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

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

Irradiance influences tea leaf (<Emphasis Type="Italic">Camellia sinensis</Emphasis> L.) photosynthesis and transpiration

Rates of net photosynthesis (P _N) and transpiration (E), and leaf temperature (T_L) of maintenance leaves of tea under plucking were affected by photosynthetic photon flux densities (PPFD) of 200–2 200 μmol m⁻² s⁻¹. P _N gradually increased with the increase of PPFD from 200 to 1 200 μmol m⁻² s⁻¹ and thereafter sharply declined. Maximum P _N was 13.95 μmol m⁻² s⁻¹ at 1 200 μmol m⁻² s⁻¹ PPFD. There was no significant variation of P _N among PPFD at 1 400–1 800 μmol m⁻² s⁻¹. Significant drop of P _N occurred at 2 000 μmol m⁻² s⁻¹. PPFD at 2 200 μmol m⁻² s⁻¹ reduced photosynthesis to 6.92 μmol m⁻² s⁻¹. PPFD had a strong correlation with T_L and E. Both T_L and E linearly increased from 200 to 2 200 μmol m⁻² s⁻¹ PPFD. T_L and E were highly correlated. The optimum T_L for maximum P _N was 26.0 °C after which P _N declined significantly. E had a positive correlation with P _N.     

Effects of the interaction between ozone and carbon dioxide on gas exchange,photosystem II and antioxidants in rice leaves

To understand the interactive effects of O₃ and CO₂ on rice leaves; gas exchange, chlorophyll (Chl) fluorescence, ascorbic acid and glutathione were examined under acute (5 h), combined exposures of O₃ (0, 0.1, or 0.3 cm³ m⁻³, expressed as O₀, O_0.1, or O_0.3, respectively), and CO₂ (400 or 800 cm³ m⁻³, expressed as C⁴⁰⁰ or C⁸⁰⁰, respectively) in natural-light gas-exposure chambers. The net photosynthetic rate (P _N), maximum (F_v/F_m) and operating (F_q′/F_m′) quantum efficiencies of photosystem II (PSII) in young (8^th) leaves decreased during O₃ exposure. However, these were ameliorated by C⁸⁰⁰ and fully recovered within 3 d in clean air (O⁰ + C⁴⁰⁰) except for the O^0.3 + C⁴⁰⁰ plants. The maximum PSII efficiency at 1,500 μmol m⁻² s⁻¹ PPFD (F_v′/F_m′) for the O^0.3 + C₄₀₀ plants decreased for all measurement times, likely because leaves with severely inhibited P _N also had a severely damaged PSII. The P _N of the flag (16^th) leaves at heading decreased under O₃ exposure, but the decline was smaller and the recovery was faster than that of the 8^th leaves. The F_q′/F_m′ of the flag leaves in the O^0.3 + C⁴⁰⁰ and O^0.3 + C⁸⁰⁰ plants decreased just after gas exposure, but the F_v/F_m was not affected. These effects indicate that elevated CO₂ interactively ameliorated the inhibition of photosynthesis induced by O₃ exposure. However, changes in antioxidant levels did not explain the above interaction.     

Developmental phase-dependent photosynthetic responses to ultraviolet-B radiation: damage, defence, and adaptation of primary leaves of wheat seedlings

Alterations in photosynthetic capacity of primary leaves of wheat seedlings in response to ultraviolet-B (UV-B; 280–320 nm; 60 μmol m⁻² s⁻¹) exposure alone and in combination with photosynthetically active radiation (PAR; 400–800 nm; 200 μmol m⁻² s⁻¹) during different phases of leaf growth and development were assessed. UV-B exposure resulted in a phase-dependent differential loss in photosynthetic pigments, photochemical potential, photosystem 2 (PS2) quantum yield, and in vivo O₂ evolution. UV-B exposure induced maximum damage to the photosynthetic apparatus during senescence phase of development. The damages were partially alleviated when UV-B exposure was accompanied by PAR. UV-B induced an enhancement in accumulation of flavonoids during all phases of development while it caused a decline in anthocyanin content during senescence. The differential changes in these parameters demonstrated the adaptation ability of leaves to UV-B stress during all phases of development and the ability was modified in UV-B+ PAR exposed samples.     

Photosynthesis of lichen symbiotic alga Trebouxia erici as affected by irradiance and osmotic stress

The relation between oxygen evolution rate (OER) and quantum yield of photochemical reactions in photosystem 2 (Φ_PS2) was examined in lichen symbiotic alga Trebouxia erici Ahmadjian (strain UTEX 911) exposed to different irradiances and osmotic stress (2 M sucrose for 60 h). Linear relationship was found between OER and Φ_PS2 in control cell suspension within irradiance range of 0 – 500 μmol m⁻² s⁻¹. Under osmotic stress, OER and Φ_PS2 were significantly reduced. Relation between OER and Φ_PS2 was curvilinear due to strong osmotically-induced inhibition of OER at high irradiance. The highest used irradiance (500 μmol m⁻² s⁻¹) was photoinhibitory for osmotically-stressed T. erici because non-photochemical quenching (NPQ) increased substantially. Energy-dependent quenching represented major part of NPQ increase. Osmotic stress led also to the reduction of capacity of photochemical processes in PS 2 (F_V/F_M) and increase in F₀/F_M. These changes indicated negative effects of osmoticum on structure and function of photosynthetic apparatus.     

The impact of ozone fumigation and fertilization on chlorophyll fluorescence of birch leaves (Betula pendula)

 The impact of ozone fumigation on chlorophyll a fluorescence parameters and chlorophyll content of birch trees grown at high and low fertilization were studied for 6-, 8-, and 12-week old leaves. Fluorescence parameters were measured with a portable fluorometer with its fibre optics tightly inserted in a gas exchange cuvette at light intensities from 0 to 220 μmol photons m⁻² s⁻¹. Ozone caused significant changes of primary photosynthetic reactions: a decrease of the quantum yield of photosystem II and an increase of non-photochemical quenching. In all leaves a biphasic light response of non-photochemical quenching was observed. Ozone fumigation shifted the onset of the second phase from a PFD of about 60 μmol m⁻² s⁻¹ to about 30 μmol m⁻² s⁻¹. While the fertilizer concentration had no influence on this character, high fertilization supply of plants partially reduced O₃-induced damage. The light responses of Ft, Fm′ and NPQ observed in birch leaves grown in O₃-free air indicate the existence of at least two different processes governing energy conversion of the photosynthetic apparatus at PS II in the range of PFD 0–200 μmol photons m⁻² s⁻¹. The first phase was attributed to a rather slowly relaxing type of non-photochemical quenching, which, at least at low PFD, is thought to be related to a state 1–2 transition. The further changes of the fluorescence parameters studied at higher PFD might be explained by an increase of energy-dependent quenching, connected with the energization of the thylakoid membrane and zeaxanthin synthesis. A major effect of ozone treatment was a lowering of PS II quantum yield. This reflects a reduction of PS II electron transport and corresponds to the reduction of CO₂-fixation observed in ozonated leaves. Received: 24 September 1996 / Accepted: 27 January 1999     

Excess irradiance causes early symptoms of senescence during leaf expansion in photoautotrophically <Emphasis Type="Italic">in vitro</Emphasis> grown tobacco plants

Photosynthetic parameters, growth, and pigment contents were determined during expansion of the fourth leaf of in vitro photoautotrophically cultured Nicotiana tabacum L. plants at three irradiances [photosynthetically active radiation (400–700 nm): low, LI 60 μmol m⁻² s⁻¹; middle, MI 180 μmol m⁻² s⁻¹; and high, HI 270 μmol m⁻² s⁻¹]. During leaf expansion, several symptoms usually accompanying leaf senescence appeared very early in HI and then in MI plants. Symptoms of senescence in developing leaves were: decreasing chlorophyll (Chl) a+b content and Chl a/b ratio, decreasing both maximum (F_V/F_M) and actual (Φ_PS2) photochemical efficiency of photosystem 2, and increasing non-photochemical quenching. Nevertheless, net photosynthetic oxygen evolution rate (P _N) did not decrease consistently with decrease in Chl content, but exhibited a typical ontogenetic course with gradual increase. P _N reached its maximum before full leaf expansion and then tended to decline. Thus excess irradiance during in vitro cultivation did not cause early start of leaf senescence, but impaired photosynthetic performance and Chl content in leaves and changed their typical ontogenetic course.     

Photosynthetically versatile thin shade leaves: A paradox of irradiance-response curves

Thick sun leaves have a larger construction cost per unit leaf area than thin shade leaves. To re-evaluate the adaptive roles of sun and shade leaves, we compared the photosynthetic benefits relative to the construction cost of the leaves. We drew photosynthetically active radiation (PAR)-response curves using the leaf-mass-based photosynthetic rate to reflect the cost. The dark respiration rates of the sun and shade leaves of mulberry (Morus bombycis Koidzumi) seedlings did not differ significantly. At irradiances below 250 μmol m⁻² s⁻¹, the shade leaves tended to have a significantly larger net photosynthetic rate (P _N) than the sun leaves. At irradiances above 250 μmol m⁻² s⁻¹, the P _N did not differ significantly. The curves indicate that plants with thin shade leaves have a larger daily CO₂ assimilation rate per construction cost than those with thick sun leaves, even in an open habitat. These results are consistently explained by a simple model of PAR extinction in a leaf. We must target factors other than the effective assimilation when we consider the adaptive roles of thick sun leaves.     

Diurnal and seasonal changes in the intensity of photosynthesis in stems of lilac (Syringa vulgaris L.)

It has been demonstrated that during the whole year the stems are photosyntheticaly active and capable of assimilating atmospheric CO₂. The intensity of photosynthesis varies. During the vegetation period the registered net photosynthesis lasted up to 13 hours per day, and in the leafless period for 2–3 hours a day. Photosynthesis was registered also at temperatures below zero (−3 °C) as a reduced CO₂ evolution in light in comparison with darkness. The maximal net photosynthesis values during the vegetation period amounted to 6 up 8 μmol (CO₂)·m⁻²·s⁻¹, and in the leafless period 0.5 – 1 μmol (CO₂)·m⁻²·s⁻¹, and they were close to being up to twice as big as the values obtained of darkness respiration. An increase of the photosynthetic activity of stems preceded the spring development of the leaves.     

Susceptibility of green leaves and green flower petals of CAM orchid <Emphasis Type="Italic">Dendrobium</Emphasis> cv. Burana Jade to high irradiance under natural tropical conditions

Photosynthetic rates of green leaves (GL) and green flower petals (GFP) of the CAM plant Dendrobium cv. Burana Jade and their sensitivities to different growth irradiances were studied in shade-grown plants over a period of 4 weeks. Maximal photosynthetic O₂ evolution rates and CAM acidities [dawn/dusk fluctuations in titratable acidity] were higher in leaves exposed to intermediate sunlight [a maximal photosynthetic photon flux density (PPFD) of 500–600 μmol m⁻² s⁻¹] than in leaves grown under full sunlight (a maximal PPFD of 1 000–1 200 μmol m⁻² s⁻¹) and shade (a maximal PPFD of 200–250 μmol m⁻² s⁻¹). However, these two parameters of GFP were highest in plants grown under the shade and lowest in full sun-grown plants. Both GL and GFP of plants exposed to full sunlight had lower predawn F_v/F_m [dark adapted ratio of variable to maximal fluorescence (the maximal photosystem 2 yield without actinic irradiation)] than those of shade-grown plants. When exposed to intermediate sunlight, however, there were no significant changes in predawn F_v/F_m in GL whereas a significant decrease in predawn F_v/F_m was found in GFP of the same plant. GFP exposed to full sunlight exhibited a greater decrease in predawn F_v/F_m compared to those exposed to intermediate sunlight. The patterns of changes in total chlorophyll (Chl) content of GL and GFP were similar to those of F_v/F_m. Although midday F_v/F_m fluctuated with prevailing irradiance, changes of midday F_v/F_m after exposure to different growth irradiances were similar to those of predawn F_v/F_m in both GL and GFP. The decreases in predawn and midday F_v/F_m were much more pronounced in GFP than in GL under full sunlight, indicating greater sensitivity in GFP to high irradiance (HI). In the laboratory, electron transport rate and photochemical and non-photochemical quenching of Chl fluorescence were also determined under different irradiances. All results indicated that GFP are more susceptible to HI than GL. Although the GFP of Dendrobium cv. Burana Jade require a lower amount of radiant energy for photosynthesis and this plant is usually grown in the shade, is not necessarily a shade plant.     

Photosynthesis of a temperate fallow C<Subscript>3</Subscript> herbaceous ecosystem: measurements and model simulations at the leaf and canopy levels

The objectives of the study were to characterize photosynthesis of temperate fallow C₃herbaceous species and examine the performance of a simple photosynthesis model (based on the Farquhar’s equations) to simulate carbon fluxes at the leaf and canopy levels. The maximum rate of carboxylation at 25°C (V _m0) was estimated for sunlit leaves using in situ gas exchange data under saturating irradiance. Throughout the seasons, leaf measurements indicate that values of V _m0 were similar for the four major species of the fallow. The rate declined from March (100 μmol m⁻² s⁻¹) to July (50 μmol m⁻² s⁻¹) and remained almost constant until November. The maximum quantum yield estimated for Potentilla reptans L. (dominant species) was 0.082 mol(CO₂) mol⁻¹(photon absorbed), similar to values already published for C₃ species. Leaf area index (LAI) increased from winter (less than 0.2 m² m⁻²) to spring (up to 4 m² m⁻²). Rates of canopy photosynthesis (measured with a canopy chamber) strongly depended on LAI and temperature, in addition to irradiance. They reached a maximum of 25 μmol m⁻² s⁻¹ and were intermediate between those published for C₄ grassland or cultivated species, and on woody species. At leaf level, simulations gave realistic predictions. At canopy level, the model had the ability to reproduce the effects of environmental and seasonal conditions. However, simulations underestimated the photosynthetic activity of the fallow canopy.     

Irradiance prior to and during desiccation improves the tolerance to excess irradiance in the desiccated state of the old forest lichen Lobaria pulmonaria

Hydrated thalli of the lichen Lobaria pulmonaria were either preconditioned to dim irradiance (DI, 5 μmol m⁻² s⁻¹) or medium irradiance (MI, 200 μmol m⁻² s⁻¹) for 6 h. After this 6 h period, the thalli were allowed to desiccate under the two respective irradiances. Thereafter, these dry lichens were exposed to high irradiance (HI, 1 000 μmol m⁻² s⁻¹) for 60 h. After this HI treatment, the maximal photochemical quantum yield (F_V/F_M) and the de-epoxidation state of xanthophyll cycle pigments (DEPS) were highest in thalli preconditioned to MI. Hence irradiance in the last hydrated period before sampling is significant for the physiological state of lichens. A standardized irradiance pre-treatment before start of experiments is recommended.     

Photosystem 2 activity of <Emphasis Type="Italic">Citrus volkameriana</Emphasis> (L.) leaves as affected by Mn nutrition and irradiance

Citrus volkameriana (L.) plants were grown for 43 d in nutrient solutions containing 0, 2, 14, 98, or 686 μM Mn (Mn₀, Mn₂, Mn₁₄, Mn₉₈, and Mn₆₈₆, respectively). To adequately investigate the combined effects of Mn nutrition and irradiance on photosystem 2 (PS2) activity, irradiance response curves for electron transport rate (ETR), nonphotochemical quenching (q_N), photochemical quenching (q_P), and real photochemical efficiency of PS2 (Φ_PS2) were recorded under 10 different irradiances (66, 96, 136, 226, 336, 536, 811, 1 211, 1 911, and 3 111 μmol m⁻² s⁻¹, I₆₆ to I₃₁₁₁, respectively) generated with the PAM-2000 fluorometer. Leaf chlorophyll content was significantly lower under Mn excess (Mn₆₈₆) compared to Mn₀; its highest values were recorded in the treatments Mn₂-Mn₉₈. However, ETR and Φ_PS2 values were significantly lower under Mn₀ compared to the other Mn treatments, when plants were exposed to irradiances ≥96 μmol m⁻² s⁻¹. Furthermore, Mn₀ plants had significantly higher values of q_N and lower values of q_P at irradiances ≤226 and ≥336 μmol m⁻² s⁻¹, respectively, than those grown under Mn₂-Mn₆₈₆. Irrespective of Mn treatment, the values of Φ_PS2 and q_N decreased, while those of q_P increased progressively by increasing irradiance from I₁₃₆ to I₃₁₁₁. Finally, Mn₂-Mn₉₈ plants were less sensitive to photoinhibition of photosynthesis (≥811 μmol m⁻² s⁻¹) than the Mn₆₈₆ (≥536 μmol m⁻² s⁻¹) and Mn₀ (≥336 μmol m⁻² s⁻¹) ones.     

Stomatal and non-stomatal limitations to photosynthesis in field-grown grapevine cultivars

Diurnal changes of photosynthesis in the leaves of grapevine (Vitis vinifera × V. labrusca) cultivars Campbell Early and Kyoho grown in the field were compared with respect to gas exchanges and actual quantum yield of photosystem 2 (Φ_PS2) in late May. Net photosynthetic rate (P_N) of the two cultivars rapidly increased in the morning, saturated at photosynthetic photon flux density (PPFD) from 1200 to 1500 μmol m⁻² s⁻¹ between 10:00 and 12:00 and slowly decreased after midday. Maximum P_N was 13.7 and 12.5 μmol m⁻² s⁻¹ in Campbell Early and Kyoho, respectively. The stomatal conductance (g_s) and transpiration rate changed in parallel with P_N, indicating that P_N was greatly affected by g_s. However, the decrease in P_N after midday under saturating PPFD was also associated with the observed depression of Φ_PS2 at high PPFD. The substantial increase in the leaf to air vapour pressure deficit after midday might also contribute to decline of g_s and P_N.     

Photosynthetic Response of Carrots to Varying Irradiances

Response to irradiance of leaf net photosynthetic rates (P _N) of four carrot cultivars: Cascade, Caro Choice (CC), Oranza, and Red Core Chantenay (RCC) were examined in a controlled environment. Gas exchange measurements were conducted at photosynthetic active radiation (PAR) from 100 to 1 000 μmol m⁻² s⁻¹ at 20 °C and 350 μmol (CO₂) mol⁻¹(air). The values of P _N were fitted to a rectangular hyperbolic nonlinear regression model. P _N for all cultivars increased similarly with increasing PAR but Cascade and Oranza generally had higher P _N than CC. None of the cultivars reached saturation at 1 000 μmol m⁻² s⁻¹. The predicted P _N at saturation (P _Nmax) for Cascade, CC, Oranza, and RCC were 19.78, 16.40, 19.79, and 18.11 μmol (CO₂) m⁻² s⁻¹, respectively. The compensation irradiance (I _c) occurred at 54 μmol m⁻² s⁻¹ for Cascade, 36 μmol m⁻² s⁻¹ for CC, 45 μmol m⁻² s⁻¹ for Oranza, and 25 μmol m⁻² s⁻¹ for RCC. The quantum yield among the cultivars ranged between 0.057–0.033 mol(CO₂) mol⁻¹(PAR) and did not differ. Dark respiration varied from 2.66 μmol m⁻² s⁻¹ for Cascade to 0.85 μmol m⁻² s⁻¹ for RCC. As P _N increased with PAR, intercellular CO₂ decreased in a non-linear manner. Increasing PAR increased stomatal conductance and transpiration rate to a peak between 600 and 800 μmol m⁻² s⁻¹ followed by a steep decline resulting in sharp increases in water use efficiency. This revised version was published online in August 2006 with corrections to the Cover Date.     

Involvement of betacyanin in chilling-induced photoinhibition in leaves of <Emphasis Type="Italic">Suaeda salsa</Emphasis>

Seeds of Suaeda salsa were cultured in dark for 3 d and betacyanin accumulation in seedlings was promoted significantly. Then the seedlings with accumulated betacyanin (C+B) were transferred to 14/10 h light/dark and used for chilling treatment 15 d later. Photosystem 2 (PS2) photochemistry, D1 protein content, and xanthophyll cycle during the chilling-induced photoinhibition (exposed to 5 °C at a moderate photon flux density of 500 μmol m⁻² s⁻¹ for 3 h) and the subsequent restoration were compared between the C+B seedlings and the control (C) ones. The maximal efficiency of PS2 photochemistry (F_v/F_m), the efficiency of excitation energy capture by open PS2 centres (F_v′/F_m′), and the yield of PS2 electron transport (Φ_PS2) of the C+B and C leaves both decreased during photoinhibition. However, smaller decreases in F_v/F_m, F_v′/F_m′, and Φ_PS2 were observed in the C+B leaves than in C ones. At the same time, the deepoxidation state of xanthophyll cycle, indicated by (A+Z)/(V+A+Z) ratio, increased rapidly but the D1 protein content decreased considerably during the photoinhibition. The increase in rate of (A+Z)/(V+A+Z) was higher but the D1 protein turnover was slower in C+B than C leaves. After photoinhibition treatment, the plants were transferred to a dim irradiation (10 μmol m⁻² s⁻¹) at 25 °C for restoration. During restoration, the chlorophyll (Chl) fluorescence parameters, D1 protein content, and xanthophyll cycle components relaxed gradually, but the rate and level of restoration in the C+B leaves was greater than those in the C leaves. The addition of betacyanins to the thylakoid solution in vitro resulted in similar changes of F_v/F_m, D1 protein content, and (A+Z)/(V+A+Z) ratio during the chilling process. Therefore, betacyanin accumulation in S. salsa seedlings may result in higher resistance to photoinhibition, larger slowing down of D1 protein turnover, and enhancement of non-radiative energy dissipation associated with xanthophyll cycle, as well as in greater restoration after photoinhibition than in the control when subjected to chilling at moderate irradiance.     

Damaging mechanisms of chilling- and salt stress to <Emphasis Type="Italic">Arachis hypogaea</Emphasis> L. leaves

To investigate damaging mechanisms of chilling and salt stress to peanut (Arachis hypogaea L.) leaves, LuHua 14 was used in the present work upon exposure to chilling temperature (4°C) accompanied by high irradiance (1,200 μmol m⁻² s⁻¹) (CH), salt stress accompanied by high irradiance (1,200 μmol m⁻² s⁻¹) (SH), and high-irradiance stress (1,200 μmol m⁻² s⁻¹) at room temperature (25°C) (NH), respectively. Additionally, plants under low irradiance (100 μmol m⁻² s⁻¹) at room temperature (25°C) were used as control plants (CK). Relative to CK and NH treatments, both the maximal photochemical efficiency of PSII (F_v/F_m) and the absorbance at 820 nm decreased greatly in peanut leaves under CH and SH stress, which indicated that severe photoinhibition occurred in peanut leaves under such conditions. Initial fluorescence (F_o), 1 − q_P and nonphotochemical quenching (NPQ) in peanut leaves significantly increased under CH- and SH stress. Additionally, the activity of superoxide dismutase (SOD), one of the key enzymes of water-water cycle, decreased greatly, the accumulation of malondialdehyde (MDA) and membrane permeability increased. These results suggested that damages to peanut photosystems might be related to the accumulation of reactive oxygen species (ROS) induced by excess energy, and the water-water cycle could not dissipate energy efficiently under the stress of CH and SH, which caused the accumulation of ROS greatly. CH and SH had similar damaging effects on peanut photosystems, except that CH has more severe effects. All the results showed that CH- and SH stress has similar damaging site and mechanisms in peanut leaves.     

A coupled model of stomatal conductance and photosynthesis for winter wheat

The model couples stomatal conductance (g _s) and net photosynthetic rate (P _N) describing not only part of the curve up to and including saturation irradiance (I _max), but also the range above the saturation irradiance. Maximum stomatal conductance (g _smax) and I _max can be calculated by the coupled model. For winter wheat (Triticum aestivum) the fitted results showed that maximum P _N (P _max) at 600 μmol mol⁻¹ was more than at 350 μmol mol⁻¹ under the same leaf temperature, which can not be explained by the stomatal closure at high CO₂ concentration because g _smax at 600 μmol mol⁻¹ was less than at 350 μmol mol⁻¹. The irradiance-response curves for winter wheat had similar tendency, e.g. at 25 °C and 350 μmol mol⁻¹ both P _N and g _s almost synchronously reached the maximum values at about 1 600 μmol m⁻² s⁻¹. At 25 °C and 600 μmol mol⁻¹ the I _max corresponding to P _max and g _smax was 2 080 and 1 575 μmol m⁻² s⁻¹, respectively.     

Differences in photosynthetic apparatus of leaves from different sides of the chestnut canopy

In crowns of chestnut trees the absorption of radiant energy is not homogeneous; leaves from the south (S) side are the most irradiated, but leaves from the east (E) and west (W) sides receive around 70 % and those from north (N) face less than 20 % of the S irradiation. Compared to the S leaves, those from the N side were 10 % smaller, their stomata density was 14 % smaller, and their laminae were 21 % thinner. N leaves had 0.63 g(Chl) m⁻², corresponding to 93 % of total chlorophyll (Chl) amount in leaves of S side. The ratios of Chl a/b were 2.9 and 3.1 and of Chl/carotenoids (Car) 5.2 and 4.8, respectively, in N and S leaves. Net photosynthetic rate (P _N) was 3.9 μmol(CO₂) m⁻² s⁻¹ in S leaves, in the E, W, and N leaves 81, 77, and 38 % of that value, respectively. Morning time (10:00 h) was the period of highest P _N in the whole crown, followed by 13:00 h (85 % of S) and 16:00 h with 59 %. Below 500 μmol m⁻² s⁻¹ of photosynthetic photon flux density (PPFD), N leaves produced the highest P _N, while at higher PPFD, the S leaves were most active. In addition, the fruits from S side were 10 % larger than those from the N side.     

Chlorophyll fluorescence imaging of photosynthetic activity in sun and shade leaves of trees

The differences in pigment levels, photosynthetic activity and the chlorophyll fluorescence decrease ratio R _Fd (as indicator of photosynthetic rates) of green sun and shade leaves of three broadleaf trees (Platanus acerifolia Willd., Populus alba L., Tilia cordata Mill.) were compared. Sun leaves were characterized by higher levels of total chlorophylls a + b and total carotenoids x + c as well as higher values for the weight ratio chlorophyll (Chl) a/b (sun leaves 3.23–3.45; shade leaves: 2.74–2.81), and lower values for the ratio chlorophylls to carotenoids (a + b)/(x + c) (with 4.44–4.70 in sun leaves and 5.04–5.72 in shade leaves). Sun leaves exhibited higher photosynthetic rates P _N on a leaf area basis (mean of 9.1–10.1 μmol CO₂ m⁻² s⁻¹) and Chl basis, which correlated well with the higher values of stomatal conductance G _s (range 105–180 mmol m⁻² s⁻¹), as compared to shade leaves (G _s range 25–77 mmol m⁻² s⁻¹; P _N: 3.2–3.7 μmol CO₂ m⁻² s⁻¹). The higher photosynthetic rates could also be detected via imaging the Chl fluorescence decrease ratio R _Fd, which possessed higher values in sun leaves (2.8–3.0) as compared to shade leaves (1.4–1.8). In addition, via R _Fd images it was shown that the photosynthetic activity of the leaves of all trees exhibits a large heterogeneity across the leaf area, and in general to a higher extent in sun leaves than in shade leaves.