Chlorophyll a Fluorescence Response of Norway Spruce Needles to the Long-Term Effect of Elevated CO2 in Relation to Their Position Within the Canopy

The long-term impact of elevated CO₂ concentration on photosynthetic activity of sun-exposed (E) versus shaded (S) foliage was investigated in a Picea abies stand (age 12 years) after three years of cultivation in adjustable-lamella-domes (ALD). One ALD is supplied with either ambient air [ca. 350 µmol(CO₂) mol^–1; AC-variant) and the second with elevated CO₂ concentration [ambient plus 350 µmol(CO₂) mol^–1; EC-variant). The pronounced vertical profile of the photosynthetically active radiation (PAR) led to the typical differentiation of the photosynthetic apparatus between the S- and E-needles in the AC-variant estimated from the irradiance-responses of various parameters of the room temperature chlorophyll (Chl) a fluorescence parameters. Namely, electron transport rate (ETR), photochemical efficiency of photosystem 2, PS2 (_PS2), irradiance-saturated values of non-photochemical quenching of minimum (SV₀) and maximum (NPQ) fluorescence levels, and photochemical fluorescence quenching (q_p) at higher irradiances were all significantly higher for E-needles as compared with the S-ones. The prolonged exposure to EC did not cause any stimulation of ETR for the E-needles but a strongly positive effect of EC on ETR was observed for the S-needles resulting in more than doubled ETR capacity in comparison with S-needles from the AC-variant. For the E-needles in EC-variant a slightly steeper reduction of the _PS2 and q_p occurred with the increasing irradiance as compared to the E-needles of AC-variant. On the contrary, the S-needles in EC variant revealed a significantly greater capacity to maintain a high _PS2 at irradiances lower than 200 µmol m^–2 s^–1 and to prevent the over-reduction of the PS2 reaction centres. Moreover, compared to the AC-variant the relation between SV₀ and NPQ exhibited a strong decrease (up to 72 %) of the SV₀-NPQ slope for the E-needles and an increase (up to 76 %) of this value for the S-needles. Hence the E- and S-foliage responded differently to the long-term impact of EC. Moreover, this exposure was responsible for the smoothing of the PAR utilisation vertical gradient in PS2 photochemical and non-photochemical reactions within the canopy.     

Long-Term Effects of Elevated CO2 on Woody Tissues Respiration of Norway Spruce Studied in Open-Top Chambers

In an open-top chamber experiment located in a mountain stand of 14-years-old Norway spruce (Picea abies [L.] Karst.), trees were continuously exposed to either ambient CO₂ concentration (A), or ambient + 350 µmol mol^–1 (E) over four growing seasons. Respiration rates of different woody parts (stem, branches, coarse roots) were measured during the last growing season. The calculated increase in the respiration rate related to a 10 °C temperature change (Q₁₀) was different in stem compared to branches and roots. Differences between the E and A variants were statistically significant only for roots in the autumn. Stem maintenance respiration (R_Ms) measured in April and November (periods of no growth activity) were not different. The stem respiration values (R_s) were recalculated to a standard temperature of 15 °C to estimate the seasonal course. The obtained R_s differed significantly between used variants during July and August. At the end of the season, R_s in E decreased slower than in A, indicating some prolongation of the physiological activity under the elevated CO₂ concentration. The total stem respiration carbon losses for the investigated growing season (May – September) were higher for A (2.32 kg(C) m^–2 season^–1) compared to E (2.12 kg(C) m^–2 season^–1). The respiration rates of the whorl branches (R_b) were lower compared with the stem respiration but not significantly different between the used variants. The root respiration rate was increased in E variant.     

Acclimation of Photosystem 2 Function of Norway Spruce Induced during First Season under Elevated CO2 in Lamellar Domes

Since July 28^th, 1997 the two experimental mini-stands of young Norway spruce [Picea abies (L.) Karst.] have been grown in lamellar domes at ambient (AC) and elevated concentrations of CO₂ [EC, i.e., ambient + 350 µmol(CO₂) mol^–1]. Before the start of exposure to EC (June 1997) the dependencies of photosystem 2 (PS2) quantum yield (Y) on irradiance, estimating the efficiency of PPFD utilisation in PS2 photochemistry, were the same for AC and EC shoots. After one month of EC simulation (August 1997), Y values were higher for EC needles as compared with the AC ones (by 1–42 %), whereas two months later (October 1997) an opposite effect was observed (decrease of Y by from 1 to 33 %). By chlorophyll a (Chl a) fluorescence induction the effects of EC on PS2 function were further characterised. During the first month a moderate improvement of PS2 function was estimated for EC needles from slightly higher potential yield of PS2 photochemistry (F_V/F_M, by 1 %) and reduced amount of inactive PS2 reaction centres (relative F_p1 level, by 15 %). However, the prolonged exposure to EC led firstly to a slight but significant decrease of F_V/F_M (by 3 %), secondly to a reduction of half time of fluorescence rise (t_1/2, by 14 %), and finally to pronounced accumulation of inactive PS2 reaction centres (by 41 %). From the gradual response of individual Chl a fluorescence parameters we suggest a probable sequence of events determining the stimulation and subsequent depression of PS2 function for Norway spruce during the first season under EC.     

Control Mechanisms of Photosynthetic Capacity Under Elevated CO2 Concentration: Evidence from Three Experiments with Norway Spruce Trees

Twelve-year-old Norway spruce (Picea abies [L.] Karst.) were exposed to ambient (AC) or elevated (EC) [ambient + 350 mol(CO₂) mol^–1] CO₂ concentration [CO₂] using the facilities of open-top-chambers (OTCs) and glass domes (GDs). A combination of gas exchange measurements and application of a biochemical model of photosynthesis were used for the evaluation of CO₂ assimilation characteristics. Morphological change was assessed on the base of specific leaf area (SLA). Nitrogen (N) content in the assimilation apparatus was considered a main factor influencing the biochemical capacity. Three experiments confirm the hypothesis that an adjustment of photosynthetic capacity under EC is controlled by the combination of biochemical, morphological, and physiological feedback mechanisms. We observed periodicity of down-regulation of photosynthetic capacity (Experiment No. 1) during the vegetation seasons. In the spring months (May–June), i.e. during the occurrence of active carbon sink associated with the formation of new foliage, up-regulation (10–35 %) of photosynthetic capacity (P _Nsat) was observed. On the contrary, in the autumn months (September–October) down-regulation (25–35 %) of P _Nsat was recorded that was mainly associated with reduced carbon sink strength and biochemical change, i.e. decrease of N status (up to 32 %) and accumulation of saccharides (up to 72 %) in leaves. Different adjustments of photosynthetic activities were observed in current (C) and one-year-old (C-1) needles exposed to EC (Experiment No. 2). Strong down-regulation of P _Nsat and the diminution of the initial stimulation of photosynthetic rate (P _Nmax) was associated with decreases of both ribulose-1,5-bisphosphate carboxylase/oxygenase carboxylation activity (by 32 %) and RuBP regeneration (by 40 %). This performance was tightly correlated with the absence of active carbon sinks, decrease of N content, and starch accumulation in C-1 needles. Finally, different responses of sun- and shade-adapted needles to EC (Experiment No. 3) were associated with the balance between morphological and biochemical changes. Observed P _Nsat down-regulation (by 22 %) of exposed needles in EC was predominantly caused by effects of both higher assimilate accumulation and stronger N dilution, resulting from higher absolute photosynthetic rates and incident irradiances in the upper canopy.     

Responses of Rice Cultivars to the Elevated CO2

The effect of CO₂ concentration elevated to 575 – 620 µmol mol^–1 on growth, tillering, grain yield, net photosynthetic rate, dark respiration rate, stomatal conductance, sugar content and protein profile of two rice (Oryza sativa L.) cultivars Pusa Basmati-1 and Pusa-677 at flowering stage was studied using open top chambers. The cultivar Pusa Basmati-1 responded more markedly for most of the growth and physiological parameters compared to Pusa-677. The increase in grain yield in Pusa Basmati-1 attributed largely to increased grain number. The increased net photosynthetic rate and greater accumulation of sugar contributed significantly to the accelerated development of leaves and tillers in both the cultivars. The reduction in the low molecular mass proteins including Rubisco and increase in high molecular mass photosystem 2 proteins was observed in both the cultivars. Additional sugars may possibly help in balancing the profile of photosynthetic proteins and sustain greater growth and productivity in rice cultivars.     

Seasonal Changes of Selected Parameters of CO2 Fixation Biochemistry of Norway Spruce Under the Long-Term Impact of Elevated CO2

Twelve-year-old Norway spruce (Picea abies [L.] Karst.) trees were exposed to ambient (AC) or elevated (EC) [ambient + 350 μmol(CO₂) mol^-1] CO₂ concentrations in open-top-chamber (OTC) experiment under the field conditions of a mountain stand. Short-term (4 weeks, beginning of the vegetation season) and long-term (4 growing seasons, end of the vegetation season) effects of this treatment on biochemical parameters of CO₂ assimilation were evaluated. A combination of gas exchange, fluorescence of chlorophyll a, and application of a mathematical model of ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBPCO) activity was used. The analysis showed that the depression of photosynthetic activity by long-term impact of elevated CO₂ was mainly caused by decreased RuBPCO carboxylation rate. The electron transport rate as well as the rate of ribulose-1,5-bisphosphate (RuBP) formation were also modified. These modifications to photosynthetic assimilation depended on time during the growing season. Changes in the spring were caused mainly by local deficiency of nitrogen in the assimilating tissue. However, the strong depression of assimilation observed in the autumn months was the result of insufficient carbon sink capacity. This revised version was published online in June 2006 with corrections to the Cover Date.     

Relationship Between Photosystem 2 Electron Transport and Photosynthetic CO2 Assimilation Responses to Irradiance in Young Apple Tree Leaves

The responses to irradiance of photosynthetic CO₂ assimilation and photosystem 2 (PS2) electron transport were simultaneously studied by gas exchange and chlorophyll (Chl) fluorescence measurement in two-year-old apple tree leaves (Malus pumila Mill. cv. Tengmu No.1/Malus hupehensis Rehd). Net photosynthetic rate (P _N) was saturated at photosynthetic photon flux density (PPFD) 600-1 100 (mol m^-2 s^-1, while the PS2 non-cyclic electron transport (P-rate) showed a maximum at PPFD 800 mol m^-2 s^-1. With PPFD increasing, either leaf potential photosynthetic CO₂ assimilation activity (F_d/F_s) and PS2 maximal photochemical activity (F_v/F_m) decreased or the ratio of the inactive PS2 reaction centres (RC) [(F_i – F_o)/(F_m – F_o)] and the slow relaxing non-photochemical Chl fluorescence quenching (q_s) increased from PPFD 1 200 mol m^-2 s^-1, but cyclic electron transport around photosystem 1 (RFp), irradiance induced PS2 RC closure [(F_s – F_o)/F_m – F_o)], and the fast and medium relaxing non-photochemical Chl fluorescence quenching (q_f and q_m) increased remarkably from PPFD 900 (mol m^-2 s^-1. Hence leaf photosynthesis of young apple leaves saturated at PPFD 800 mol m^-2 s^-1 and photoinhibition occurred above PPFD 900 mol m^-2 s^-1. During the photoinhibition at different irradiances, young apple tree leaves could dissipate excess photons mainly by energy quenching and state transition mechanisms at PPFD 900-1 100 mol m^-2 s^-1, but photosynthetic apparatus damage was unavoidable from PPFD 1 200 mol m^-2 s^-1. We propose that Chl fluorescence parameter P-rate is superior to the gas exchange parameter P _N and the Chl fluorescence parameter F_v/F_m as a definition of saturation irradiance and photoinhibition of plant leaves.     

Photosynthetic Gas Exchange and Chlorophyll a Fluorescence of Three Wild Soybean Species in Response to NaCl Treatments

Responses of photosynthetic gas exchange and chlorophyll (Chl) a fluorescence of three wild soybeans, Glycine soja, G. tomentella, and G. tabacina occurring in different habitats of Taiwan, to four NaCl treatments, 0S, LS, MS, and HS (i.e. 0, 17, 51, and 85 mM NaCl) were compared. In G. soja following exposure to NaCl treatment for one month, the photon saturated photosynthetic rate (P _N), the ratio of variable to maximum fluorescence (F_v/F_m), the quantum yield of photosystem 2 (_PS2), and the electron transport rate (ETR) decreased dramatically. These reductions increased with increasing concentration of NaCl treatment. Plants of MS and HS treatments did not survive after extending the treatment to two months. Reductions in P _N, _PS2, and ETR (but not in F_v/F_m) were found in G. tabacina after two months of exposure to MS and HS treatments, but the reduction was not as severe as that in G. soja. In G. tomentella, significant reductions in P _N and g _s were found only in HS plants after two months of treatment, but no significant differences in F_v/F_m, _PS2, and ETR were found among plants of the four treatments. Thus the three wild soybeans in Taiwan have differentiated in their photosynthetic susceptibility to salinity, G. tomentella being the least susceptible, G. soja the most sensitive, and G. tabacina the intermediate. Different mechanisms are attributed to the inhibition effect of salinity on photosynthesis of the three species.     

Modifications in Photosynthetic Pigments and Chlorophyll Fluorescence in 20-Year-Old Pine Trees after a Four-Year Exposure to Carbon Dioxide and Temperature Elevation

Changes in pigment composition and chlorophyll (Chl) fluorescence parameters were studied in 20 year-old Scots pine (Pinus sylvestris L.) trees grown in environment-controlled chambers and subjected to ambient conditions (CON), doubled ambient CO₂ concentration (EC), elevated temperature (ambient +2−6 °C, ET), or a combination of EC and ET (ECT) for four years. EC did not significantly alter the optimal photochemical efficiency of photosystem 2 (PS2; F_v/F_m), or Chl a+b content during the main growth season (days 150–240) but it reduced F_v/F_m and the Chl a+b content and increased the ratio of total carotenoids to Chl a+b during the ‘off season’. By contrast, ET significantly enhanced the efficiency of PS2 in terms of increases in F_v/F_m and Chl a+b content throughout the year, but with more pronounced enhancement in the ‘off season’. The reduction in F_v/F_m during autumn could be associated with the CO₂-induced earlier yellowing of the leaves, whereas the temperature-stimulated increase in the photochemical efficiency of PS2 during the ‘off season’ could be attributed to the maintenance of a high sink capacity. The pigment and fluorescence responses in the case of ECT showed a similar pattern to that for ET, implying the importance of the temperature factor in future climate changes in the boreal zone. This revised version was published online in August 2006 with corrections to the Cover Date.     

Response of Photosynthesis to Radiation and Intercellular CO2 Concentration in Sun and Shade Shoots of Norway Spruce

Functional differentiation of assimilation activity of sun versus shade foliage was analysed in a Norway spruce monoculture stand (age 15 years). The investigated stand density (leaf area index 8.6) and crown structure led to variation in the photosynthetically active photon flux density (PPFD) within the crowns of the sampled trees. At the saturating PPFD, the maximum rate of CO₂ uptake (P _Nmax) of exposed shoots (E-shoots) was 1.7 times that of the shaded shoots (S-shoots). The apparent quantum yield (α) of E-shoots was 0.9 times that of the S-shoots. A lower ability to use excess energy at high PPFD in photosynthesis was observed in the S-layer. The CO₂- and PPFD-saturated rate of CO₂ uptake (P _Nsat) of the E-shoots was 1.12 times and the carboxylation efficiency (τ) 1.6 times that of the S-shoots. The CO₂-saturated rate of ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBPCO) carboxylation (V_Cmax) and of actual electron transport (J_amax) in the S-needles amounted to 89 and 95 % of V_Cmax and J_amax in the E-needles. Thus, in addition to the irradiation conditions and thus limitation by low J_a, the important limitation of photosynthesis in shade needles is due to carboxylation. This limitation of photosynthesis is accompanied by lower stomatal conductance. This revised version was published online in June 2006 with corrections to the Cover Date.     

Chlorophyll Fluorescence Parameters: The Definitions,Photosynthetic Meaning,and Mutual Relationships

Chlorophyll fluorescence parameters (Chl FPs) derived from the slow (long-term) induction kinetics of modulated Chl a fluorescence are reviewed and analysed with respect to their application in photosynthesis research. Only four mutually independent Chl FPs, calculated from values of five essential Chl fluorescence (ChlF) yields, are distinguished as the basic ones. These are: the maximum quantum yield of PS2 photochemistry (_{P O}), the photochemical quenching of variable ChlF (q_P), the non-photochemical quenching of variable ChlF (q_N), and the relative change of minimum ChlF (q_O). _{P O} refers to the dark-adapted state of a thylakoid membrane, q_P, q_N and q_O characterise the light-adapted state. It is demonstrated that all other Chl FPs can be determined using this quartet of parameters. Moreover, three FPs related to the non-radiative energy dissipation within thylakoid membranes are evaluated, namely: the non-photochemical ChlF quenching (NPQ), the complete non-photochemical quenching of ChlF (q_CN), and the effective quantum yield of non-photochemical processes in PS2 (_N). New FPs, the total quenching of variable ChlF (q_TV) and the absolute quenching of ChlF (q_A) which allow to quantify co-action of the photochemical and non-photochemical processes during a light period are defined and analysed. The interpretation of Chl FPs and recommendations for their application in the photosynthesis research are also given. Some alternative FPs used in the laboratory practice have only an approximate character and can lead to incorrect conclusions if applied to stressed plants. They are reviewed and compared with the standard ones. All formulae and conclusions discussed herein are verified using experimental values obtained on young seedlings of the Norway spruce (Picea abies [L.] Karst.).     

Photosynthetic Responses of Tropical Trees to Short-Term Exposure to Ozone

Saplings of the tropical trees Tibouchina pulchra (Cham.) Cogn., Caesalpinia echinata Lam., and Psidium guajava L. cv. Paluma were exposed in open-top chambers with charcoal filtered air and measurements of gas exchange and chlorophyll fluorescence were made before (t₁) and after exposure to non-filtered air plus O₃ (t₂), simulating 6-h peaks of O₃ similar to those observed in São Paulo city (SE Brazil, reaching an AOT40 of 641 nmol mol^–1). After the fumigation, the net photosynthetic rate, stomatal conductance, transpiration rate, and F_v/F_m were reduced (p<0.05) for the three species. C. echinata was the most sensitive species and P. guajava cv. Paluma the most resistant.     

Gas Exchange Responses to CO2 Concentration Instantaneously Elevated in Flag Leaves of Winter Wheat Cultivars Released in Different Years

Three winter wheat (Triticum aestivum L.) cultivars, representatives of those widely cultivated in Beijing over the past six decades, were grown in the same environmental conditions. Net photosynthetic rate (P _N) per unit leaf area and instantaneous water use efficiency (WUE) of flag leaves increased with elevated CO₂ concentration. With an increase in CO₂ concentration from 360 to 720 µmol mol^–1, P _N and WUE of Jingdong 8 (released in 1990s and having the highest yield) increased by 173 and 81 %, while those of Nongda 139 (released in 1970s) increased by 88 and 66 %, and Yanda 1817 (released in 1945, with lowest yield) by 76 and 65 %. Jingdong 8 had the highest P _N and WUE values under high CO₂ concentration, but Yanda 1817 showed the lowest P _N. Stomatal conductance (g _s) of Nongda 139 and Yanda 1817 declined with increasing CO₂ concentration, but g _s of Jingdong 8 firstly went down and then up as the CO₂ concentration further increased. Intercellular CO₂ concentration (C _i) of Jingdong 8 and Nongda 139 increased when CO₂ concentration elevated, while that of Yanda 139 increased at the first stage and then declined. Jingdong 8 had the lowest C _i of the three wheat cultivars, and Yanda 1817 had the highest C _i value under lower CO₂ concentrations. However, Jingdong 8 had the highest P _N and lowest C _i at the highest CO₂ concentration which indicates that its photosynthetic potential may be high.     

Diurnal Changes of Gas Exchange,Chlorophyll Fluorescence,and Stomatal Aperture of Hybrid Poplar Clones Subjected to Midday Light Stress

Diurnal changes in net photosynthetic rate (P _N), chlorophyll (Chl) fluorescence, and stomatal aperture of several hybrid poplar clones subjected to midday light stress were measured in July and August of 1996. Midday depression of P _N, photosystem 2 (PS2) efficiency, stomatal conductance (g _s), and stomatal aperture was observed in all clones, though at differing rates among them. Non-uniform stomatal closure occurred at noon and at other times, requiring a modification of intercellular CO₂ concentration (C ₁). A linear relationship was found between g _s and stomatal aperture. More than half of the photons absorbed by PS2 centre dissipated thermally when subjected to light stress at noon. There was a linear relationship between the rate of PS2 photochemical electron transport (PxPFD) and P _N. There was a consensus for two fluorescence indicators (1 – q_P/q_N and (F_m' – F)/F_m') in assessment of susceptibility of photoinhibition in the clones. According to P _N, Chl fluorescence, and stomatal aperture, we conclude that midday depression of photosynthesis can be attributed to both stomatal and non-stomatal limitations.     

Acclimation of Two Distinct Plant Species,Spring Barley and Norway Spruce,to Combined Effect of Various Irradiance and CO2 Concentration During Cultivation in Controlled Environment

The short-term acclimation (10-d) of Norway spruce [Picea abies (L.) Karst] to elevated CO₂ concentration (EC) in combination with low irradiance (100 mol m^–2 s^–1) resulted in stimulation of CO₂ assimilation (by 61 %), increased total chlorophyll (Chl) content (by 17 %), significantly higher photosystem 2 (PS2) photochemical efficiency (F_v/F_m; by 4 %), and reduced demand on non-radiative dissipation of absorbed excitation energy corresponding with enhanced capacity of photon utilisation within PS2. On the other hand, at high cultivation irradiance (1 200 mol m^–2 s^–1) both Norway spruce and spring barley (Hordeum vulgare L. cv. Akcent) responded to EC by reduced photosynthetic capacity and prolonged inhibition of F_v/F_m accompanied with enhanced non-radiative dissipation of absorbed photon energy. Norway spruce needles revealed the expressive retention of zeaxanthin and antheraxanthin (Z+A) in darkness and higher violaxanthin (V) convertibility (yielding even 95 %) under all cultivation regimes in comparison with barley plants. In addition, the non-photochemical quenching of minimum Chl a fluorescence (SV₀), expressing the extent of non-radiative dissipation of absorbed photon energy within light-harvesting complexes (LHCs), linearly correlated with V conversion to Z+A very well in spruce, but not in barley plants. Finally, a key role of the Z+A-mediated non-radiative dissipation within LHCs in acclimation of spruce photosynthetic apparatus to high irradiance alone and in combination with EC was documented by extremely high SV₀ values, fast induction of non-radiative dissipation of absorbed photon energy, and its stability in darkness.     

Developmental Change in CO2 Compensation Concentrations in Spartina alterniflora Results from Sigmoidal Photosynthetic CO2 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 () 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 did not result from changing respiration, but rather from a sigmoidal response of net photosynthetic rate (P _N) 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 P _N 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.     

Gas Exchange of Carrot Leaves in Response to Elevated CO2 Concentration

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

Chlorophyll a Fluorescence Emission,Xanthophyll Cycle Activity,and Net Photosynthetic Rate Responses to Ozone in Some Foliose and Fruticose Lichen Species

The lichens Parmelia quercina, Parmelia sulcata, Evernia prunastri, Hypogymnia physodes, and Anaptychia ciliaris were exposed to ozone (O₃) in controlled environment cuvettes designed to maintain the lichens at optimal physiological activity during exposure. Measurements of gas exchange, modulated chlorophyll (Chl) fluorescence, and pigment analysis were conducted before and after exposure to 300 mm³ (O₃) m^–3, 4 h per d for 14 d. No changes in the efficiency of photosystem 2 (PS2) photochemistry, the reduction state of Q_A, or the electron flow through PS2, measured by Chl fluorescence, were detected in any of the five lichen species studied. Additionally, neither photosynthetic CO₂ assimilation nor xanthophyll cycle activity or photosynthetic pigment concentration were affected by high O₃ concentrations. Thus the studied lichen species have significant capacities to withstand oxidative stresses induced by high concentration of O₃.     

Photosynthetic Assimilation of Sun versus Shade Norway Spruce [Picea abies (L.) Karst] Needles Under the Long-Term Impact of Elevated CO2 Concentration

The long-term impact of elevated concentration of CO₂ on assimilation activity of sun-exposed (E) versus shaded (S) foliage was investigated in a Norway spruce stand [Picea abies (L.) Karst, age 14 years] after three years of cultivation in two domes with adjustable windows (DAW). One DAW was supplied with ambient air [AC, ca. 350 µmol(CO₂) mol^–1) and the second with elevated CO₂ concentration [EC = AC plus 350 µmol(CO₂) mol^–1]. The pronounced vertical profile of the photosynthetic photon flux density (PPFD) led to the typical differentiation of the photosynthetic apparatus between the shaded and sun needles. Namely, photon-saturated values of maximal net photosynthetic rate (P _Nmax) and apparent quantum yield () were significantly higher/lower for E-needles as compared with the S-ones. The prolonged exposure to EC was responsible for the apparent assimilatory activity stimulation observed mainly in deeply shaded needles. The degree of this stimulation decreases in the order: S-needles dense part > S-needles sparse part > E-needles dense part > E-needles sparse part. In exposed needles some signals on a manifestation of the acclimation depression of the photosynthetic activity were found. The long-term effect of EC was responsible for the decrease of nitrogen content of needles and for its smoother gradient between E- and S-needles. The obtained results indicate that the E- and S-foliage respond differently to the long-term impact of EC.     

Single-Turnover Flashes to Saturate the QA Reduction in a Leaf were Generated by the Light-Emitting Diodes from a Double Modulation Kinetic Chlorophyll Fluorometer

The earlier developed double-modulation chlorphyll (Chl) fluorometer was modified for measurements with intact leaves of higher plants. The Chl fluorometer is based on a non-periodic modulation of both actinic and measuring flashes. In addition, continuous orange actinic and far-red radiation were produced by separate arrays of light-emitting diodes (LEDs). Programmable timing of the flashes allows to cover a wide dynamic range from microseconds to minutes. We have demonstrated that the LEDs can produce single-turnover flashes that saturate Q_A reduction of intact leaves of Glyceria maxima and shoots of Picea abies. This revised version was published online in August 2006 with corrections to the Cover Date.