Simulated influence of altitude on photosynthetic CO2 uptake potential in plants

Abstract. A simulation of the quantitative influence of altitude on photosynthetic CO₂ uptake capability (A^P) included the effects of predicted changes (1) in air temperature (lapse rate) and (2) leaf temperature, (3) ambient pressure and CO₂ concentration, and (4) the diffusion coefficient for CO² in air. When a dry lapse rate (0.01°C m⁻¹) in air temperature was simulated, significant declines (up to 14%) in A^P were predicted from sea level to 4km altitude. A moist lapse rate of 0.003°C m⁻¹ resulted in less than a 4% decrease in A^P over the same altitude range. When natural leaf temperatures (predicted from heat balance analyses) were simulated, A^P was significantly greater (∼20%) than when leaf temperatures were considered equal to air temperature for all lapse conditions. There was virtually no change in A^P with altitude when predicted leaf temperatures and moist lapse conditions were simulated. There was a significant (∼10%) increase in A^P with altitude when leaf temperature was held constant at 30°C (regardless of altitude) under moist lapse conditions. Future studies evaluating the effects of elevation on photosynthesis could benefit from the above considerations of the effects of natural leaf temperature regimes and prevailing lapse conditions on CO₂ uptake potential.     

Drought-induced shifts in daily CO2 uptake patterns for leafy cacti

For cacti with persistent, relatively large leaves, most shoot CO₂ uptake under well-watered conditions occurs by the leaves using the C₃ pathway. For three species in the primitive subfamily Pereskioideae, droughts of 7 or 14 days decreased leaf daytime net CO₂ uptake by an average of 49 and 88%, respectively; these species always had a net CO₂ release at night by the leaves and both at night and during the day by the stems. For three leafy species in subfamily Opuntioideae, 7 and 14 days of drought reduced leaf daytime net CO₂ uptake by 90 and 100%, respectively. Although drought reduced the total CO₂ uptake over 24 h, the average percentage occurring at night by the leaves of these species increased from 5% under wet conditions to 71% after 7 days of drought to 99% after 14 days of drought. For two of the three species of Opuntioideae, 7 days of drought caused the small net CO₂ uptake by the sterns to shift from the daytime to the nighttime, while for the third species drought caused a reduction of its stem nocturnal net CO₂ uptake. Thus, shifts from predominantly daytime to predominantly nighttime net CO₂ uptake can be induced by drought for the leaves and the stems of leafy cacti in subfamily Opuntioideae, indicating a high degree of biochemical versatility.     

Hypometabolism in torpid goldsinny wrasse subjected to rapid reductions in seawater temperature

Goldsinny Ctenolabrus rupestris were subjected to rapid, environmentally realistic, reductions in temperature at 2° C increments from 10 to 4° C over a 3-day period in full-strength sea water. In separate experiments, oxygen uptake measurements and ultrasound recordings of heart rate and opercular motion were carried out at regular intervals over the same temperature regime. Mean oxygen uptake rates fell from 0.042 to 0.028 ml O₂ g⁻¹ h⁻¹ between 10 and 6° C respectively (Q₁₀=2.71). Between 6 and 4° C mean rates decreased from 0.028 to 0.008 ml O₂ g⁻¹ h⁻¹ (Q₁₀=542). Mean opercular motion and heart beat rates decreased from 49.5 and 60.3 beats min⁻¹ respectively at 10° C to 18.7 and 18.0 beats min⁻¹ respectively at 4° C. Most goldsinny subjected to 4° C were observed in a torpid state and would not react to external stimulation. Opercular motion was erratic at 4° C and would at times cease altogether for periods up to 1.3 min duration. Heart movement was diffcult to detect at 4° C and may also have ceased for prolonged periods. Q₁₀ values for opercular motion and heart beat rates recorded between 6 and 4° C were 6.39 and 24.52 respectively compared with values of 2.42 and 2.93 respectively recorded between 10 and 8° C. Such large depressions in metabolism appear not to have been reported previously for a marine fish species. No goldsinny mortalities were recorded at any temperature. The possibility that hypometabolic torpor is an adaptive strategy for goldsinny survival at low environmental temperatures is discussed.     

Diel patterns of CO2 exchange for epiphytic cacti differing in succulence

Net CO: uptake over 24-h periods was examined for 6 species of epiphytic cacti. which as a group number about 120 species that vary considerably in stem succulence. Nocturnal CO, uptake as a percentage of the total daily CO, uptake increased from 0% for Rhipsalis houlletiana to 14% for Epipliyl/urn pittieri to 21% for E. oxypetnluin to 51% for R. rhornbea to 87% for Schlumbergera truncata to 98% for Cryplocercus anthonyamis. For R. houlletiana and S. truncata, maximum net CO, uptake occurred at a photosynthetically active radiation of only 200 μmol m⁻² s⁻¹, as is typical for shade plants. Short-term drought of R. rhombea caused a shift favoring nocturnal CO, uptake. For the 6 species examined, increased reliance on nocturnal CO, uptake characteristic of Crassulacean acid metabolism (CAM) was closely correlated with increases in stem thickness, fresh weight per unit area, water mass per unit area, and to a somewhat lesser extent with another measure of succulence, water mass per unit mass of chlorophyll.     

Gas exchange and carbon isotope composition of Ananas comosus in response to elevated CO2 and temperature

Ananas comosus L. (Merr.) (pineapple) was grown at three day/night temperatures and 350 (ambient) and 700 (elevated) μ mol mol^–1 CO₂ to examine the interactive effects of these factors on leaf gas exchange and stable carbon isotope discrimination ( Δ ,‰). All data were collected on the youngest mature leaf for 24 h every 6 weeks. CO₂ uptake (mmol m^–2 d^–1) at ambient and elevated CO₂, respectively, were 306 and 352 at 30/20 °C, 175 and 346 at 30/25 °C and 187 and 343 at 35/25 °C. CO₂ enrichment enhanced CO₂ uptake substantially in the day in all environments. Uptake at night at elevated CO₂, relative to that at ambient CO₂, was unchanged at 30/20 °C, but was 80% higher at 30/25 °C and 44% higher at 35/25 °C suggesting that phosphoenolpyruvate carboxylase was not CO₂-saturated at ambient CO₂ levels and a 25 °C night temperature. Photosynthetic water use efficiency (WUE) was higher at elevated than at ambient CO₂. Leaf Δ -values were higher at elevated than at ambient CO₂ due to relatively higher assimilation in the light. Leaf Δ was significantly and linearly related to the fraction of total CO₂ assimilated at night. The data suggest that a simultaneous increase in CO₂ level and temperature associated with global warming would enhance carbon assimilation, increase WUE, and reduce the temperature dependence of CO₂ uptake by A. comosus .     

Effects of development, temperature and salinity on metabolism in eggs and yolk-sac larvae of milkfish, Chanos chanos (Forsskål)

Oxygen uptake rates and yolk-inclusive dry weiGhts were measured during the egg and yolk-sac larval stages of milkfish, Chanos chanos (Forsskal). Oxygen uptake by eggs and yolk-sac larvae was measured to assess the effects of four salinities (20,25,30,35 ppt) at 28°C. The effects of three temperatures (23,28,33°C) on oxygen uptake by yolk-sac larvae were determined at a salinity of 35 ppt. Dry weights were measured throughout embryonic development at 28°C and the yolk-sac stage at 23.28 and 33°C.
Oxygen uptake rates of eggs increased more than fivefold during embryogenesis (0.07±0.03 to 0.40 ± 03 μl O₂ egg ⁻¹ h ⁻¹;blastula to prehatch stage). Larval oxygen uptake did not change with age but was affected by rearing temperature (0.33 ± 0.08, 0.44 ± 0.07 and 0.63 ± 0.13 μl O₂ larva ⁻¹ h⁻¹ at 23, 28 and 33°C, respectively; Q₁₀= 1.93). Acute temperature changes from 28 to 33°C caused significant increases in oxygen uptake by embryos (Q ₁₀= 1.69–3.58) and yolk-sac larvae (Q ₁₀=2.55). Salinity did not affect metabolic rates.
Dry weight of eggs incubated at 28°C decreased 13% from fertilization to hatching. Incubation temperatures from 23–33°C did not affect dry weights at hatching. Rearing temperatures significantly affected the rate of larval yolk absorption (Q ₁₀= 2.25).     

Kinetic changes with temperature of phosphoenolpyruvate carboxylase from a CAM plant

Abstract. Purified and crude phosphoenolpyruvate carboxylase from the CAM plant Kalanchoë daigremontiana Hamet et Perrier ( Bryophyllum diagremontianum ) was assayed at temperatures between 10 and 45° C. The optimum temperature of the enzyme activity changed with substrate availability and effector concentration in the assay. l -malate inhibited the enzyme activity and lowered the optimum temperature. Glucose-6-phosphate raised the optimum temperature to 43°C. K _m values for phosphoenolpyruvate increased with assay temperature from 0.12 mol m^-3 at 15° C to 0.36 mol^m−3 at 35° C. Inhibition by malate increased with temperature and acidity of the assay. In the crude enzyme 50% of control activity was inhibited by 1.65 mol m^-3 malate at 15° C and by 0.5 mol m^-3 at 35° C (at pH 7.0). With purification malate sensitivity was lost ( K _i values for malate at least 10 times higher). The shift in optimum temperatures for PEP-carboxylase activity thus results from changes in the kinetic parameters with temperature and allosteric effectors. The often low optimum temperatures for CO₂ fixation observed in nature may thus be the result of substrate and effector concentrations in the cytoplasm and the antagonistic effect of temperature on substrate affinity and effector efficiency on phosphoenolpyruvate carboxylase.     

A comparison of low temperature growth vs low temperature shifts to induce resistance to photoinhibition in spinach (Spinacia oleracea)

Plants of Spinacia oleracea L. cv. Savoy grown under cold-hardening (5°C) and nonhardening (16°C) conditions were exposed to a photoinhibitory irradiance of 1300 μmol rrr^: m-² S-¹ 5°C for 12 h. Plants grown at 5°C exhibited a greater resistance to photoinhibition at low temperature in comparison to plants grown at 16°C as measured by the photochemical efficiency of photosyslem II. In contrast, tuily expanded leaves of plants grown at 16°C and then shifted to 5°C for 10 days did not exhibit increased resistance to photoinhibition. This was observed irrespective of the phoioperiod experienced during the shift to a lower temperature. Furthermore, spinach grown at 16°C and subsequently exposed to a stepped, daily decrease in temperature from 16 to 1°C over 10 days w ith a concomitant reduction in photoperiod. also did not exhibit any change in susceptibility to photoinhibition. Thus, a decrease in photoperiod accompanied by either an abrupt or stepped low temperature shift cannot induce increased resistance to photoinhibition. This confirms the hypothesis that growth and development at cold-hardening temperature are absolute requirements for the acquisition of resistance to photoinhibition at low temperature.     

C4 photosynthesis in Cyperus longus L., a species occurring in temperate climates

Abstract. Cyperus longus L. , which has a widespread but disjunct distribution throughout Europe and extends northwards into Britain, was found to be a C₄ species based upon its Kranz leaf anatomy, low CO₂ compensation point and the labelling of malate as an early product of ¹⁴CO₂ fixation. The photosynthetic characteristics of C. longus are similar to many other C₄ species with a high maximum rate of photosynthesis (> 1.5 mg CO₂ m ⁻² s ⁻¹) and a relatively high temperature optimum (30–35°C), but unlike many C₄ species the rate of photosynthesis does not decline rapidly below the optimum temperature and a substantial rate (0.6 mgCO₂ m⁻²s⁻¹)occursat 15°C. Leaf extension is very slow at 15°C and shows a curvilinear response to temperatures between 15 and 25°C. Leaves extend at a rate of almost 4 cm d⁻¹ at 25°C.     

Photosynthetic response of Eragrostis tef to temperature

Photosynthetic characteristics of leaves of tef, Eragrostis tef (Zucc.) Trotter, plants, grown at 25/15°C (day/night), were measured at temperatures from 18 to 48°C. The highest carbon exchange rates (CER) occurred between 36 and 42°C. and averaged 27 μmol m⁻² s⁻¹. At lower or higher temperatures, CER was reduced, but the availability of CO₂ to the mesophyll, measured as internal CO₂ concentration, was highest when temperatures were above or below the optimum for CER. In addition, CER and stomatal conductance were not correlated, but residual conductance was highly correlated with CER (r = 0.98). In additional experiments, relative ¹³C composition for leaf tissue grown at 25, 35 and 45°C averaged -14.4 per mille, confirming that tef is a C₄ grass species. Dry matter accumulation was higher at 35 than at 25, and lowest at 45°C. Leaf CER rates increased hyperbolically with increased light when measured from 0 to 2000 μmol m⁻² s⁻¹ PPFD. The highest CER, 31.8 μ-mol m_-2 s⁻¹, occurred at 35°C and 2000 μmol m⁻² s⁻¹ PPFR. At high light, CER at 25 and 35°C were nearly equal because of higher stomatal conductance at 25°C. Residual conductance was, however, clearly highest at 35°C compared to 25 and 45°C treatments. Stomatal conductance and residual conductance were not correlated in either set of experiments, yet residual conductance was always highest when temperatures were between 35 and 42°C across experiments, suggesting that internal leaf photosynthetic potential was highest across that temperature range.     

The effect of temperature fluctuations on oxygen consumption and ammonia excretion of underyearling Lake Inari Arctic charr

Underyearling Lake Inari Arctic charr Salvelinus alpinus were acclimated to 11·0) C for 3 weeks, and then one group was maintained at 11·0) C and others were exposed to 14·4) Cconst, 17·7) C_const or a diel fluctuating temperature of 14·3° C ± 1° C (14·3° C_fluc). Routine rates of oxygen consumption and ammonia excretion were measured over 10 days before the temperature change and over 31 days following the change. Measurements were made on fish that were feeding and growing. The temperature increase produced an immediate increase in oxygen consumption. There was then a decline over the next few days, suggesting that thermal acclimation was rapid. For groups exposed to constant temperature there was an increase in oxygen consumption ( M _accl, mg kg⁻¹ h⁻¹) with increasing temperature ( T ), the relationship being approximated by an exponential model: M _accl= 46·53e0·^{086 T} . At 14·3° C_fluc oxygen consumption declined during the 3–4 days following the temperature shift, but remained higher than at 14·4° C_const. This indicates that small temperature fluctuations have some additional influences that increase metabolic rate. Ammonia excretion rates showed diel variations. Excretion was lower at 11° C_const than at other temperatures, and increases in temperature had a significant effect on ammonia excretion rate. Fluctuating (14·3° C_fluc) temperature did not influence ammonia excretion relative to constant temperature (14·4° C_const).     

The influence of the rate of temperature change on the activation of dormant seeds of Rumex obtusifolius L.

1. One temperature shift from 20 to 30°C in darkness induces 30–40% germination in Rumex obtusifolius seeds. The same germination percentages are found with heat treatment varying between 1 and 6h duration, indicating that the total heat sum of the temperature shift is not important.
2. Germination is greatly enhanced by three consecutive heat shifts of 1h at 30°C separated by 1h periods at 20°C.
3. The seeds are activated to a small extent after a slow warming (+2°Ch^–1) from 20 to 30°C, followed by incubation for 1h at 30°C. Germination is much higher after rapid heating (+10°Ch^–1) to 30°C, followed by 1h incubation at this temperature. Repeated fast heating treatments on four consecutive days enhances germination. Moderately rapid heatings (+3·3°Ch^–1) give intermediate results.
4. The rate of cooling does not influence the germination percentage. Cooling alone cannot induce germination.
5. Heating alone from 15 to 25°C without cooling also activates germination. In this temperature range the seeds are more activated by rapid warming than by slow warming.
6. The ecological relevance of the response to different warming rate is discussed. The insensitivity of seeds to a slow warming might keep deeply buried seeds in a dormant stage.     

The effect of temperature and mass on routine metabolism in Sarotherodon (Tilapia)mossambicus (Peters)

A reappraisal of oxygen uptake by Sarotherodon mossambicus was undertaken using a continuous flow respirometer. Measurements were obtained over the temperature range 16°C–37°C for fish weighing between 10 g and 150 g. Oxygen uptake was converted to energy equivalents ( Q _ox) using the value 13.68 J mg O₂^–1and the routine metabolic energy expenditure can be described by the equation E =0.0086 t ^{2 0783} M ^{0 652} where E is the energy requirement for routine metabolism expressed in J h^-1, t the temperature in °C and M the mass in g.     

Photoinhibition of photosynthesis in Lemna gibba as induced by the interaction between light and temperature. I. Photosynthesis in vivo

The floating angiosperm Lemna gibba L. was exposed for 2 h to various combinations of photosynthetic photon flux densities and temperature. The extent of photoinhibition of photosynthesis was assayed by measuring the net CO₂ uptake before and after a photoinhibitory treatment, and the time course for photoinhibition was studied. It was found that the maximum quantum yield and the light-saturated rate of CO₂ uptake were affected by the interaction between light and temperature during the photoinhibitory treatment. At a constant photon flux density of 650 μmol m⁻² s⁻¹ the extent of photoinhibition increased with decreasing temperature showing that even a chilling-resistant plant like L. gibba is much more susceptible to photoinhibition at chilling temperatures. About 60% photoinhibition of the quantum yield for CO₂ uptake could be obtained either by a high photon flux density of 1 750 μmol m⁻² s⁻¹ and 25°C or by a moderate photon flux density of 650 μmol m⁻² s⁻¹ and 3°C. The time courses of recovery from 60% photoinhibition produced by either of these two treatments were similar, indicating that the nature of the photoinhibition was intrinsically similar. The extent of photoinhibition was related to the amount of light absorbed in excess to what could be handled by photosynthesis at that temperature. The vital importance of photosynthesis in alleviating photoinhibition is discussed.     

Effects of temperature and [His7]-corazonin on the body darkening in Locusta migratoria

Abstract.  [His⁷]-corazonin is a neuropeptide that induces dark coloration in locusts. This study examined the effect of temperature on body colour in the migratory locust, Locusta migratoria . L. Injection of a low dose (1 pmol) of [His⁷]-corazonin caused albino nymphs to develop dark coloration at 25–34 °C, but little darkening occurred at 38 and 42 °C. However, injection of a high dose (10 pmol) induced darkening even at these high temperatures. Transfer of nymphs injected with 1 pmol of [His⁷]-corazonin from 30 to 42 °C, and vice versa, indicated that temperature influenced darkening at any time after injection. Measurements of the luminance of the pronotum were made using commercially available computer software to follow continuous changes in darkening of the body. The body colour of nymphs injected with [His⁷]-corazonin was reddish-brown at 25 °C, blackish at 30 and 34 °C, and greyish or whitish at 38 and 42 °C. High temperature also suppressed darkening in a normal (pigmented) strain. Most nymphs transferred from 30 to 42 °C during the first three stadia developed a light colour in the fifth stadium without the striking black patterns that are typically manifested in gregarious nymphs at lower temperatures. Such individuals developed black patterns in the fifth stadium when injected with [His⁷]-corazonin at a mid stage of the previous stadium. These results indicate that high temperature may induce light body coloration by suppressing the release of [His⁷]-corazonin in normal locusts.     

The role of temperature in determining the stimulation of CO2 assimilation at elevated carbon dioxide concentration in soybean seedlings

Soybean ( Glycine max cv. Clark) was grown at both ambient (ca 350 μmol mol⁻¹) and elevated (ca 700 μmol mol⁻¹) CO₂ concentration at 5 growth temperatures (constant day/night temperatures of 20, 25, 30, 35 and 40°C) for 17–22 days after sowing to determine the interaction between temperature and CO₂ concentration on photosynthesis (measured as A, the rate of CO₂ assimilation per unit leaf area) at both the single leaf and whole plant level. Single leaves of soybean demonstrated increasingly greater stimulation of A at elevated CO₂ as temperature increased from 25 to 35°C (i.e. optimal growth rates). At 40°C, primary leaves failed to develop and plants eventually died. In contrast, for both whole plant A and total biomass production, increasing temperature resulted in less stimulation by elevated CO₂ concentration. For whole plants, increased CO₂ stimulated leaf area more as growth temperature increased. Differences between the response of A to elevated CO₂ for single leaves and whole plants may be related to increased self-shading experienced by whole plants at elevated CO₂ as temperature increased. Results from the present study suggest that self-shading could limit the response of CO₂ assimilation rate and the growth response of soybean plants if temperature and CO₂ increase concurrently, and illustrate that light may be an important consideration in predicting the relative stimulation of photosynthesis by elevated CO₂ at the whole plant level.     

Effect of temperature and dissolved oxygen concentration on the metabolic rate of the turbot and the relationship between metabolic scope and feeding demand

Turbot Scophthalmus maximus maximum oxygen uptake following feeding and exhaustive exercise increased from 107 mg O2 kg⁻¹ h⁻¹ at 6° C to c . 218 mg O2 kg⁻¹ h⁻¹ at 18° C, then increased slightly from 18 to 22° C to 224 mg O2 kg⁻¹ h⁻¹. Standard oxygen uptake increased exponentially as a function of temperature from 11 mg O2 kg ⁻¹ h⁻¹ at 6° C to 66 mg O2 kg⁻¹ h⁻¹ at 22° C. Gradual reduction in oxygen concentration to 87–90% air saturation at 6, 10. 18° C and <80% at 14 and 22° C limited the maximum metabolic rate but, supersaturation (>100% saturation) had little effect. Metabolic scope attained a maximum of 176 mg O₂ kg⁻¹ h⁻¹ at 18° C. Interpolation of the results showed that this value changed little between 16 and 20° C. It is suggested that this temperature range is optimal for turbot of c . 500 g. A comparison with a previous study on feeding demand in intensive farming conditions showed a linear relationship between appetite and metabolic scope. It is concluded that the ability of a fish to supply energy (including the energy requirement of digestive metabolism) above a standard level is a limiting factor in the manifestation of its feeding demand.     

Reappraisal of the effect of temperature on the growth kinetics of Aeromonas salmonicida

The effect of temperature (1–34 °C ) on the maximum specific growth rate of Aeromonas salmonicida could not be described by the classical growth models ; for some strains, two optimal temperatures at 23 °C and 30 °C were observed, as well as an unexpected increase in the pseudolag time above 27 °C. This could be explained by the presence of two subsets, notably S-layer⁺ and S-layer⁻ sub-populations. The A⁻ cells had higher growth parameters (T_opt and μ_opt) than the A⁺ cells and were selected by subcultures above 30 °C. Yet the relative proportion of A⁺ cells did not explain all the variation of μ_max versus temperature, and the growth kinetics of an Aer. salmonicida isolate remained unpredictable.     

A desk study of the relationship between temperature and hatching time for the eggs of five species of salmonid fishes

SUMMARY. Information on temperature (T°C) and time from fertilization to 50% hatch ( D days) for five species of salmonid fishes has been used to assess several mathematical models relating D and T . No single equation gave the best fit to all five data sets. The power law with temperature correction (α), log₁₀₁ D = log₁₀ a + b log₁₀ ( T - α) and the quadratic, log₁₀ D = log₁₀ a + bT + b ₁ T ² (where a, b, b ₁, and α are constants), each accounted for over 97 % of the variance of D and were good fits to the observed data points for all five species. There was little difference between the predictions obtained from these two equations within the range of observed temperatures. Therefore, the simpler power-law model is preferred. However, there were substantial within-species differences between values of D predicted from extrapolations of the two models from 2 or 3°C down to 0°C. When more data for low temperatures become available it will be possible to make a more objective choice of model.     

Growth and photosynthetic response of three soybean cultivars to simultaneous increases in growth temperature and CO2

Three soybean ( Glycine max L. Merr.) cultivars (Maple Glen, Clark and CNS) were exposed to three CO₂ concentrations (370, 555 and 740 μmol mol⁻¹) and three growth temperatures (20/15°, 25/20° and 31/26°C, day/night) to determine intraspecific differences in single leaf/whole plant photosynthesis, growth and partitioning, phenology and final biomass. Based on known carboxylation kinetics, a synergistic effect between temperature and CO₂ on growth and photosynthesis was predicted since elevated CO₂ increases photosynthesis by reducing photorespiration and photorespiration increases with temperature. Increasing CO₂ concentrations resulted in a stimulation of single leaf photosynthesis for 40–60 days after emergence (DAE) at 20/15°C in all cultivars and for Maple Glen and CNS at all temperatures. For Clark, however, the onset of flowering at warmer temperatures coincided with the loss of stimulation in single leaf photosynthesis at elevated CO₂ concentrations. Despite the season-long stimulation of single leaf photosynthesis, elevated CO₂ concentrations did not increase whole plant photosynthesis except at the highest growth temperature in Maple Glen and CNS, and there was no synergistic effect on final biomass. Instead, the stimulatory effect of CO₂ on growth was delayed by higher temperatures. Data from this experiment suggest that: (1) intraspecific variation could be used to select for optimum soybean cultivars with future climate change; and (2) the relationship between temperature and CO₂ concentration may be expressed differently at the leaf and whole plant levels and may not solely reflect known changes in carboxylation kinetics.