Impact of CO2 Enrichment and Variable Nitrogen Supplies on Composition and Partitioning of Essential Nutrients of Wheat

This study was conducted to determine effects of nitrogen supply (75 and 150 kg(N) ha⁻¹) and CO₂ enrichment on partitioning of macro and micro nutrients in wheat (Triticum aestivum L. cv. HD-2285). Plants were grown from seedling emergence to maturity inside open top chambers under ambient CO₂ (CA, 350 ± 50 μmol mol⁻¹) and elevated CO₂ (CE, 600 ± 50 μmol mol⁻¹). Leaves, stems and roots of the same physiological age were analyzed for carbon, nitrogen, calcium, copper, iron, zinc and manganese content at 40, 60 and 90 d after germination. C, Cu, Mn and Zn content was higher in the stem, leaves and roots on dry mass basis under CE than CA. However, N and Fe contents decreased in CE grown plants. Ca content was unaffected due to CE and variable N supplies. This revised version was published online in July 2006 with corrections to the Cover Date.     

Short-term effect of elevated CO2 concentration and high irradiance on the antioxidant enzymes in bean plants

The effect of short-term exposure to elevated CO₂ concentration and high irradiance on the activity of superoxide dismutase (SOD), ascorbate peroxidase (APX), guaiacol peroxidases (GPX) and catalase (CAT), and on the extent of the lipid peroxidation was studied in bean (Phaseolus vulgaris L.) plants. Plants were exposed for 4 d (8 h a day) to irradiance of 100 (LI) or 1000 (HI) μmol m⁻² s⁻¹ at ambient (CA, 350 μmol mol⁻¹) or elevated (CE, 1300 μmol mol⁻¹) CO₂ concentration. Four-day exposure to CE increased the leaf dry mass in HI plants and RuBPC activity and chlorophyll content in LI plants. Total soluble protein content, leaf dry matter and RuBPC activity were higher in HI than in LI plants, although the HI and CE increased the contents of malonyldialdehyde and H₂O₂. Under CA, exposure to HI increased the activity of APX and decreased the total SOD activity. Under CE, HI treatment also activated APX and led to reduction of both, SOD and GPX, enzymes activities. CE considerably reduced the CAT activity at both irradiances, possibly due to suppressed rate of photorespiration under CE conditions.     

The effect of elevated CO2 concentration on leaf chlorophyll and nitrogen contents in rice during post-flowering phases

The effect of elevated CO₂ concentration (CE) on leaf chlorophyll (Chl) and nitrogen (N) contents and photosynthetic rate (P_N) was evaluated during the post-flowering stages of rice grown at CE (570 ± 50 μmol mol⁻¹) in open top chamber (OTC), at ambient CO₂ concentration (∼ 365 μmol mol⁻¹) in OTC and at open field. Thirty-five day old seedlings were transplanted in OTCs or in field and allowed to grow till maturity. Chl and N contents were highest at the time of flowering and thereafter it started to decline. The rate of decline in Chl and N contents was faster in plants grown under CE mostly in later part of growth. Irrespective of treatment difference, flag leaf contained the highest amount of Chl and N than penultimate and third leaf. The higher P_N was observed in leaves under CE than in the leaves in other two growing conditions. Considering growth stage, P_N was the highest at flowering which reduced at the later part of growth due to degradation of Chl and N content of the leaf. Under CE it was 40.02 μmol m⁻² s⁻¹ at flowering and it reduced to only 14.77 μmol m⁻² s⁻¹ at maturity stage. The beneficial effect of CE in increasing leaf P_N may be maintained by applying extra dose of nitrogen at the later stages of plant growth.     

Diurnal Changes in Photosynthesis,Sugars, and Nitrogen of Wheat and Mungbean Grown Under Elevated CO2 Concentration

The diurnal changes in leaf net photosynthetic rate (P _N) and sugar and nitrogen contents in wheat [Triticum aestivum (L.) cv. HD 2285] and mungbean [Vigna radiata (L.) Wilczek cv. PS 16] were analysed under ambient, AC [350±25 µmol mol^–1] and elevated, EC [600±50 µmol mol^–1] CO₂ concentrations. In both mungbean and wheat P _N of AC- and EC-grown plants compared at the same CO₂ concentration showed that P _N was higher under EC. However, increased P _N in EC-plants declined in the afternoon and approached P _N of AC-plants. Depression in P _N, however, was less in mungbean compared with the large depression in wheat. Greater down regulation of P _N in wheat was associated with the accumulation of large amount of sugars and low nitrogen content in wheat leaves. Mungbean leaves accumulated mostly starch under EC and the difference in N content in AC- and EC-plants was relatively less than in wheat.     

Effects of elevated CO<Subscript>2</Subscript> and nitrogen on wheat growth and photosynthesis

The effects of nitrogen [75 and 150 kg (N) ha⁻¹] and elevated CO₂ on growth, photosynthetic rate, contents of soluble leaf proteins and activities of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) and nitrate reductase (NR) were studied on wheat (Triticum aestivum L. cv. HD-2285) grown in open top chambers under either ambient (AC) or elevated (EC) CO₂ concentration (350 ± 50, 600 ± 50 μmol mol⁻¹) and analyzed at 40, 60 and 90 d after sowing. Plants grown under EC showed greater photosynthetic rate and were taller and attained greater leaf area along with higher total plant dry mass at all growth stages than those grown under AC. Total soluble and Rubisco protein contents decreased under EC but the activation of Rubisco was higher at EC with higher N supply. Nitrogen increased the NR activity whereas EC reduced it. Thus, EC causes increased growth and P_N ability per unit uptake of N in wheat plants, even if N is limiting.     

Does the Response of Perennial Ryegrass to Elevated CO2 Concentration Depend on the Form of the Supplied Nitrogen?

To test whether different nitrogen form (nitrate or ammonium) in substrate can alter the response to elevated partial pressure of CO₂ (pCO₂) plants of perennial ryegrass (Lolium perenne cv. Bastion) were grown from seeds in growth chambers under pCO₂ of either 35 Pa (ambient, CA) or 70 Pa (elevated, CE) in a hydroponic system (with nutrient and pH control) for 24 d. Nitrogen was supplied as ammonium, nitrate or an equimolar mixture of both N forms. Under CE plants grew faster than their counterparts under CA during the first 14 d but after 23 d of cultivation stimulation disappeared. Despite the strong positive effect of mixed forms of N on plant growth, the beneficial effect of CE was similar to that in the other two N treatments. However, the almost alike final growth response to CE had different underlying mechanisms in different N treatments. Plants supplied with nitrate as a sole source of nitrogen had lower leaf mass ratio but much higher specific leaf area compared to plants supplied with ammonium. The decrease in the content of leaf organic N (per unit of structural dry mass) under CE was found only in leaves of plants supplied with ammonium on day 14. Nevertheless, the available form of N evidently contributes to changes of leaf N content under CE. The high levels of N and non-structural saccharides in plants supplied with ammonium at CE suggest that the CO₂ response of these plants was controlled by factors other than amount of available carbon and nitrogen.     

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.     

Effects of elevated CO<Subscript>2</Subscript> applied to potato roots on the anatomy and ultrastructure of leaves

The root system of potato (Solanum tuberosum L. cv. Favorita) plants was treated with different O₂ and CO₂ concentrations for 35 d in aeroponic culture. Under 21 or 5 % O₂ in the root zones, the thickness of leaves and palisade parenchyma significantly increased at 3 600 μmol(CO₂) mol⁻¹ in the root zone, compared with CO₂ concentration 380 μmol mol⁻¹ or low CO₂ concentration (100 μmol mol⁻¹). In addition, smaller cells of palisade tissue, more intercellular air spaces and partially two layers of palisade cells were observed in the leaves with root-zone CO₂ enrichment. Furthermore, there was a significant increase in the size of chloroplasts and starch grains, and the number of starch grains per chloroplast due to elevated CO₂ only under 21 % O₂. In addition, a significant decline in the thickness of grana and the number of lamellas, but no significant differences in the number of grana per chloroplast were observed under elevated CO₂ concentration. The accumulation of starch grains in the chloroplast under elevated CO₂ concentration could change the arrangement of grana thylakoids and consequently inhibited the absorption of sun radiation and photosynthesis of potato plants.     

Plant responses to past concentrations of CO2

The influence of recent historical changes in atmospheric CO₂ have been investigated by two methods: 1, the responses of leaf development and physiology as indicated by leaves stored in herbaria and 2, by investigating the differential growth responses of populations originating from naturally different CO₂ concentrations. Herbarium leaves indicate that stomatal density and leaf nitrogen have decreased over the last 150 to 200 years, while water use efficiency, estimated from leaf ¹³C and historical measurements of climate, has increased.Natural populations ofBoehmeria cylindrica were found growing at sites, in Florida, with CO₂ mole fractions varying naturally from 350 µmol mol^-1 to 505 µmol mol^-1. Plants were grown in the controlled environment, using seeds originating from populations occurring in the different CO₂ mole fractions. Plants from the different ambient CO₂ mole fractions showed different rates of growth and different non-linear responses of the shoot to root ratio in response to changes in the CO₂ mole fraction from 350 to 675 µmol mol^-1.The proposal that plants originating from high altitude will whow greater stimulations of growth with an increase in CO₂, than plants from low altitude, was rejected in experiments which simulated the atmospheric pressure at altitudes of 0 and 2000m, at CO₂ mole fractions of 350 and 700 µmol mol^-1 and on populations ofPlantago major originating from altitudes of 0 and 3335 m.     

Modelling Net Photosynthetic Rate of Winter Wheat in Elevated Air CO2 Concentrations

Winter wheat plants were grown in open top chambers either at 365 µmol mol^–1 (AC) or at 700 µmol mol^–1 (EC) air CO₂ concentrations. The photosynthetic response of flag leaves at the beginning of flowering and on four vertical leaf levels at the beginning of grain filling were measured. Net photosynthetic rates (P _N) were higher at both developmental phases in plants grown at EC coupled with larger leaf area and photosynthetic pigment contents. The widely accepted Farquhar net photosynthesis model was parameterised and tested using several observed data. After parameterisation the test results corresponded satisfactorily with observed values under several environmental conditions.     

Photosynthetic acclimation to elevated CO2 in wheat cultivars

Wheat (T. aestivum) cvs. Kalyansona and Kundan grown under atmospheric (CA) and elevated CO₂ concentrations (650±50 cm³ m^-3 - CE) in open top chambers were examined for net photosynthetic rate (PN), stomatal limitation (l _s) of P _N, ribulose-1,5-bisphosphate carboxylase (RuBPC) activity, and saccharide content of the leaves. The P _N values of both CA- and CE-grown plants compared at the same CO₂ concentration showed a down regulation under CE at the post-anthesis stage. The negative acclimation of P _N appeared to be due to both stomatal and mesophyll components, and the RuBPC activity got also adjusted. There was a decrease in activation state of RuBPC under CE. In connection with this, an increased accumulation of saccharides in wheat leaf under CE was observed. Kalyansona, owing to its larger sink potential in terms of the number of grains, showed a greater enhancement under CE in both post-ear emergence dry matter production and grain yield. Under CE, this cultivar also showed a lower down regulation of P _N than Kundan.     

Contrasting Effects of Carbon Dioxide and Irradiance on the Acclimation of Photosynthesis in Developing Soybean Leaves

Leaves developed at high irradiance (I) often have higher photosynthetic capacity than those developed at low I, while leaves developed at elevated CO₂ concentration [CO₂] often have reduced photosynthetic capacity compared with leaves developed at lower [CO₂]. Because both high I and elevated [CO₂] stimulate photosynthesis of developing leaves, their contrasting effects on photosynthetic capacity at maturity suggest that the extra photosynthate may be utilized differently depending on whether I or [CO₂] stimulates photosynthesis. These experiments were designed to test whether relationships between photosynthetic income and the net accumulation of soluble protein in developing leaves, or relationships between soluble protein and photosynthetic capacity at full expansion differed depending on whether I or [CO₂] was varied during leaf development. Soybean plants were grown initially with a photosynthetic photon flux density (PPFD) of 950 µmol m^–2 s^–1 and 350 µmol [CO₂] mol^–1, then exposed to [CO₂] ranging from 135 to 1400 µmol mol^–1 for the last 3 d of expansion of third trifoliolate leaves. These results were compared with experiments in which I was varied at a constant [CO₂] of 350 µmol mol^–1 over the same developmental period. Increases in area and dry mass over the 3 d were determined along with daily photosynthesis and respiration. Photosynthetic CO₂ exchange characteristics and soluble protein content of leaves were determined at the end of the treatment periods. The increase in leaflet mass was about 28 % of the dry mass income from photosynthesis minus respiration, regardless of whether [CO₂] or I was varied, except that very low I or [CO₂] increased this percentage. Leaflet soluble protein per unit of area at full expansion had the same positive linear relationship to photosynthetic income whether [CO₂] or I was varied. For variation in I, photosynthetic capacity varied directly with soluble protein per unit area. This was not the case for variation in [CO₂]. Increasing [CO₂] reduced photosynthetic capacity per unit of soluble protein by up to a factor of 2.5, and photosynthetic capacity exhibited an optimum with respect to growth [CO₂]. Thus CO₂ did not alter the relationship between photosynthetic income and the utilization of photosynthate in the net accumulation of soluble protein, but did alter the relationship between soluble protein content and photosynthetic characteristics in this species.     

Control of steady-state photosynthesis in sunflowers growing in enhanced CO2

Control coefficients were used to describe the degree to which ribulose bisphosphate carboxylase/oxygenase (Rubisco) limits the steady-state rate of CO₂ assimilation in sunflower leaves from plants grown at high (800 μmol mol⁻¹) and low (350 μmol mol⁻¹) CO₂. The magnitude of a control coefficient is approximately the percentage change in the flux that would result from a 1% rise in enzyme active site concentration. In plants grown at low CO₂, leaves of different ages varied considerably in their photosynthetic capacities. In a saturating light flux and an ambient CO₂ concentration of 350 μmol mol⁻¹, the Rubisco control coefficient was about 0.7 in all leaves, indicating that Rubisco activity largely limited the assimilation flux. The Rubisco control coefficient for leaves grown at 350 μmol mol⁻¹ CO₂ dropped to about zero when the ambient CO₂ concentration was raised to 800 μmol mol⁻¹. In relatively young, fully expanded leaves of plants grown at high CO₂, the Rubisco control coefficient was also about 0.7 at a saturating light flux and at the CO₂ concentration at which the plants were grown (800 μmol mol⁻¹). This apparently resulted from a decrease in the concentration of Rubisco active sites. In older leaves, however, the control coefficient was about 0.2. Because, on the whole, Rubisco activity still largely limits the assimilation flux in plants grown at high CO₂, the kinetics of this enzyme can still be used to model photosynthesis under these conditions. The relatively high Rubisco control coefficient under enhanced CO₂ indicates that the young sunflower leaves have the capacity to acclimate their photosynthetic biochemistry in a way consistent with an optimal use of protein resources.     

Photosynthetic acclimation and photosynthate partitioning in soybean leaves in response to carbon dioxide enrichment

Photosynthetic rates and photosynthate partitioning were studied in three-week-old soybean [Glycine max (L.) Merr. cv. Williams] plants exposed to either ambient (35 Pa) or elevated (70 Pa) CO₂ in controlled environment chambers. Ambient CO₂-grown plants also were given a single 24 h treatment with 70 Pa CO₂ 1 d prior to sampling. Photosynthetic rates of ambient CO₂-grown plants initially increased 36% when the measurement CO₂ was doubled from 35 to 70 Pa. Photosynthetic rates of the third trifoliolate leaf, both after 1 and 21 d of elevated CO₂ treatment, were 30 to 45% below those of ambient CO₂-grown plants when measured at 35 Pa CO₂. These reduced photosynthetic rates were not due to increased stomatal resistance and were observed for 2 to 8 h after plants given 1 d of CO₂ enrichment were returned to ambient CO₂. Initial and total ribulose 1,5-bisphosphate carboxylase/oxygenase (Rubisco) activities, percent activation, Rubisco protein, soluble protein and leaf chlorophyll content were similar in all CO₂ treatments. Quantum yields of photosynthesis, determined at limiting irradiances and at 35 Pa CO₂, were 0.049±0.003 and 0.038±0.005 mol CO₂ fixed per mol quanta for ambient and elevated CO₂-grown plants, respectively (p<0.05). Leaf starch and sucrose levels were greater in plants grown at 70 than at 35 Pa CO₂. Starch accumulation rates during the day were greater in ambient CO₂-grown plants than in plants exposed to elevated CO₂ for either 1 or 21 d. However, the percentage of C partitioned to starch relative to total C fixed was unaffected by 1 d of CO₂ enrichment. The above results showed that both photosynthetic and starch accumulation rates of soybean leaflets measured at 35 Pa CO₂ were temporarily reduced after 1 and 21 d of CO₂ enrichment. The biochemical mechanism affecting these responses was not identified.Abbreviations SLW- specific leaf weight (g m^–2) - Rubisco- ribulose 1,5-bisphosphate carboxylase/oxygenase - Rul- 5bisP, ribulose 1,5 bisphosphate - DAP- days after planting - SAR- starch accumulation rate - C_i- intercellular CO₂ concentration     

Growth and Photosynthetic Carbon Metabolism in Tobacco Plants under an Oscillating CO2 Concentration in the Atmosphere

Abstract: The hypothesis for the present work was that photosynthetic acclimation to increased atmospheric CO₂ in Nicotiana tabacum could be prevented by an oscillating supply of CO₂. This was tested by growing half of the plants (for the 20 day period after sowing) at 700 μmol mol^‐1 CO₂ (S+ plants) and half at 350 μmol mol^‐1 CO₂ (S‐ plants) and thereafter switching them every 48 h from high to low CO₂ and vice versa. These plants were compared with plants continuously kept (from sowing onwards) at 350 μmol mol^‐1 CO₂ (C‐ plants) and 700 μmol mol^‐1 CO₂ (C+ plants). Switching plants from high to low CO₂ and vice versa (S+ and S‐) did not improve plant growth efficiency, as hypothesized. The extra carbon fixed by the leaves under increased CO₂ in the atmosphere, supplied either continuously or intermittently, was mostly stored as starch and not used to build additional structural biomass. The differences in final plant biomass, observed between S+ and S‐ plants, are explained by the CO₂ concentration in the atmosphere during the first 20 days after sowing, the oscillation in CO₂ supply thereafter is playing a smaller role in this response. Switching plants from high to low CO₂ and vice versa, also did not prevent down‐regulation of photosynthesis, despite lower leaf sugar concentrations than in C+ plants. Nitrate concentration decreased dramatically in C+, S+ and S‐ plants. The leaf C/N ratio was highest in C+ plants (ranging from 8 to 13), intermediate in S+ and S‐ plants (from 7 to 11) and lowest in C‐ plants (from 6 to 8). This supports the view that the balance between carbohydrates and nitrogen may have a triggering role in plant response under elevated CO₂. Carbon export rates by the leaves seem to be independent of total carbon assimilation, suggesting a sink limiting effect on tobacco growth and phototsynthesis under elevated CO₂.     

Is photosynthetic acclimation to free-air CO<Subscript>2</Subscript> enrichment (FACE) related to a strong competition for the assimilatory power between carbon assimilation and nitrogen assimilation in rice leaf?

Net photosynthetic rate (P _N) of leaves grown under free-air CO₂ enriched condition (FACE, about 200 μmol mol⁻¹ above ambient air) was significantly lower than P _N of leaves grown at ambient CO₂ concentration (AC) when measured at CO₂ concentration of 580 μmol mol⁻¹. This difference was found in rice plants grown at normal nitrogen supply (25 g m⁻²; NN-plants) but not in plants grown at low nitrogen supply (15 g m⁻²; LN-plants). Namely, photosynthetic acclimation to FACE was observed in NN-plants but not in LN-plants. Different from the above results measured in a period of continuous sunny days, such photosynthetic acclimation occurred in NN-plants, however, it was also observed in LN-plants when P _N was measured before noon of the first sunny day after rain. Hence strong competition for the assimilatory power between nitrogen (N) and carbon (C) assimilations induced by an excessive N supply may lead to the photosynthetic acclimation to FACE in NN-plants. The hypothesis is supported by the following facts: FACE induced significant decrease in both apparent photosynthetic quantum yield (Φ_c) and ribulose-1,5-bisphosphate (RuBP) content in NN-plants but not in LN-plants.     

Effects of nitrogen supply and elevated carbon dioxide on construction cost in leaves of Pinus taeda (L.) seedlings

Seedlings of loblolly pine (Pinus taeda L.) were grown under varying conditions of soil nitrogen and atmospheric carbon dioxide availability to investigate the interactive effects of these resources on the energetic requirements for leaf growth. Increasing the ambient CO₂ partial pressure from 35 to 65 Pa increased seedling growth only when soil nitrogen was high. Biomass increased by 55% and photosynthesis increased by 13% after 100 days of CO₂ enrichment. Leaves from seedlings grown in high soil nitrogen were 7.0% more expensive on a g glucose g^–1 dry mass basis to produce than those grown in low nitrogen, while elevated CO₂ decreased leaf cost by 3.5%. Nitrogen and CO₂ availability had an interactive effect on leaf construction cost expressed on an area basis, reflecting source-sink interactions. When both resources were abundant, leaf construction cost on an area basis was relatively high (81.8±3.0 g glucose m^–2) compared to leaves from high nitrogen, low CO₂ seedlings (56.3±3.0 g glucose m^–2) and low nitrogen, low CO₂ seedlings (67.1±2.7 g glucose m^–2). Leaf construction cost appears to respond to alterations in the utilization of photoassimilates mediated by resource availability.     

Photosynthetic responses to elevated CO2and O3 in field-grown potato(Solanum tuberosum)

Potato plants (Solanum tuberosum L. cv. Bintje) were grown to maturity in open-top chambers under three carbon dioxide (CO₂; ambient and 24 h d⁻¹ seasonal mean concentrations of 550 and 680 μmol mol⁻¹) and two ozone levels (O₃; ambient and an 8 h d⁻¹ seasonal mean of 50 nmol mol⁻¹). Chlorophyll content, photosynthetic characteristics, and stomatal responses were determined to test the hypothesis that elevated atmospheric CO₂ may alleviate the damaging influence of O₃ by reducing uptake by the leaves. Elevated O₃ had no detectable effect on photosynthetic characteristics, leaf conductance, or chlorophyll content, but did reduce SPAD values for leaf 15, the youngest leaf examined. Elevated CO₂ also reduced SPAD values for leaf 15, but not for older leaves; destructive analysis confirmed that chlorophyll content was decreased. Leaf conductance was generally reduced by elevated CO₂, and declined with time in the youngest leaves examined, as did assimilation rate (A). A generally increased under elevated CO₂, particularly in the older leaves during the latter stages of the season, thereby increasing instantaneous transpiration efficiency. Exposure to elevated CO₂ and/or O₃ had no detectable effect on dark-adapted fluorescence, although the values decreased with time. Analysis of the relationships between assimilation rate and intercellular CO₂ concentration and photosynthetically active photon flux density showed there was initially little treatment effect on CO₂-saturated assimilation rates for leaf 15. However, the values for plants grown under 550 μmol mol⁻¹ CO₂ were subsequently greater than in the ambient and 680 μmol mol⁻¹ treatments, although the beneficial influence of the former treatment declined sharply towards the end of the season. Light-saturated assimilation was consistently greater under elevated CO₂, but decreased with time in all treatments. The values decreased sharply when leaves grown under elevated CO₂ were measured under ambient CO₂, but increased when leaves grown under ambient CO₂ were examined under elevated CO₂. The results obtained indicate that, although elevated CO₂ initially increased assimilation and growth, these beneficial effects were not necessarily sustained to maturity as a result of photosynthetic acclimation and the induction of earlier senescence.     

Estimation of the whole-plant CO2 compensation point of tobacco (Nicotiana tabacum L.)

The whole-plant CO₂ compensation point (Γ_plant) is the minimum atmospheric CO₂ level required for sustained growth. The minimum CO₂ requirement for growth is critical to understanding biosphere feedbacks on the carbon cycle during low CO₂ episodes; however, actual values of Γ_plant remain difficult to calculate. Here, we have estimated Γ_plant in tobacco by measuring the relative leaf expansion rate at several low levels of atmospheric CO₂, and then extrapolating the leaf growth vs. CO₂ response to estimate CO₂ levels where no growth occurs. Plants were grown under three temperature treatments, 19/15, 25/20 and 30/25°C day/night, and at CO₂ levels of 100, 150, 190 and 270 μmol CO₂ mol⁻¹ air. Biomass declined with growth CO₂ such that Γ_plant was estimated to be approximately 65 μmol mol⁻¹ for plants grown at 19/15 and 30/25°C. In the first 19 days after germination, plants grown at 100 μmol mol⁻¹ had low growth rates, such that most remained as tiny seedlings (canopy size <1 cm²). Most seedlings grown at 150 μmol mol⁻¹ and 30/25°C also failed to grow beyond the small seedling size by day 19. Plants in all other treatments grew beyond the small seedling size within 3 weeks of planting. Given sufficient time (16 weeks after planting) plants at 100 μmol mol⁻¹ eventually reached a robust size and produced an abundance of viable seed. Photosynthetic acclimation did not increase Rubisco content at low CO₂. Instead, Rubisco levels were unchanged except at the 100 and 150 μmol mol⁻¹ where they declined. Chlorophyll content and leaf weight per area declined in the same proportion as Rubisco, indicating that leaves became less expensive to produce. From these results, we conclude that the effects of very low CO₂ are most severe during seedling establishment, in large part because CO₂ deficiency slows the emergence and expansion of new leaves. Once sufficient leaf area is produced, plants enter the exponential growth phase and acquire sufficient carbon to complete their life cycle, even under warm conditions (30/25°C) and CO₂ levels as low as 100 μmol mol⁻¹.     

Elevated CO2 and plant nitrogen-use: is reduced tissue nitrogen concentration size-dependent?

Plants often respond to elevated atmospheric CO₂ levels with reduced tissue nitrogen concentrations relative to ambient CO₂-grown plants when comparisons are made at a common time. Another common response to enriched CO₂ atmospheres is an acceleration in plant growth rates. Because plant nitrogen concentrations are often highest in seedlings and subsequently decrease during growth, comparisons between ambient and elevated CO₂-grown plants made at a common time may not demonstrate CO₂-induced reductions in plant nitrogen concentration per se. Rather, this comparison may be highlighting differences in nitrogen concentration between bigger, more developed plants and smaller, less developed plants. In this study, we directly examined whether elevated CO₂ environments reduce plant nitrogen concentrations independent of changes in plant growth rates. We grew two annual plant species. Abutilon theophrasti (C₃ photosynthetic pathway) and Amaranthus retroflexus (C₄ photosynthetic pathway), from seed in glass-sided growth chambers with atmospheric CO₂ levels of 350 mol·mol^–1 or 700 mol·mol^–1 and with high or low fertilizer applications. Individual plants were harvested every 2 days starting 3 days after germination to determine plant biomass and nitrogen concentration. We found: 1. High CO₂-grown plants had reduced nitrogen concentrations and increased biomass relative to ambient CO₂-grown plants when compared at a common time; 2. Tissue nitrogen concentrations did not vary as a function of CO₂ level when plants were compared at a common size; and 3. The rate of biomass accumulation per rate of increase in plant nitrogen was unaffected by CO₂ availability, but was altered by nutrient availability. These results indicate that a CO₂-induced reduction in plant nitrogen concentration may not be due to physiological changes in plant nitrogen use efficiency, but is probably a size-dependent phenomenon resulting from accelerated plant growth.