High-light damage in air-dry thalli of the old forest lichen Lobaria pulmonaria-interactions of irradiance, exposure duration and high temperature

High-light damage in air-dry thalli of Lobaria pulmonaria were measured in the laboratory as reductions in maximal PSII efficiency (Fv/Fm) after a 48 h recovery in a hydrated state at low light to account for permanent damage. Thalli treated with the lowest light dose (90 mol photons m^-2) recovered normal Fv/Fm-values with increasing irradiances (400-700 nm) up to 1000 mol photons m^-2 s^-1. Doubling this dose lowered the threshold level for damage from 1000 to 320 mol photons m^-2 s^-1, and reduced Fv/Fm at 1000 mol photons m^-2 s^-1 by more than 50%. A second doubling of the dose to 360 mol photons m^-2 caused damage at 200 mol photons m^-2 s^-1, and a nearly complete cessation of PSII efficiency occurred at 1000 mol photons m^-2 s^-1. No reciprocity of irradiance and duration of illumination for PSII function was found. The measured time-dependent decrease in Fv/Fm was remarkably similar for the naturally coupled, but artificially separated, light and temperature factors. Therefore, the damage of high light on desiccated L. pulmonaria seemed to be an additive effect of high irradiance and high temperatures. Air-dry thalli were highly heat susceptible, being affected already at temperatures around 40C. Logging operations in forests are likely to raise the solar radiation at remaining lichen sites to destructive levels.Keywords: Lichens, high-light damage, heat stress, poikilohydric organisms, reciprocity.      

Light screening in lichen cortices can be quantified by chlorophyll fluorescence techniques for both reflecting and absorbing pigments

Lichens, representing mutualistic symbioses between photobionts and mycobionts, often accumulate high concentrations of secondary compounds synthesized by the fungal partner. Light screening is one function for cortical compounds being deposited as crystals outside fungal hyphae. These compounds can non-destructively be extracted by 100% acetone from air-dry living thalli. Extraction of atranorin from Physcia aipolia changed the lichen colour from pale grey to green in the hydrated state, whereas acetone-rinsed and control thalli were all pale grey when dry. Removal of parietin from Xanthoria parietina changed the colour of desiccated thalli from orange to grey. Colour changes were quantified by reflectance measurements. By a new chlorophyll fluorescence method, screening was assessed as the decrease in incident irradiance (PAR) necessary to reach identical effective quantum yields of PSII (Φ_PSII) in acetone-rinsed and control thalli. Thereby, we estimated a screening efficiency due to cortical atranorin crystals at 61, 38, and 40% of blue, green and red light, respectively, whereas parietin screened 81, 27 and 1% of these wavelength ranges. Removal of atranorin caused similar levels of increased photoinhibition for P. aipolia in blue, green and red light, whereas parietin-deficient thalli of X. parietina exhibited increased photoinhibition with decreasing wavelengths. Atranorin possibly prevents water from entering the spaces between the hyphae in the cortex. The air-filled cavities with white atranorin crystals reflect excess light, whereas the yellow compound parietin absorbs excess light. Thereby, both atranorin and parietin play significant photoprotective roles for symbiotic green algae, but with compound-specific screening mechanisms.     

Response of the Photosynthetic Apparatus of the Diatom Thallassiosira weisflogii to High Irradiance Light

The response of the photosynthetic apparatus to high irradiance illumination (440–2200 W/m²) was studied in the diatom Thallassiosira weisflogii by fluorescence methods. Changes in the photosynthetic apparatus were monitored by measuring characteristics of chlorophyll fluorescence F ₀, F _m, F _v/F _m, and qN for several hours after illumination of the alga with high-intensity light. Incubation of the alga with 2 mM DTT, an inhibitor of de-epoxidase of carotenoids in the diadinoxanthin cycle, led to a decrease in the nonphotochemical quenching of chlorophyll fluorescence and a drop in the F _v/F _m ratio, a characteristic that reflects the quantum efficiency of the functioning of the photosynthetic apparatus. Light-induced absorption changes associated with transformations of carotenoids of diadinoxanthin cycle were recorded in vivo in algal suspensions in the absence and in the presence of DTT. Using the microfluorometric method, we measured cell distribution over the efficiency of the primary processes of photosynthesis (F _v/F _m) after illumination. We found cells with a high tolerance of their photosynthetic apparatus to photooxidative damage. The relatively high tolerance of a portion of the cell population to high-light illumination can be related to light-induced transformation of carotenoids and to the functioning of other protective systems of the photosynthetic apparatus in diatoms.     

Mating systems in representatives of Parmeliaceae, Ramalinaceae and Physciaceae (Lecanoromycetes, lichen-forming ascomycetes)

The progeny of meiosis of eight Parmeliaceae, two Ramalinaceae and seven Physciaceae were subjected to fingerprint analysis using RAPD-PCR applied to single spore isolates. The sample set included common and widespread rarely fertile species (Parmelia sulcata, Pseudevernia furfuracea, Physcia tenella), local to common, infrequently fertile species (Melanelixia glabra, Parmelina tiliacea, Xanthoparmelia conspersa, X. stenophylla, Anaptychia runcinata, Diploicia canescen, Physconia distorta), local to rare, infrequently or regularly fertile species with declining distributions (Parmelina carporrhizans, P. quercina, Ramalina fastigiata, R. fraxinea, Anaptychia ciliaris), and local to common, regularly fertile species (Physcia aipolia, P. stellaris). All species turned out to be heterothallic, polymorphisms among RAPD markers ranging from 10–87 %. The significance of these findings for population genetics and conservation biology, and potential reasons for infrequent ascoma formation in some of the species are discussed.     

Net photosynthetic response patterns of the basidiomycete lichen Cora pavonia (Web.) E. Fries from the tropical volcano La Soufrière (Guadeloupe)

Summary The response of net photosynthesis to temperature, moisture, and light was examined in thalli of the tropical basidiomycete lichen Cora pavonia from recent lahar flows on the volcanic summit La Soufrière (Guadeloupe, French West Indies). Although thalli of C. pavonia are typically exposed to only low light intensities and isothermal temperature conditions under prevailing cloud/shroud conditions on La Soufrière, their photosynthetic response matrix reveals an unexpected breadth of response. The temperature optimum of net photosynthetic uptake in C. pavonia rises from 6°C at a photon flux area density of 25 mol m^–2 s^–1 PAR to 27°C at 1000 mol m^–2 s^–1 PAR, with rates of maximal net photosynthetic uptake exceeding 25 mg CO₂ g^–1 h^–1. Net photosynthesis was optimal at thallus moisture contents of 250 to 350 percent water content by weight, declining only slightly in fully saturated thalli. These response patterns pose an apparent paradox, as on most days they will act to severely restrict net photosynthetic uptake by thalli of C. pavonia on La Soufrière. This paradox is discussed in context of those selective pressures faced by lichen thalli in later successional stages as well as those imposed by brief periods of atypical weather conditions.     

Curling during desiccation protects the foliose lichen Lobaria pulmonaria against photoinhibition

This study aims to assess the photoprotective potential of desiccation-induced curling in the light-susceptible old forest lichen Lobaria pulmonaria by using chlorophyll fluorescence imaging. Naturally curled thalli showed less photoinhibition-induced limitations in primary processes of photosynthesis than artificially flattened specimens during exposures to 450 μmol m⁻² s⁻¹ in the laboratory after both 12- (medium dose treatment) and 62-h duration (high dose treatment). Thallus areas shaded by curled lobes during light exposure showed unchanged values of measured chlorophyll fluorescence parameters (F _V/F _M, Φ_{PS II}), whereas non-shaded parts of curled thalli, as well as the mean for the entire flattened thalli, showed photoinhibitory limitation after light treatments. Furthermore, the chlorophyll fluorescence imaging showed that the typical small-scale reticulated ridges on the upper side of L. pulmonaria caused a spatial, small-scale reduction in damage due to minor shading. Severe dry-state photoinhibition readily occurred in flattened and light-treated L. pulmonaria, although the mechanisms for such damage in a desiccated and inactive stage are not well known. Natural curling is one strategy to reduce the chance for serious photoinhibition in desiccated L. pulmonaria thalli during high light exposures.     

Changes of fluorescence and xanthophyll pigments during dehydration in the resurrection plantSelaginella lepidophylla in low and medium light intensities

The changes in photosynthetic efficiency and photosynthetic pigments during dehydration of the resurrection plantSelaginella lepidophylla (from the Chiuhahuan desert, S.W. Texas, USA) were examined under different light conditions. Changes in the photosynthetic efficiency were deduced from chlorophyll a fluorescence measurements (F_o, F_m, and F_v) and pigment changes were measured by HPLC analysis. A small decrease in F_v/F_m was seen in hydrated stems in high light (650 μmol photons·m⁻²·s⁻¹) but not in low light (50 μmol photons·m⁻²·s⁻¹). However, a pronounced decline in F_v/F_m was observed during dehydration in both light treatments, after one to two hours of dehydration. A rise in F_o was observed only after six to ten hours of dehydration. Concomitant with the decrease in photosynthetic efficiency during dehydration a rise in the xanthophyll zeaxanthin was observed, even in low-light treatments. The increase in zeaxanthin can be related to previously observed photoprotective non-photochemical quenching of fluorescence in dehydrating stems ofS. lepidophylla. We hypothesize that under dehydrating conditions even low light levels become excessive and zeaxanthin-related photoprotection is engaged. We speculate that these processes, as well as stem curling and self shading (Eickmeier et al. 1992), serve to minimize photoinhibitory damage toS. lepidophylla during the process of dehydration.     

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.     

Reduced photoinhibition with stem curling in the resurrection plant Selaginella lepidophylla

Summary Selaginella lepidophylla, the resurrection plant, curls dramatically during desiccation and the hypothesis that curling may help limit bright light-induced damage during desiccation and rehydration was tested under laboratory conditions. Restraint of curling during desiccation at 25° C and a constant irradiance of 2000 mol m^–2 s^]t-1 significantly decreased PSII and whole-chain electron transport and the F_v/F_m fluorescence yield ratio following rehydration relative to unrestrained plants. Normal curling during desiccation at 37.5°C and 200 mol m^–2 s^–1 irradiance did not fully protect against photoinhibition or chlorophyll photooxidation indicating that some light-induced damage occurred early in the desiccation process before substantial curling. Photosystem I electron transport was less inhibited by high-temperature, high-irradiance desiccation than either PSII or whole-chain electron transport and PSI was not significantly affected by restraint of curling during desiccation at 25°C and high irradiance. Previous curling also helped prevent photoinhibition of PSII electron transport and loss of whole-plant photosynthetic capacity as the plants uncurled during rehydration at high light. These results demonstrate that high-temperature desiccation exacerbated photoinhibition, PSI was less photoinhibited than PSII or whole-chain electron transport, and stem curling ameliorated bright light-induced damage helping to make rapid recovery of photosynthetic competence possible when the plants are next wetted.     

Photosynthetic response of Ulva rotundata to light and temperature during emersion on an intertidal sand flat

Summary We have investigated the diurnal response of photosynthesis and variable photosystem II (PSII) chlorophyll fluorescence at 77 K for thalli of the chlorophyte macroalga, Ulva rotundata, grown in outdoor culture and transplanted to an intertidal sand flat in different seasons. The physiological response in summer indicated synergistic effects of high PFD and aerial exposure, the latter probably attributable to temperature, which usually increased by 8 to 10° C during midday emersion. Except at extreme emersed temperatures in summer (38° C), the light-saturated photosynthesis rate (P_m) did not decline at midday. In contrast, light-limited quantum yield of photosynthetic O₂ exchange () and the ratio of variable to maximum fluorescence yield (F_v/F_m) reversibly declined during midday low tides in all seasons. Shade-grown thalli exhibited a fluorescence response suggestive of greater photodamage to PSII, whereas sun-grown thalli had greater photoprotective capacity. The fluorescence decline was smaller when high tide occurred at midday, and was delayed during morning cloudiness. These results suggest that the diurnal response to PFD in this shallow water species is modified by tidal and meteorological factors. U. rotundata has a great capacity for photoprotection which allows it to tolerate and even thrive in the harsh intertidal environment.Abbreviations F_o instantaneous yield of chlorophyll fluorescence - F_m maximum yield of fluorescence - F_v variable yield (F_m–F_o) of fluorescence - PFD photon flux density (400–700 nm) - P_m light-saturated rate of photosynthesis - PSH photosystem II - Q_A electron acceptor of PSII - light-limited quantum yield of photosynthesis     

Photosynthetic acclimation to different light levels in the brown marine macroalga, <Emphasis Type="Italic">Hizikia fusiformis</Emphasis> (Sargassaceae,Phaeophyta)

Hizikia fusiformis thalli experience dynamic incident light conditions during the period of growth. The present study was designed to examine how changing photon irradiance affects the photosynthesis both in the short and long terms by culturing H. fusiformis under three different light levels: 35 μmol photons m^-2 s^-1 (low light, LL), 85 μmol photons m^-2 s^-1 (intermediate light, IL), and 165 μmol photons m^-2 s^-1 (high light, HL). A similar relative growth rate was observed between IL- and HL-grown algae, but the growth rate was significantly reduced in LL-grown algae. The photosynthetic rates (P _n) measured at their respective growth light levels were found to be lowest in the thalli grown at LL and highest at HL. However, LL-grown algae exhibited much higher P _n in comparison with IL- and the HL-grown thalli at the same measuring photosynthetic photon flux density, indicating the photosynthetic acclimation to low growth light in H. fusiformis. The photosynthesis–light curves showed that LL-grown algae had a highest light-saturating maximum P _n (P _max) in comparison with IL- or HL-grown algae when the photosynthetic rates were expressed on the biomass basis. However, P _max was highest in HL-grown algae compared to IL- or LL-grown algae when the rates were normalized to chlorophyll a. The photosynthesis–inorganic carbon (Ci) response curves were also significantly affected by the growth light conditions. The highest value of apparent photosynthetic conductance occurred in LL-grown algae while the lowest value in HL-grown algae. Additionally, the activity of external carbonic anhydrase (CA) tended to increase while the total CA activity inclined to decrease in H. fusiformis thalli when the growth light level altered from 35 to 165 μmol photons per square meter per second. The external CA inhibitors showed a higher inhibition in HL-grown algae compared with LL-grown algae. It was proposed that photosynthetic acclimation to low light condition in H. fusiformis was achieved through an increase in the number of reaction centers and increased capacities of electron transport and of Ci transport within cells. The ability of photosynthetic acclimation to low light confers H. fusiformis thalli to overcome the environmental low light condition as a result of the attenuation of seawater or self-shading through enhancing its photosynthetic performance and carbon assimilation necessary for growth.     

Ecological and physiological investigations in continental Antarctic cryptogams

Summary Microclimate and CO₂ exchange of the lichen Usnea sphacelata were measured during summen on a hill near Casey Station, Bailey Peninsula, Wilkes Land, Antarctica. Within a period of 52 days (November 10 until December 31, 1985), 8 diurnal courses of net photosynthesis were measured in naturally snow-covered lichen thalli, and 9 diurnal courses in thalli experimentally sprayed with melt water. Photosynthetic performance of a light-form of Usnea sphacelata was compared with that of a shade-form. Net photosynthesis was reversibly depressed in snow-covered lichen thalli of both forms when irradiance was higher than 600 mol m^–2 s^–1 photosynthetic active radiation (PAR), the depression persisting several hours after a period of strong light. These responses suggest photoinhibition. Models of photosynthesis were established for the light-form by non-linear regressions with field data from water-sprayed thalli (Model W) and field data measured in snow-covered lichens (SNO I, SNO II). Model SNO I is based on median values of photosynthetic rates and SNO II on maximum values for each light/temperature combination. Photosynthetic rates were calculated using model W; the results showed values approximately three times higher than measured in the field with naturally moistened thalli. Photosynthetic rates according to model SNO II fitted the data of naturally moistened lichens measured during the day, before strong light (> 600 mol m^–2s^–1 PAR) caused reversible decrease of net photosynthesis. Model SNO I fitted the data measured during and after a phase of strong irradiance. Model SNO I demonstrated that light stress was highest at temperatures below 2 °C. This study has shown that long-term calculation of the photosynthetic productivity must take into account decreases in net photosynthesis rate caused by strong light, as well as effects of water content and temperature. For the investigated period of the austral summer, a carbon production of 3.44 gm^–2 was estimated for U. sphacelata.     

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.     

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     

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.     

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)     

Photoinhibition of photosynthesis under natural conditions in ivy (Hedera helix L.) growing in an understory of deciduous trees

We investigated to what extent south-exposed leaves (E-leaves) of the evergreen ivy (Hedera helix L.) growing in the shadow of two deciduous trees suffered from photoinhibition of photosynthesis when leaf-shedding started in autumn. Since air temperatures drop concomitantly with increase in light levels, changes in photosynthetic parameters (apparent quantum yield, _i and maximal photosynthetic capacity of O₂ evolution, P_max; chlorophyll-a fluorescence at room temperature) as well as pigment composition were compared with those in north-exposed leaves of the same clone (N-leaves; photosynthetic photon flux density PPFD< 100 mol · m^–2 · s^–2) and phenotypic sun leaves (S-leaves; PPFD up to 2000 mol · m^–2 · s^–1).In leaves exposed to drastic light changes during winter (E-leaves) strong photoinhibition of photosynthesis could be observed as soon as the incident PPFD increased in autumn. In contrast, in N-leaves the ratio of variable fluorescence to maximum fluorescence (F_V/F_Mm) and _i did not decline appreciably prior to severe frosts (up to -12° C) in January. At this time, _i was reduced to a similar extent in all leaves, from about 0.073 mol O₂ · mol^–1 photons before stress to about 0.020. Changes in _i were linearly correlated with changes in f_v/f_m (r = 0.955). The strong reduction in F_V/F_M on exposure to stress was caused by quenching in F_M. The initial fluorescence (F₀), however, was also quenched in all leaves. The diminished fluorescence yield was accompanied by an increase in zeaxanthin content. These effects indicate that winter stress in ivy primarily induces an increase in non-radiative energy-dissipation followed by photoinhibitory damage of PSII. Although a pronounced photooxidative bleaching of chloroplast pigments occurred in January (especially in E-leaves), photosynthetic parameters recovered completely in spring. Thus, the reduction in potential photosynthetic yield in winter may be up to three times greater in leaves subjected to increasing light levels than in leaves not exposed to a changing light environment.Abbreviations and Symbols F₀, F_M initial and maximal fluorescence yield when all PSII centres are open and closed - F_V variable fluorescence (F_M-F₀) - P_max maximal photosynthetic capacity at 1000 umol · m^–2 · s^–1 PPFD and CO₂ saturation - PPFD photosynthetic photon flux density - _i apparent quantum yield of photosynthetic O₂ evolution - E-leaves, N-leaves shade leaves exposed, not exposed to drastic light changes during winter - S-leaves sun leaves from an open ivy stand Dedicated to Professor Otto Härtel on the occasion of his 80th birthdayThis work was supported by the Austrian Fonds zur Förderung der wissenschaftlichen Forschung.     

Xanthophyll cycle and light stress in nature: uniform response to excess direct sunlight among higher plant species

Photosystem II (PS II) efficiency, nonphotochemical fluorescence quenching, and xanthophyll cycle composition were determined in situ in the natural environment at midday in (i) a range of differently angled sun leaves ofEuonymus kiautschovicus Loesener and (ii) in sun leaves of a wide range of different plant species, including trees, shrubs, and herbs. Very different degrees of light stress were experienced by these leaves (i) in response to different levels of incident photon flux densities at similar photosynthetic capacities amongEuonymus leaves and (ii) as a result of very different photosynthetic capacities among species at similar incident photon flux densities (that were equivalent to full sunlight). ForEuonymus as well as the interspecific comparison all data fell on one single, close relationship for changes in intrinsic PSII efficiency, nonphotochemical fluorescence quenching, or the levels of zeaxanthin + antheraxanthin in leaves, respectively, as a function of the actual level of light stress. Thus, the same conversion state of the xanthophyll cycle and the same level of energy dissipation were observed for a given degree of light stress independent of species or conditions causing the light stress. Since all increases in thermal energy dissipation were associated with increases in the levels of zeaxanthin + antheraxanthin in these leaves, there was thus no indication of any form of xanthophyll cycle-independent energy dissipation in any of the twenty-four species or varieties of plants examined in their natural environment. It is also concluded that transient diurnal changes in intrinsic PSII efficiency in nature are caused by changes in the efficiency with which excitation energy is delivered from the antennae to PSII centers, and are thus likely to be purely photoprotective. Consequently, the possibility of quantifying the allocation of absorbed light into PSII photochemistry versus energy dissipation in the antennae from changes in intrinsic PSII efficiency is explored.Abbreviations A antheraxanthin - F actual level of fluorescence - F_a, F _o minimal fluorescence in the absence, presence of thylakoid energization - F_m, F _m maximal fluorescence in the absence, presence of thylakoid energization - F_m, - F)/F _m actual PSII efficiency ( = percent of absorbed light utilized in PSII photochemistry) - F_v/F_m, F _v /F_m/ PSII efficiency of open centers in the absence, presence of thylakoid energization - NPQ nonphotochemical fluorescence quenching - F_m/F _m - 1; q_p quenching coefficient for photochemical quenching - V violaxanthin - Z zeaxanthin     

Mineral nutrient deficiency increases the sensitivity of photosynthesis to sulphur dioxide in needles of a coniferous tree,Abies nordmanniana

Summary Abies nordmanniana (Stev.) Spach was cultivated in rooting media either rich in nutrients (control) or low in magnesium (low Mg) or low in magnesium and nitrogen (low Mg-N), respectively. Intact, attached needles were exposed, in the light (460 mol photons m^-2 s^-1), to an atmosphere containing 1 ppm SO₂ for 5 h. Measurements of light- and CO₂-saturated rates of photosynthetic O₂ evolution, A _max, were performed before and after SO₂ treatments. In needles from well fertilized plants, A _max was high (about 50 mol m^-2 s^-1) and was not affected by SO₂. Needles from low-Mg and low-Mg-N plants had lower photosynthetic rates and showed a marked decline in A _max in response to the SO₂ treatment. Stomatal conductance was similar in the three groups of plants during SO₂ treatments.Abbreviations A _max photosynthetic capacity (CO₂- and light-saturated rate of O₂ evolution) - DW dry weight - F_o yield of dark level fluorescence - F_M maximum yield of fluorescence, induced in a pulse of saturating light - F_v yield of variable fluorescence (= F_M–F_O) - FW fresh weight; g, conductance to water vapor transfer     

Transgenic tobacco with suppressed zeaxanthin formation is susceptible to stress-induced photoinhibition

Tobacco (Nicotiana tabacum cv. Xanthi) transformed with the antisense construct of tobacco violaxanthin de-epoxidase was analyzed for responses in growth chambers to both short and long-term stress treatments. Following a short-term (2 or 3 h) high-light treatment, antisense plants had a greater reduction in F_v/F_m relative to wild-type, indicating a greater susceptibility to photoinhibition. The responses of antisense plants to long-term stress were examined in two separate experiments, one with high light alone and the other wherein high light and water stress were combined. In the light-stress experiment, plants were grown at 1300 mol photons m^–2 s^–1 under a 12 h photoperiod. In the light and water-stress experiment, plants were grown under moderately high light of 900 mol photons m^–2 s^–1, under a 16 h photoperiod, in combination with water stress. Both conditions caused formation of high antheraxanthin and zeaxanthin levels in wild-type plants but not in antisense plants. In both cases, antisense plants showed significant reductions in F_v/F_m and total leaf-pigment content relative to wild-type. The data demonstrate a critical photoprotective function of the xanthophyll cycle-dependent energy dissipation in tobacco exposed suddenly to high amounts of excess light over extended times.This revised version was published online in October 2005 with corrections to the Cover Date.