Changes in Chlorophyll Fluorescence in Maize Plants with Imposed Rapid Dehydration at Different Leaf Ages

A comparison of the effects of a rapidly imposed water deficit with different leaf ages on chlorophyll a fluorescence and gas exchange was performed in maize (Zea mays L.) plants. The relationships between photosynthesis and leaf relative turgidity (RT) and ion leakage were further investigated. Leaf dehydration substantially decreased net photosynthetic rate (A) and stomatal conductance (G _s), particularly for older leaves. With dehydration time, F _v /F _m maintained a relatively stable level for youngest leaves but significantly decreased for the older leaves. The electron transport rate (ETR) sharply decreased with intensifying dehydration and remained at lower levels during continuous dehydration. The photochemical quenching of variable chlorophyll fluorescence (q _P) gradually decreased with dehydration intensity for the older leaves but increased for the youngest leaves, whereas dehydration did not affect the nonphotochemical chlorophyll fluorescence quenching (NPQ) for the youngest leaves but remarkably decreased it for the older leaves. The leaf RT was significantly and positively correlated with its F _v /F _m, ETR, and q _P, and the leaf ion leakage was significantly and negatively correlated with F _v /F _m and NPQ. Our results suggest that the photosynthetic systems of young and old leaves decline at different rates when exposed to rapid dehydration.     

'Photoinhibition' of Heliconia under natural tropical conditions: the importance of leaf orientation for light interception and leaf temperature

The influence of irradiance on photosynthesis under natural conditions was studied in aseasonal Singapore using three Heliconia taxa: H. rostrata, H. psittacorum × H. spathocircinata cv. Golden Torch and H. psittacorum cv. Tay. When grown under full sunlight, all three heliconias exhibited reduced phatosynthetic capacities and lowered chlorophyll content per leaf area as compared with those grown under intermediate and deep shade. A marked decrease in the chlorophyll fluorescence F_v/Fm ratio and an increase in photochemical quenching (1- q_p) and non-photochemical quenching (q_N) were observed in upper leaves of plants grown under full sunlight. Increases in q_N suggest that ‘photoinhibition’ (decreases in F_v/F_m) in Heliconia grown under natural tropical conditions are probably due to photoprotective energy dissipation processes. The quantum yield, the maximum photosynthetic rate, F_v/F_m and the chlorophyll content of upper leaves were lower than those of lower leaves on the same plants grown under full sunlight. Similarly, lower values were obtained for the tip (sun) portion than for the base (shaded) portion of the leaves. The changes in F_v/F_m and in the levels of (1 –q_p) in leaves grown under intermediate and deep shade were negligible in plants during the course of day. However, there was a steep decrease in F_v/F_m and an increase in the levels of (1 –q_p), along with an increase in incident light in the sun leaves. The lowest F_v/F_m and the highest level of (1 –q_p) indicated minimum PSII efficiency at midday in full sun. These results indicate that, in Heliconia, the top leaves (particularly leaf tips) experienced sustained decreases in PSII efficiency upon exposure to full sunlight. Although all three taxa exhibited sustained decreases in photosynthetic capacity in full sunlight, the sun leaves of ‘Tay’ showed higher photosynthetic capacity than those of the other two taxa. This could be due, at least in part, to the vertical leaf angle and smaller lamina area. When the upright leaves of ‘Tay’ were constrained to a horizontal angle, they exhibited lower PSII efficiency (F_vIF_m ratio), while horizontal leaves of ‘Rostrata’ and ‘Golden Torch’ inclined lo near-vertical angles showed increased efficiency. Thus, an increase in leaf angle helps to achieve a reduction in the sustained decrease in PSII efficiency by decreasing the levels of incident sunlight and subsequently the leaf temperature.     

Short-term responses of photosynthetic membrane lipids and photochemical efficiency in plants of <Emphasis Type="Italic">Phaseolus vulgaris</Emphasis> and <Emphasis Type="Italic">Vigna unguiculata</Emphasis> submitted to high irradiance

Primary leaves of young plants of common bean (Phaseolus vulgaris cv. Carioca and Negro Huasteco) and cowpea (Vigna unguiculata Walp cv. Epace 10) were exposed to high irradiance (HI) of 2 000 μmol m⁻² s⁻¹ for 10, 20, and 30 min. The initial fluorescence (F₀) was nearly constant in response to HI in each genotype except for Carioca. A distinct reduction of maximum fluorescence (F_m) was clearly observed in stressed genotypes of beans after 20 min followed by a slight recovery for the longer stress times. In common bean, the maximum quantum yield (F_v/F_m) was reduced slowly from 10 to 30 min of HI. In cowpea, only a slight reduction of F_v/F_m was observed at 20 min followed by recovery to normal values at 30 min. HI resulted in changes in the photochemical (q_P) and non-photochemical (q_N) quenching in both species, but to a different extent. In cowpea plants, more efficiency in the use of the absorbed energy under photoinhibitory conditions was related to increase in q_P and decrease in q_N. In addition, lipid peroxidation changed significantly in common bean genotypes with an evident increase after 20 min of HI. Hence the photosynthetic apparatus of cowpea was more tolerant to HI than that of common bean and the integrity of cowpea cell membranes was apparently maintained under HI.     

Photoinhibitory responses of the epiphytic bromeliad Guzmania monostachia during the dry season in Trinidad maintain photochemical integrity under adverse conditions

The C₃–CAM epiphytic bromeliad Guzmania monostachia var. monostachia may be exposed to high incident photosynthetically active radiation (PAR) during the dry season in Trinidad, and resultant variations in photochemical efficiency have been investigated for ‘exposed’ (receiving ～100% incident PAR), ‘semi-exposed’ (～60% PAR) and shaded populations under natural conditions. The more succulent leaves of the plants growing fully exposed within the canopy had higher overall CAM activity (measured as ΔH⁺, the dawn-dusk titratable acidity), a smaller proportion of chlorenchyma and lower total chlorophyll content. There was a gradation of morphological and physiological characteristics between these and shaded leaves. Diurnal time-courses of photosynthetic light responses (as O₂ evolution) showed marked variations in apparent quantum yield (AQY) and light-saturated rates for both exposed and semi-exposed populations, dependent on incident PAR during the day. Similar measurements of photosystem II fluorescence characteristics showed that F_v/F_m declined from 0·70 to 0·42 at midday for exposed plants (on a day when total incident PAR was 44 mol photon m^?2), indicating non-photochemical quenching (qNP) of photosynthesis. However, in contrast to AQY determinations, F_v/F_m recovered during the afternoon. The decrease in F_v/F_m was reduced from 0·72 to 0·64 under 24 mol photon m^?2 d^?1. The long–term recovery of photo-synthetic efficiency was assessed for exposed plants placed under three shading regimes (60, 30 and 3% of incident PAR) over a 17-d period. During this time, total chlorophyll content increased from 228 to 515 and 585 μg g^?1 fresh weight (for 3 and 30%, respectively) and chlorophyll a:b declined. While AQY recovery was much longer under the lowest PAR (17d), under 30% PAR both AQY and F_v/F_m had recovered after 2d shading. The differences between timing of recovery for F_v/F_m during diurnal time courses and in the long term suggest that, while quenching associated with PSII recovers rapidly, enzyme activation and/or protein synthesis of other photosynthetic components may be limiting under low PAR. However, it is suggested that the occurrence of qNP on a daily basis may preclude long-term photoinhibitory damage under natural conditions during the dry season.     

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.     

Enhanced thermotolerance of photosystem II in salt-adapted plants of the halophyte<Emphasis Type="Italic"> Artemisia anethifolia</Emphasis>

Thermotolerance of photosystem II (PSII) in leaves of salt-adapted Artemisia anethifolia L. plants (100–400 mM NaCl) was evaluated after exposure to heat stress (30–45°C) for 30 min. After exposure to 30°C, salt adaptation had no effects on the maximal efficiency of PSII photochemistry (F_v/F_m), the efficiency of excitation capture by open PSII centers (F_v/F_m), or the actual PSII efficiency (_PSII). After pretreatment at 40°C, there was a striking difference in the responses of F_v/F_m, F_v/F_m and _PSII to heat stress in non-salt-adapted and salt-adapted leaves. Leaves from salt-adapted plants maintained significantly higher values of F_v/F_m, F_v/F_m and _PSII than those from non-salt-adapted leaves. The differences in F_v/F_m, F_v/F_m and _PSII between non-salt-adapted and salt-adapted plants persisted for at least 12 h following heat stress. These results clearly show that thermotolerance of PSII was enhanced in salt-adapted plants. This enhanced thermotolerance was associated with an improvement in thermotolerance of the PSII reaction centers, the oxygen-evolving complexes and the light-harvesting complex. In addition, we observed that after exposure to 42.5°C for 30 min, non-salt-adapted plants showed a significant decrease in CO₂ assimilation rate while in salt-adapted plants CO₂ assimilation rate was either maintained or even increased to some extent. Given that photosynthesis is considered to be the physiological process most sensitive to high-temperature damage and that PSII appears to be the most heat-sensitive part of the photosynthetic apparatus, enhanced thermotolerance of PSII may be of significance for A. anethifolia, a halophyte plant, which grows in the high-salinity regions in the north of China, where the air temperature in the summer is often as high as 45°C.     

Effect of daily photon receipt on the susceptibility of dwarf bean (Phaseolus vulgaris L.) leaves to photoinhibition of photosynthesis

Bean (Phaseolus vulgaris L.) plants were grown at two light periods of 8 and 13 h with a similar photon flux density (PFD) giving a daily photon receipt (DPR) of 17.9 and 38.2 mol · m–2, respectively. Shoot growth and leaf area development were followed at regular intervals and diurnal whole-plant photosynthesis measured. Single mature trifoliate leaves were exposed to photoinhibitory treatments at PFDs of 800 and 1400 mol · m–2 · s–1 and at temperatures of 12 and 20°C. Chlorophyll fluorescence and photon yields were measured at regular intervals throughout each treatment. Plants grown in 13 h had significantly greater leaf areas than those grown in 8 h. There were no differences in maximum rates of photosynthesis, photon yields and only minor but significant differences in F_v/F_m for plants in the two treatments, showing photosynthetic characteristics were dependent on PFD but not DPR. A significant decline in photosynthesis and F_v/F_m occurred over the 13-h but little change in photosynthesis for plants in the 8 h, indicating some feedback inhibition of photosynthesis was occurring. Plants grown in 8 h were consistently more susceptible to photoinhibition of photosynthesis at all treatments than 13-h plants. Nevertheless, photoinhibition was exacerbated by increases in PFD, and by decreases in temperature for leaves from both treatments. However, for plants from the 8-h day, exposing leaves to 12°C and 1400 mol · m–2 · s–1 caused photo-oxidation and severe bleaching but no visible damage on leaves from 13-h-grown plants. Closure of the photosystem II reaction-centre pool was partially correlated with increasing extents of photoinhibition but the relationship was similar for plants from both treatments. There remains no clear explanation for their wide differences in susceptibility to photoinhibition.Abbreviations and Symbols DPR daily photon receipt - F₀ and F_m initial and maximal fluorescence - F_v/F_m fluorescence ratio in dark-treated leaves - F/F_m intrinsic efficiency of PSII during illumination - PFD photon flux density - _i photon yield (incident basis) - _psi quantum yield of PSII electron transport - P_max maximum rate of photosynthesis - q_N non-photochemical quenching coefficient - q_P photochemical quenching coefficient Many thanks to my colleague William Laing who spent a considerable effort in developing the programme to run the photosynthesis apparatus. I am also indebted to one reviewer with whom I corresponded to resolve some issues in the paper. This project was funded by the New Zealand Foundation for Research, Science and Technology.     

PSII photochemistry, thermal energy dissipation, and the xanthophyll cycle in Kalanchoë daigremontiana exposed to a combination of water stress and high light

Kalanchoë daigremontiana, a CAM plant grown in a greenhouse, was subjected to severe water stress. The changes in photosystem II (PSII) photochemistry were investigated in water‐stressed leaves. To separate water stress effects from photoinhibition, water stress was imposed at low irradiance (daily peak PFD 150 μmol m^?2 s^?1). There were no significant changes in the maximal efficiency of PSII photochemistry (F_v/F_m), the traditional fluorescence induction kinetics (OIP) and the polyphasic fluorescence induction kinetics (OJIP), suggesting that water stress had no direct effects on the primary PSII photochemistry in dark‐adapted leaves. However, PSII photochemistry in light‐adapted leaves was modified in water‐stressed plants. This was shown by the decrease in the actual PSII efficiency (Φ_PSII), the efficiency of excitation energy capture by open PSII centres (F_v′/F_m′), and photochemical quenching (q_P), as well as a significant increase in non‐photochemical quenching (NPQ) in particular at high PFDs. In addition, photoinhibition and the xanthophyll cycle were investigated in water‐stressed leaves when exposed to 50% full sunlight and full sunlight. At midday, water stress induced a substantial decrease in F_v/F_m which was reversible. Such a decrease was greater at higher irradiance. Similar results were observed in Φ_PSII, q_P, and F_v′/F_m′. On the other hand, water stress induced a significant increase in NPQ and the level of zeaxanthin via the de‐epoxidation of violaxanthin and their increases were greater at higher irradiance. The results suggest that water stress led to increased susceptibility to photoinhibition which was attributed to a photoprotective process but not to a photodamage process. Such a photoprotection was associated with the enhanced formation of zeaxanthin via de‐epoxidation of violaxanthin. The results also suggest that thermal dissipation of excess energy associated with the xanthophyll cycle may be an important adaptive mechanism to help protect the photosynthetic apparatus from photoinhibitory damage for CAM plants normally growing in arid and semi‐arid areas where they are subjected to a combination of water stress and high light.     

Protective Effects of Exogenous Antioxidants and Phenolic Compounds on Photosynthesis of Wheat Leaves under High Irradiance and Oxidative Stress

Infiltration of methyl viologen (MV, source of O₂ ^–) and Na-diethyldithiocarbamate (DDC, inhibitor of SOD) into wheat leaves resulted in the accumulation of active oxygen species and photo-oxidative damage to photosynthetic apparatus under both moderate and high irradiance. Exogenous antioxidants, ascorbate (ASA) and mannitol, scavenged active oxygen efficiently, protected the photosynthetic system from MV and DDC induced oxidative damage, and maintained high F_v/F_m [maximal photochemical efficiency of photosystem 2 (PS2) while all PS2 reaction centres are open], F_m/F₀ (another expression for the maximal photochemical efficiency of PS2), _PS2 (actual quantum yield of PS2 under actinic irradiation), q_P (photochemical quenching coefficient), P _N (net photosynthetic rate), and lowered q_NP (non-photochemical quenching coefficient) of the leaves kept under high irradiance and oxidative stress. Phenolic compounds used in these experiments, catechol (Cat), resorcinol (Res), and tannic acid (Tan), had similar anti-oxidative activity and protective effect on photosynthetic apparatus as ASA and mannitol. The anti-oxidative activity and the protective effect of phenolic compounds increased with increase in their concentration from 100 to 300 g m^–3. The number and the position of hydroxyl group in phenolic molecules seemed to influence their antioxidative activity.     

Dithiothreitol,an inhibitor of violaxanthin de-epoxidation,increases the susceptibility of leaves ofNerium oleander L. to photoinhibition of photosynthesis

Leaves ofNerium oleander L. plants, which had been previously kept in a shaded glasshouse for at least two months, were fed 1 mM dithiothreitol (DTT) through their petioles, either for 12h in darkness (overnight) or for 2h in low light (28 μmol photons·m⁻²·s⁻¹), in each case followed by a 3-h exposure to high light (1260 μmol photons·m⁻²·s⁻¹). During exposure to high light, violaxanthin became converted to zeaxanthin in control leaves, to which water had been fed, whereas zeaxanthin did not accumulate in leaves treated with DTT. Total carbon gain was not reduced by DTT during the photoinhibitory treatment. Exposure to high light led to a decrease in the photochemical efficiency of photosystem II, measured as the ratio of variable over maximum fluorescence emission,F _v/F _M, at both 298 K and 77K. The decrease was much more pronounced in the presence of DTT, mainly owing to a sustained increase in the instantaneous fluorescence,F _o. By contrast, in the control leaves,F _o determined immediately after the high-light treatment showed a transient decrease below theF _o value obtained before the onset of the photoinhibitory treatment (i.e. after 12 h dark adaptation), followed by a rapid return (within seconds) to this original level ofF _o during the following recovery period in darkness. Incubation of leaves with DTT led to large, sustained decreases in the photon-use efficiency of photosynthetic O₂ evolution by bright light, whilst the capacity of photosynthetic O₂ evolution at light and CO₂ saturation was less affected. In the control leaves, only small reductions in the photon yield and in the photosynthetic capacity were observed. These findings are consistent with previous suggestions that zeaxanthin, formed in the xanthophyll cycle by de-epoxidation of violaxanthin, is involved in protecting the photosynthetic apparatus against the adverse effects of excessive light.     

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.     

In situ monitoring of chlorophyll fluorescence to assess the synergistic effect of low temperature and high irradiance stresses inSpirulina cultures grown outdoors in photobioreactors

A chlorophyll fluorescence technique was applied to anin situ study on the effects of low temperature and high light stresses onSpirulina cultures grown outdoors in controlled tubular photobioreactors at high (1.1 g L^–1) and low (0.44 g L^–1) biomass concentrations. Diurnal changes in PSII photochemistry (F _v/F _m) after 15 min of darkness, or in the light (dF/F _m), and non-photochemical (qN) quenching were measured using a portable, pulse-amplitude-modulated fluorometer. The depression of theF _v/F _m ratio ofSpirulina cultures grown outdoors at 25°C (i.e. 10°C below optimum for growth) and 0.44 g L^–1, reached 30% at the middle of the day. At the same time of the day thedF/F _m ratio showed a reduction of up to 52%. The depression of bothF _v/F _m anddF/F _m was lower in the cultures grown at 1.1 g L^–1. Photoinhibition reduced the daily productivity of the culture grown at 0.44 g L^–1 and 25°C by 33% with respect to that grown at 35°C. Changes in the growth yields of the cultures grown under different temperatures and growth rates correlate well with analogous changes in photon yield (dF/F _m). Simple measurements of photochemical yield (F _v/F _m) can be used to test the physiological status ofSpirulina cultures. The results indicate that the saturating pulse fluorescence technique, when usedin situ, is a powerful tool for assessment of the photosynthetic characteristics of outdoor cultures ofSpirulina.     

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

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

Is the bark of shining gum (Eucalyptus nitens) a sun or a shade leaf?

Photoinhibition and pigment composition of green stem tissues of field-grown adult Eucalyptus nitens were measured on clear spring days with low morning temperatures—conditions that cause photoinhibition in leaves of many plant species. The sun-exposed (north-facing) bark contained less chlorophyll a+b (531 vs 748 mol m^–2), neoxanthin (29 vs 41), and -carotene (54 vs 73), more xanthophyll cycle pigments per unit surface area and per unit chlorophyll (71 vs 53 mol m^–2 and 141 vs 66 mmol mol^–1 chlorophyll), and less lutein per unit chlorophyll (239 vs 190) than the shaded (southern) side. Maximum electron flow rates were 60 mol m^–2 s^–1 on the sun-exposed side, and about 10 mol m^–2 s^–1 on the shaded side. F_v/F_m was always lower than 0.8 on the sun-exposed side and the de-epoxidation state (DEPS) of the xanthophyll cycle was dominated by zeaxanthin in midday samples. F_v/F_m increased quickly after darkening, but DEPS recovered more slowly to 40% overnight. This suggested that rapidly reversible pH-dependent quenching was responsible for the bulk of changes in PS II efficiency. F_v/F_m remained below 0.8 overnight, which may well be indicative of photo-damage to PSII. In contrast, DEPS of the shaded side was lower, and F_v/F_m was higher, than for the sun-exposed side. We conclude that E. nitens chlorenchyma on the sun-exposed stem side has a photosynthetic pigment composition similar to sun leaves and it experiences significant photoinhibition in the field.     

CO2 assimilation, xanthophyll cycle pigments and PSII efficiency in pumpkin plants as affected by ozone fumigation

CO₂ assimilation, xanthophyll cycle pigments and PSII efficiency were analyzed in two different ages of pumpkin leaves (Cucurbita pepo L. cv. Ambassador) exposed to 150 nmol mol^-1 of ozone (5 days, 5 h day^-1). Gas-exchange measurements revealed a reduction in CO₂ assimilation and stomatal conductance, accompanied by an increase in the intercellular CO₂ concentration both in young and in mature leaves as compared to their respective controls. In both leaves, F₀ remained unchanged, while F_m and the F_v/F_m ratio decreased after O₃ fumigation, indicating that ozone may induce an alteration in the capability of photosystem II (PSII) to reduce the primary acceptor Q_A. In the mature leaves the photochemical quenching (q_p) was significantly lowered by the pollutant, but this was not the case in the young leaves where q_p did not change. In both mature and young ozonated pumpkin leaves, the development of non-photochemical quenching caused a decrease in the PSII photochemical rate, as shown by the correlation between F_v/F_m and the de-epoxidation state of dark-adapted leaves. Decreases in the F_v/F_m ratio are generally attributed to damage to the PSII reaction centre, apart from the down-regulation of the capacity of PSII electron transport. While in young ozonated leaves the decrease in the F_v/F_m ratio was not associated with damage to the D1 protein, in mature ozonated pumpkin leaves, the decrease in the F_v/F_m was accompanied by a significant decline in the D1 content. In conclusion, ozone exposure induces alterations in the light reactions of photosynthesis in both young and mature leaves. However, in young leaves the engagement of the xanthophyll cycle appears to counteract ozone effects against the photosynthetic apparatus as demonstrated by the absence of damage to the D1 protein. On the other hand, the loss of D1 protein in mature fumigated leaves suggests that the activation of the xanthophyll cycle is not sufficient to prevent photoinhibition, probably because a physiological state of senescence adds to the oxidative stress.     

Role of calcium ion in protection against heat and high irradiance stress-induced oxidative damage to photosynthesis of wheat leaves

Cross stress of heat and high irradiance (HI) resulted in the accumulation of active oxygen species and photo-oxidative damage to photosynthetic apparatus of wheat leaves during grain development. Pre-treatment with calcium ion protected the photosynthetic system from oxidative damage by reducing O^-. ₂ production, inhibiting lipid peroxidation, and retarding electrolyte leakage from cell. Therefore, high F_v/F_m [maximal photochemical efficiency of photosystem 2 (PS2) while all PS2 reaction centres are open], F_m/F₀ (another expression for the maximal photochemical efficiency of PS2), Φ_PS2 (actual quantum yield of PS2 under actinic irradiation), q_P (photochemical quenching coefficient), and P _N (net photosynthetic rate) were maintained, and lower q_NP (non-photochemical quenching coefficient) of the leaves was kept under heat and HI stress. EGTA (a chelant of calcium ion) and LaCl₃ (a blocker of Ca²⁺ channel in cytoplasmic membrane) had the opposite effect. Thus Ca ion may help protect the photosynthetic system of wheat leaves from oxidative damage induced by the cross stress of heat and HI.     

Rehydration of Sugar Beet Plants after Water Stress: Effect of Cytokinins

The possibility to improve the recovery of sugar beet plants after water stress by application of synthetic cytokinins N⁶-benzyladenine (BA) or N⁶-(m-hydroxybenzyl)adenosine (HBA) was tested. Relative water content (RWC), net photosynthetic rate (P_N), transpiration rate (E), stomatal conductance (g_s), chlorophyll (Chl) a and Chl b contents, and photosystem 2 efficiency characterized by variable to maximal fluorescence ratio (F_v/F_m) were measured in control plants, in water-stressed plants, and after rehydration (4, 8, 24, and 48 h). Water stress markedly decreased parameters of gas exchange, but they started to recover soon after irrigation. Application of BA or HBA to the substrate or sprayed on leaves only slightly stimulated recovery of P_N, E, and g_s in rehydrated plants, especially during the first phases of recovery. Chl contents decreased only under severe water stress and F_v/F_m ratio was not significantly affected by water stress applied. Positive effects of BA or HBA application on Chl content and F_v/F_m ratio were mostly not observed.     

Simultaneous gas exchange and fluorescence measurements indicate differences in the response of sunflower,bean and maize to water stress

Gas exchange and fluorescence measurements of attached leaves of water stressed bean, sunflower and maize plants were carried out at two light intensities (250 mol quanta m^-2s^-1 and 850 mol quanta m^-2s^-1). Besides the restriction of transpiration and CO₂ uptake, the dissipation of excess light energy was clearly reflected in the light and dark reactions of photosynthesis under stress conditions. Bean and maize plants preferentially use non-photochemical quenching for light energy dissipation. In sunflower plants, excess light energy gave rise to photochemical quenching. Autoradiography of leaves after photosynthesis in ¹⁴CO₂ demonstrated the occurrence of leaf patchiness in sunflower and maize but not in bean. The contribution of CO₂ recycling within the leaves to energy dissipation was investigated by studies in 2.5% oxygen to suppress photorespiration. The participation of different energy dissipating mechanisms to quanta comsumption on agriculturally relevant species is discussed.Abbreviations F_o minimal fluorescence - F_m maximal fluorescence - F_p peak fluorescence - g leaf conductance - P_N net CO₂ uptake - q_N coefficient of non-photochemical quenching - q_P coefficient of photochemical quenching     

Performance of active Photosystem II centers in photoinhibited pea leaves

Effects of photoinhibition on photosynthesis in pea (Pisum sativum L.) leaves were investigated by studying the relationship between the severity of a photoinhibitory treatment (measured as F_v/F_m) and several photoacoustic and chlorophyll a fluorescence parameters. Because of the observed linear relationship between the decline of F_v/F_m and the potential oxygen evolution rate determined by the photoacoustic method, the parameter F_v/F_m was used as an indicator for the severity of photoinhibition. Our analysis revealed that part of the Photosystem II (PS II) reaction centers is inactive in oxygen evolution and is also less sensitive to photoinhibition. Correcting the parameter q_P (fraction of open PS II reaction centers) for inactive PS II centers unveiled a strong increase of q_P in severely inhibited pea leaves, indicating that the inactivated active centers do no longer contribute to q_P and that photoinhibition has an all or none effect on PS II centers. Analysis of q_E (energy quenching) demonstrated its initial increase possibly associated with dephosphorylation of LHC II. Analysis of q_I (photoinhibition dependent quenching) showed that the half-time of recovery of q_I increases steeply below an F_v/F_m of 0.65. This increase of the relaxation half-time corresponds with a decrease of the electron transport rate J and tentatively indicates that the supply of ATP, needed for the recovery, starts to decrease. The data indicate the necessity of correcting for inactive centers in order to make valuable conclusions about effects of photoinhibition on photosynthetic parameters.