Leaf gas exchange,chlorophyll fluorescence and growth responses of <Emphasis Type="Italic">Melaleuca alternifolia</Emphasis> seedlings to flooding and subsequent recovery

Periodic flooding of trees in tropical floodplains and reservoirs where water levels fluctuate is a common phenomenon. The effects of flooding and subsequent recovery on gas exchange, chlorophyll fluorescence and growth responses of Melaleuca alternifolia seedlings, a tall shrub species used in floodplain and reservoir forest restoration in southern China, were studied during a grow season (from March to December in 2007). M. alternifolia seedlings were flooded for 180 days, drained and left to recover for another 60 days. Survival rates of the seedlings were 100% during the 180-day flooding period. Chlorophyll (Chl) content, net photosynthetic rate (P _N), stomatal conductance (g _s), and transpiration rate (E) of the flooded seedlings were all significantly lower than those of the control. Significant reductions of photochemical quenching coefficient (q_p) and increases of nonphotochemical quenching (NPQ) in the flooded seedlings were observed. However, there were no significant differences in the maximal quantum yield of PSII photochemistry (F_v/F_m) between treatments. All seedlings survived during the two-month recovery period after the flooded treatment was drained, and the biomass and height of the recovered seedlings approached those of the control at the end of the experiment. During the first-month recovery period, Chl content, P _N, g _s and E in the recovered seedlings were all obviously low, then increased gradually and rose to the levels similar to the control by the end of the experiment. Quenching analysis revealed significant reductions of q_p and increments of NPQ in the recovered seedlings at the beginning of draining, and a nearly complete recovery for both parameters by the end of the experiment. However, F_v/F_m of the recovered seedlings did not differ significantly from the control during the recovery period. Our study demonstrated that M. alternifolia seedlings can survive and grow through 180 days of flooding with a subsequent 60-day recovery period in drained conditions, indicating that seedlings of this species would be suitable for afforestation in areas exposed to intermittent flooding.     

Photoprotective function of betacyanin in leaves of <Emphasis Type="Italic">Amaranthus cruentus</Emphasis> L. under water stress

The photoprotective function of leaf betacyanin in water-stressed Amaranthus cruentus plants was examined by comparing leaves of two strains which differ significantly in the amount of betacyanin. At 0, 1, and 2 days after the imposed water stress, leaves were subjected to high-light (HL) treatment to assess their photosynthetic capacity and photoinhibition susceptibility. The water stress equally reduced leaf relative water content (RWC), gas-exchange rate and chlorophyll (Chl) contents in both leaves, indicating that the severity of water stress was comparable between the strains. Consequently, the extent of photoinhibition after the HL treatment increased in both strains as water stress developed; however, it was significantly greater in acyanic leaves than in betacyanic leaves, suggesting lower photoinhibition susceptibility in the betacyanic strain. The betacyanic leaves also exhibited approximately 30% higher values for photochemical quenching coefficient (q_P) during the period of water stress despite the nonphotochemical quenching coefficient (q_N) did not differ significantly between the strains. These results may be partially explained by the increased amount of leaf betacyanin under water stress. Moreover, a decrease in Chl content in betacyanic leaves might have enhanced light screening effect of betacyanin by increasing relative abundance of betacyanin to Chl molecule. In addition, reduced Chl content increased light penetrability of leaves. As a result, the extent of photoinhibition at the deeper tissue was exacerbated and the Chl fluorescence emitted from these tissues was more readily detected, facilitating assessment of photoinhibition at deeper tissues where the effect of betacyanic light screening is considered to be most apparent. Our results demonstrated that leaf betacyanin contributes to total photoprotective capacity of A. cruentus leaves by lowering excitation pressure on photosystem II (PSII) via attenuation of potentially harmful excess incident light under water stress.     

Lanthanum improves salt tolerance of maize seedlings

In this study, the effects of lanthanum were investigated on contents of pigments, chlorophyll (Chl) fluorescence, antioxidative enzymes, and biomass of maize seedlings under salt stress. The results showed that salt stress significantly decreased the contents of Chl and carotenoids, maximum photochemical efficiency of PSII (F_v/F_m), photochemical quenching (q_P), and quantum efficiency of PSII photochemistry (Φ_PSII), net photosynthetic rate (P_N), and biomass. Salt stress increased nonphotochemical quenching (q_N), the activities of ascorbate peroxidase, catalase, superoxide dismutase, glutathione peroxidase, and the contents of malondialdehyde and hydrogen peroxide compared with control. Pretreatment with lanthanum prior to salt stress significantly enhanced the contents of Chl and carotenoids, F_v/F_m, q_P, q_N, Φ_PSII, P_N, biomass, and activities of the above antioxidant enzymes compared with the salt-stressed plants. Pretreatment with lanthanum also significantly reduced the contents of malondialdehyde and hydrogen peroxide induced by salt stress. Our results suggested that lanthanum can improve salt tolerance of maize seedlings by enhancing the function of photosynthetic apparatus and antioxidant capacity.     

Use of a SPAD meter to estimate chlorophyll content in Eugenia uniflora L. leaves as affected by contrasting light environments and soil flooding

In three separate experiments, the effectiveness of a SPAD-502 portable chlorophyll (Chl) meter was evaluated for estimating Chl content in leaves of Eugenia uniflora seedlings in different light environments and subjected to soil flooding. In the first experiment, plants were grown in partial or full sunlight. In the second experiment plants were grown in full sunlight for six months and then transferred to partial sunlight or kept in full sunlight. In the third experiment plants were grown in a shade house (40% of full sunlight) for six months and then transferred to partial shade (25–30% of full sunlight) or full sunlight. In each experiment, plants in each light environment were either flooded or not flooded. Non-linear regression models were used to relate SPAD values to leaf Chl content using a combination of the data obtained from all three experiments. There were no significant effects of flooding treatments or interactions between light and flooding treatments on any variable analyzed. Light environment significantly affected SPAD values, chlorophyll a (Chl a), chlorophyll b (Chl b), and total chlorophyll [Chl (a+b)] contents in Experiment I (p≤0.01) and Experiment III (p≤0.05). The relationships between SPAD values and Chl contents were very similar among the three experiments and did not appear to be influenced by light or flooding treatments. There were high positive exponential relationships between SPAD values and Chl (a+b), Chl a, and Chl b contents.     

Changes in the photosynthetic characteristics and photosystem stoichiometries in wild-type and Chl <Emphasis Type="Italic">b</Emphasis>-deficient mutant rice seedlings under various irradiances

By using a wild-type rice (Oryza sativa L. cv. Norin No. 8) and the chlorophyll (Chl) b-deficient mutant derived from Norin No. 8 (chlorina 11), the present study monitored the oxygen evolution, contents of Chl a and b, β-carotene, and lutein in leaf and the contents of cytochrome f, and the reaction centres of photosystem I (PSI) and photosystem II (PSII) in thylakoids. The oxygen evolution, maximal quantum yield of PSII (F_v/F_m) and Chl concentration remained constant in both Norin No. 8 and chlorina 11 under 5 and 2% of full sunlight for six days. On the other hand, on the thylakoid level, the PSII reaction centre of chlorina 11 was more stable even under high irradiance, while approximately 40% decrease in levels of the PSII reaction centre occurred under 2% of full sunlight for six days. However, under such conditions, by regulating the stoichiometry of active PSII and PSI centres, the light absorption balance in both rice types was adjusted between the two photosystems. The present study attempted to examine whether the light absorption balance between PSII and PSI is altered to effectively conduct photosynthesis in the wild-type and Chl b-deficient mutant rice seedlings.     

Role of xanthophyll cycle-mediated photoprotection in <Emphasis Type="Italic">Arbutus unedo</Emphasis> plants exposed to water stress during the Mediterranean summer

We analyzed the response of potted strawberry tree (Arbutus unedo L.) seedlings exposed to water stress by withholding water for 10 d (WS). Leaf water potential, net CO₂ assimilation, and stomatal conductance decreased with increasing water deficit. A 30 % reduction of chlorophyll (Chl) content in the antenna complexes was observed in WS-plants. Simultaneously, a decline of photochemical efficiency (F_v/F_m) occurred as a result of an excess of solar radiation energy when carbon assimilation was limited by stomata closure due to soil water deficit. The non-photochemical quenching of Chl fluorescence (Φ_NPQ) significantly increased, as well as the leaf contents of zeaxanthin (Z) and antheraxanthin (A) at the expense of violaxanthin during the WS-period. Elevated predawn contents of de-epoxidized xanthophyll cycle components were associated with a sustained lowering of predawn photosystem 2 efficiency; this suggested an engagement of Z+A in a state primed for energy dissipation. Thus, the ability of strawberry trees to maintain the functionality of the xanthophyll cycle during the Mediterranean summer is an efficient mechanism to prevent irreversible damages to the photosynthetic machinery through thermal energy dissipation in the antenna and the reduction in photochemical efficiency.     

Physiological behaviors of <Emphasis Type="Italic">Acer mono</Emphasis> under drought and low light

The seedlings of Acer mono Maxim. were exposed to two watering regimes (well watered (100% of field capacity) and drought (30% of field capacity)) and two light levels (high light (100% of full sunlight) and low light (15% of full sunlight)) in a greenhouse to assess growth, photosynthesis, and foliar nutrient traits of woody plants under drought and low light. Drought significantly reduced growth and gas exchange characteristics of A. mono, including net photosynthetic rate (P _N), stomatal conductance (g _s), intercellular CO₂ concentration (C_i), and photosynthetic nitrogen use efficiency (PNUE). Also, drought reduced relative water content (RWC) and foliar C and N concentrations, but increased the C/N ratio. P _N and C concentration were lower under drought and low light than in any other treatment, indicating that deep shade might seriously decrease C assimilation. However, the negative effect induced by drought was alleviated by improving RWC and maintaining C balance, and therefore low-light seedlings accumulated more biomass than those under high light when they were exposed to drought. Our results showed that trade-off and facilitation effects of drought and low light might be complementary and alter in different species.     

Method for resolution and quantification of components of the non-photochemical quenching (<Emphasis Type="Italic">q</Emphasis><Subscript>N</Subscript>)

A new method of the chlorophyll (Chl) a fluorescence quenching analysis is described, which allows the calculation of values of (at least) three components of the non-photochemical quenching of the variable Chl a fluorescence (q _N) using a non-linear regression of a multi-exponential function within experimental data. Formulae for coefficients of the “energy”-dependent (ΔpH-dependent) quenching (q _E), the state-transition quenching (q _T) and the photo/inhibitory quenching (q _I) of Chl a fluorescence were found on the basis of three assumptions: (i) the dark relaxation kinetics of q _N, as well as of all its components, is of an exponential nature, (ii) the superposition principle is valid for individual Chl a fluorescence quenching processes and (iii) the same reference fluorescence level (namely the maximum variable Chl a fluorescence yield in the dark-adapted state, F _V) is used to define both q _N and its components. All definitions as well as the algorithms for analytical recognition of the q _N components are theoretically clarified and experimentally tested. The described theory results in a rather simple equation allowing to compute values for all q _N components (q _E, q _T, q _I) as well as the half-times of relaxation (τ_1/2) of corresponding quenching processes. It is demonstrated that under the above assumptions it holds: q _N = q _E + q _T + q _I. The theoretically derived equations are tested, and the results obtained are discussed for non-stressed and stressed photosynthetically active samples. Semi-empirical formulae for a fast estimation of values of the q _N components from experimental data are also given.     

Morphological plasticity,photosynthesis and chlorophyll fluorescence of <Emphasis Type="Italic">Athyrium pachyphlebium</Emphasis> at different shade levels

Athyrium pachyphlebium C. is a popular ornamental fern with considerable shade tolerance. The aim of this study was to investigate how the mature sporophytes acclimate to different light levels and to obtain an optimal light environment for their growth both in natural forest canopy and in urban landscapes. Plant growth and morphology, photosynthetic light-response curves and chlorophyll (Chl) fluorescence were measured at four different light levels (45% full sunlight, 30%, 20% and 8%). As the light intensities declined from 45% to 20%, seedling height, crown growth, foliage number and plant lifespan increased significantly. Seedlings grown at 20% light level were vigorous with great ornamental value. Plants grown in deep shade (8% light) showed severe symptoms of lodging and in 45% full sun, the plants showed highlight-stress symptoms. Seedlings in high light levels exhibited a higher light-saturated photosynthetic rate (P _max), light compensation point (LCP), light saturation point (LSP) and a reduced ability for nonphotochemical quenching (NPQ) of excess light than those in low light levels. However, seedlings in low light exhibited greater efficiency in absorbing and utilizing light energy, characterized by higher chlorophyll b (Chl b) and electron transport rate (ETR). These results indicated that a light level of about 20% full sun appeared to be optimal for A. pachyphlebium when both physiological and morphological performance in the landscape were considered.     

Photosynthetic performance of <Emphasis Type="Italic">Lycoris radiata</Emphasis> var. <Emphasis Type="Italic">radiata</Emphasis> to shade treatments

The effects of shade on the growth, leaf photosynthetic characteristics, and chlorophyll (Chl) fluorescence parameters of Lycoris radiata var. radiata were determined under differing irradiances (15, 65, and 100% of full irradiance) within pots. The HI plants exhibited a typical decline in net photosynthetic rate (P _N) during midday, which was not observed in MI- and LI plants. This indicated a possible photoinhibition in HI plants as the ratio of variable to maximum fluorescence (F_v/F_m) value was higher and the minimal fluorescence (F₀) was lower in the, and LI plants. Diurnal patterns of stomatal conductance (g _s) and transpiration rate (E) were remarkably similar to those of P _N at each shade treatments, and the intercellular CO₂ concentration (C _i) had the opposite change trend. Under both shading conditions, the light saturation point, light compensation point and photon-saturated photosynthetic rate (P _max) became lower than those under full sunlight, and it was the opposite for the apparent quantum yield (AQY). The higher the level of shade, the lower the integrated daytime carbon gain, stomatal and epidermis cell densities, specific leaf mass (SLM), bulb mass ratio (BMR), leaf thickness, and Chl a/b ratio. In contrast, contents of Chls per dry mass (DM), leaf area ratio (LAR), leaf mass ratio (LMR), leaf length, leaf area and total leaf area per plant increased under the same shade levels to promote photon absorption and to compensate for the lower radiant energy. Therefore, when the integrated daytime carbon gain, leaf area and total leaf area per plant, which are the main factors determining the productivity of L. radiata var. radiata plant, were taken into account together, this species may be cultivated at about 60∼70% of ambient irradiance to promote its growth.     

Photosystem 2 photochemistry and pigment composition of a yellow mutant of rice (<Emphasis Type="Italic">Oryza sativa</Emphasis> L.) under different irradiances

A yellow leaf colouration mutant (named ycm) generated from rice T-DNA insertion lines was identified with less grana lamellae and low thylakoid membrane protein contents. At weak irradiance [50 μmol(photon) m⁻² s⁻¹], chlorophyll (Chl) contents of ycm were ≈20 % of those of WT and Chl a/b ratios were 3-fold that of wild type (WT). The leaf of ycm showed lower values in the actual photosystem 2 (PS2) efficiency (Φ_PS2), photochemical quenching (q_P), and the efficiency of excitation capture by open PS2 centres 1 (F_v′/F_m′) than those of WT, except no difference in the maximal efficiency of PS2 photochemistry (F_v/F_m). With progress in irradiance [100 and 200 μmol(photon) m⁻² s⁻¹], there was a change in the photosynthetic pigment stoichiometry. In ycm, the increase of total Chl contents and the decrease in Chl a/b ratio were observed. Φ_PS2, q_P, and F_v′/F_m′ of ycm increased gradually along with the increase of irradiance but still much less than in WT. The increase of xanthophyll ratio [(Z+A)/(V+A+Z)] associated with non-photochemical quenching (q_N) was found in ycm which suggested that ycm dissipated excess energy through the turnover of xanthophylls. No significant differences in pigment composition were observed in WT under various irradiances, except Chl a/b ratio that gradually decreased. Hence the ycm mutant developed much more tardily than WT, which was caused by low photon energy utilization independent of irradiance.     

Effects of nitrogen supply on photosynthetic and anatomical changes in current-year needles of <Emphasis Type="Italic">Pinus koraiensis</Emphasis> seedlings grown under two irradiances

We investigated the responses of photon-saturated photosynthesis rate (P _sat) and its simultaneous acclimation of anatomy and nitrogen use patterns of current needles of Korean pine (Pinus koraiensis) seedlings grown under factorial combinations of two nitrogen levels and irradiances. Although N supply resulted in a significant increase of N content in needles under both irradiances, the increase of P _sat tended to be suppressed only in shade (S). The significant increase of P _sat in full sunlight (O) was associated with the increase of ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBPCO) and chlorophyll (Chl) contents. In contrast, small increase of Chl content and no increase of RuBPCO content were found in S (90 % cut of full irradiance), which would result in a small increase of P _sat. This result suggests that extra N is stocked in needles under shade for the growth in next season. With N supply, a significant decrease of specific leaf area (SLA) was detected only in O. This decrease of SLA was due to the increase of density of needle. Furthermore, the increase of needle density was not due to the increased number and size of mesophyll cells, but the increased density of each mesophyll cell. Therefore, although SLA changed in O, the change did not involve anatomical adaptation to use increased N effectively, at least observable by light microscopy. Hence, even though the SLA would change, N deposition will improve the photosynthetic capacity of Korean pine seedlings, not through the development of needle anatomy but through improvement of the allocation of N in both irradiances.     

Effects of copper and cadmium on photosynthesis in cucumber cotyledons

The effects of 20 and 50 μM concentrations of Cu and Cd on photosynthesis in cucumber (Cucumis sativus L.) cotyledons were studied by the measurements of gas exchange characteristics, chlorophyll (Chl) fluorescence parameters, photosynthetic pigment contents, and two Calvin cycle enzymes activities: glyceraldehyde-3-phosphate dehydrogenase (GAPDH) and 3-phosphoglyceric acid kinase (PGK). To minimize indirect metal action, seedlings were treated with metals in the stage of green, fully developed cotyledons. The metals reached the cotyledon tissue after 48 h of treatments, though symptoms of metal action were not visible at that time. The effect of metals on the light phase of the photosynthesis parameters such as potential efficiency of photosystem 2 (PS2; F_v/F_m), and photochemical and nonphotochemical quenching of Chl fluorescence (q_P and q_NP) was negligible. In contrast, a decrease of PS2 quantum efficiency (Φ_PS2) was much more noticeable. Changes in the pigment contents were slight, as only 50 μM Cd decreased Chl a and b contents in small extent. On the contrary, metals in both concentrations drastically decreased (50 and more % of control) the net photosynthetic rate and the stomatal conductance, but not the internal CO₂ concentration. The activities of both GAPDH and PGK were also decreased by metals, although the effect on PGK was more prominent, particularly on its potential activity (dithiothreitol in extraction and incubation media). Hence Cu and Cd affected the synthesis of enzyme proteins rather than they influenced their modifications. The effects of both metals on most of the measured photosynthesis parameters were similar, but the accumulation of Cd in the cotyledons was significantly higher than Cu accumulation. Thus Cu was more toxic for the photosynthesis of cucumber cotyledons than Cd.     

Effect of Different Sugars on Photosynthesis and Chlorophyll Fluorescence in Photoautotrophic Tomato Suspension Cell Cultures

The effects of metabolisable sugars sucrose and glucose along with non-metabolisable isomers of sucrose palatinose and turanose were tested. Rate of oxygen evolution (P), electron transport rate (ETR), and photochemical quenching (q_p) showed substantial decrease after 24 and 48 h by glucose and sucrose treatments, whereas there was no effect on all these parameters by the treatment with palatinose and turanose. Also the F_v/F_m ratio remained constant through the time of studies revealing that the maximal photochemical capacity of the cells was unchanged. Non-photochemical quenching (q_N) showed a decrease compared to the control values by all the treatments. Hence P and Chl fluorescence parameter were affected only by those sugars which are used in the metabolic pathways and not by sugar analogues.     

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.).     

Enhancement of susceptivity to photoinhibition and photooxidation in rice chlorophyll <Emphasis Type="Italic">b</Emphasis>-less mutants

Two rice chlorophyll (Chl) b-less mutants (VG28-1, VG30-5) and the respective wild type (WT) plant (cv. Zhonghua No. 11) were analyzed for the changes in Chl fluorescence parameters, xanthophyll cycle pool, and its de-epoxidation state under exposure to strong irradiance, SI (1 700 μmol m⁻² s⁻¹). We also examined alterations in the chloroplast ultrastructure of the mutants induced by methyl viologen (MV) photooxidation. During HI (0–3.5 h), the photoinactivation of photosystem 2 (PS2) appeared earlier and more severely in Chl b-less mutants than in the WT. The decreases in maximal photochemical efficiency of PS2 in the dark (F_v/F_m), quantum efficiency of PS2 electron transport (Φ_PS2), photochemical quenching (q_P), as well as rate of photochemistry (P_rate), and the increases in de-epoxidation state (DES) and rate of thermal dissipation of excitation energy (D_rate) were significantly greater in Chl b-mutants compared with the WT plant. A relatively larger xanthophyll pool and 78–83 % conversion of violaxanthin into antheraxanthin and zeaxanthin in the mutants after 3.5 h of HI was accompanied with a high ratio of inactive/total PS2 (0.55–0.73) and high 1–q_P (0.57–0.68) which showed that the activities of the xanthophyll cycle were probably insufficient to protect the photosynthetic apparatus against photoinhibition. No apparent difference of chloroplast ultrastructure was found between Chl b-less mutants and WT plants grown under low, LI (180 μmol m⁻² s⁻¹) and high, HI (700 μmol m⁻² s⁻¹) irradiance. However, swollen chloroplasts and slight dilation of thylakoids occurred in both mutants and the WT grown under LI followed by MV treatment. These typical symptoms of photooxidative damage were aggravated as plants were exposed to HI. Distorted and loose scattered thylakoids were observed in particular in the Chl b-less mutants. A greater extent of photoinhibition and photooxidation in these mutants indicated that the susceptibility to HI and oxidative stresses was enhanced in the photosynthetic apparatus without Chl b most likely as a consequence of a smaller antenna size.     

How to correctly determine the different chlorophyll fluorescence parameters and the chlorophyll fluorescence decrease ratio R<Subscript>Fd</Subscript> of leaves with the PAM fluorometer

This contribution is a practical guide to the measurement of the different chlorophyll (Chl) fluorescence parameters and gives examples of their development under high-irradiance stress. From the Chl fluorescence induction kinetics upon irradiation of dark-adapted leaves, measured with the PAM fluorometer, various Chl fluorescence parameters, ratios, and quenching coefficients can be determined, which provide information on the functionality of the photosystem 2 (PS2) and the photosynthetic apparatus. These are the parameters F_v, F_m, F₀, F_m′, F_v′, NF, and ΔF, the Chl fluorescence ratios F_v/F_m, F_v/F₀, ΔF/F_m′, as well as the photochemical (q_P) and non-photochemical quenching coefficients (q_N, q_CN, and NPQ). q_N consists of three components (q_N = q_E + q_T + q_I), the contribution of which can be determined via Chl fluorescence relaxation kinetics measured in the dark period after the induction kinetics. The above Chl fluorescence parameters and ratios, many of which are measured in the dark-adapted state of leaves, primarily provide information on the functionality of PS2. In fully developed green and dark-green leaves these Chl fluorescence parameters, measured at the upper adaxial leaf side, only reflect the Chl fluorescence of a small portion of the leaf chloroplasts of the green palisade parenchyma cells at the upper outer leaf half. Thus, PAM fluorometer measurements have to be performed at both leaf sides to obtain information on all chloroplasts of the whole leaf. Combined high irradiance (HI) and heat stress, applied at the upper leaf side, strongly reduced the quantum yield of the photochemical energy conversion at the upper leaf half to nearly zero, whereas the Chl fluorescence signals measured at the lower leaf side were not or only little affected. During this HL-stress treatment, q_N, q_CN, and NPQ increased in both leaf sides, but to a much higher extent at the lower compared to the upper leaf side. q_N was the best indicator for non-photochemical quenching even during a stronger HL-stress, whereas q_CN and NPQ decreased with progressive stress even though non-photochemical quenching still continued. It is strongly recommended to determine, in addition to the classical fluorescence parameters, via the PAM fluorometer also the Chl fluorescence decrease ratio R_Fd (F_d/F_s), which, when measured at saturation irradiance is directly correlated to the net CO₂ assimilation rate (_{P N}) of leaves. This R_Fd-ratio can be determined from the Chl fluorescence induction kinetics measured with the PAM fluorometer using continuous saturating light (cSL) during 4–5 min. As the R_Fd-values are fast measurable indicators correlating with the photosynthetic activity of whole leaves, they should always be determined via the PAM fluorometer parallel to the other Chl fluorescence coefficients and ratios.     

Chlorophyll content and photosystem II efficiency in soybean exposed to supplemental ultraviolet-B radiation

Chlorophyll (Chl) a fluorescence parameters and rapid light curves of soybean [Glycine max (L.) Merrill] were measured by pulse amplitude modulation fluorometry. Measurements were taken during different stages of soybean growth under field conditions with 20% enhancement in ultraviolet-B (UV-B) radiation. Results showed that supplemental UV-B radiation decreased Chl contents by 5.5% (P=0.048), 8.7% (P=0.046), and 10.5% (P=0.005) in seedling, in branching-flowering, and in pod-setting stages, respectively. In the branching-flowering and pod-setting stages, maximum quantum yield of photosystem (PS) II photochemistry (F_v/F_m) decreased by 6.1% (P=0.001) and 3.0% (P=0.009), respectively. Supplemental UV-B radiation significantly decreased the effective quantum yield (Y). The photosynthetic capacity at light saturation (P_m) also decreased in both the seedling and branching-flowering stages by 28.9% (P=0.007) and 15.5% (P=0.041), respectively. However, Y and Pm showed no significant difference in the trefoil and pod-setting stages with and without the UV treatment. The light saturation parameter (E _k) decreased by 21.1% (P=0.000) and 23.2% (P=0.029) in the trefoil and seedling stages, respectively. Moreover, the initial slope (α) decreased by 21.1% (P=0.001) in the branching-flowering stage. Nonphotochemical quenching (NPQ) in the seedling stage and photochemical quenching coefficient (q_p) in the branching-flowering stage decreased significantly under UV-B treatments. The results of the present study suggest that supplemental UV-B radiation adversely affected Chl content and electron transport activity in PSII and consequently decreased the photosynthetic efficiency of soybean plants.     

Responses of photosynthetic parameters of <Emphasis Type="Italic">Mikania micrantha</Emphasis> and <Emphasis Type="Italic">Chromolaena odorata</Emphasis> to contrasting irradiance and soil moisture

Photosynthetic parameters were measured in two invasive weeds, Mikania micrantha and Chromolaena odorata, grown in soil under full, medium, and low irradiance and full, medium, and low water supply. Both species showed significantly higher net photosynthetic rate, quantum yield of PS 2 photochemistry and photochemical quenching coefficient under high than low irradiance. For M. micrantha, low irradiance caused decreased chlorophyll content (Chl), Chl a/b ratio and maximum photochemical efficiency of PS 2 (F_v/F_m), while drought decreased Chl content and F_v/F_m and increased nonphotochemical quenching (NPQ). However, these parameters were much less affected in C. odorata except that Chl content and NPQ slightly increased under drought and high irradiance. High irradiance increased xanthophyll pools in both species, especially M. micrantha under combination with drought.