Comparison of photosynthetic damage from arthropod herbivory and pathogen infection in understory hardwood saplings

Arthropods and pathogens damage leaves in natural ecosystems and may reduce photosynthesis at some distance away from directly injured tissue. We quantified the indirect effects of naturally occurring biotic damage on leaf-level photosystem II operating efficiency (Φ_PSII) of 11 understory hardwood tree species using chlorophyll fluorescence and thermal imaging. Maps of fluorescence parameters and leaf temperature were stacked for each leaf and analyzed using a multivariate method adapted from the field of quantitative remote sensing. Two tree species, Quercus velutina and Cercis canadensis, grew in plots exposed to ambient and elevated atmospheric CO₂ and were infected with Phyllosticta fungus, providing a limited opportunity to examine the potential interaction of this element of global change and biotic damage on photosynthesis. Areas surrounding damage had depressed Φ_PSII and increased down-regulation of PSII, and there was no evidence of compensation in the remaining tissue. The depression of Φ_PSII caused by fungal infections and galls extended >2.5 times further from the visible damage and was ∼40% more depressed than chewing damage. Areas of depressed Φ_PSII around fungal infections on oaks growing in elevated CO₂ were more than 5 times larger than those grown in ambient conditions, suggesting that this element of global change may influence the indirect effects of biotic damage on photosynthesis. For a single Q. velutina sapling, the area of reduced Φ_PSII was equal to the total area directly damaged by insects and fungi. Thus, estimates based only on the direct effect of biotic agents may greatly underestimate their actual impact on photosynthesis.     

Potential mechanisms of low-temperature tolerance of C<Subscript>4</Subscript> photosynthesis in<Emphasis Type="Italic"> Miscanthus</Emphasis> ×<Emphasis Type="Italic"> giganteus</Emphasis>: an in vivo analysis

Miscanthus × giganteus (Greef & Deuter ex Hodkinson & Renvoize) is unique among C₄ species in its remarkable ability to maintain high photosynthetic productivity at low temperature, by contrast to the related C₄ NADP-malic enzyme-type species Zea mays L. In order to determine the in vivo physiological basis of this difference in photosynthesis, water vapor and CO₂ exchange and modulated chlorophyll fluorescence were simultaneously monitored on attached leaf segments from plants grown and measured at 25/20°C or 14/11°C (day/night temperature). Analysis of the response of photosynthesis to internal CO₂ concentration suggested that ribulose bisphosphate carboxylase/oxygenase (Rubisco) and/or pyruvate orthophosphate dikinase (PPDK) play a more important role in determining the response to low temperature than does phosphoenolpyruvate carboxylase (PEPc). For both species at both temperatures, the linear relationship between operating efficiency of whole-chain electron transport through photosystem II (_PSII) and the efficiency of CO₂ assimilation (_CO2) was unchanged and had a zero intercept, suggesting the absence of non-photosynthetic electron sinks. The major limitation at low temperature could not be solely at Rubisco or at any other point in the Calvin cycle, since this would have increased leakage of CO₂ to the mesophyll and increased _PSII/_CO2. This in vivo analysis suggested that maintenance of high photosynthetic rates in M. × giganteus at low temperature, in contrast to Z. mays, is most likely the result of different properties of Rubisco and/or PPDK, reduced susceptibility to photoinhibition, and the ability to maintain high levels of leaf absorptance during growth at low temperature.     

Responses of leaf photosynthesis,pigments and chlorophyll fluorescence within canopy position in a boreal grass (<Emphasis Type="Italic">Phalaris arundinacea</Emphasis> L.) to elevated temperature and CO<Subscript>2</Subscript> under varying water regimes

The effects of elevated growth temperature (ambient + 3.5°C) and CO₂ (700 μmol mol⁻¹) on leaf photosynthesis, pigments and chlorophyll fluorescence of a boreal perennial grass (Phalaris arundinacea L.) under different water regimes (well watered to water shortage) were investigated. Layer-specific measurements were conducted on the top (younger leaf) and low (older leaf) canopy positions of the plants after anthesis. During the early development stages, elevated temperature enhanced the maximum rate of photosynthesis (P _max) of the top layer leaves and the aboveground biomass, which resulted in earlier senescence and lower photosynthesis and biomass at the later periods. At the stage of plant maturity, the content of chlorophyll (Chl), leaf nitrogen (N_L), and light response of effective photochemical efficiency (Φ_PSII) and electron transport rate (ETR) was significantly lower under elevated temperature than ambient temperature in leaves at both layers. CO₂ enrichment enhanced the photosynthesis but led to a decline of N_L and Chl content, as well as lower fluorescence parameters of Φ_PSII and ETR in leaves at both layers. In addition, the down-regulation by CO₂ elevation was significant at the low canopy position. Regardless of climate treatment, the water shortage had a strongly negative effect on the photosynthesis, biomass growth, and fluorescence parameters, particularly in the leaves from the low canopy position. Elevated temperature exacerbated the impact of water shortage, while CO₂ enrichment slightly alleviated the drought-induced adverse effects on P _max. We suggest that the light response of Φ_PSII and ETR, being more sensitive to leaf-age classes, reflect the photosynthetic responses to climatic treatments and drought stress better than the fluorescence parameters under dark adaptation.     

Photosynthetic alterations of pea leaves infected systemically by pea enation mosaic virus: A coordinated decrease in efficiencies of CO2 assimilation and photosystem II photochemistry

We have investigated photosynthetic changes of fully expanded pea leaves infected systemically by pea enation mosaic virus (PEMV) that often attacks legumes particularly in northern temperate regions. A typical compatible virus–host interaction was monitored during 40 post-inoculation days (dpi). An initial PEMV-induced decrease in photosynthetic CO₂ assimilation was detected at 15 dpi, when the virus appeared in the measured leaves. This decrease was not induced by stomata closure and corresponded with a decrease in the efficiency of photosystem II photochemistry (Φ_PSII). Despite of a slight impairment of oxygen evolution at this stage, PSII function was not primarily responsible for the decrease in Φ_PSII. Chlorophyll fluorescence imaging revealed that Φ_PSII started to decrease from the leaf tip to the base. More pronounced symptoms of PEMV disease appeared at later stages, when a typical mosaic and enations appeared in the infected leaves and oxidative damage of cell membranes was detected. From 30 dpi, a degradation of photosynthetic pigments accelerated, stomata were closing and corresponding pronounced decline in CO₂ assimilation was observed. A concomitant photoprotective responses, i.e. an increase in non-photochemical quenching and accumulation of de-epoxidized xanthophylls, were also detected. Interestingly, alternative electron sinks in chloroplasts were not stimulated by PEMV infection, which is in contradiction to earlier reports dealing with virus-induced plant stresses. The presented results show that the PEMV-induced alterations in mature pea leaves accelerated leaf senescence during which a decrease in Φ_PSII took place in coordinated manner with an inhibition of CO₂ assimilation.     

Cold Tolerance of the Photosynthetic Apparatus: Pleiotropic Relationship between Photosynthetic Performance and Specific Leaf Area of Maize Seedlings

The objective of this study was to elucidate the genetic relationship between the specific leaf area (SLA) and the photosynthetic performance of maize (Zea mays L.) as dependent on growth temperature. Three sets of genotypes: (i) 19 S₅ inbred lines, divergently selected for high or low operating efficiency of photosystem II (Φ_PSII) at low temperature, (ii) a population of 226 F_2:3 families from the cross of ETH-DL3 × ETH-DH7, and (iii) a population of 168 F_2:4 families from the cross of Lo964 × Lo1016 were tested at low (15/13 °C day/night) or at optimal (25/22 °C day/night) temperature. The latter cross was originally developed to study QTLs for root traits. At 15/13 °C the groups of S₅ inbred lines selected for high or low Φ_PSII differed significantly for all the measured traits, while at optimal temperature the groups differed only with regard to leaf greenness (SPAD). At low temperature, the SLA of these inbred lines was negatively correlated with Φ_PSII (r = − 0.56, p < 0.05) and SPAD (r = − 0.80, p < 0.001). This negative relationship was confirmed by mapping quantitative trait loci (QTL) in the two mapping populations. A co-location of three QTLs for SLA with QTLs for photosynthesis-related traits was detected in both populations at 15/13 °C, while co-location was not detected at 25/22 °C. The co-selection of SLA and Φ_PSII in the inbred lines and the co-location of QTL for SLA, SPAD, and Φ_PSII at 15/13 °C in the QTL populations strongly supports pleiotropy. There was no evidence that selecting for high Φ_PSII at low temperature leads to a constitutively altered SLA.     

QTLs for early vigor of tropical maize

A strong photosynthetic performance and rapid leaf development, are important indicators of vigorous early growth. The aim of this study was to (1) evaluate the tropical maize (Zea mays L.) inbred lines CML444 and SC-Malawi for their photosynthetic performance at different growth stages and (2) assess quantitative trait loci (QTL) of photosynthesis-related traits in their 236 recombinant inbred lines at the heterotrophic growth stage. CML444 had a higher leaf chlorophyll (SPAD) content than SC-Malawi. Ten QTLs were found for the quantum efficiency of photosystem II (Φ_PSII; four), SPAD (three) and the specific leaf area (SLA; three). The relevance of seedling QTLs for Φ_PSII, SPAD and SLA for yield formation is emphasized by seven collocations (bins 5.01, 7.03, 8.05) with QTLs for kernel number and grain yield under field conditions. QTLs for SPAD at the V2 and at the reproductive stage did not collocate, indicating differences in the genetic control of SPAD at different growth stages. Knowing which loci affect SLA, SPAD and Φ_PSII simultaneously and which do not will help to optimize light harvest by the canopy.     

Changes in electron transport,superoxide dismutase and ascorbate peroxidase isoenzymes in chloroplasts and mitochondria of cucumber leaves as influenced by chilling

In order to clarify the relationship between chill-induced disturbance in photosynthetic, respiratory electron transport and the metabolism of reactive oxygen species (ROS), leaf gas exchange, chlorophyll fluorescence quenching, respiration, and activities of superoxide dismutase (SOD) and ascorbate peroxidase (APX) were investigated in chloroplasts and mitochondria of cucumber (Cucumis sativus) leaves subjected to a chill (8 °C) for 4 d. Chilling decreased net photosynthetic rate (P _N) and quantum efficiency of photosystem 2 (Φ_PS2), but increased the ratio of Φ_PS2 to the quantum efficiency of CO₂ fixation (ΦCO₂) and non-photochemical quenching (NPQ) in cucumber leaves. While chilling inhibited the activity of cytochrome respiration pathway, it induced an increase of alternative respiration pathway activity and the reduction level of Q-pool. Chilling also significantly increased O₂ ^• production rate, H₂O₂ content, and SOD and APX activities in chloroplasts and mitochondria. There was a more significant increase in SOD and APX activities in chloroplasts than in mitochondria with the increase of membrane-bound Fe-SOD and tAPX in chloroplasts being more significant than other isoenzymes. Taken together, chilling inhibited P _N and cytochrome respiratory pathway but enhanced the photosynthetic electron flux to O₂ and over-reduction of respiratory electron transport chain, resulting in ROS accumulation in cucumber leaves. Meanwhile, chilling resulted in an enhancement of the protective mechanisms such as thermal dissipation, alternative respiratory pathway, and ROS-scavenging mechanisms (SODs and APXs) in chloroplasts and mitochondria.     

Spectral reflectance indices and pigment functions during leaf ontogenesis in six subtropical landscape plants

Pigment combinations are regulated during leaf ontogenesis. To better understand pigment function, alterations in chlorophyll, carotenoid and anthocyanin concentrations were investigated during different leaf development stages in six subtropical landscape plants, namely Ixora chinensis Lam, Camellia japonica Linn, Eugenia oleina Wight, Mangifera indica L., Osmanthus fragrans Lowr and Saraca dives Pierre. High concentrations of anthocyanin were associated with reduced chlorophyll in juvenile leaves. As leaves developed, the photosynthetic pigments (chlorophyll and carotenoid) of all six species increased while anthocyanin concentration declined. Chlorophyll fluorescence imaging of Φ_PSII (effective quantum yield of PSII) and of NPQ (non-photochemical fluorescence quenching) and determination of electron transport rate-rapid light curve (RLC) showed that maximum ETR (leaf electron transport rate), Φ_PSII and the saturation point in RLC increased during leaf development but declined as they aged. Juvenile leaves displayed higher values of NPQ and Car/Chl ratios than leaves at other developmental stages. Leaf reflectance spectra (400–800 nm) were measured to provide an in vivo non-destructive assessment of pigments in leaves during ontogenesis. Four reflectance indices, related to pigment characters, were compared with data obtained quantitatively from biochemical analysis. The results showed that the ARI (anthocyanin reflectance index) was linearly correlated to anthocyanin concentration in juvenile leaves, while a positive correlation of Chl NDI (chlorophyll normalized difference vegetation index) to chlorophyll a concentration was species dependent. Photosynthetic reflectance index was not closely related to Car/Chl ratio, while a structural-independent pigment index was not greatly altered by leaf development or species. Accordingly, it is suggested that the high concentration of anthocyanin, higher NPQ and Car/Chl ratio in juvenile leaves are important functional responses to cope with high radiation when the photosynthetic apparatus is not fully developed. Another two leaf reflectance indices, ARI and Chl NDI, are valuable for in vivo pigment evaluation during leaf development.     

Diurnal cycle of chlorophyll fluorescence in <Emphasis Type="Italic">Phalaenopsis</Emphasis>

Chlorophyll (Chl) fluorescence of warm day/cool night temperature exposed Phalaenopsis plants was measured hourly during 48 h to study the simultaneous temperature and irradiance response of the photosynthetic physiology. The daily pattern of fluorescence kinetics showed abrupt changes of photochemical quenching (q_P), non-photochemical quenching (NPQ) and quantum yield of photosystem II electron transport (Φ_PSII) upon transition from day to night and vice versa. During the day, the course of Φ_PSII and NPQ was related to the air temperature pattern, while maximum quantum efficiency of PSII photochemistry (F_v/F_m) revealed a rather light dependent response. Information on these daily dynamics in fluorescence kinetics is important with respect to meaningful data collection and interpretation.     

Photoinhibitory damage is modulated by the rate of photosynthesis and by the photosystem II light-harvesting chlorophyll antenna size

We investigated the effect of photosynthetic electron transport and of the photosystem II (PSII) chlorophyll (Chl) antenna size on the rate of PSII photoinhibitory damage. To modulate the rate of photosynthesis and the light-harvesting capacity in the unicellular chlorophyte Dunaliella salina Teod., we varied the amount of inorganic carbon in the culture medium. Cells were grown under high irradiance either with a limiting supply of inorganic carbon, provided by an initial concentration of 25 mM NaHCO₃, or with supplemental CO₂ bubbled in the form of 3% CO₂ in air. The NaHCO₃-grown cells displayed slow rates of photosynthesis and had a small PSII light-harvesting Chl antenna size (60 Chl molecules). The half-time of PSII photodamage was 40 min. When switched to supplemental CO₂ conditions, the rate of photodamage was retarded to a t_1/2 = 70 min. Conversely, CO₂-supplemented cells displayed faster rates of photosynthesis and a larger PSII light-harvesting Chl antenna size (500 Chl molecules). They also showed a rate of photodamage with t_1/2 = 40 min. When depleted of CO₂, the rate of photodamage was accelerated (t_1/2  = 20 min). These results indicate that the in-vivo susceptibility to photodamage is modulated by the rate of forward electron transport through PSII. Moreover, a large Chl antenna size enhances the rate of light absorption and photodamage and, therefore, counters the mitigating effect of forward electron transport. We propose that under steady-state photosynthesis, the rate of light absorption (determined by incident light intensity and PS Chl antenna size) and the rate of forward electron transport (determined by CO₂ availability) modulate the oxidation/reduction state of the primary PSII acceptor Q_A, which in turn defines the low/high probability for photodamage in the PSII reaction center. Received: 14 August 1997 / Accepted: 26 September 1997     

The mechanisms contributing to photosynthetic control of electron transport by carbon assimilation in leaves

Photosynthetic control describes the processes that serve to modify chloroplast membrane reactions in order to co-ordinate the synthesis of ATP and NADPH with the rate at which these metabolites can be used in carbon metabolism. At low irradiance, optimisation of the use of excitation energy is required, while at high irradiance photosynthetic control serves to dissipate excess excitation energy when the potential rate of ATP and NADPH synthesis exceed demand. The balance between pH, ATP synthesis and redox state adjusts supply to demand such that the [ATP]/[ADP] and [NADPH]/[NADP⁺] ratios are remarkably constant in steady-state conditions and modulation of electron transport occurs without extreme fluctuations in these pools.Abbreviations FBPase Fructose-1,6-bisphosphatase - PS I Photosystem I - PS II Photosystem II - Pi inorganic phosphate - PGA glycerate 3-phosphate - PQ plastoquinone - Q_A the bound quinone electron acceptor of PS II - q_P Photochemical quenching of chlorophyll fluorescence associated with the oxidation of Q_A - q_N non-photochemical quenching of chlorophyll fluorescence - q_E non-photochemical quenching associated with the high energy state of the membrane - RuBP ribulose-1,5-bisphosphate - TP triose phosphate - intrinsic quantum yield of PS II - quantum yield of electron transport - quantum yield of CO₂ assimilation     

Regulation of photosynthetic electron-transport in Phaseolus vulgaris L., as determined by room-temperature chlorophyll a fluorescence

The regulation of photosystem II (PSII) by light-, CO₂-, and O₂-dependent changes in the capacity for carbon metabolism was studied. Estimates of the rate of electron transport through PSII were made from gas-exchange data and from measurements of chlorophyll fluorescence. At subsaturating photon-flux density (PFD), the rate of electron transport was independent of O₂ and CO₂. Feedback on electron transport was observed under two conditions. At saturating PFD and low partial pressure of CO₂, p(CO₂), the rate of electron transport increased with p(CO₂). However, at high p(CO₂), switching from normal to low p(O₂) did not affect the net rate of photosynthetic CO₂ assimilation but the rate of electron-transport decreased by an amount related to the change in the rate of photorespiration. We interpret these effects as 1) regulation of ribulose-1,5-bisphosphatecarboxylase (RuBPCase, EC 4.1.1.39) activity to match the rate of electron transport at limiting PFD, 2) regulation of electron-transport rate to match the rate of RuBPCase at low p(CO₂), and 3) regulation of the electron-transport rate to match the capacity for starch and sucrose synthesis at high p(CO₂) and PFD. These studies provide evidence that PSII is regulated so that the capacity for electron transport is matched to the capacity for other processes required by photosynthesis, such as ribulose-bisphosphate carboxylation and starch and sucrose synthesis. We show that at least two mechanisms contribute to the regulation of PSII activity and that the relative engagement of these mechanisms varies with time following a step change in the capacity for ribulose-bisphosphate carboxylation and starch and sucrose synthesis. Finally, we take advantage of the relatively slow activation of deactivated RuBPCase in vivo to show that the activation level of this enzyme can limit the rate of electron transport as evidenced by increased feedback on PSII following a step change in p(CO₂). As RuBPCase as activated, the feedback on PSII declined.Abbreviations and symbols J_C electron-transport rate calculated from CO₂-assimilation measurements - J_F electron-transport rate calculated from fluorescence parameters - PFD photon-flux density - q_E energy-dependent quenching - PSII photosystem II - q_Q Q-dependent quenching - QY quantum yield - RuBPCase ribulose-1,5-bisphosphate carboxylase (EC 4.1.1.39) C.I.W. publication No. 1015     

Caragana korshinskii seedlings maintain positive photosynthesis during short-term,severe drought stress

Seedling performance may determine plant distribution, especially in water-limited environments. Plants of Caragana korshinskii commonly grow in arid and semiarid areas in northwestern China, and endure water shortage in various ways, but little is known about their performance when water shortage occurs at early growth stages. The water relations, photosynthetic activity, chlorophyll (Chl) content and proline accumulation were determined in 1-year-old seedlings growing in a 1:1 mixture of Loess soil and Perlite and subjected to (1) a water deficit for 20 days and (2) kept adequately watered throughout. The water deficit induced low (−6.1 MPa) predawn leaf water potentials (LWP), but did not induce any leaf abscission. Stomatal conductance (g _s), leaf transpiration rate (E), and net photosynthetic rate (P _N) decreased immediately following the imposition of the water deficit, while the maximal photochemical efficiency of photosystem II (PSII) (F_v/F_m) and the effective quantum yield of PSII (Φ_PSII) decreased 15 days later. An early and rapid decrease in g _s, reduced E, increased Chl (a+b) loss, increased the apparent rate of photochemical transport of electrons through PSII (ETR)/P _N, as well as a gradual increase in non-photochemical quenching of fluorescence (NPQ) and proline may have contributed to preventing Φ_PSII from photodamage. C. korshinskii seedlings used a stress-tolerance strategy, with leaf maintenance providing a clear selective advantage, considering the occasional rainfall events during the growing season.     

The response of electron transport mediated by active NADPH dehydrogenase complexes to heat stress in the cyanobacterium Synechocystis 6803

The electron-transport machinery in photosynthetic membranes is known to be very sensitive to heat. In this study, the rate of electron transport (ETR) driven by photosystem I (PSI) and photosystem II (PSII) during heat stress in the wild-type Synechocystis sp. strain PCC 6803 (WT) and its ndh gene inactiva-tion mutants ΔndhB (M55) and ΔndhD1/ndhD2 (D1/D2) was simultaneously assessed by using the novel Dual-PAM-100 measuring system. The rate of electron transport driven by the photosystems (ETR_PSs) in the WT, M55, and D1/D2 cells incubated at 30℃ and at 55℃ for 10 min was compared. Incubation at 55℃ for 10 min significantly inhibited PSII-driven ETR (ETR^_PSII) in the WT, M55 and D1/D2 cells, and the ex-tent of inhibition in both the M55 and D1/D2 cells was greater than that in the WT cells. Further, PSI-driven ETR (ETR_PSI) was stimulated in both the WT and D1/D2 cells, and this rate was increased to a greater extent in the D1/D2 than in the WT cells. However, ETR_PSI was considerably inhibited in the M55 cells. Analysis of the effect of heat stress on ETR_PSs with regard to the alterations in the 2 active NDH-1 complexes in the WT, M55, and D1/D2 cells indicated that the active NDH-1 supercomplex and medi-umcomplex are essential for alleviating the heat-induced inhibition of ETRPSII and for accelerating the heat-induced stimulation of ETR_PSI, respectively. Further, it is believed that these effects are most likely brought about by the electron transport mediated by each of these 2 active NDH-1 complexes.     

ATP and NADPH as the driving force of carbon reduction in leaves in relation to thylakoid energization by light

Carbon assimilation of spinach (Spinacia oleracea L.) leaves was measured in the presence of 2000l· l^–1CO₂ and 2% O₂ in the gas phase to suppress photorespiratory reactions and to reduce stomatal diffusion resistance. Simultaneously, membrane parameters such as modulated chlorophyll fluorescence, oxidation of P₇₀₀ in the reaction centre of photosystem I, and apparent changes in absorbance at 535 nm were recorded. After light-regulated enzymes were activated at a high irradiance, illumination was changed. About 3 min later (to maintain the previous activation state of enzymes), leaves were shock-frozen and freeze-dried. Chloroplasts were isolated nonaqueously and analysed for ATP, ADP, inorganic phosphate, NADPH and NADP. Observations made under the chosen conditions differed in some important aspects from those commonly observed when leaves are illuminated in air. (i) Not only assimilation, but also the phosphorylation potential [ATP]/([ADP]·[Pi]) increased hyperbolically with irradiance towards saturation. In contrast, the ratio of NADPH to NADP did not change much as irradiances increased from low to high photon flux densities. When ATP, the phosphorylation potential and the assimilatory force, F_A (the product of phosphorylation potential and NADPH/NADP ratio), were plotted against assimilation, ATP increased relatively less than assimilation, whereas the phosphorylation potential increased somewhat more steeply than assimilation did. A linear relationship existed between assimilation and F_A at lower irradiances. The assimilatory force F_A increased more than assimilation did when irradiances were very high. Differences from previous observations, where F_A was under some conditions higher at low than at high rates of carbon assimilation, are explained by differences in flux resistances caused not only by stomatal diffusion resistance but also by differences in the activity of light-regulated enzymes, (ii) The relationship between P₇₀₀ oxidation and a fast absorption change with a maximum close to 520 nm on one hand and carbon assimilation on the other hand was largely linear under the specific conditions of the experiments. A similar linear relationship existed also between the quantum efficiency of electron flow through photosystem II and the quantum efficiency of photosystem I electron transport. (iii) Whereas the increase in non-photochemical fluorescence quenching, q_N, was similar to the increase in assimilation, the relationship between light scattering and assimilation was distinctly sigmoidal. Light scattering appeared to be a better indicator of control of photosystem II activity under excessive irradiation than q_N. (iv) The results are discussed in relation to the relative significance of chloroplast levels of ATP and NADPH and of the assimilatory force F_A in driving carbon assimilation. From the observations, the proton/electron (H⁺/e^–) ratio of linear electron transport is suggested to be 3 and the H⁺/ATP ratio to be 4 in leaves. An H⁺/e^– ratio of 3 implies the existence of an obligatory Q-cycle in leaves.Abbreviations F_A assimilatory force - F_o fluorescence after long dark adaptation - F_m maximum fluorescence level - F_s steady-state fluorescence - PGA 3-phosphoglycerate - PFD photon flux density - P₇₀₀ (P₇₀₀+) electron-donor pigment in the reaction center of PSI (its oxidized form) - Q_A primary quinone acceptor of PSII - q_P photochemical quenching - q_N non-photochemical quenching - _PSII relative quantum efficiency of energy conversation at the level of photosystem II - _PSI relative quantum efficiency of photosystem II This research was supported by the Sonderforschungsbereich 251 of the University of Würzburg and the Stiftung Volkswagenwerk. U.G. is a member of the Graduate College of the Julius-von-Sachs Institut für Biowissenschaften, University of Würzburg, being on leave from Tartu University, Tartu, Estonia. The authors are grateful to Prof. A. Laisk, Chair of Plant Physiology, Tartu University, for stimulating discussions.     

The Response of Antioxidant Enzymes in Cellular Organelles in Cucumber (Cucumis sativus L.) Leaves to Methyl Viologen-induced Photo-oxidative Stress

In order to clarify the response of antioxidant systems in various cellular organelles to photo-oxidative stress, the activities of superoxide dismutase (SOD) and enzymes of the ascorbate–glutathione (AsA-GSH) cycle were investigated in chloroplasts, mitochondria and cytosol of cucumber leaves subjected to methyl viologen (MV) treatment. Photo-oxidation by MV resulted in significant reductions in net photosynthetic rate (Pn) and increases in the ratio of the quantum efficiency of photosystem II (PSII), Φ_PSII to that of the quantum efficiency of CO₂ fixation (ΦCO₂), followed by increased activities of SOD, and a general increase of AsA-GSH cycle enzymes in chloroplasts, mitochondria and cytosol. These increases were however, most significant in chloroplasts. There were also significant increases in dehydroascorbate (DHA), reduced glutathione (GSH), and oxidized glutathione (GSSG) except that the content of ascorbate (AsA) in chloroplasts and cytosol was slightly decreased and little effected, respectively. However, GSSG in mitochondria and GSH in cytosol were little influenced by the MV treatment. The activity of ascorbate oxidase (AO) in these organelles was independent of the MV treatment while the activity of l-galactono-1,4- lactone dehydrogenase (GLDH) in mitochondria was slightly inhibited by MV treatment. These results indicate that disturbance of electron transport in chloroplasts by MV influenced the metabolism of whole cell by a crosstalk signaling system and that the AsA-GSH cycle played a primary role in sustaining the levels of AsA.     

Effects of groundwater depth variation on photosynthesis and photoprotection of <Emphasis Type="Italic">Elaeagnus angustifolia</Emphasis> L.

Physiological and photosynthetic responses were investigated at three different depths of groundwater (DGW: 1.4, 2.4, and 3.4 m) in Elaeagnus angustifolia L., a locally adapted tree to the arid region in northwest China. Predawn leaf water potential and chlorophyll content declined gradually with the increasing DGW, whereas there was little effect on predawn variable-to-maximum chlorophyll fluorescence ratio F _v/F _m and leaf carotenoid compositions (xanthophyll cycle pool, neoxanthin, lutein, and β-carotene). Net photosynthetic rate (P _n), quantum yield of PSII electron transport (Φ_PSII), stomatal conductance (Gs), and intercellular CO₂ concentration (Ci) declined obviously; however, P _n decreased more than Φ_PSII at deeper DGW. The photoinhibition of PSII at all three DGW occurred at midday in summer and increased as DGW increased. The ΔpH-dependent thermal dissipation and the level of de-epoxidation of the xanthophyll cycle at all three DGW reached their maxima at midday with the increase of light intensity. However, the fraction of functional PSII and light intensity at deeper DGW (2.4, 3.4 m) showed a negative correlation. This correlation suggested that most of violaxanthin was converted into zeaxanthin at midday, and the reversible inactivation of partial PSII reaction centers took place at deeper DGW. These results together suggest that both the xanthophyll cycle-dependent thermal dissipation and the reversible inactivation of partial PSII might have played important roles in avoiding the excess light-induced energy damage in leaves of this tree species at deeper DGW.     

Stomatal development and associated photosynthetic performance of capsicum in response to differential light availabilities

The mechanisms of capsicum growth in response to differential light availabilities are still not well elucidated. Hereby, we analyzed differential light availabilities on the relationship between stomatal characters and leaf growth, as well as photosynthetic performance. We used either 450–500 μmol m⁻² s⁻¹ as high light (HL) or 80–100 μmol m⁻² s⁻¹ as low light (LL) as treatments for two different cultivars. Our results showed that the stomatal density (SD) and stomatal index (SI) increased along with the leaf area expansion until the peak of the correlation curve, and then decreased. SD and SI were lower under the LL condition after three days of leaf expansion. For both cultivars, downregulation of photosynthesis and electron transport components was observed in LL-grown plants as indicated by lower light- and CO₂-saturated photosynthetic rate (P _max and RuBP_max), quantum efficiency of photosystem II (PSII) photochemistry (Φ_PSII), electron transport rate (ETR) and photochemical quenching of fluorescence (q_p). The observed inhibition of the photosynthesis could be explained by the decrease of SD, SI, Rubisco content and by the changes of the chloroplast. The low light resulted in lower total biomass, root/shoot ratio, and the thickness of the leaf decreased. However, the specific leaf area (SLA) and the content of leaf pigments were higher in LL-treatment. Variations in the photosynthetic characteristics of capsicum grown under different light conditions reflected the physiological adaptations to the changing light environments.     

Photorespiration is Essential for the Protection of the Photosynthetic Apparatus of C3 Plants Against Photoinactivation Under Sunlight

CO₂ assimilation, transpiration and modulated chlorophyll fluorescence of leaves of Chenopodium bonus-henricus (L.) were measured in the laboratory and, at a high altitude location, in the field. Direct calibration of chlorophyll fluorescence parameters against carbon assimilation in the presence of 1 or 0.5% oxygen (plus CO₂) proved necessary to calculate electron transport under photorespiratory conditions in individual experiments. Even when stomata were open in the field, total electron transport was two to three times higher in sunlight than indicated by net carbon gain. It decreased when stomata were blocked by submerging leaves under water or by forcing them to close in air by cutting the petiole. Even under these conditions, electron transport behind closed stomata approached 10 nmol electrons m^?2 leaf area s^?1 at temperatures between 25 and 30 °C. No photoinactivation of photosystem II was indicated by fluorescence analysis after a day's exposure to full sunlight. Only when leaves were submerged in ice was appreciable photoinactivation noticeable after 4 h exposure to sunlight. Even then almost full recovery occurred overnight. Electron transport behind blocked stomata was much decreased when leaves were darkened for 70 min (in order to deactivate light-regulated enzymes of the Calvin cycle) before exposure to full sunlight. Brief exposure of leaves to HCN (to inhibit photoassimilation and photorespiration) also decreased electron transport drastically compared to electron transport in unpoisoned leaves with blocked stomata. Non-photochemical fluorescence quenching and reduction of Q_A, the primary electron acceptor of photosystem II was increased by HCN-poisoning. Very similar observations were made when glyceraldehyde was used instead of HCN to inhibit photosynthesis and photorespiration. In HCN-poisoned leaves, residual electron transport increased linearly with temperature and showed early light saturation revealing characteristics of the Mehler reaction. During short exposure of these leaves to photon flux densities equivalent to 25% of sunlight, no or only little photoinactivation of photosystem II was observed. However, prolonged exposure to sunlight caused inactivation even though non-photochemical quenching of chlorophyll fluorescence was extensive. Simultaneously, oxidation of cellular ascorbate and glutathione increased. Inactivation of photosystem II was reversible in dim light and in the dark only after short times of exposure to sunlight. Glyceraldehyde was very similar to HCN in increasing the sensitivity of photosystem II in leaves to sunlight. We conclude from the observations that the electron transport permitted by the interplay of photoassimilatory and photorespiratory electron transport is essential to prevent the photoinactivation of photosynthetic electron transport. The Mehler and Asada reactions, which give rise to strong nonphotochemical fluorescence quenching, are insufficient to protect the chloroplast electron transport chain against photoinactivation.