CO2 Compensation Concentration Research Articles

Inorganic carbon acquisition in the acid-tolerant alga Chlorella kessleri.

The ability of the freshwater alga, Chlorella kessleri, to maintain a carbon concentrating mechanism when grown at acid pH was investigated. The alga grows over the pH range 4.0-9.0 and was found to take up bicarbonate and CO2 actively when grown at pH 6.0. However, when grown at acid pH (below 5.5), it does not have active CO2 uptake. The acidotolerant species maintained an internal pH of 6.1-7.5 over the external pH range 4.5-7.5, thus the pH difference between the cell interior and the external medium was large enough to allow for the diffusive uptake of CO2 at acid external pH. Mass spectrometric monitoring of O2 and CO2 fluxes by suspensions of C. kessleri, grown at acid pH, and maintained at pH 7.5 showed that the rates of O2 evolution did not exceed those of CO2 uptake. The final CO2 compensation concentrations of 14.0-17.7 µM reached by photosynthetic cells were above the CO2 equilibrium concentration in the external medium, indicating a lack of active CO2 uptake at acid pH. Chlorella kessleri accumulated CO2 with internal concentrations that were 9.9, 18.7 and 22.7-fold that of the external medium for cells grown, respectively, at pH 4.5, 5.0 and 5.5. The ability of C. kessleri cells to accumulate high intracellular concentrations of inorganic carbon at acid pH would provide a sufficiently high concentration of CO2 at the active site of Rubisco thus allowing the alga to maintain growth rates similar to those at alkaline pH.

Structural and biochemical characterization of the C₃-C₄ intermediate Brassica gravinae and relatives, with particular reference to cellular distribution of Rubisco.

On the basis of its CO2 compensation concentration, Brassica gravinae Ten. has been reported to be a C3–C4 intermediate. This study investigated the structural and biochemical features of photosynthetic metabolism in B. gravinae. The cellular distribution of ribulose 1,5-bisphosphate carboxylase/oxygenase (Rubisco) was also examined in B. gravinae, B. napus L. (C3), Raphanus sativus L. (C3), and Diplotaxis tenuifolia (L.) DC. (C3–C4) by immunogold electron microscopy to elucidate Rubisco expression during the evolution from C3 to C3–C4 intermediate plants. The bundle sheath (BS) cells of B. gravinae contained centrifugally located chloroplasts as well as centripetally located chloroplasts and mitochondria. Glycine decarboxylase P-protein was localized in the BS mitochondria. Brassica gravinae had low C4 enzyme activities and high activities of Rubisco and photorespiratory enzymes, suggesting that it reduces photorespiratory CO2 loss by the glycine shuttle. In B. gravinae, the labelling density of Rubisco was higher in the mesophyll chloroplasts than in the BS chloroplasts. A similar cellular pattern was found in other Brassicaceae species. These data demonstrate that, during the evolution from C3 to C3–C4 intermediate plants, the intercellular pattern of Rubisco expression did not change greatly, although the amount of chloroplasts in the BS cells increased. It also appears that intracellular variation in Rubisco distribution may occur within the BS cells of B. gravinae.

PHOTOSYNTHETIC INORGANIC CARBON ACQUISITION IN AN ACID-TOLERANT, FREE-LIVING SPECIES OF COCCOMYXA (CHLOROPHYTA)(1).

The processes of CO2 acquisition were characterized for the acid-tolerant, free-living chlorophyte alga, CPCC 508. rDNA data indicate an affiliation to the genus Coccomyxa, but distinct from other known members of the genus. The alga grows over a wide range of pH from 3.0 to 9.0. External carbonic anhydrase (CA) was detected in cells grown above pH 5, with the activity increasing marginally from pH 7 to 9, but most of the CA activity was internal. The capacity for HCO3 (-) uptake of cells treated with the CA inhibitor acetazolamide (AZA), was investigated by comparing the calculated rate of uncatalyzed CO2 formation with the rate of photosynthesis. Active bicarbonate transport occurred in cells grown in media above pH 7.0. Monitoring CO2 uptake and O2 evolution by membrane-inlet mass spectrometry demonstrated that air-grown cells reduced the CO2 concentration in the medium to an equilibrium concentration of 15 μM, but AZA-treated cells caused a drop in extracellular CO2 concentration to a compensation concentration of 27 μM at pH 8.0. CO2 -pulsing experiments with cells in the light indicated that the cells do not actively take up CO2 . An internal pool of unfixed inorganic carbon was not detected at the CO2 compensation concentration, probably because of the lack of active CO2 uptake, but was detectable at times before compensation point was reached. These results indicate that this free-living Coccomyxa possesses a CO2 -concentrating mechanism (CCM) due to an active bicarbonate-uptake system, unlike the Coccomyxa sp. occurring in symbiotic association with lichens.

Screening of certain mangroves for photosynthetic carbon metabolic pathway

The mangroves Rhizophora lamarkii, Ceriops roxburghiana, Bruguiera gymnorrhiza, Aegiceras corniculatum, and Lumnitzera racemosa were screened for their carbon metabolic pathways by measuring net photosynthetic rate (P N), 13C discrimination rate, leaf anatomy, titratable acidity, and activities of phosphoenolpyruvate carboxylase, NADH-malate dehydrogenase, alanine aminotransferase, aspartate aminotransferase, and pyruvate phosphate dikinase. The tested mangroves had a well developed succulence, opening of stomata during day time and closure in the night hours, and absence of diurnal fluctuation of organic acids in their leaves which excludes the possibility of these species being CAM plants. Moreover, the leaf anatomy had not exhibited Kranz syndrome. The high values of discrimination against 13C, low P N, high CO2 compensation concentration, and the activities of aminotransferases in the direction of alanine formation suggest that the species may follow C3 mode of carbon metabolic pathway.

Growth and some physiological attributes of pea (Pisum sativum L.) as affected by salinity.

The effects of salt stress were studied on growth and physiology of pea (Pisum sativum L. cv. Green Arrow) in a pot study. Pea plants were treated with NaCl at 0, 10, 30, 50 and 70 mM in Hoagland solution. Plants were harvested after 21 days for measurements of physiological parameters. The highest NAR and RGR were found in 10 mM NaCl. However, in 70 mM NaCl, RGR and RLGR were significantly decreased in respect of other concentrations of NaCl. In 50 and 70 mM NaCl, chlorophylls contents and photosynthetic rate, were significantly decreased and CO2 compensation concentration and respiration rate increased in comparison with control. In 10 and 30 mM NaCl gas exchanges and chlorophyll contents were not significantly decrease in respect of control. Results indicated that Pisum sativum L. cv. Green Arrow can tolerate to 70 mM NaCl, also growth of plants in 10 and 30 mM NaCl was better than that of those in 0 mM NaCl.

Intrinsic changes in photosynthetic parameters of carrot leaves under increasing CO<sub>2</sub> concentrations and soil moisture regimes

A controlled growth chamber experiment was conducted to investigate the short-term water use and photosynthetic responses of 30-d-old carrot seedlings to the combined effects of CO2 concentration (50–1 050 µmol mol –1 ) and moisture deficits (–5, –30, –55, and –70 kPa). The photosynthetic response data was fitted to a non-rectangular hyperbola model. The estimated parameters were compared for effects of moisture deficit and elevated CO2 concentration (EC). The carboxylation efficiency (α) increased in response to mild moisture stress (–30 kPa) under EC when compared to the unstressed control. However, moderate (–55 kPa) and extreme (–70 kPa) moisture deficits reduced α under EC. Maximum net photosynthetic rate (PNmax) did not differ between mild water deficit and unstressed controls under EC. Moderate and extreme moisture deficits reduced PNmax by nearly 85 % compared to controls. Dark respiration rate (RD) showed no consistent response to moisture deficit. The CO2 compensation concentration (Γ) was 324 µmol mol –1 for –75 kPa and ranged 63–93 µmol mol –1 for other moisture regimes. Interaction between moisture deficit and EC was noticed for PN, ratio of intercellular and ambient CO2 concentration (Ci/Ca), stomatal conductance (gs), and transpiration rate (E). PN was maximum and Ci/Ca was minimum at –30 kPa moisture deficit and at Ca of 350 µmol mol –1 . The gs and E showed an inverse relationship at all moisture deficit regimes and EC. Water use efficiency (WUE) increased with moisture deficit up to –55 kPa and declined thereafter. EC showed a positive influence towards sustaining PN and increasing WUE only under mild moisture stress, and no beneficial effects of EC were noticed at moderate or extreme moisture deficits.

δ<sup>13</sup>C values, photosynthetic pathways, and plant functional types for some plants from saline meadows, Northeastern China

Based on stable carbon isotope ratio (δ13C) measurements, photosynthetic pathway types were determined for 61 species in 54 genera and 24 families of flowering plants from the saline meadows of Northeastern China. Of these total vascular plants, 18 species in 17 genera from 6 families were found to have C4 photosynthesis; 43 species in 38 genera from 20 families had C3 photosynthesis. Six dicotyledonous species exhibited C4 pathway, 12 monocotyledonous species were found with C4 photosynthesis. The dicotyledonous C4 species had relative greater mean δ13C value and less total carbon content than both monocotyledonous C4 and C3 species. Most dicotyledonous C4 species were annual forbs and halophytes. Some C4 species had been previously documented, but their δ13C values varied remarkably from those of the present study. Even though there are some fluctuations for the δ13C values of some C4 species, δ13C value was still more reliable for C3 and C4 identification than the use of the enzyme ratio method and of low CO2 compensation concentration.

Mechanisms of inorganic carbon acquisition in twoEuglenaspecies

The mechanism of inorganic carbon uptake was examined in Euglena gracilis Klebs. and the acidophilic species Euglena mutabilis Schmitz. Both species, whether grown in acidic (pH 3.5) or alkaline (pH 7.5) media lack external carbonic anhydrase. Acid-grown E. gracilis was shown to have no capacity for bicarbonate transport, but transport was induced on transfer to alkaline medium (pH 7.5) in the light over a period of 8 h. In contrast, acid-grown E. mutabilis appears to have no capacity for bicarbonate transport even at neutral pH. The overall internal pH of the cells was determined by equilibration with14C-labelled benzoic acid over the pH range 3.5–5.0 and with14C-labelled 5,5-dimethyloxazolidine-2,4-dione over the range pH 5.5–7.5. The acidophilic species maintains an internal pH range of 6.6–6.8 in an external pH range of 3.5–5.5, whereas the acid-tolerant species E. gracilis maintains a neutral internal pH in an external pH range of 3.5–7.5. Measurement, by mass spectrometry, of the fluxes of CO2and O2in photosynthesizing cells at pH 3.5 demonstrated a rapid uptake of CO2by both species that was completely blocked by iodoacetamide, an inhibitor of CO2fixation. Uptake of CO2by E. gracilis, grown at pH 7.5, was not completely inhibited by iodoacetamide and O2evolution was sustained when the cells reached the CO2compensation concentration, indicating a direct uptake of bicarbonate. These data indicate that the acidophilic species, E. mutabilis, takes up CO2by diffusion, whereas the acid-tolerant species, E. gracilis, takes up CO2by diffusion at acid pH levels but has some capacity for active bicarbonate uptake when grown at alkaline pH levels.Key words: acidophilic alga, acidotolerant alga, bicarbonate uptake, CO2uptake, Euglena gracilis, Euglena mutabilis, internal pH.

Gas Exchange of Carrot Leaves in Response to Elevated CO<sub>2</sub> Concentration

Short-term responses of four carrot (Daucus carota) cultivars: Cascade, Caro Choice (CC), Oranza, and Red Core Chantenay (RCC) to CO2 concentrations (Ca) were studied in a controlled environment. Leaf net photosynthetic rate (PN), intercellular CO2 (Ci), stomatal conductance (gs), and transpiration rate (E) were measured at Ca from 50 to 1 050 μmol mol-1. The cultivars responded similarly to Ca and did not differ in all the variables measured. The PN increased with Ca until saturation at 650 μmol mol-1 (Ci= 350-400 μmol mol-1), thereafter PN increased slightly. On average, increasing Ca from 350 to 650 and from 350 to 1 050 μmol mol-1 increased PN by 43 and 52 %, respectively. The PNvs.Ci curves were fitted to a non-rectangular hyperbola model. The cultivars did not differ in the parameters estimated from the model. Carboxylation efficiencies ranged from 68 to 91 μmol m-2 s-1 and maximum PN were 15.50, 13.52, 13.31, and 14.96 μmol m-2 s-1 for Cascade, CC, Oranza, and RCC, respectively. Dark respiration rate varied from 2.80 μmol m-2 s-1 for Oranza to 3.96 μmol m-2 s-1 for Cascade and the CO2 compensation concentration was between 42 and 46 μmol mol-1. The gs and E increased to a peak at Ca= 350 μmol mol-1 and then decreased by 17 and 15 %, respectively when Ca was increased to 650 μmol mol-1. An increase from 350 to 1 050 μmol mol-1 reduced gs and E by 53 and 47 %, respectively. Changes in gs and PN maintained the Ci:Ca ratio. The water use efficiency increased linearly with Ca due to increases in PN in addition to the decline in E at high Ca. Hence CO2 enrichment increases PN and decreases gs, and can improve carrot productivity and water conservation.

Developmental Change in CO<sub>2</sub> Compensation Concentrations in Spartina alterniflora Results from Sigmoidal Photosynthetic CO<sub>2</sub> Responses

We investigated seasonal patterns of photosynthetic responses to CO2 concentrations in Spartina alterniflora Loisel, an aerenchymous halophyte grass, from a salt marsh of the Bay of Fundy (NB, Canada), and from plants grown from rhizome in controlled-environment chambers. From late May to August, CO2 compensation concentrations (Γ) of field-grown leaves varied between 2.5–10.7 cm3(CO2) m−3, with a mean of 5.4 cm3(CO2) m−3. From September onwards field leaves showed CO2 compensation concentrations from 6.6–21.1 cm3(CO2) m−3, with a mean of 13.1 cm3 m−3 well into the C3–C4 intermediate range. The seasonal variability in Γ did not result from changing respiration, but rather from a sigmoidal response of net photosynthetic rate (PN) to applied CO2 concentration, found in all tested leaves but which became more pronounced late in the season. One explanation for the sigmoidal response of PN to external CO2 concentration could be internal delivery of CO2 from roots and rhizomes to bundle sheath cells via the aerenchyma, but the sigmoidal responses in S. alterniflora persisted out to the tips of leaves, while the aerenchyma extend only to mid-leaf. The sigmoidicity persisted when CO2 response curves were measured from low to high CO2, or from high to low CO2, and even when prolonged acclimation times were used at each CO2 concentration.

Nonstomatal limitations are responsible for drought-induced photosynthetic inhibition in four C4 grasses.

• Here, the contribution of stomatal and nonstomatal factors to photosynthetic inhibition under water stress in four tropical C4 grasses was investigated (Panicum coloratum, Bothriochloa bladhii, Cenchrus ciliaris and Astrebla lappacea). • Plants were grown in well watered soil, and then the effects of soil drying were measured on leaf gas exchange, chlorophyll a fluorescence and water relations. • During the drying cycle, leaf water potential (Ψleaf ) and relative water content (RWC) decreased from c. -0.4 to -2.8MPa and 100-40%, respectively. The CO2 assimilation rates (A) and quantum yield of PSII (ΦPSII ) of all four grasses decreased rapidly with declining RWC. High CO2 concentration (2500µll-1 ) had no effect on A or ΦPSII at any stage of the drying cycle. Electron transport capacity and dark respiration rates were unaltered by drought. The CO2 compensation concentrations of P. coloratum and C. ciliaris rose sharply when leaf RWC fell below 70%. In P. coloratum, 5% CO2 did not prevent the decline of O2 evolution rates under water stress. • We conclude that inhibition of photosynthesis in the four C4 grasses under water stress is dependent mainly on biochemical limitations.

Photosynthesis of Cockspur [Echinochloa crus-galli (L.) Beauv.] at Sites of Naturally Elevated CO<sub>2</sub> Concentration

High abundance of cockspur (Echinochloa crus-galli) at the geothermal carbon dioxide spring area in StaveĹĄinci indicates that this species is able to grow under widely varying CO2 concentrations. Living cockspur plants can even be found very close to gas-releasing vents where growth is significantly reduced. Plant height correlated well with CO2 exposure. The δ13C value of the CO2 spring air was -3.9 ‰ and δ13C values of high-, medium-, and low-CO2 plants were -10.14, -10.44, and -11.95 ‰, respectively. Stomatal response directly followed the prevailing CO2 concentrations, with the highest reduction of stomatal conductance in high CO2 concentration grown plants. Analysis of the curves relating net photosynthetic rate to intercellular CO2 concentration (PN-Ci curves) revealed higher CO2 compensation concentration in plants growing at higher CO2 concentration. This indicates adjustment of respiration and photosynthetic carbon assimilation according to the prevailing CO2 concentrations during germination and growth. There was no difference in other photosynthetic parameters measured.

Comparison of Some Photosynthetic Characters Between Two Hybrid Rice Combinations Differing in Yield Potential

Photosynthetic characteristics of two hybrid rice combinations, Peiai 64S/E32 and Shanyou 63, were compared at the panicle differentiation stage. As compared with Shanyou 63, the new combination Peiai 64S/E32 showed a significantly higher net photosynthetic rate (PN), apparent quantum yield of carbon assimilation (Φc), carboxylation efficiency (CE), and photorespiratory rate (RP) as well as leaf chlorophyll content, but a significantly lower dark respiration rate (RD) and compensation irradiance (Ic). It also showed a slightly higher photochemical efficiency (Fv/Fm and ΔF/Fm') of photosystem 2, a lower non-photochemical quenching (qN), and a similar CO2 compensation concentration (Γ) as compared to Shanyou 63.

Photosynthetic Response of Wheat to Stress Induced by Puccinia Recondita and Post-Infection Drought

To quantify photosynthetic response of wheat to the combination of a fungal brown rust infection and a post-infection drought, four treatments were compared: no stress (control), fungal stress (FS), water stress (WS), and twofold stress (WS×FS). Predawn leaf water potential (Ψwp) was similar in FS and WS treatments over a 3-week period. In the WS treatment, net photosynthetic rate (P N) and stomata CO2 conductance (g s) diminished concomitantly with a constant intercellular CO2 concentration (C i) close to 200 µmol mol−1. In the FS treatment, a reduction of P N occurred with an increase in respiration rate (doubling of the CO2 compensation concentration) and in C i but with no water loss modification. Healthy leaves of infected plants (FS) showed a reduction of P N as well, with constant g s and increased C i. In the twofold stress treatment (WS×FS), leaves showed reduced P N in relation to the lower Ψwp. Deleterious effects of both drought and fungal infection on the final area of leaves and dry matter were additive.

Responses of Two Olive Tree (Olea Europaea L.) Cultivars to Elevated CO<sub>2</sub> Concentration in the Field

Five-year-old plants of two olive cultivars (Frantoio and Moraiolo) grown in large pots were exposed for 7 to 8 months to ambient (AC) or elevated (EC) CO2 concentration in a free-air CO2 enrichment (FACE) facility. Exposure to EC enhanced net photosynthetic rate (PN) and decreased stomatal conductance, leading to greater instantaneous transpiration efficiency. Stomata density also decreased under EC, while the ratio of intercellular (Ci) to atmospheric CO2 concentration and chlorophyll content did not differ, except for the cv. Moraiolo after seven months of exposure to EC. Analysis of the relationship between photosynthesis and Ci indicated no significant change in carboxylation efficiency of ribulose-1,5-bisphosphate carboxylase/oxygenase after five months of exposure to EC. Based on estimates derived from the PN-Ci relationship, there were no apparent treatment differences in daytime respiration, CO2 compensation concentration, CO2-saturated photosynthetic rate, or photosynthetic rate at the mean Ci, but there was a reduction in stomata limitation to PN at EC. Thus 5-year-old olive trees did not exhibit down regulation of leaf-level photosynthesis in their response to EC, though some indication of adjustment was evident for the cv. Frantoio with respect to the cv. Moraiolo.

Effects of Cadmium and Gibberellin on Growth and Photosynthesis of Glycine Max

The effects of 0, 2.5, 5.0, and 10.0 mg(Cd2+) m-3 [Cd(NO3)2×4 H2O] and 0 and 10.0 mg m-3 gibberellin on certain parameters of photosynthesis and growth in soybean (Glycine max L. cv. Pershing) plants were studied. With increasing Cd2+ concentration in the Hoagland nutrient solution, the contents of chlorophyll and CO2 compensation concentration decreased. The addition of 10 mg m-3 gibberellin reduced the negative effects of Cd2+ in shoot and root growth. With increasing of Cd2+ concentration in the culture medium, the dry matter production in both the roots and shoots decreased as shown by the decline in growth rate (PGR), net assimilation rate (NAR), and leaf area ratio. The addition of gibberellin caused a partial elimination of the Cd effects on the roots and shoots and the PGR and NAR and it increased leaf area and length of stem.

Clonal Differences in Photosynthesis in Hevea Brasiliensis Müll. Arg.

The objective of the present investigation was to examine the extent of variations in single leaf net photosynthetic rate (PN) and its relative dependence on stomatal conductance (gs) and the mesophyll capacity to fix carbon in 12 clones of the natural rubber plant. There were significant variations in PN measured at low and saturating photon flux density (PFD); the extent of variation was larger at low than at saturating PFD. The compensation irradiance (CI) and apparent quantum yield of CO2 assimilation (φc) calculated from the PN/PFD response curves showed significant variations among the clones. PN at low irradiance was positively correlated with φc. Thus a clone with large PN at low irradiance, high φc, and low CI may tolerate shade better and thus produce a high tree stand per hectare. A strong positive correlation existed between PN saturated with radiant energy (Psat) and carboxylation efficiency (CE) estimated from the response curves of PN on intercellular CO2 concentration (Ci), but gs showed a poor correlation with Psat High CO2 compensation concentration (Γ) led to low CE in Hevea clones. A clone with large Psat, high CE, low gs, and low Γ is the one in which photosynthesis is more dependent on the mesophyll factors than stomata. Such a clone may produce relatively high biomass and maintain high water use efficiency.

The effects of NaCl and CaCl2 on photosynthesis and growth of alfalfa plants

The effects of 0, 30, 60, and 90 mM NaCl, and 0 and 5 mM CaCl2 on certain parameters of photosynthesis and growth in alfalfa (Medicago sativa L. cv. Ghara yonjeh) plants were studied. The increasing NaCl concentration in the Hoagland nutrient solution decreased the contents of chlorophylls and the net photosynthetic rate, and increased the rate of respiration (RD) and CO2 compensation concentration in the leaves of treated plants. The contents of carotenoids (Car) were not significantly affected. The addition of 5 mM CaCl2 enhanced the RD and increased the Car contents in treated leaves. With the NaCl concentration in the culture medium increasing, the dry matter production in both root and shoot decreased, as well as the relative growth rate (RGR), net assimilation rate (NAR), and leaf area ratio (LAR). The addition of CaCl2 caused a partial elimination of the NaCl effects on the root and shoot, RGR and NAR, and it decreased the LAR.

A STUDY ON THE RELATIONSHIP BETWEEN LEAF CONDUCTANCE, CO2 CONCENTRATION AND CARBOXYLATION RATE IN VARIOUS SPECIES

Leaf conductance gL is strongly influenced by environmental factors like CO2, irradiance and air humidity. According to Ball et al. (1987), gL is correlated with an index calculated as the product of net CO2 exchange rate A and ambient water vapour concentration Wa, divided by ambient CO2 concentration ca. However, this empirical model does not apply to high values of gL observed at ca below CO2 compensation concentration Γ. Therefore, we applied modified indices in which A is replaced by estimates for the rate of carboxylation. Such estimates, P1 and P2, were determined by adding to A the quotient of Γ and the sum of gas phase resistance rg and intracellular resistance for CO2 exchange ri, P1 = A+Γ/(rg + ri), or the quotient of Γ and ri, P2 = A + Γ/ri. If P2 is chosen, ca in the Ball index has to be replaced by the intercellular CO2 concentration ci. By using the modified indices P1·Wa/ca and P2·Wa/ci, we analysed data from the C3 species Nicotiana tabacum and Nicotiana plumbaginifolia, the C3–C4 intermediate species Diplotaxis tenuifolia, and the C4 species Zea mays. The data were collected at widely varying levels of irradiance and CO2 concentration. For all species uniform relationships between gL and the new indices were found for the whole range of CO2 concentrations below and above Γ. Correlations between gL and P1·Wa/ca were closer than those between gL and P2·Wa/ci because P1/ca implicitly contains gL. Highly significant correlations were also obtained for the relationships between gL and the ratios P1/ca and P2/ci.

Chloroplastic Carbon Dioxide Concentration in Norway Spruce (Picea Abies [L.] Karst.) needles relates to the position within the crown

Differences between sun (E) and shaded (S) foliage were studied in a Norway spruce (Picea abies [L.] Karst.) stand. Response curves describing the dependence of the CO2 assimilation rate (PN) on the CO2 concentration at the catalytic site of ribulose-1,5-bisphosphate carboxylase/oxygenase, RuBPCO (PN-Cc) were estimated using the simultaneous measurements of chlorophyll fluorescence and leaf gas exchange. Higher PN, higher electron transport (Ja), higher carboxylation capacity (Vc), and higher RuBPCO activity (τ) for sun acclimated needles was found. The S-needles had higher portion of internal limitation and higher CO2 compensation concentration (Γ) than the E-needles. Because higher degree of limitation of photosynthesis by carboxylation was ascertained, it can be assumed that photosynthesis in shade foliage is limited mainly by lower carboxylation capacity and by low chloroplastic CO2 concentration