which in turn yielded higher protein content. Droughted plants showed changes in water status and in foliar nitrogen metabolism. Leaf water potential (Ψ_w) and nitrogen assimilation rates decreased after the onset of water deprivation. NR_act and NR_max activity declined rapidly in response to drought. Similarly, drought decreased GS whereas NAD-GDH rose. Moreover, protein content fell dramatically in parallel with decreased leaf Ψ_w. In contrast, elevated CO₂ reduced the water stress effect on both nitrate reduction and ammonia assimilation coincident with a less-steep decrease in Ψ_w. On the other hand, Ψ_w practically reached control levels after 3 days of rewatering. In parallel with the recovery of plant water status, nitrogen metabolism was also restored. Thus, both NR_act and NR_max activities were restored to about 75-90% of control levels when water supply was restored; the GS activity reached 80-90% of control values; and GDH activities and protein content were similar to those of control plants. The recovery was always faster and slightly higher in plants grown under elevated CO₂ conditions compared to those grown in ambient CO₂, but midday Ψ_w dropped to similar values under both CO₂ conditions. The results suggest that elevated CO₂ improves nitrogen metabolism in droughted plants by maintaining better water status and enhanced photosynthesis performance, allowing superior nitrate reduction and ammonia assimilation. Ultimately, elevated CO₂ mitigates many of the effects of drought on nitrogen metabolism and allows more rapid recovery following water stress.     

Effects of elevated CO2 on growth and carbon/nutrient balance in the deciduous woody shrub Lindera benzoin (L.) Blume (Lauraceae)

We examined the effects of elevated CO₂ on growth and carbon/nutrient balance in a natural population of the deciduous temperate zone shrub Lindera benzoin. Our data concern whole plant, leaf, and stem growth for the first two seasons of a long-term field experiment in which CO₂ levels were manipulated in situ. In addition to growth parameters, we evaluated changes in leaf and stem chemistry, including total nitrogen, nonstructural carbohydrates, and total phenolics. Over the course of this study, L. benzoin appeared to respond to elevated CO₂ primarily by physiological and biochemical changes, with only a slight enhancement in aboveground growth (ramet height). Positive effects on aboveground growth were primarily evident in young (nonreproductive) ramets. Our results suggest that nitrogen limitation may have constrained plants to allocate carbohydrates produced in response to elevated CO₂ primarily to storage and belowground growth, and perhaps to increased secondary chemical production, rather than to increased stem and leaf growth. We discuss our results in terms of changes in carbon/nutrient balance induced by elevated CO₂, and provide predictions for future changes in this system based upon constraints imposed by intrinsic and extrinsic factors and their potential effects on the reallocation of stored reserves.     

Salinity and Nitrogen Effects on Photosynthesis, Ribulose-1,5-Bisphosphate Carboxylase and Metabolite Pool Sizes in Phaseolus vulgaris L

Salinity (100 millimolar NaCl) was found to reduce photosynthetic capacity independent of stomatal closure in Phaseolus vulgaris. This reduction was shown to be a consequence of a reduction in the efficiency of ribulose-1,5-bisphosphate (RuBP) carboxylase (RuBPCase) rather than a reduction in the leaf content of photosynthetic machinery. In control plants, photosynthesis became RuBP-limited at approximately 1.75 moles RuBP per mole 2-carboxyarabinitol bisphosphate binding sites. Salinization caused the RuBP pool size to reach this limiting value for CO₂ fixation at much lower values of intercellular CO₂. Plants grown at low nitrogen and ± NaCl became RuBP limited at similar RuBP pool sizes as the high nitrogen-grown plants. At limiting RuBP pool sizes and equal values of intercellular CO₂ photosynthetic capacity of salt-stressed plants was less than control plants. This effect of salinity on RuBPCase activity could not be explained by deactivation of the enzyme or inhibitor synthesis. Thus, salinity reduced photosynthetic capacity by reducing both the RuBP pool size by an effect on RuBP regeneration capacity and RuBPCase activity by an unknown mechanism when RuBP was limiting.     

The effect of free air carbon dioxide enrichment (FACE) and soil nitrogen availability on the photosynthetic capacity of wheat

A simple system for free air carbon dioxide enrichment (FACE) was recently developed and it is here briefly described. Such a MiniFACE system allowed the elevation of CO₂ concentration of small field plots avoiding the occurrence of large spatial and temporal fluctuations. A CO₂ enrichment field experiment was conducted in Italy in the season 1993–1994 with wheat (cv. Super-dwarf Mercia). A randomized experimental design was used with the treatment combination CO₂ × soil N, replicated twice. Gas exchange measurements showed that photosynthetic capacity was significantly decreased in plants exposed to elevated CO₂ and grown under nitrogen deficiency. Photosynthetic acclimation was, in this case, due to the occurrence of reduced rates of rubP saturated and rubP regeneration limited photosynthesis. Gas exchange measurements did not instead reveal any significant effect of elevated CO₂ on the photosynthetic capacity of leaves of plants well fertilized with nitrogen, in spite of a transitory negative effect on rubP regeneration limited photosynthesis that was detected to occur in the central part of a day with high irradiance. It is concluded that the levels of nitrogen fertilization will play a substantial role in modulating CO₂ fertilization effects on growth and yields of wheat crops under the scenario of future climate change.     

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.