Partitioning of photosynthetic electron flow between CO2 and O2 reduction in a C3 leaf (Phaseolus vulgaris L.) at different CO2 concentrations and during drought stress

Photosystem II chlorophyll fluorescence and leaf net gas exchanges (CO₂ and H₂O) were measured simultaneously on bean leaves (Phaseolus vulgaris L.) submitted either to different ambient CO₂ concentrations or to a drought stress. When leaves are under photorespiratory conditions, a simple fluorescence parameter F/ F_m (B. Genty et al. 1989, Biochem. Biophys. Acta 990, 87–92; F = difference between maximum, F_m, and steady-state fluorescence emissions) allows the calculation of the total rate of photosynthetic electron-transport and the rate of electron transport to O₂. These rates are in agreement with the measurements of leaf O₂ absorption using ¹⁸O₂ and the kinetic properties of ribulose-1,5bisphosphate carboxylase/oxygenase. The fluorescence parameter, F/F_m, showed that the allocation of photosynthetic electrons to O₂ was increased during the desiccation of a leaf. Decreasing leaf net CO₂ uptake, either by decreasing the ambient CO₂ concentration or by dehydrating a leaf, had the same effect on the partitioning of photosynthetic electrons between CO₂ and O₂ reduction. It is concluded that the decline of net CO₂ uptake of a leaf under drought stress is only due, at least for a mild reversible stress (causing at most a leaf water deficit of 35%), to stomatal closure which leads to a decrease in leaf internal CO₂ concentration. Since, during the dehydration of a leaf, the calculated internal CO₂ concentration remained constant or even increased we conclude that this calculation is misleading under such conditions.Abbreviations Ca, Ci ambient, leaf internal CO₂ concentrations - F_m, F_o, F_s maximum, minimal, steady-state fluorescence emission - F_v variable fluorescence emission - PPFD photosynthetic photon flux density - q_p, q_N photochemical, non-photochemical fluorescence quenching - Rubisco ribulose-1,5-bisphosphate carboxylase/oxygenase     

Quantum yields for uptake of carbon dioxide in C3 vascular plants of contrasting habitats and taxonomic groupings

The maximum quantum yields (_a,c) for CO₂ uptake in low-oxygen atmospheres were determined for 11 species of C₃ vascular plants of diverse taxa, habitat and life form using an Ulbricht-sphere leaf chamber. Comparisons were also made between tissues of varied age within species. The species examined were Psilotum nudum (L.) P. Beauv., Davallia bullata Wall. ex Hook., Cycas revoluta Thunb., Araucaria heterophylla (Salisb.) Franco, Picea abies (L.) Karst., Nerium oleander L., Ruellia humilis Nutt., Pilea microphylla (L.) Karst., Beaucarnea stricta Lem., Oplismenus hirtellus (L.) P. Beauv. and Poa annua L. Quantum yields were calculated from the initial slopes of the response of CO₂ uptake to the quantity of photons absorbed in conditions of diffuse lighting. Regression analysis of variance of the initial slopes of the response of CO₂ uptake to photon absorption failed to show any statistically significant differences between age classes within species or between the mature photosynthetic organs of different species. The constancy of _a,c was apparent despite marked variation in the light-saturated rates of CO₂ uptake within and between species. The mean _a,c was 0.093±0.003 for 11 species. By contrast, surface absorptance varied markedly between species from 0.90 to 0.60, producing proportional variation in the quantum yield calculated on an incidentlight basis. The ratio of variable to maximum fluorescence emission at 695 nm for the same tissues also failed to show any statistically significant variation between species, with a mean of 0.838±0.008. Mean values of _a,c reported here for C₃ species, in the absence of photorespiration, are higher than reported in previous surveys of vascular plants, but consistent with recent estimates of the quantum yields of O₂ evolution.Abbreviations and Symbols A rate of CO₂ uptake per unit projected area (mol · m^–2 · s^–1) - F_m the maximum fluorescence emission at 695 nm in saturating excitation light when closure of PSII reaction centres is maximal (relative units) - F_o the ground fluorescence at 695 nm when all PSII reaction centres are assumed open (relative units) - F_v the difference between F_m and F_o - J_Q rate of CO₂ uptake by the sample (nmol · s^–1) - J_Q rate of photon absorption by the sample (nmol · s^–1) - Q absorbed photon flux per unit of projected area (nmol · m^–2 · s^–1) - ₁ the light absorptance of photosynthetic organs (dimensionless) - s₁ and s'₁ the total and projected surface areas of the photosynthetic organs examined (m²) - _a,c and _i,c the quantum yields for CO₂ uptake on an absorbed- and incident-light basis, respectively (dimensionless) - _a,o the quantum yield for O₂ evolution on an absorbed-light basis (dimensionless) This work was supported by grant PI7179-BIO, FWF, Austria to H.B-N. and by a British Council travel award to S.P.L. This work was completed under the auspices of U.S. Department of Energy under Contract No. DE-AC02-76CH00016. We also thank Dr. K.J. Parkinson of PP Systems, Hitchin, UK for the loan of a prototype of a commercial integrating-sphere leaf chamber developed from our design.     

Effects of <Emphasis Type="Italic">in vitro</Emphasis>rooting environments and irradiance on growth and photosynthesis of strawberry plantlets during acclimatization

Strawberry (Fragaria ananassaDuch. cv. Fengxiang) plantlets were cultured under two in vitroenvironments for rooting, and then acclimatized under two ex vitroirradiance conditions. At the end of rooting stage plant height, fresh weight and specific leaf area of T1-plants grown under high sucrose concentration (3 sucrose), low photosynthetic photon flux density (30 mol m^–2 s^–1) and normal CO₂ concentration (350–400 l l^–1) were significantly higher than those of T2-plantlets grown under low sucrose concentration (0.5), high photosynthetic photon flux density (90 mol m^–2 s^–1) and elevated CO₂ concentration (700–800 l l^–1). But T2-plantlets had higher net photosynthetic rate (Pn), effective photochemical quantum yield of PSII (_PSII), effective photosynthetic electron transport rate (ETR), photochemical quenching (q_P) and ratio of chlorophyll fluorescence yield decrease (Rfd). After transfer, higher irradiance obviously promoted the growth of plantlets and was beneficial for the development of photosynthetic functions during acclimatization. T2-plantlets had higher fresh weight, leaf area, _PSII and ETR under higher ex vitroirradiance condition.     

Adenine nucleotides and the xanthophyll cycle in leaves

The relationships among the leaf adenylate energy charge, the xanthophyll-cycle components, and photosystem II (PSII) fluorescence quenching were determined in leaves of cotton (Gossypium hirsutum L. cv. Acala) under different leaf temperatures and different intercellular CO₂ concentrations (C_i). Attenuating the rate of photosynthesis by lowering the C_i at a given temperature and photon flux density increased the concentration of high-energy adenylate phosphate bonds (adenylate energy charge) in the cell by restricting ATP consumption (A.M. Gilmore, O. Björkman 1994, Planta 192, 526–536). In this study we show that decreases in photosynthesis and increases in the adenylate energy charge at steady state were both correlated with decreases in PSII photo-chemical efficiency as determined by chlorophyll fluorescence analysis. Attenuating photosynthesis by decreasing C_i also stimulated violaxanthin-de-epoxidation-dependent nonradiative dissipation (NRD) of excess energy in PSII, measured by nonphotochemical fluorescence quenching. However, high NRD levels, which indicate a large trans-thylakoid proton gradient, were not dependent on a high adenylate energy charge, especially at low temperatures. Moreover, dithiothreitol at concentrations sufficient to fully inhibit violaxanthin de-epoxidation and strongly inhibit NRD, affected neither the increased adenylate energy charge nor the decreased PSII photo-chemical efficiency that result from inhibiting photosynthesis. The build-up of a high adenylate energy charge in the light that took place at low C_i and low temperatures was accompanied by a slowing of the relaxation of non-photochemical fluorescence quenching after darkening. This slowly relaxing component of nonphotochemical quenching was also correlated with a sustained high adenylate energy charge in the dark. These results indicate that hydrolysis of ATP that accumulated in the light may acidify the lumen and thus sustain the level of NRD for extended periods after darkening the leaf. Hence, sustained nonphotochemical quenching often observed in leaves subjected to stress, rather than being indicative of photoinhibitory damage, apparently reflects the continued operation of NRD, a photoprotective process.Abbreviations A antheraxanthin - adenylate kinase (myokinase), ATP:AMPphosphotransferase - C_i intercellular CO₂ concentration - DPS de-epoxidation state of violaxanthin, ([Z+A]/[V+A+Z]) - DTT dithiothreitol - pH trans-thylakoid proton gradient - [2ATP+ADP] - F steady-state fluorescence in the presence of NRD - F_M maximal fluorescence in the absence of NRD - F_M maximal fluorescence in the presence of NRD - NRD nonradiative energy dissipation - PET photosynthetic electron transport rate - PFD photon flux density - _PSII photon yield of PSII photochemistry at the actual reduction state in the light or dark - Q_A the primary electron acceptor of PSII - [ATP+ADP+AMP] - SV_N Stern-Volmer nonphotochemical quenching - V violaxanthin - Z zeaxanthin We thank Connie Shih for skillful assistance in growing plants and for conducting HPLC analyses. A Carnegie Institution Fellowship to A.G. is also gratefully acknowledged.     

Inhibition of photosynthesis of sunflower leaves by an endogenous solute and interdependence of different photosynthetic reactions

The relationship between components of non-photochemical quenching of chlorophyll fluorescence yield (q_NP) and dissipation of excessive excitation energy was determined in cotton leaves using concurrent measurements of fluorescence and gas-exchange at 2% and 20% O₂ under a range of photon flux densities and CO₂ pressures. A nearly stoichiometric relationship was obtained between dissipation of energy not used in photosynthetic CO₂ fixation or photorespiration and q_NP provided that a component, probably associated with state transitions, was not included in q_NP. Although two distinct components of q_NP were resolved on the basis of their relaxation kinetics, both components appear effective in energy dissipation. The photon yield of open photosystem-II reaction centers decreased linearly with increases in q_NP, indicating that much of the energy dissipation occurs in the pigment bed. However, increases in q_NP appear dependent on the redox state of these centers. The results are discussed in relation to current hypotheses of the molecular basis of non-radiative energy dissipation. It is concluded that determinations of q_NP can provide a quantitative measure of the dissipation of excessive excitation energy if precautions are taken to ensure that the maximum fluorescence yield is measured under conditions that provide complete closure of the photosystem-II reaction centers. It is also concluded that such dissipation can prevent photoinhibitory damage in cotton leaves even under extreme conditions where as much as 80% of the excitation energy is excessive.Abbreviations and symbols F _M, F _O, F _V, F _S fluorescence yield when all PSII centers are closed, when all centers are open, F_M-F_O, at steady state in the light - PFD photon flux density (photon fluence rate) - P(CO₂) sum of rates of CO₂ uptake and dark respiration - P(ET) sum of P(CO₂) and rate of oxygenation - PSI, PSII photosystem I, II - q_NP, q_P non-photochemical, photochemical fluorescence quenching - Q the acceptor for PSII - Q _r/Q _t the fraction of reduced Q or closed PSII centers - _r/ _t intrinsic photon yield of CO₂ fixation in the absence of photorespiration of O₂ evolution - _a P(ET)/PFD (absorbed light) C.I.W. Publication No. 1016     

Evaluation of the role of State transitions in determining the efficiency of light utilisation for CO2 assimilation in leaves

Wheat leaves were exposed to light treatments that excite preferentially Photosystem I (PS I) or Photosystem II (PS II) and induce State 1 or State 2, respectively. Simultaneous measurements of CO₂ assimilation, chlorophyll fluorescence and absorbance at 820 nm were used to estimate the quantum efficiencies of CO₂ assimilation and PS II and PS I photochemistry during State transitions. State transitions were found to be associated with changes in the efficiency with which an absorbed photon is transferred to an open PS II reaction centre, but did not correlate with changes in the quantum efficiencies of PS II photochemistry or CO₂ assimilation. Studies of the phosphorylation status of the light harvesting chlorophyll protein complex associated with PS II (LHC II) in wheat leaves and using chlorina mutants of barley which are deficient in this complex demonstrate that the changes in the effective antennae size of Photosystem II occurring during State transitions require LHC II and correlate with the phosphorylation status of LHC II. However, such correlations were not found in maize leaves. It is concluded that State transitions in C₃ leaves are associated with phosphorylation-induced modifications of the PS II antennae, but these changes do not serve to optimise the use of light absorbed by the leaf for CO₂ assimilation.Abbreviations Fm, Fo, Fv maximal, minimal and variable fluorescence yields - Fm, Fv maximal and variable fluorescence yields in a light adapted state - LHC II light harvesting chlorophyll a/b protein complex associated with PS II - q_P photochemical quenching - A₈₂₀ light-induced absorbance change at 820 nm - _{PS I}, _{PS II} relative quantum efficiencies of PS I and PS II photochemistry - _{CO 2} quantum yield of CO₂ assimilation     

Effect of temperature on net CO2 assimilation and photosystem II quantum yield of electron transfer of French bean (Phaseolus vulgaris L.) leaves during drought stress

The effect of leaf temperature on stomatal conductance and net CO₂ uptake was studied on French bean (Phaseolus vulgaris L.) using either dehydrated attached leaves (25–40% water deficit) or cut leaves supplied with 10^–4 M abscisic acid (ABA) solution to the transpiration stream. Decreasing leaf temperature caused stomatal opening and increased net CO₂ uptake (which was close to zero at around 25° C) to a level identical to that of control leaves (without water deficit) at around 15° C. (i) The ABA effect on stomatal closure was modulated by temperature and, presumably, ABA is at least partly responsible for stomatal closure of french bean submitted to a drought stress. (ii) For leaf temperatures lower than 15° C, net CO₂ uptake was no longer limited by water deficit even on very dehydrated leaves. This shows that dehydrated leaves retain a substantial part of their photosynthetic capacity which can be revealed at normal CO₂ concentrations when stomata open at low temperature. In contrast to leaves fed with ABA, decreasing the O₂ concentration from 21% to 1% O₂ did not increase either the rate of net CO₂ uptake or the thermal optimum for photosynthesis of dehydrated leaves. The quantum yield of PSII electron flow (measured by F/F_m) was lower in 1% O₂ than in 21% O₂ for each leaf pretreatment given (non-dehydrated leaves, dehydrated leaves, and leaves fed with ABA) even within a temperature range in which leaf photosynthesis at normal CO₂ concentration was the same in these two O₂ concentrations. It is concluded that this probably indicates an heterogeneity of photosynthesis, since this difference in quantum yield disappears when using high CO₂ concentrations during measurements.Abbreviations and Symbols ABA abscisic acid - F_m maximum chlorophyll fluorescence - F difference between steady-state chlorophyll fluorescence and F_m - PPFD photosynthetic photon flux density We would like to thank Dr. J.-M. Briantais (Laboratoire d'écologie végétale, Orsay, France) for help during fluorescence measurements and Ms. J. Liebert for technical assistance.     

Short-term CO2 exchange response to temperature,irradiance, and CO2 concentration in strawberry

Relative importance of short-term environmental interaction and preconditioning to CO₂ exchange response was examined in Fragaria ananasa (strawberry, cv. Quinault). Tests included an orthogonal comparison of 15 to 60-min and 6 to 7-h exposures to different levels of temperature (16 to 32°C), photosynthetically active radiation (PAR, 200 to 800 E m² s^-1), and CO₂ (300 to 600 l/l) on successive days of study. Plants were otherwise maintained at 21°C, 300 E m² s^-1 PAR and 300–360 l/l CO₂ as standard conditions. Treatment was restricted to the mean interval of 14 h daily illumination and the first 3–4 days of each test week over a 12-week cultivation period. CO₂ exchange rates were followed with each step-change in environmental level including ascending/descending temperature/PAR within a test period, initial response at standard conditions on successive days of testing, and measurement at reduced O₂. Response generally supported prior concepts of leaf biochemical modeling in identifying CO₂ fixation as the major site of environmental influence, while overall patterns of whole plant CO₂ exchange suggested additional effects for combined environmental factors and preconditioning. These included a positive interaction between temperature and CO₂ concentration on photosynthesis at high irradiance and a greater contribution by dark respiration at lower PAR than previously indicated. The further importance of estimating whole plant CO₂ exchange from repetitive tests and measurements was evidenced by a high correlation of response to prior treatment both during the daily test period and on consecutive days of testing.Abbreviations C₃ plant a plant in which the product of CO₂ fixation is a 3-carbon acid (3-phosphoglyceric acid) - IRGA intra-red gas analyzer - PAR photosynthetically active radiation - RH relative humidity - RuBisCO ribulose-1,5-bisphosphate carboxylase/oxygenase Reference to a company and/or product named by the Department is only for purposes of information and does not imply approval or recommendation of the product to the exclusion of others which may also be suitable.     

Changes in Lipid Metabolism during Adaptation of the Dunaliella salina Photosynthetic Apparatus to High CO2 Concentration

The effects of CO₂ on the content and composition of lipid fatty acids (FA) and on the photosynthetic characteristics of unicellular halophilic green alga Dunaliella salina (known to be susceptible to CO₂ stress) were investigated. It was shown that even one-day-long increase in the CO₂ concentration (from 2 to 10%) provoked an increase in the total amount of FA on the dry weight basis by 30%. After 7-day-long growth at 10% CO₂, this value was 2.7-fold higher than that at 2% CO₂. The difference in the FA content and composition indicated the activation of FA synthesis de novo and inhibition of their elongation and desaturation, as well as the increase in the relative content of saturated FA at 10% CO₂. It was demonstrated that, after one-day-long CO₂ stress, the MGDG/DGDG ratio increased fourfold without change in the sum of their FA, which indicates the increase in the proportion of lipids predisposed to micellar (hexagonal phase) but not lamellar structure formation. Under short-term CO₂ stress, the ratio of 3/6 FA increased and the content of E-16:113 FA in phosphatidylglycerols increased sharply. The drop in protein content especially in the photosystem I (PSI) preparations, as well as diminishing the ratio of F ₇₀₀-to-F ₆₈₆ nm fluorescence (F ₇₀₀/F ₆₈₆) under short-term CO₂ stress argued for the significant damage to PSI. The reversibility of these changes at more prolonged treatment (7 and 10 days) demonstrated that D. salina cells could restore the functional activity of PSI. The lower level of F ₇₀₀/F ₆₈₆, chlorophyll a (Chla)/Chlb, and 3/6 FA ratio in line with the higher level of E-16:113 in the cells growing for a long time at the high CO₂ concentration is characteristic for the new structural and functional state of the photosynthetic apparatus providing for the effective photosynthesis of D. salina under these conditions.     

Photoinhibition of photosynthesis in intact kiwifruit (Actinidia deliciosa) leaves: Effect of temperature

Photoinhibition of photosynthesis was induced in attached leaves of kiwifruit grown in natural light not exceeding a photon flux density (PFD) of 300 mol·m^-2·s^-1, by exposing them to a PFD of 1500 mol·m^-2·s^-1. The temperature was held constant, between 5 and 35° C, during the exposure to high light. The kinetics of photoinhibition were measured by chlorophyll fluorescence at 77K and the photon yield of photosynthetic O₂ evolution. Photoinhibition occurred at all temperatures but was greatest at low temperatures. Photoinhibition followed pseudo first-order kinetics, as determined by the variable fluorescence (F _v) and photon yield, with the long-term steady-state of photoinhibition strongly dependent on temperature wheareas the observed rate constant was only weakly temperature-dependent. Temperature had little effect on the decrease in the maximum fluorescence (F _m) but the increase in the instantaneous fluorescence (F _o) was significantly affected by low temperatures in particular. These changes in fluorescence indicate that kiwifruit leaves have some capacity to dissipate excessive excitation energy by increasing the rate constant for non-radiative (thermal) energy dissipation although temperature apparently had little effect on this. Direct photoinhibitory damage to the photosystem II reaction centres was evident by the increases in F _o and extreme, irreversible damage occurred at the lower temperatures. This indicates that kiwifruit leaves were most susceptible to photoinhibition at low temperatures because direct damage to the reaction centres was greatest at these temperatures. The results also imply that mechanisms to dissipate excess energy were inadequate to afford any protection from photoinhibition over a wide temperature range in these shade-grown leaves.Abbreviations and symbols fluorescence yield correction coefficient - F _o, F _m, F _v instantaneous, maximum, variable fluorescence - K _D, K _F, K _P, K _T rate constants for non-radiative energy dissipation, fluorescence, photochemistry, energy transfer to photosystem I - PFD photon flux density - PSI, II photosystem I, II - _i photon yield of photosynthesis (incident light)     

Light and the maintenance of photosynthetic competence in leaves of Populus balsamifera L. during short-term exposures to high concentrations of sulfur dioxide

Leaves of Populus balsamifera grown under full natural sunlight were treated with 0, 1, or 2 l SO₂·1^-1 air under one of four different photon flux densities (PFD). When the SO₂ exposures took place in darkness or at 300 mol photons·m^-2·s^-1, sulfate accumulated to the levels predicted by measurements of stomatal conductance during SO₂ exposure. Under conditions of higher PFD (750 and 1550 mol·m^-2·s^-1), however, the predicted levels of accumulated sulfate were substantially higher than those obtained from anion chromatography of the leaf extracts. Light-and CO₂-saturated capacity as well as the photon yield of photosynthetic O₂ evolution were reduced with increasing concentration of SO₂. At 2 l SO₂·1^-1 air, the greatest reductions in both photosynthetic, capacity and photon yield occurred when the leaves were exposed to SO₂ in the dark, and increasingly smaller reductions in each occurred with increasing PFD during SO₂ exposure. This indicates that the inhibition of photosynthesis resulting from SO₂ exposure was reduced when the exposure occurred under conditions of higher light. The ratio F _v/F _M (variable/maximum fluorescence emission) for photosyntem II (PSII), a measure of the photochemical efficiency of PSII, remained unaffected by exposure of leaves to SO₂ in the dark and exhibited only moderate reductions with increasing PFD during the exposure, indicating that PSII was not a primary site of damage by SO₂. Pretreatment of leaves with SO₂ in the dark, however, increased the susceptibility of PSII to photoinhibition, as such pretreated leaves exhibited much greater reductions inF _V/F _M when transferred to moderate or high light in air than comparable control leaves.Abbreviations and symbols A₁₂₀₀ photosynthetic capacity (CO₂-saturated rate of O₂ evolution at 1200 mol photons·m^-2·s^-1) - F_o instantaneous fluorescence emission - F_M maximum fluorescence emission - F_V variable fluorescence emission - PFD photon flux density (400–700 nm) - PSII photosystem II     

Photon yield of O2 evolution and chlorophyll fluorescence characteristics at 77 K among vascular plants of diverse origins

Photon yields of oxygen evolution at saturating CO₂ were determined for 44 species of vascular plants, representing widely diverse taxa, habitats, life forms and growth conditions. The photonyield values on the basis of absorbed light ( ^a) were remarkably constant among plants possessing the same pathway of photosynthetic CO₂ fixation, provided the plants had not been subjected to environmental stress. The mean ^a value ±SE for 37 C₃ species was 0.106±0.001 O₂·photon^-1. The five C₄ species exhibited lower photon yields and greater variation than the C₃ species ( ^a=0.0692±0.004). The ^a values for the two Crassulaceanacid-metabolism species were similar to those of C₃ species. Leaf chlorophyll content had little influence on ^a over the range found in normal, healthy leaves. Chlorophyll fluorescence characteristics at 77 K were determined for the same leaves as used for the photon-yield measurements. Considerable variation in fluorescence emission both at 692 nm and at 734 nm, was found 1) among the different species; 2) between the upper and lower surfaces of the same leaves; and 3) between sun and shade leaves of the same species. By contrast, the ratio of variable to maximum fluorescence emission at 692 nm (F_v/F_M, 692) remained remarkably constant (The mean value for the C₃ species was 0.832±0.004). High-light treatments of shade leaves resulted in a reduction in both ^a and the F_v/F_M, 692 ratio. The extent of the reductions increased with time of exposure to bright light. A linear relationship was obtained when ^a was plotted against F_v/F_M, 692. The results show that determinations of the photon yield of O₂ evolution and the F_v/F_M, 692 ratio can serve as excellent quantitative measures of photoinhibition of overall photosynthetic energy-conversion system and of photochemistry of photosystem II, respectively. This is especially valuable in field work where it is often impossible to obtain appropriate controls.Abbreviations and symbols CAM Crassulacean acid metabolism - PFD photon flux density (photon fluence rate) - PSI, PSII photosystem I, II - F_o, F_M, F_v instantaneous, maximum, variable fluorescence emission - absorptance - ^a photon yield (absorbed light) - ⁱ photon yield (incident light) C.I.W.-D.P.B. Publication No. 923     

Recycling of respiratory CO2 during Crassulacean acid metabolism: alleviation of photoinhibition in Pyrrosia piloselloides

The regulation of Crassulacean acid metabolism (CAM) in the fern Pyrrosia piloselloides (L.) Price was investigated in Singapore on two epiphytic populations acclimated to sun and shade conditions. The shade fronds were less succulent and had a higher chlorophyll content although the chlorophyll a:b ratio was lower and light compensation points and dark-respiration rates were reduced. Dawn-dusk variations in titratable acidity and carbohydrate pools were two to three times greater in fronds acclimated to high photosynthetically active radiation (PAR), although water deficits were also higher than in shade fronds. External and internal CO₂ supply to attached fronds of the fern was varied so as to regulate the magnitude of CAM activity. A significant proportion of titratable acidity was derived from the refixation of respiratory CO₂ (27% and 35% recycling for sun and shade populations, respectively), as measured directly under CO₂-free conditions. Starch was shown to be the storage carbodydrate for CAM in Pyrrosia, with a stoichiometric reduction of C₃-skeleton units in proportion to malic-acid accumulation. Measurements of photosynthetic O₂ evolution under saturating CO₂ were used to compare the light responses of sun and shade fronds for each CO₂ supply regime, and also following the imposition of a photoinhibitory PAR treatment (1600 mol·m^-2·s^-1 for 3 h). Apparent quantum yield declined following the high-PAR treatment for sun- and shade-adapted plants, although for sun fronds CAM activity derived from respiratory CO₂ prevented any further reduction in photosynthetic efficiency. Recycling of respiratory CO₂ by shade plants could only partly prevent photoinhibitory damage. These observations provide experimental evidence that respiratory CO₂ recycling, ubiquitous in CAM plants, may have developed so as to alleviate photoinhibition.Abbreviations and symbols CAM Crassulacean acid metabolism - F_M maximal photosystem II fluorescence - F_T terminal steady-state fluorescence - PAR photosynthetically active radiation, 400–700 nm - H⁺ (dawn-dusk) variation in titratable acidity     

Effect of dissolved inorganic carbon on oxygen evolution and uptake by Chlamydomonas reinhardtii suspensions adapted to ambient and CO2-enriched air

Mass spectrometric measurements of ¹⁶O₂ and ¹⁸O₂ isotopes were used to compare the rates of gross O₂ evolution (E₀), O₂ uptake (U₀) and net O₂ evolution (NET) in relation to different concentrations of dissolved inorganic carbon (DIC) by Chlamydomonas reinhardtii cells grown in air (air-grown), in air enriched with 5% CO₂ (CO₂-grown) and by cells grown in 5% CO₂ and then adapted to air for 6h (air-adapted).At a photon fluence rate (PFR) saturating for photosynthesis (700 mol photons m^-2 s^-1), pH=7.0 and 28°C, U₀ equalled E₀ at the DIC compensation point which was 10M DIC for CO₂-grown and zero for air-grown cells. Both E₀ and U₀ were strongly dependent on DIC and reached DIC saturation at 480 M and 70 M for CO₂-grown and air-grown algae respectively. U₀ increased from DIC compensation to DIC saturation. The U₀ values were about 40 (CO₂-grown), 165 (air-adapted) and 60 mol O₂ mg Chl^-1 h^-1 (air-grown). Above DIC compensation the U₀/E₀ ratios of air-adapted and air-grown algae were always higher than those of CO₂-grown cells. These differences in O₂ exchange between CO₂- and air-grown algae seem to be inducable since air-adapted algae respond similarly to air-grown cells.For all algae, the rates of dark respiratory O₂ uptake measured 5 min after darkening were considerably lower than the rates of O₂ uptake just before darkening. The contribution of dark respiration, photorespiration and the Mehler reaction to U₀ is discussed and the energy requirement of the inducable CO₂/HCO₃ ^- concentrating mechanism present in air-adapted and air-grown C. reinhardtii cells is considered.Abbreviations DIC dissolved inorganic carbon - DCMU 3-(3,4-dichlorophenyl)-1,1-dimethylurea - E₀ rate of photosynthetic gross O₂ evolution - PCO photosynthetic carbon oxidation - PFR photon fluence rate - PS I photosystem I - PS II photosystem II - U₀ rate of O₂ uptake in the light - MS mass spectrometer     

Acclimation of Arabidopsis thaliana to the light environment: changes in photosynthetic function

Arabidopsis thaliana (L.) Heynh. cv. Landsberg erecta was grown under light regimes of differing spectral qualities, which results in differences in the stoichiometries of the two photosynthetic reaction centres. The acclimative value of these changes was investigated by assessing photosynthetic function in these plants when exposed to two spectrally distinct actinic lights. Plants grown in an environment enriched in far-red light were better able to make efficient use of non-saturating levels of actinic light enriched in long-wavelength red light. Simultaneous measurements of chlorophyll fluorescence and absorption changes at 820 nm indicated that differences between plants grown under alternative light regimes can be ascribed to imbalances in excitation of photosystems I and II (PSI, PSII). Measurements of chlorophyll fluorescence emission and excitation spectra at 77 K provided strong evidence that there was little or no difference in the composition or function of PSI or PSII between the two sets of plants, implying that changes in photosynthetic stoichiometry are primarily responsible for the observed differences in photosynthetic function.Abbreviations Chl chlorophyll - FR far-red light - HF highirradiance FR-enriched light (400 mol·m^–2·s^–1, RFR = 0.72) - HW high-irradiance white light (400 mol·m^–2 1·1 s^–1RFR = 1.40) - LHCI, LHCII light-harvesting complex of PSI, PSII - qO quenching of dark-level chlorophyll fluorescence - qN non-photochemical quenching of variable chlorophyll fluorescence - qP photochemical quenching of variable chlorophyll fluorescence - R red light - Rubisco ribulose-1,5-bisphosphate carboxylase/oxygenase We thank Dr. Sasha Ruban for assistance with the 77 K fluorescence measurements and for helpful discussions. This work was supported by Natural Environment Research Council Grant GR3/7571A.     

Photoinhibition of photosynthesis in intact kiwifruit (Actinidia deliciosa) leaves: Changes in susceptibility to photoinhibition and recovery during the growth season

Kiwifruit (Actinidia deliciosa (A. Chev.) C.F. Liang et A.R. Ferguson) plants grown in an outdoor enclosure were exposed to the natural conditions of temperature and photon flux density (PFD) over the growing season (October to May). Temperatures ranged from 14 to 21° C while the mean monthly maximum PFD varied from 1000 to 1700 mol · m^–2 · s^–1, although the peak PFDs exceeded 2100 mol · m^–2 · s^–1. At intervals, the daily variation in chlorophyll fluorescence at 692 nm and 77K and the photon yield of O₂ evolution in attached leaves was monitored. Similarly, the susceptibility of intact leaves to a standard photoinhibitory treatment of 20° C and a PFD of 2000 mol · m^–2 · s^–1 and the ability to recover at 25° C and 20 mol · m^–2 · s^–2 was followed through the season. On a few occasions, plants were transferred either to or from a shade enclosure to assess the suceptibility to natural photoinhibition and the capacity for recovery. There were minor though significant changes in early-morning fluorescence emission and photon yield throughout the growing season. The initial fluorescence, F_o, and the maximum fluorescence, F_m, were, however, significantly and persistently different from that in shade-grown kiwifruit leaves, indicative of chronic photoinhibition occurring in the sun leaves. In spring and autumn, kiwifruit leaves were photoinhibited through the day whereas in summer, when the PFDs were highest, no photoinhibition occurred. However, there was apparently no non-radiative energy dissipation occurring then also, indicating that the kiwifruit leaves appeared to fully utilize the available excitation energy. Nevertheless, the propensity for kiwifruit leaves to be susceptible to photoinhibition remained high throughout the season. The cause of a discrepancy between the severe photoinhibition under controlled conditions and the lack of photoinhibition under comparable, natural conditions remains uncertain. Recovery from photoinhibition, by contrast, varied over the season and was maximal in summer and declined markedly in autumn. Transfer of shade-grown plants to full sun had a catastrophic effect on the fluorescence characteristics of the leaf and photon yield. Within 3 d the variable fluorescence, F_v, and the photon yield were reduced by 80 and 40%, respectively, and this effect persisted for at least 20 d. The restoration of fluorescence characteristics on transfer of sun leaves to shade, however, was very slow and not complete within 15 d.Abbreviations and Symbols F_o, F_m, F_v initial, maximum, variable fluorescence - F_i F_v at t = 0 - F F_v at t = - PFD photon flux density - PSII photosystem II - leaf absorptance ratio - (_a photon yield of O₂ evolution (absorbed basis) - _{i a} at t = 0 - _a at t = We thank Miss Linda Muir and Amanda Yeates for their technical assistance in this study.     

Leaf cavity CO2 concentrations and CO2 exchange in onion,Allium cepa L.

Onion (Allium cepa L.) plants were examined to determine the photosynthetic role of CO₂ that accumulates within their leaf cavities. Leaf cavity CO₂ concentrations ranged from 2250 L L^–1 near the leaf base to below atmospheric (<350 L L^–1) near the leaf tip at midday. There was a daily fluctuation in the leaf cavity CO₂ concentrations with minimum values near midday and maximum values at night. Conductance to CO₂ from the leaf cavity ranged from 24 to 202 mol m^–2 s^–1 and was even lower for membranes of bulb scales. The capacity for onion leaves to recycle leaf cavity CO₂ was poor, only 0.2 to 2.2% of leaf photosynthesis based either on measured CO₂ concentrations and conductance values or as measured directly by ¹⁴CO₂ labeling experiments. The photosynthetic responses to CO₂ and O₂ were measured to determine whether onion leaves exhibited a typical C₃-type response. A linear increase in CO₂ uptake was observed in intact leaves up to 315 L L^–1 of external CO₂ and, at this external CO₂ concentration, uptake was inhibited 35.4±0.9% by 210 mL L^–1 O₂ compared to 20 mL L^–1 O₂. Scanning electron micrographs of the leaf cavity wall revealed degenerated tissue covered by a membrane. Onion leaf cavity membranes apparently are highly impermeable to CO₂ and greatly restrict the refixation of leaf cavity CO₂ by photosynthetic tissue.Abbreviations C_a external CO₂ concentration - C_i intercellular CO₂ concentration - CO₂ compensation concentration - PPFR photosynthetic photon fluence rate     

Growth,photosynthesis and photorespiration of Lemna gibba: response to variations in CO2 and O2 concentrations and photon flux density

Dry weight and Relative Growth Rate of Lemna gibba were significantly increased by CO₂ enrichment up to 6000 l CO₂ l^–1. This high CO₂ optimum for growth is probably due to the presence of nonfunctional stomata. The response to high CO₂ was less or absent following four days growth in 2% O₂. The Leaf Area Ratio decreased in response to CO₂ enrichment as a result of an increase in dry weight per frond. Photosynthetic rate was increased by CO₂ enrichment up to 1500 l CO₂ l^–1 during measurement, showing only small increases with further CO₂ enrichment up to 5000 l CO₂ l^–1 at a photon flux density of 210 mol m^–2 s^–1 and small decreases at 2000 mol m^–1 s^–1. The actual rate of photosynthesis of those plants cultivated at high CO₂ levels, however, was less than the air grown plants. The response of photosynthesis to O₂ indicated that the enhancement of growth and photosynthesis by CO₂ enrichment was a result of decreased photorespiration. Plants cultivated in low O₂ produced abnormal morphological features and after a short time showed a reduction in growth.     

Effects of extreme high temperature,drought and elevated CO2 on photosynthesis of the Mojave Desert evergreen shrub,Larrea tridentata

The interaction of extreme temperature events with future atmospheric CO₂ concentrations may have strong impacts on physiological performance of desert shrub seedlings, which during the critical establishment phase often endure temperature extremes in conjunction with pronounced drought. To evaluate the interaction of drought and CO₂ on photosynthesis during heat stress, one-year-old Larrea tridentata[DC] Cov. seedlings were exposed to nine days of heat with midday air temperature maxima reaching 53 °C under three atmospheric CO₂ concentrations (360, 550 and 700 mol mol^–1) and two water regimes (well-watered and droughted). Photosynthetic gas exchange, chlorophyll fluorescence and water potential responses were measured prior to, during and one week following the high temperature stress event. Heat stress markedly decreased net photosynthetic rate (A _net), stomatal conductance (g _s), and the photochemical efficiency of photosystem II (F _v/F _m) in all plants except for well-watered L. tridentata grown in 700 mol mol^–1 CO₂. A _net and g _s remained similar to pre-stress levels in these plants. In droughted L. tridentata, A _net was ca. 2× (in 550 mol mol^–1 CO₂) to 3× (in 700 mol mol^–1 CO₂) higher than in ambient-CO₂-grown plants, while g _s and F _v/F _m were similar and low in all CO₂ treatments. Following heat stress, g _s in all well-watered plants rose dramatically, exceeding pre-stress levels by up to 100%. In droughted plants, g _s and A _net rose only in plants grown at elevated CO₂ following release from heat. This recovery response was strongest at 700 mol mol^–1 CO₂, which returned to A _net and g _s values similar to pre-heat following several days of recovery. Extreme heat diminished the photosynthetic down-regulation response to growth at elevated CO₂ under well-watered conditions, similar to the action of drought. Ambient-CO₂-grown L. tridentata did not show significant recovery of photosynthetic capacity (A _\max and CE) after alleviation of temperature stress, especially when exposed to drought, while plants exposed to elevated CO₂ appeared to be unaffected. These findings suggest that elevated CO₂ could promote photosynthetic activity during critical periods of seedling establishment, and enhance the potential for L. tridentata to survive extreme high temperature events.