Consequences of growth at two carbon dioxide concentrations and two temperatures for leaf gas exchange in Pascopyrum smithii (C3) and Bouteloua gracilis (C4)*

Continually rising atmospheric CO₂ concentrations and possible climatic change may cause significant changes in plant communities. This study was undertaken to investigate gas exchange in two important grass species of the short-grass steppe, Pascopyrum smithii (western wheat-grass), C₃, and Bouteloua gracilis (blue grama), C4, grown at different CO₂ concentrations and temperatures. Intact soil cores containing each species were extracted from grasslands in north-eastern Colorado, USA, placed in growth chambers, and grown at combinations of two CO₂ concentrations (350 and 700 μmol mol⁻¹) and two temperature regimes (field average and elevated by 4°C). Leaf gas exchange was measured during the second, third and fourth growth seasons. All plants exhibited higher leaf CO₂ assimilation rates (A) with increasing measurement CO₂ concentration, with greater responses being observed in the cool-season C₃ species P. smithii. Changes in the shape of intercellular CO₂ response curves of A for both species indicated photosynthetic acclimation to the different growth environments. The photosynthetic capacity of P. smithii leaves tended to be reduced in plants grown at high CO₂ concentrations, although A for plants grown and measured at 700μmol mol⁻¹ CO₂ was 41% greater than that in plants grown and measured at 350 μmol mol⁻¹ CO₂. Low leaf N concentration may have contributed to photosynthetic acclimation to CO₂. A severe reduction in photosynthetic capacity was exhibited in P. smithii plants grown long-term at elevated temperatures. As a result, the potential response of photosynthesis to CO₂ enrichment was reduced in P. smithii plants grown long-term at the higher temperature.     

Effects of elevated carbon dioxide concentration on photosynthesis and growth of small birch plants (Betula pendula Roth.) at optimal nutrition

Small birch plants (Betula pendula Roth.) were grown from seed for periods of up to 70d in a climate chamber at optimal nutrition and at present (350 μmol mol^?1) or elevated (700 μmol mol^?1) concentrations of atmospheric CO₂. Nutrients were sprayed over the roots in Ingestad-type units. Relative growth rate and net assimilation rate were slightly higher at elevated CO₂, whereas leaf area ratio was slightly lower. Smaller leaf area ratio was associated with lower values of specific leaf area. Leaves grown at elevated CO₂ had higher starch concentrations (dry weight basis) than leaves grown at present levels of CO₂. Biomass allocation showed no change with CO₂, and no large effects on stem height, number of side shoots and number of leaves were found. However, the specific root length of fine roots was higher at elevated CO₂. No large difference in the response of carbon assimilation to intercellular CO₂ concentration (A/C_i curves) were found between CO₂ treatments. When measured at the growth environments, the rates of photosynthesis were higher in plants grown at elevated CO₂ than in plants grown at present CO₂. Water use efficiency of single leaves was higher in the elevated treatment. This was mainly attributable to higher carbon assimilation rate at elevated CO₂. The difference in water use efficiency diminished with leaf age. The small treatment difference in relative growth rate was maintained throughout the experiment, which meant that the difference in plant size became progressively greater. Thus, where plant nutrition is sufficient to maintain maximum growth, small birch plants may potentially increase in size more rapidly at elevated CO₂.     

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

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

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

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

Infection with the parasitic angiosperm Striga hermonthica influences the response of the C3 cereal Oryza sativa to elevated CO2

Upland rice (Oryza sativa L.) was grown at both ambient (350 μmol mol^?1) and elevated (700 μmol mol^?1) CO₂ in either the presence or absence of the root hemi‐parasitic angiosperm Striga hermonthica (Del) Benth. Elevated CO₂ alleviated the impact of the parasite on host growth: biomass of infected rice grown at ambient CO₂ was 35% that of uninfected, control plants, while at elevated CO₂, biomass of infected plants was 73% that of controls. This amelioration occurred despite the fact that O. sativa grown at elevated CO₂ supported both greater numbers and a higher biomass of parasites per host than plants grown at ambient CO₂. The impact of infection on host leaf area, leaf mass, root mass and reproductive tissue mass was significantly lower in plants grown at elevated as compared with ambient CO₂. There were significant CO₂ and Striga effects on photosynthetic metabolism and instantaneous water‐use efficiency of O. sativa. The response of photosynthesis to internal [CO₂] (A/C_i curves) indicated that, at 45 days after sowing (DAS), prior to emergence of the parasites, uninfected plants grown at elevated CO₂ had significantly lower CO₂ saturated rates of photosynthesis, carboxylation efficiencies and ribulose‐1,5‐bisphosphate carboxylase/oxygenase (Rubisco; EC 4.1.1.39) contents than uninfected, ambient CO₂‐grown O. sativa. In contrast, infection with S. hermonthica prevented down‐regulation of photosynthesis in O. sativa grown at elevated CO₂, but had no impact on photosynthesis of hosts grown at ambient CO₂. At 76 DAS (after parasites had emerged), however, infected plants grown at both elevated and ambient CO₂ had lower carboxylation efficiencies and Rubisco contents than uninfected O. sativa grown at ambient CO₂. The reductions in carboxylation efficiency (and Rubisco content) were accompanied by similar reductions in nitrogen concentration of O. sativa leaves, both before and after parasite emergence. There were no significant CO₂ or infection effects on the concentrations of soluble sugars in leaves of O. sativa, but starch concentration was significantly lower in infected plants at both CO₂ concentrations. These results demonstrate that elevated CO₂ concentrations can alleviate the impact of infection with Striga on the growth of C₃ hosts such as rice and also that infection can delay the onset of photosynthetic down‐regulation in rice grown at elevated CO₂.     

How is the relationship between the C4 cereal Sorghum bicolor and the C3 root hemi-parasites Striga hermonthica and Striga asiatica affected by elevated CO2?

The C₄ cereal Sorghum bicolor was grown under either ambient (350 μmol mol^?1) or elevated (700 μmol mol^?1) [CO₂] in either the presence or absence of the C₃ obligate root hemi-parasites Striga hermonthica or S. asiatica. Both uninfected and infected sorghum plants were taller and had greater biomass, photosynthetic rates, water-use efficiencies and leaf areas under elevated compared with ambient [CO₂]. There was no evidence of any downregula-tion of photosynthesis in sorghum grown at elevated [CO₂]. Biomass of infected sorghum was lower under both ambient and elevated [CO₂], and although infected plants were larger under elevated [CO₂] the relative impact of infection on host biomass was either the same (S. asiatica) or only slightly less (S. hermonthica) than under ambient [CO₂]. In contrast, biomass of S. hermonthica and S. asiatica per host was lower under elevated than ambient [CO₂], although rates of photosynthesis were higher at elevated [CO₂] and parasite stomatal conductance was not responsive to [CO₂]. Parasites emerged above-ground and flowered earlier under ambient compared with elevated [CO₂]. It appears that the mechanism(s) by which the parasites affect host growth is (are) relatively insensitive to increased atmospheric [CO₂], although the parasites themselves were adversely affected by growth at elevated [CO₂].     

Interactive effects of nitrogen and phosphorus on the acclimation potential of foliage photosynthetic properties of cork oak,Quercus suber,to elevated atmospheric CO2 concentrations

Leaf gas-exchange and chemical composition were investigated in seedlings of Quercus suber L. grown for 21 months either at elevated (700 μmol mol^–1) or normal (350 μmol mol^–1) ambient atmospheric CO₂ concentrations, [CO₂], in a sandy nutrient-poor soil with either ‘high’ N (0.3 mol N m^–3 in the irrigation solution) or with ‘low’ N (0.05 mol N m^–3) and with a constant suboptimal concentration of the other macro- and micronutrients. Although elevated [CO₂] yielded the greatest total plant biomass in ‘high’ nitrogen treatment, it resulted in lower leaf nutrient concentrations in all cases, independent of the nutrient addition regime, and in greater nonstructural carbohydrate concentrations. By contrast, nitrogen treatment did not affect foliar N concentrations, but resulted in lower phosphorus concentrations, suggesting that under lower N, P use-efficiency in foliar biomass production was lower. Phosphorus deficiency was evident in all treatments, as photosynthesis became CO₂ insensitive at intercellular CO₂ concentrations larger than ≈ 300 μmol mol^–1, and net assimilation rates measured at an ambient [CO₂] of 350 μmol mol^–1 or at 700 μmol mol^–1 were not significantly different. Moreover, there was a positive correlation of foliar P with maximum Rubisco (Ribulose-1,5-bisphosphate carboxylase/oxygenase) carboxylase activity (V_cmax), which potentially limits photosynthesis at low [CO₂], and the capacities of photosynthetic electron transport (J_max) and phosphate utilization (P_max), which are potentially limiting at high [CO₂]. None of these potential limits was correlated with foliar nitrogen concentration, indicating that photosynthetic N use-efficiency was directly dependent on foliar P availability. Though the tendencies were towards lower capacities of potential limitations of photosynthesis in high [CO₂] grown specimens, the effects were statistically insignificant, because of (i) large within-treatment variability related to foliar P, and (ii) small decreases in P/N ratio with increasing [CO₂], resulting in balanced changes in other foliar compounds potentially limiting carbon acquisition. The results of the current study indicate that under P-deficiency, the down-regulation of excess biochemical capacities proceeds in a similar manner in leaves grown under normal and elevated [CO₂], and also that foliar P/N ratios for optimum photosynthesis are likely to increase with increasing growth CO₂ concentrations. Symbols: A, net assimilation rate (μmol m^–2 s^–1); A_max, light-saturated A (μmol m^–2 s^–1); α, initial quantum yield at saturating [CO₂] and for an incident Q (mol mol^–1); [CO₂], atmospheric CO₂ concentration (μmol mol^–1); C_i, intercellular CO₂ concentration (μmol mol^–1); C_a, CO₂ concentration in the gas-exchange cuvette (μmol mol^–1); F_B, fraction of leaf N in ‘photoenergetics’; F_L, fraction of leaf N in light harvesting; F_R, fraction of leaf N in Rubisco; Γ*, CO₂ compensation concentration in the absence of R_d (μmol mol^–1); J_max*, capacity for photosynthetic electron transport; J_mc, capacity for photosynthetic electron transport per unit cytochrome f (mol e^–[mol cyt f]^–1 s^–1); K_c, Michaelis-Menten constant for carboxylation (μmol mol^–1); K_o, Michaelis-Menten constant for oxygenation (mmol mol^–1); M_A, leaf dry mass per area (g m^–2); O, intercellular oxygen concentration (mmol mol^–1); [P_i], concentration of inorganic phosphate (mM); P_max*, capacity for phosphate utilization; Q, photosynthetically active quantum flux density (μmol m^–2 s^–1); R_d*, day respiration (CO₂ evolution from nonphotorespiratory processes continuing in the light); Rubisco, ribulose-1,5-bisphosphate carboxylase/oxygenase; RUBP, ribulose-1,5-bisphosphate; T_l, leaf temperature (°C); U_TPU*, rate of triose phosphate utilization; V_cmax*, maximum Rubisco carboxylase activity; V_cr, specific activity of Rubisco (μmol CO₂[g Rubisco]^–1 s^–1] *given in either μmol m^–2 s^–1 or in μmol g^–1 s^–1 as described in the text.     

Direct and acclimatory responses of stomatal conductance to elevated carbon dioxide in four herbaceous crop species in the field

In order to separate the net effect of growth at elevated [CO₂] on stomatal conductance (g_s) into direct and acclimatory responses, mid‐day values of g_s were measured for plants grown in field plots in open‐topped chambers at the current ambient [CO₂], which averaged 350 μmol mol^?1 in the daytime, and at ambient + 350 μmol mol^?1[CO₂] for winter wheat, winter barley, potato and sorghum. The acclimatory response was determined by comparing g_s measured at 700 μmol mol^?1[CO₂] for plants grown at the two [CO₂]. The direct effect of increasing [CO₂] from 350 to 700 μmol mol^?1 was determined for plants grown at the lower concentration. Photosynthetic rates were measured concurrently with g_s. For all species, growth at the higher [CO₂] significantly reduced g_s measured at 700 μmol mol^?1[CO₂]. The reduction in g_s caused by growth at the higher [CO₂] was larger for all species on days with low leaf to air water vapour pressure difference for a given temperature, which coincided with highest conductances and also the smallest direct effects of increased [CO₂] on conductance. For barley, there was no other evidence for stomatal acclimation, despite consistent down‐regulation of photosynthetic rate in plants grown at the higher [CO₂]. In wheat and potato, in addition to the vapour pressure difference interaction, the magnitude of stomatal acclimation varied directly in proportion to the magnitude of down‐regulation of photosynthetic rate through the season. In sorghum, g_s consistently exhibited acclimation, but there was no down‐regulation of photosynthetic rate. In none of the species except barley was the direct effect the larger component of the net reduction in g_s when averaged over measurement dates. The net effect of growth at elevated [CO₂] on mid‐day g_s resulted from unique combinations of direct and acclimatory responses in the various species.     

Leaf senescence of Quercus myrtifolia as affected by long-term CO2 enrichment in its native environment

The long‐term effects of elevated (ambient plus 350 μmol mol^?1) atmospheric CO₂ concentration (C_a) on the leaf senescence of Quercus myrtifolia Willd was studied in a scrub‐oak community during the transition from autumn (December 1997) to spring (April 1998). Plants were grown in large open‐top chambers at the Smithsonian CO₂ Research Site, Merritt Island Wildlife Refuge, Cape Canaveral, Florida. Chlorophyll (a + b) concentration, Rubisco activity and N concentration decreased by 75%, 82%, and 52%, respectively, from December (1997) to April (1998) in the leaves grown at ambient C_a. In contrast, the leaves of plants grown at elevated C_a showed no significant decrease in chlorophyll (a + b) concentration or Rubisco activity, and only a 25% reduction in nitrogen. These results indicate that leaf senescence was delayed during this period at elevated C_a. Delayed leaf senescence in elevated C_a had important consequences for leaf photosynthesis. In elevated C_a the net photosynthetic rate of leaves that flushed in Spring 1997 (last year's leaves) and were 13 months old was not different from fully‐expanded leaves that flushed in 1998, and were approximately 1 month old (current year's leaves). In ambient C_a the net photosynthetic rate of last year's leaves was 54% lower than for current year's leaves. When leaves were fully senesced, nitrogen concentration decreased to about 40% of the concentration in non‐senesced leaves, in both CO₂ treatments. In April, net photosynthesis was 97% greater in leaves grown in elevated C_a than in those grown at ambient. During the period when elevated C_a delayed leaf senescence, more leaves operating at higher photosynthetic rate would allow the ecosystem dominated by Q. myrtifolia to gain more carbon at elevated C_a than at ambient C_a.     

Stomatal acclimation to increased CO2 concentration in a Florida scrub oak species Quercus myrtifolia Willd

Native scrub‐oak communities in Florida were exposed for three seasons in open top chambers to present atmospheric [CO₂] (approx. 350 μmol mol^?1) and to high [CO₂] (increased by 350 μmol mol^?1). Stomatal and photosynthetic acclimation to high [CO₂] of the dominant species Quercus myrtifolia was examined by leaf gas exchange of excised shoots. Stomatal conductance (g_s) was approximately 40% lower in the high‐ compared to low‐[CO₂]‐grown plants when measured at their respective growth concentrations. Reciprocal measurements of g_s in both high‐ and low‐[CO₂]‐grown plants showed that there was negative acclimation in the high‐[CO₂]‐grown plants (9–16% reduction in g_s when measured at 700 μmol mol^?1), but these were small compared to those for net CO₂ assimilation rate (A, 21–36%). Stomatal acclimation was more clearly evident in the curve of stomatal response to intercellular [CO₂] (c_i) which showed a reduction in stomatal sensitivity at low c_i in the high‐[CO₂]‐grown plants. Stomatal density showed no change in response to growth in high growth [CO₂]. Long‐term stomatal and photosynthetic acclimation to growth in high [CO₂] did not markedly change the 2·5‐ to 3‐fold increase in gas‐exchange‐derived water use efficiency caused by high [CO₂].     

Leaf photosynthesis and carbohydrate dynamics of soybeans grown throughout their life-cycle under Free-Air Carbon dioxide Enrichment

A lower than theoretically expected increase in leaf photosynthesis with long‐term elevation of carbon dioxide concentration ([CO₂]) is often attributed to limitations in the capacity of the plant to utilize the additional photosynthate, possibly resulting from restrictions in rooting volume, nitrogen supply or genetic constraints. Field‐grown, nitrogen‐fixing soybean with indeterminate flowering might therefore be expected to escape these limitations. Soybean was grown from emergence to grain maturity in ambient air (372 µmol mol^?1[CO₂]) and in air enriched with CO₂ (552 µmol mol^?1[CO₂]) using Free‐Air CO₂ Enrichment (FACE) technology. The diurnal courses of leaf CO₂ uptake (A) and stomatal conductance (g_s) for upper canopy leaves were followed throughout development from the appearance of the first true leaf to the completion of seed filling. Across the growing season the daily integrals of leaf photosynthetic CO₂ uptake (A′) increased by 24.6% in elevated [CO₂] and the average mid‐day g_s decreased by 21.9%. The increase in A′ was about half the 44.5% theoretical maximum increase calculated from Rubisco kinetics. There was no evidence that the stimulation of A was affected by time of day, as expected if elevated [CO₂] led to a large accumulation of leaf carbohydrates towards the end of the photoperiod. In general, the proportion of assimilated carbon that accumulated in the leaf as non‐structural carbohydrate over the photoperiod was small (< 10%) and independent of [CO₂] treatment. By contrast to A′, daily integrals of PSII electron transport measured by modulated chlorophyll fluorescence were not significantly increased by elevated [CO₂]. This indicates that A at elevated [CO₂] in these field conditions was predominantly ribulose‐1,5‐bisphosphate (RubP) limited rather than Rubisco limited. There was no evidence of any loss of stimulation toward the end of the growing season; the largest stimulation of A′ occurred during late seed filling. The stimulation of photosynthesis was, however, transiently lost for a brief period just before seed fill. At this point, daytime accumulation of foliar carbohydrates was maximal, and the hexose:sucrose ratio in plants grown at elevated [CO₂] was significantly larger than that in plants grown at current [CO₂]. The results show that even for a crop lacking the constraints that have been considered to limit the responses of C₃ plants to rising [CO₂] in the long term, the actual increase in A over the growing season is considerably less than the increase predicted from theory.     

Modification of the response of photosynthetic productivity to rising temperature by atmospheric CO2 concentrations: Has its importance been underestimated?

Abstract. Climate change will include correlated increases in temperature and atmospheric CO₂ concentration (C_a). Rising temperatures will increase the ratio of photorespiratory loss of carbon to photosynthetic gain, whilst rising C_a will have an opposing effect. The mechanism of these effects at the level of carboxylation in C₃ photosynthesis are quantitatively well understood and provide a basis for models of the response of leaf and canopy carbon exchange to climate change. The principles of such a model are referred to here and used to quantitatively examine the implications of concurrent increase in temperature and C_a. Simulations of leaf photosynthesis show the increase, with elevation of C_a from 350 to 650 μmol mol^-1, in light saturated rates of CO₂ uptake (A_sat) and maximum quantum yields (φ) to rise with temperature. An increase in C_a from 350 to 650 μmol mol^-1 can increase A_sat by 20% at 10°C and by 105% at 35°C, and can raise the temperature optimum of A_sat by 5°C. This pattern of change agrees closely with experimental data. At the canopy level, simulations also suggest a strong interaction of increased temperature and CO₂ concentration. Predictions are compared with the findings of long-term field studies. The principles used here suggest that elevated C_a will alter both the magnitude of the response of leaf and canopy carbon gain to rising temperature, and sometimes, the direction of response. Findings question the value of models for predicting plant production in response to climate change which ignore the direct effects of rising C_a and the modifications that rising C_a imposes on the temperature response of net CO₂ exchange.     

The impact of elevated CO2 on growth and photosynthesis in Agrostis canina L. ssp. monteluccii adapted to contrasting atmospheric CO2 concentrations

 The aim of this study was to characterise growth and photosynthetic capacity in plants adapted to long-term contrasting atmospheric CO₂ concentrations (C _a). Seeds of Agrostis canina L. ssp. monteluccii were collected from a natural CO₂ transect in central-western Italy and plants grown in controlled environment chambers at both ambient and elevated CO₂ (350 and 700 μmol mol⁻¹) in nutrient-rich soil. Seasonal mean C _a at the source of the plant material ranged from 610 to 451 μmol CO₂ mol⁻¹, derived from C₄ leaf stable carbon isotope discrimination (δ¹³C). Under chamber conditions, CO₂ enrichment stimulated the growth of all populations. However, plants originating from elevated C _a exhibited higher initial relative growth rates (RGRs) irrespective of chamber CO₂ concentrations and a positive relationship was found between RGR and C _a at the seed source. Seed weight was positively correlated with C _a, but differences in seed weight were found to explain no more than 34% of the variation in RGRs at elevated CO₂. Longer-term experiments (over 98 days) on two populations originating from the extremes of the transect (451 and 610 μmol CO₂ mol⁻¹) indicated that differences in growth between populations were maintained when plants were grown at both 350 and 700 μmol CO₂ mol⁻¹. Analysis of leaf material revealed an increase in the cell wall fraction (CWF) in plants grown at elevated CO₂, with plants originating from high C _a exhibiting constitutively lower levels but a variable response in terms of the degree of lignification. In vivo gas exchange measurements revealed no significant differences in light and CO₂ saturated rates of photosynthesis and carboxylation efficiency between populations or with CO₂ treatment. Moreover, SDS-PAGE/ LISA quantification of leaf ribulose bisphosphate carboxylase/oxygenase (Rubisco) showed no difference in Rubisco content between populations or CO₂ treatments. These findings suggest that long-term adaptation to growth at elevated CO₂ may be associated with a potential for increased growth, but this does not appear to be linked with differences in the intrinsic capacity for photosynthesis. Received: 16 August 1996 / Accepted: 19 October 1996     

Response of small birch plants (Betula pendula Roth.) to elevated CO2 and nitrogen supply

Small birch plants were grown for up to 80 d in a climate chamber at varied relative addition rates of nitrogen in culture solution, and at ambient (350 μmol mol^-1) or elevated (700 μmol mol^-1) concentrations of CO₂. The relative addition rate of nitrogen controlled relative growth rate accurately and independently of CO₂ concentration at sub-optimum levels. During free access to nutrients, relative growth rate was higher at elevated CO₂. Higher values of relative growth rate and net assimilation rate were associated with higher values of plant N-concentration. At all N-supply rates, elevated CO₂ resulted in higher values of net assimilation rate, whereas leaf weight ratio was independent of CO₂. Specific leaf area (and leaf area ratio) was less at higher CO₂ and at lower rates of N-supply. Lower values of specific leaf area were partly because of starch accumulation. Nitrogen productivity (growth rate per unit plant nitrogen) was higher at elevated CO₂. At sub-optimal N-supply, the higher net assimilation rate at elevated CO₂ was offset by a lower leaf area ratio. Carbon dioxide did not affect root/shoot ratio, but a higher fraction of plant dry weight was found in roots at lower N-supply. In the treatment with lowest N-supply, five times as much root length was produced per amount of plant nitrogen in comparison with optimum plants. The specific fine root length at all N-supplies was greater at elevated CO₂. These responses of the root system to lower N-supply and elevated CO₂ may have a considerable bearing on the acquisition of nutrients in depleted soils at elevated CO₂. The advantage of maintaining steady-state nutrition in small plants while investigating the effects of elevated CO₂ on growth is emphasized.     

Effect of CO2 enrichment on non-structural carbohydrates in leaves, stems and ears of spring wheat

Field-grown spring wheat (Triticum aestivum L. cv. Dragon) was exposed to ambient and elevated CO₂ concentrations (1.5 and 2 times ambient) in open-top chambers. Contents of non-structural carbohydrates were analysed enzymatically in leaves, stems and ears six times during the growing season. The impact of elevated CO₂ on wheat carbohydrates was non-significant in most harvests. However, differences in the carbohydrate contents due to elevated CO₂ were found in all plant compartments. Before anthesis, at growth stage (GS) 30 (the stem is 1 cm to the shoot apex), the plants grown in elevated CO₂ contained significantly more water soluble carbohydrates (WSC), fructans, starch and total non-structural carbohydrates (TNC) in the leaves in comparison with the plants grown in ambient CO₂. It is hypothesised that the plants from the treatments with elevated CO₂ were sink-limited at GS30. After anthesis, the leaf WSC and TNC contents of the plants from elevated CO₂ started to decline earlier than those of the plants from ambient CO₂. This may indicate that the leaves of plants grown in the chambers with elevated CO₂ senesced earlier. Elevated CO₂ accelerated grain development: 2 weeks after anthesis, the plants grown in elevated CO₂ contained significantly more starch and significantly less fructans in the ears compared to the plants grown in ambient CO₂. Elevated CO₂ had no effect on ear starch and TNC contents at the final harvest. Increasing the CO₂ concentration from 360 to 520 μmol mol^?1 had a larger effect on wheat non-structural carbohydrates than the further increase from 520 to 680 μmol mol^?1. The results are discussed in relation to the effects of elevated CO₂ on yield and yield components.     

Atmospheric carbon dioxide and fertilizer nitrogen effects on radiation interception by rice 1

In order to predict the potential impacts of global change, it is important to understand the impact of increasing global atmospheric [CO₂] on the growth and yield of crop plants. The objectives of this study were to determine the interaction of N fertilization rates and atmospheric [CO₂] on radiation interception and radiation-use efficiency of rice (Oryza sativa L. cv. IR72) grown under tropical field conditions. Rice plants were grown inside open top chambers in a lowland rice field at the International Rice Research Institute in the Philippines at ambient (about 350 μmol mol^-1) or elevated (about 600 μmol mol^-1 during the 1993 wet season and 700 μmol mol^-1 during the 1994 dry season) in combination with three levels of applied N (0, 50 or 100 kg N ha^-1 in the wet season; 0, 90 or 200 kg N ha-1 in the dry season). Light interception was not directly affected by [CO₂], but elevated [CO₂] indirectly increased light interception through increasing total absorbed N. Plant N requirement for radiation interception was similar for rice grown under ambient [CO₂] or elevated [CO₂] treatments. The conversion efficiency of intercepted radiation to dry matter, radiation-use efficiency (RUE), was about 35% greater at elevated [CO₂] than at ambient [CO₂]. The relationship between leaf N and RUE was curvilinear. At ambient [CO₂], RUE was fairly stable across levels of leaf N, but leaf N less than about 2.5% resulted in lower RUE for plants grown with elevated [CO₂] than for plant grown at ambient [CO₂]. Decreased leaf N with increased [CO₂], therefore decreased RUE of rice plants grown at elevated [CO₂]. When predicting responses of rice to elevated [CO₂], RUE should be adjusted with a decrease in leaf N. This revised version was published online in June 2006 with corrections to the Cover Date.     

A meta-analysis of leaf gas exchange and nitrogen in trees grown under elevated carbon dioxide

The response of trees to rising atmospheric CO₂ concentration ([CO₂]) is of concern to forest ecologists and global carbon modellers and is the focus of an increasing body of research work. I review studies published up to May 1994, and several unpublished works, which reported at least one of the following: net CO₂ assimilation (A), stomatal conductance (g_s), leaf dark respiration (R_d) leaf nitrogen or specific leaf area (SLA) in woody plants grown at <400 μmol mol^?1 CO₂ or at 600–800 μmol mol^?1 CO₂. The resulting data from 41 species were categorized according to growth conditions (unstressed versus stressed), length of CO₂ exposure, pot size and exposure facility [growth chamber (GC), greenhouse (GH), or open-top chamber (OTC)] and interpreted using meta-analytic methods. Overall, A showed a large and significant increase at elevated [CO₂] but length of CO₂ exposure and the exposure facility were important modifiers of this response. Plants exposed for < 50 d had a significantly greater response, and those from GCs had a significantly lower response than plants from longer exposures or from OTC studies. Negative acclimation of A was significant and general among stressed plants, but in unstressed plants was influenced by length of CO₂ exposure, the exposure facility and/or pot size. Growth at elevated [CO₂] resulted in moderate reductions in gs in unstressed plants, but there was no significant effect of CO₂ on gs in stressed plants. Leaf dark respiration (mass or area basis) was reduced strongly by growth at high [CO₂] > while leaf N was reduced only when expressed on a mass basis. This review is the first meta-analysis of elevated CO₂ studies and provides statistical confirmation of several general responses of trees to elevated [CO₂]. It also highlights important areas of continued uncertainty in our understanding of these responses.     

Effects of elevated CO2 on growth,photosynthesis and respiration of sweet chestnut (Castanea sativa Mill.)

Two year old sweet chestnut seedlings (Castanea sativa Mill) were grown in pots at ambient (350 µmol·mol^–1) and double (700 µmol·mol^–1) atmospheric CO₂ concentration in constantly ventilated greenhouses during entire growing seasons. CO₂ enrichment caused either no significant change or a decrease in shoot response, depending on yearly weather conditions. Similarly, leaf area was either reduced or unchanged under elevated CO₂. However, when grown under controlled conditions in a growth chamber, leaf area was enlarged with elevated CO₂.The CO₂ exchanges of whole plants were measured during the growing season. In elevated CO₂, net photosynthetic rate was maximum in May and then decreased, reaching the level of the control at the end of the season. End of night dark respiration of enriched plants was significantly lower than that of control plants; this difference decreased with time and became negligible in the fall. The original CO₂ level acted instantaneously on the respiration rate: a double concentration in CO₂ decreased the respiration of control plants and a reduced concentration enhanced the respiration of enriched plants. The carbon balance of a chestnut seedling may then be modified in elevated CO₂ by increased carbon inputs and decreased carbon outputs.     

Photosynthetic downregulation in leaves of the Japanese white birch grown under elevated CO2 concentration does not change their temperature‐dependent susceptibility to photoinhibition

To determine the effects of elevated CO₂ concentration ([CO₂]) on the temperature‐dependent photosynthetic properties, we measured gas exchange and chlorophyll fluorescence at various leaf temperatures (15, 20, 25, 30, 35 and 40°C) in 1‐year‐old seedlings of the Japanese white birch (Betula platyphylla var. japonica), grown in a phytotron under natural daylight at two [CO₂] levels (ambient: 400 µmol mol^?1 and elevated: 800 µmol mol^?1) and limited N availability (90 mg N plant^?1). Plants grown under elevated [CO₂] exhibited photosynthetic downregulation, indicated by a decrease in the carboxylation capacity of Rubisco. At temperatures above 30°C, the net photosynthetic rates of elevated‐CO₂‐grown plants exceeded those grown under ambient [CO₂] when compared at their growth [CO₂]. Electron transport rates were significantly lower in elevated‐CO₂‐grown plants than ambient‐CO₂‐grown ones at temperatures below 25°C. However, no significant difference was observed in the fraction of excess light energy [(1 ? q_P)× F_v′/F_m′] between CO₂ treatments across the temperature range. The quantum yield of regulated non‐photochemical energy loss was significantly higher in elevated‐CO₂‐grown plants than ambient, when compared at their respective growth [CO₂] below 25°C. These results suggest that elevated‐CO₂‐induced downregulation might not exacerbate the temperature‐dependent susceptibility to photoinhibition, because reduced energy consumption by electron transport was compensated for by increased thermal energy dissipation at low temperatures.