Variation in photoinhibition among Sasa senanensis, Quercus mongolica, and Acer mono in the understory of a deciduous broad-leaved forest exposed to canopy gaps caused by typhoons

To elucidate mechanisms for tolerating sudden increases in light intensity following canopy gap formation, we investigated susceptibility to photoinhibition in the evergreen clonal plant bamboo, Sasa senanensis, and two deciduous broadleaf woody plants, Quercus mongolica, and Acer mono. We measured pre-dawn photochemical efficiency of photosystem II (F _v /F _m) in plants exposed to canopy gaps and in shade-grown plants through the month following gap formation. Photoinhibition (indicated by decreased F _v /F _m) was smallest in S. senanensis and largest in A. mono. S. senanensis had the highest area-based net CO₂ assimilation rate (A _area) and electron transport rate (ETR) under high light conditions. This species also had the highest leaf mass per area (LMA) and leaf nitrogen content per area (N _area). Higher values of LMA and N _area under shade conditions probably contribute to circumvent photoinhibition through maintenance of a higher ETR capacity. Q. mongolica, a gap-dependent species, had properties intermediate between S. senanensis and A. mono; it appeared less susceptible to photoinhibition than the shade-tolerant A. mono. None of the species examined had increased photosynthetic capacity 1 month after gap formation, indicating that shade-grown leaves were unable to fully acclimate to increased light.     

Interactions between water stress, sun-shade acclimation, heat tolerance and photoinhibition in the sclerophyll Heteromeles arbutifolia

Gas exchange and chlorophyll fluorescence techniques were used to evaluate the acclimation capacity of the schlerophyll shrub Heteromeles arbutifolia M. Roem. to the multiple co-occurring summer stresses of the California chaparral. We examined the influence of water, heat and high light stresses on the carbon gain and survival of sun and shade seedlings via a factorial experiment involving a slow drying cycle applied to plants grown outdoors during the summer. The photochemical efficiency of PSII exhibited a diurnal, transient decrease (δF/F_m′) and a chronic decrease or photoinhibition (F_v/F_m) in plants exposed to full sunlight. Water stress enhanced both transient decreases of δF/F_m’and photoinhibition. Effects of decreased δF/F_m’and F_v/F_m on carbon gain were observed only in well-watered plants since in water-stressed plants they were overidden by stomatal closure. Reductions in photochemical efficiency and stomatal conductance were observed in all plants exposed to full sunlight, even in those that were well-watered. This suggested that H. arbutifolia sacrificed carbon gain for water conservation and photoprotection (both structurally via shoot architecture and physiologically via down-regulation) and that this response was triggered by a hot and dry atmosphere together with high PFD, before severe water, heat or high PFD stresses occur. We found fast adaptive adjustments of the thermal stability of PSII (diurnal changes) and a superimposed long-term acclimation (days to weeks) to high leaf temperatures. Water stress enhanced resistance of PSII to high temperatures both in the dark and over a wide range of PFD. Low PFD protected photochemical activity against inactivation by heat while high PFD exacerbated damage of PSII by heat. The greater interception of radiation by horizontally restrained leaves relative to the steep leaves of sun-acclimated plants caused photoinhibition and increased leaf temperature. When transpirational cooling was decreased by water stress, leaf temperature surpassed the limits of chloroplast thermostability. The remarkable acclimation of water-stressed plants to high leaf temperatures proved insufficient for the semi-natural environmental conditions of the experiment. Summer stresses characteristic of Mediterranean-type climates (high leaf temperatures in particular) are a potential limiting factor for seedling survival in H. arbutifolia, especially for shade seedlings lacking the crucial structural photoprotection provided by steep leaf angles.     

Chronic photoinhibition in seedlings of tropical trees

Seedlings of five canopy species of tropical trees from Costa Rica and Puerto Rico were grown in full shade (midday range of photosynthetic photon flux density [PPFD], 100–140 μmol m^?2 s^?1), partial shade (midday PPFD, 400–600 μmol m^?2 s^?1) and full sun (midday PPFD, 1 500–1 800 μmol m^?2 s^?1) for 3 months. The species were Ochroma lagopus (Bombacaceae), a pioneer species; Inga edulis (Fabaceae), found in secondary forest; and Dipteryx panamensis (Fabaceae), Hampea appendiculata (Malvaceae), and Manilkara bidentata (Sapotaceae), three species characteristic of primary forest. After the plants were placed in the dark overnight, chlorophyll fluorescence characteristics were measured for recently expanded and mature leaves. The ratio of variable fluorescence to maximum fluorescence (F_v/F_m) was used to estimate the degree of chronic photoinhibition. Only individuals of one species, Dipteryx panamensis, showed significant depression of F_v/F_m after long-term exposure to full sun. The depression was highly correlated with quantum yield of O₂ evolution which also declined after exposure to full sun. The decline may have been related to foliar N concentration. Although all plants were supplied with ample nutrients, foliar N did not increase significantly for Dipteryx seedlings in full sun, whereas it did for Ochroma and Inga. Leaf age affected F_v/F_m only in the cases of Manilkara, where it was slightly lower in recently expanded leaves, and of Dipteryx where it interacted with the effects of light regime. We conclude that chronic photoinhibition is not common in seedlings of canopy trees of tropical rain forests except when availability of mineral nutrients may be limiting.     

Photoinhibition of photosynthesis in intact kiwifruit (Actinidia deliciosa) leaves: Effect of light during growth on photoinhibition and recovery

Photoinhibition of photosynthesis was induced in intact kiwifruit (Actinidia deliciosa (A. Chev.) C. F. Liang et A. R. Ferguson) leaves grown at two photon flux densities (PFDs) of 700 and 1300 mol·m^-2·s^-1 in a controlled environment, by exposing the leaves to PFD between 1000 and 2000 mol·m^-2·s^-1 at temperatures between 10 and 25°C; recovery from photoinhibition was followed at the same range of temperatures and at a PFD between 0 and 500 mol·m^-2·s^-1. In either case the time-courses of photoinhibition and recovery were followed by measuring chlorophyll fluorescence at 692 nm and 77K and by measuring the photon yield of photosynthetic O₂ evolution. The initial rate of photoinhibition was lower in the high-light-grown plants but the long-term extent of photoinhibition was not different from that in low-light-grown plants. The rate constants for recovery after photoinhibition for the plants grown at 700 and 1300 mol·m^-2·s^-1 or for those grown in shade were similar, indicating that differences between sun and shade leaves in their susceptibility to photoinhibition could not be accounted for by differences in capacity for recovery during photoinhibition. Recovery following photoinhibition was increasingly suppressed by an increasing PFD above 20 mol·m^-2·s^-1, indicating that recovery in photoinhibitory conditions would, in any case, be very slow. Differences in photosynthetic capacity and in the capacity for dissipation of non-radiative energy seemed more likely to contribute to differences in susceptibility to photoinhibition between sun and shade leaves of kiwifruit.Abbreviations and symbols F _o , F _m , F _v instantaneous, maximum, variable fluorescence - F _v /F _m fluorescence ratio - F _i =F _v at t=0 - F F _v at t= - K _D rate constant for photochemistry - k(F _p ) first-order rate constant for photoinhibition - k(F _r ) first-order rate constant for recovery - PFD photon flux density - PSII photosystem II - _i photon yield of O₂ evolution (incident light)     

ACCLIMATION OF EMILIANIA HUXLEYI (PRYMNESIOPHYCEAE) TO PHOTON FLUX DENSITY1

Growth rate, pigment composition, and noninvasive chl a fluorescence parameters were assessed for a noncalcifying strain of the prymnesiophyte Emiliania huxleyi Lohman grown at 50, 100, 200, and 800 μmol photons·m^?2·s^?1. Emiliania huxleyi grown at high photon flux density (PFD) was characterized by increased specific growth rates, 0.82 d^?1 for high PFD grown cells compared with 0.38 d^?1 for low PFD grown cells, and higher in vivo chl a specific attenuation coefficients that were most likely due to a decreased pigment package, consistent with the observed decrease in cellular photosynthetic pigment content. High PFD growth conditions also induced a 2.5‐fold increase in the pool of the xanthophyll cycle pigments diadinoxanthin and diatoxanthin responsible for dissipation of excess energy. Dark‐adapted maximal photochemical efficiency (F_v/F_m) remained constant at around 0.58 for cells grown over the range of PFDs, and therefore the observed decline, from 0.57 to 0.33, in the PSII maximum efficiency in the light‐adapted state, (F_v′/F_m′), with increasing growth PFD was due to increased dissipation of excess energy, most likely via the xanthophyll cycle and not due to photoinhibition. The PSII operating efficiency (F_q′/F_m′) decreased from 0.48 to 0.21 with increasing growth PFD due to both saturation of photochemistry and an increase in nonphotochemical quenching. The changes in the physiological parameters with growth PFD enable E. huxleyi to maximize rates of photosynthesis under subsaturating conditions and protect the photosynthetic apparatus from excess energy while supporting higher saturating rates of photosynthesis under saturating PFDs.     

High-light effects on CO2 fixation gradients across leaves

Chlorophyll fluorescence and internal patterns of ¹⁴CO₂ fixation were measured in sun and shade leaves of spinach after treatment with various light intensities. When sun leaves were irradiated with 2000μmol m^?2 s^?1 for 2h, F_V/F_M decreased by about 15%, but ¹⁴CO₂ fixation was unaffected, whereas shade leaves exhibited a 21% decrease in F_v/F_M and a 25% decrease in ¹⁴CO₂ fixation. Irradiation of sun and shade leaves with 4000μmol m^?1 for 4 h decreased F_V/F_M by 30% in sun leaves and 40% in shade leaves, while total ¹⁴CO₂ fixation decreased by 41% in sun leaves and 55% in shade leaves. After light treatment, gradients of CO₂ fixation across leaves were determined by measuring ¹⁴CO₂ fixed in paradermal leaf sections after a 10s pulse of ¹⁴CO₂. Gradients of ¹⁴CO₂ fixation in control sun and shade leaves were identified when expressed on a relative basis and normalized for leaf depth. Treatment of leaves with 2000 μmol PAR m^?2 s^?1 for 2h did not after patterns of carbon fixation across sun leaves, but slightly altered the pattern in shade leaves. In contrast, treatment of sun and shade leaves with 4000μmol m^?2 s^?1 for 4h decreased carbon fixation more in the palisade mesophyll cells than in the spongy mesophyll cells of sun and shade leaves, and fixation in medial tissue of shade leaves was dramatically decreased compared to the adaxial and abaxial tissue. The interaction between leaf anatomy and biochemical parameters involved in tolerance to photoinhibition in spinach is discussed.     

Mechanistic differences in photoinhibition of sun and shade plants

Leaf discs of the shade plant Tradescantia albiflora Kunth grown at 50 μmol · m^?2 · s^?1, and the facultative sun/shade plant Pisum sativum L. grown at 50 or 300 μmol · m^?2, s^?1, were photoinhibited for 4 h in 1700 μmol photons m^?2 · s^?1 at 22° C. The effects of photoinhibition on the following parameters were studied: i) photosystem II (PSII) function; ii) amount of D1 protein in the PSII reaction centre; iii) dependence of photoinhibition and its recovery on chloroplast-encoded protein synthesis; and, iv) the sensitivity of photosynthesis to photoinhibition in the presence or absence of the carotenoid zeaxanthin. We show that: i) despite different sensitivities to photoinhibition, photoinhibition in all three plants occurred at the reaction centre of PSII; ii) there was no correlation between the extent of photoinhibition and the degradation of the D1 protein; iii) the susceptibility to photoinhibition by blockage of chloroplas-tencoded protein synthesis was much less in shade plants than in plants acclimated to higher light; and iv) inhibition of zeaxanthin formation increased the sensitivity to photoinhibition in pea, but not in the shade plant Tradescantia. We suggest that there are mechanistic differences in photoinhibition of sun and shade plants. In sun plants, an active repair cycle of PSII replaces photoinhibited reaction centres with photochemically active ones, thereby conferring partial protection against photoinhibition. However, in shade plants, this repair cycle is less important for protection against photoinhibition; instead, photoinhibited PSII reaction centres may confer, as they accumulate, increased protection of the remaining connected, functional PSII centres by controlled, nonphotochemical dissipation of excess excitation energy.     

Diurnal changes in the photochemical efficiency of the symbiotic dinoflagellates (Dinophyceae) of corals: photoprotection, photoinactivation and the relationship to coral bleaching

The photochemical efficiency of symbiotic dinoflagellates within the tissues of two reef‐building corals in response to normal and excess irradiance at water temperatures < 30 °C were investigated using pulse amplitude modulated (PAM) chlorophyll fluorescence techniques. Dark‐adapted F_v/F_m showed clear diurnal changes, decreasing to a low at solar noon and increasing in the afternoon. However, F_v/F_m also drifted downwards at night or in prolonged darkness, and increased rapidly during the early morning twilight. This parameter also increased when the oxygen concentration of the water holding the corals was increased. Such changes have not been described previously, and most probably reflect state transitions associated with PQ pool reduction via chlororespiration. These unusual characteristics may be a feature of an endosymbiotic environment, reflective of the well‐documented night‐time tissue hypoxia that occurs in corals. F_v/F_m decreased to 0·25 in response to full sunlight in shade‐acclimated (shade) colonies of Stylophora pistillata, which is considerably lower than in light‐acclimated (sun) colonies. In sun colonies, the reversible decrease in F_v/F_m was caused by a lowering of F_m and F_o suggesting photoprotection and no lasting damage. The decrease in F_v/F_m, however, was caused by a decrease in F_m and an increase in F_o in shade colonies suggesting photoinactivation and long‐term cumulative photoinhibition. Shade colonies rapidly lost their symbiotic algae (bleached) during exposure to full sunlight. This study is consistent with the hypothesis that excess light leads to chronic damage of symbiotic dinoflagellates and their eventual removal from reef‐building corals. It is significant that this can occur with high light conditions alone.     

Photoinhibition of photosynthesis in intact kiwifruit (Actinidia deliciosa) leaves: effect of growth temperature on photoinhibition and recovery

Intact leaves of kiwifruit (Actinidia deliciosa (A. Chev.) C.F. Liang et A.R. Ferguson) from plants grown in a range of controlled temperatures from 15/10 to 30/25°C were exposed to a photon flux density (PFD) of 1500 μmol·m⁻²·s⁻¹ at leaf temperatures between 10 and 25°C. Photoinhibition and recovery were followed at the same temperatures and at a PFD of 20 μmol·m⁻²·s⁻¹, by measuring chlorophyll fluorescence at 77 K and 692 nm, by measuring the photon yield of photosynthetic O₂ evolution and light-saturated net photosynthetic CO₂ uptake. The growth of plants at low temperatures resulted in chronic photoinhibition as evident from reduced fluorescence and photon yields. However, low-temperature-grown plants apparently had a higher capacity to dissipate excess excitation energy than leaves from plants grown at high temperatures. Induced photoinhibition, from exposure to a PFD above that during growth, was less severe in low-temperature-grown plants, particularly at high exposure temperatures. Net changes in the instantaneous fluorescence,F ₀, indicated that little or no photoinhibition occurred when low-temperature-grown plants were exposed to high-light at high temperatures. In contrast, high-temperature-grown plants were highly susceptible to photoinhibitory damage at all exposure temperatures. These data indicate acclimation in photosynthesis and changes in the capacity to dissipate excess excitation energy occurred in kiwifruit leaves with changes in growth temperature. Both processes contributed to changes in susceptibility to photoinhibition at the different growth temperatures. However, growth temperature also affected the capacity for recovery, with leaves from plants grown at low temperatures having moderate rates of recovery at low temperatures compared with leaves from plants grown at high temperatures which had negligible recovery. This also contributed to the reduced susceptibility to photoinhibition in low-temperature-grown plants. However, extreme photoinhibition resulted in severe reductions in the efficiency and capacity for photosynthesis.     

Photoinhibition of Photosynthesis in Sun and Shade Grown Leaves of Grapevine (Vitis vinifera L.)

The degree of photoinhibition of sun and shade grown leaves of grapevine was determined by means of the ratio of variable to maximum chlorophyll (Chl) fluorescence (F_v/F_m) and electron transport measurements. The potential efficiency of photosystem 2 (PS2), F_v/F_m, markedly declined under high irradiance (HI) in shade leaves with less than 10 % of F₀ level. In contrast, F_v/F_m ratio declined with about 20 % increase of F₀ level in sun leaves. In isolated thylakoids, the rate of whole chain and PS2 activity in HI shade and sun leaves was decreased by about 60 and 40 %, respectively. A smaller inhibition of photosystem 1 (PS1) activity was also observed in both leaf types. In the subsequent dark incubation, fast recovery was observed in both leaf types that reached maximum PS2 efficiencies similar to non-photoinhibited control leaves. The artificial exogenous electron donors DPC, NH₂OH, and Mn²⁺ failed to restore the HI-induced loss of PS2 activity in sun leaves, while DPC and NH₂OH were significantly restored in shade leaves. Hence HI in shade leaves inactivates on the donor side of PS2 whereas it does at the acceptor side in sun leaves, respectively. Quantification of the PS2 reaction centre protein D1 and the 33 kDa protein of water splitting complex following HI-treatment of leaves showed pronounced differences between shade and sun leaves. The marked loss of PS2 activity in HI leaves was due to the marked loss of D1 protein of the PS2 reaction centre protein and the 33 kDa protein of the water splitting complex in sun and shade leaves, respectively.     

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.     

GROWTH AND PHOTOSYNTHESIS OF ULVA ROTUNDATA (CHLOROPHYTA) AS A FUNCTION OF TEMPERATURE AND SQUARE WAVE IRRADIANCE IN INDOOR CULTURE1

Temperature and photon flux density (PFD) vary independently in estuaries, e.g. high PFD may occur at any temperature, so it is necessary to consider synergistic effects of these factors on algal growth. Because natural PFD is highly variable and daylength changes confound seasonal temperature cycles, it is easier to interpret factorial experiments in controlled laboratory conditions. Clonal Ulva rotundata Blid. (Chlorophyta) has been studied extensively in outdoor culture. In this study it was maintained indoors under square wave photoperiods at five PFDs and three temperatures. Growth rate, photqsynthetic light response (P-I) curves, and photosystem II chlorophyll fluorescence properties were measured at the growth temperature following acclimation. Interactions between PFD and growth temperature were strongly indicated in all physiological parameters measured. Greatest PFD response occurred at the highest temperature, and the largest temperature response occurred at the highest PFD. Light-saturated photosynthesis (P_m) dark respiration (R_d), and light-limited quantum yield (Φ_m) were sufficient to describe acclimation status. The light-saturation parameter (I_k) was redundant and potentially misleading. Although U. rotundata exhibits a great amplitude of photoacclimation, it apparently has little capacity for temperature acclimation compared to the kelp, Laminaria saccharina, for which published data indicate similar photosynthetic rates over a broad range of growth temperatures. Diurnal variation of P_m and R_d at a growth PFD of ～ 1700 ± 200 μmol photons · m^?2· s^?1 was similar to the pattern observed previously in outdoor culture, suggesting endogenous control of these parameters. Quantum yield and the ratio of variable to maximum chlorophyll fluorescence (F_v/F_m), which were depressed in midday sunlight exceeding ～ 1500 μmol photons · m^?2· s^?1, were relatively invariant through the day in indoor culture, indicating that these parameters are controlled primarily by instantaneous PFD. Growth and fluorescence data are also presented for some other macroalgae for comparative purposes.     

Effects of photoinhibition on whole-plant carbon gain assessed with a photosynthesis model

A canopy photosynthesis model was modified to assess the effect of photoinhibition on whole‐plant carbon gain. Photoinhibitory changes in maximum quantum yield of photosystem II (F_v/F_m) could be explained solely from a parameter (Lflux) calculated from the light micro‐environment of the leaves. This relationship between F_v/F_m and the intercepted cumulative light dose, integrated and equally weighted over several hours was incorporated into the model. The effect of photoinhibition on net photosynthesis was described through relationships between photoinhibition and the shaping parameters of the photosynthetic light‐response curve (quantum use efficiency, convexity, and maximum capacity). This new aspect of the model was then validated by comparing measured field data (diurnal courses of F_v/F_m) with simulation results. Sensitivity analyses revealed that the extent of photoinhibitory reduction of whole‐plant photosynthesis was strongly dependent on the structural parameters (LAI and leaf angle). Simulations for a Mediterranean evergreen oak, Quercus coccifera, under climatic conditions which cause mild photoinhibition revealed a daily loss of 7·5–8·5% of potential carbon gain in the upper sunlit canopy layers, a 3% loss in the bottom canopy, and an overall loss of 6·1%. Thus, this canopy photoinhibition model (CANO‐PI) allows the quantitative evaluation of photoinhibition effects on primary production.     

PHOTOINHIBITION OF MECHANICALLY STIMULABLE BIOLUMINESCENCE IN THE AUTOTROPHIC DINOFLAGELLATE CERATIUM FUSUS (PYRROPHYTA)1

Ceratium fusus (Ehrenb.) Dujardin was exposed to light of different wavelengths and photon flux densities (PFDs) to examine their effects on mechanically stimulable bioluminescence (MSL). Photoinhibition of MSL was proportional to the logarithm of PFD. Exposure to I μmol photons·m^?2s^?1 of broadband blue light (ca. 400–500 nm) produced near-complete photoinhibition (≥90% reduction in MSL) with a threshold at ca. 0.01 μmol photons·m^?2·s^?1. The threshold of photoinhibition was ca. an order of magnitude greater for both broadband green (ca. 500–580 nm) and red light (ca. 660–700 nm). Exposure to narrow spectral bands (ca. 10 nm half bandwidth) from 400 and 700 nm at a PFD of 0.1 μmol photons·m^?2·s^?1 produced a maximal response of photoinhibition in the blue wavelengths (peak ca. 490 nm). A photoinhibition response (≥ 10%) in the green (ca. 500–540 nm) and red wavelengths (ca. 680 nm) occurred only at higher PFDs (1 and 10 μmol photons·m^?2·s^?1). The spectral response is similar to that reported for Gonyaulax polyedra Stein and Pyrocystis lunula Schütt and unlike that of Alexandrium tamarense (Lebour) Balech et Tangen. The dinoflagellate's own bioluminescence is two orders of magnitude too low to result in self-photoinhibition. The quantitative relationships developed in the laboratory predict photoinhibition of bioluminescence in populations of C. fusus in the North Atlantic Ocean.     

'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.     

Nitrogen supply as a factor influencing photoinhibition and photosynthetic acclimation after transfer of shade-grown Solanum dulcamara to bright light

We have compared the ability of shadegrown clones of Solamum dulcamara L. from shade and sun habitats to acclimate to bright light, as a function of nitrogen nutrition before and after transfer to bright light. Leaves of S. dulcamara grown in the shade with 0.6 mM NO ₃ ^- have similar photosynthetic properties as leaves of plants grown with 12.0 mM NO ₃ ^- . When transferred to bright light for 1–2 d the leaves of these plants show substantial photoinhibition which is characterized by about 50% decrease in apparent quantum yield and a reduction in the rate of photosynthesis in air at light saturation. Photoinhibition of leaf photosynthesis is associated with reduction in the variable component of low-temperature fluorescence emission, and with loss of in-vitro electron transport, especially of photosystem II-dependent processes.We find no evidence for ecotypic differentiation in the potential for photosynthetic acclimation among shade and sun clones of S. dulcamara, or of differentiation with respect to nitrogen requirements for acclimation. Recovery from photoinhibition and subsequent acclimation of photosynthesis to bright light only occurs in leaves of plants provided with 12.0 mM NO ₃ ^- . In these, apparent quantum yield is fully restored after 14 d, and photosynthetic acclimation is shown by an increase in light-saturated photosynthesis in air, of light-and CO₂-saturated photosynthesis, and of the initial slope of the CO₂-response curve. The latter changes are highly correlated with changes in ribulose-bisphosphate-carboxylase activity in vitro. Plants supplied with 0.6 mM NO ₃ ^- show incomplete recovery of apparent quantum yield after 14 d, but CO₂-dependent leaf photosynthetic parameters return to control levels.Symbols and abbreviations F_o initial level of fluorescence at 77 K - F_m maximum level of fluorescence at 77 K - F_v variable components of fluorescence at 77 K (F_v=F_m-F_o) - PSI, PSII photosystem I and II, respectively - RuBP ribulose-1,5-bisphosphate - RuBPCase ribulose-1,5-bisphosphate carboxylase-oxygenase (EC 4.1.1.39)     

Photosynthetic and stomatal responses of Avena sativa (poaceae) to a variable light environment

The net photosynthetic (A), stomatal conductance to water vapor (g), water use efficiency (WUE = A/transpiration), and leaf water potential (ψ) responses of cultivated oats (A vena saliva) were determined under an experimental regime of alternating full sun (photosynthetic photon flux density, PPFD > 1,700 μmol · m^-2-sec^-1) and shade (300-400 μmol · m^-2 · sec^-1 PPFD). Less extensive measurements were made on winter wheat (Triticum aestivum) to test the generality of crop species' responses. The rates of stomatal opening/closing after changes in PPFD in A vena and Triticum were compared with previously determined rates for native grasses and forbs to assess how domestication might have altered stomatal dynamics. Characteristics of Avena under alternating sun and shade were 1) rapid fluctuations in A between full sun (~27 μmol · m^-2 · sec^-1) and shade rates (~12 -13 μmol · m ²sec ¹); 2) slower changes in g, causing progressive stomatal closure during the measurement sequence and possible stomatal limitation of A; 3) no change in leaf ψ; and 4) a net reduction in WUE. Triticum had similar sun/shade A, g, and WUE dynamics, except that Triticum stomata returned to full-sun g between shade periods. The rates of change of g in Avena and Triticum were lower than for some desert and subalpine native species, but were similar to rates for species in adjoining native habitat. The basic stomatal dynamics of Avena and Triticum may typify many cultivated C, species, and these data indicate that crop stomatal behavior has not diverged significantly from that of native species.     

Species-specific acclimation to strong shade modifies susceptibility of conifers to photoinhibition

Photoinhibitory processes in the photosynthetic apparatus of the seedlings of Abies alba (Mill.), Picea abies (Karst.), and Pinus mugo (Turra) growing under strong shade (5 % of full solar irradiance) or full irradiance conditions were investigated in winter and spring using chlorophyll a fluorescence techniques. The extent of photoinhibition in needles as indicated by a decrease in maximum quantum yield of PS II photochemistry (F_v/F_m) depended on species, air temperature and acclimation to the light environment. Unexpectedly, shade-tolerant Abies alba was less affected by low-temperature photoinhibition compared to the other species. F_v/F_m recovered with increasing air temperature. During winter, the seedlings of Picea abies growing in shade showed higher F_v/F_m than those from full light. Non-photochemical quenching of fluorescence (NPQ) measured at the same levels of actinic light was higher in needles acclimated to full light except for Abies alba in February. Photosynthetic performance in term of ETR (apparent electron transfer rate) was also higher in full light-acclimated needles. In April, at ambient temperature, recovery of PS II efficiency from the stress induced by illumination with saturating light was faster in the needles of Picea abies than in those of Abies alba. The shade-acclimated needles of Abies alba and Picea abies showed greater down-regulation of PS II induced by high light stress.     

Photoinhibition of photosynthesis in Minquartia guianensis and Swietenia macrophylla inferred by monitoring the initial fluorescence

We assessed the effect of the exposure to full sunlight (5, 35, and 120 min, i.e. T₅, T₃₅, and T₁₂₀) on fluorescence parameters of two young tropical trees, Swietenia macrophylla, a gap-demanding species, and Minquartia guianensis, a shade tolerant species. Fluorescence parameters (F₀, F_m, F_v/F_m) were recorded before treatments and after the transition to low irradiance (LI). Recovery from photoinhibition (measured as F_v/F_m) was monitored for 24 h at LI. In Swietenia, an almost complete restoration of the F_v/F_m values occurred in T₅ and T₃₅ plants, when a rise in F₀ was observed after the transition to LI. This was inferred as indicative of dynamic photoinhibition. T₁₂₀ led to a decline in F₀ in Minquartia, but not in Swietenia. The plants of both species were unable to recovery from photoinhibition after 24 h at LI, when F₀ declined or remained unchanged. This was interpreted as indicative of chronic photoinhibition. Compared with Swietenia, Minquartia was more susceptible to photoinhibition, as indicated by lower F_v/F_m values.     

High resistance to low-temperature photoinhibition in two alpine, snowbank species

At high elevation, the combination of low temperature and high solar irradiance may be particularly conducive to the low‐temperature photoinhibition of photosynthesis (LTP). Microclimate and photosynthesis were measured in situ in Caltha leptosepala and Erythronium grandiflorum, alpine perennials that may experience even lower temperatures and higher sunlight (PFD, photon flux density) than other alpine plants due to their proximity to snowbanks. Light‐saturated CO₂ assimilation (A_sat) and chlorophyll a fluorescence were measured for naturally frosted and non‐frosted plants that also experienced either natural or experimentally‐lowered sunlight. Following several hours of full sunlight exposure, A_sat in both species was not different in leaves that experienced either frostless nights, shading (ca. 1200 versus 2000 μmol m^?2 s^?1 PFD), or the combination of frostless nights and shade, compared to leaves that experienced frost followed by full sunlight. However, increases in the maximum efficiency of photosystem II photochemistry occurred following either frostless nights (change in F_v/F_m=5.3%; P<0.001), experimental shade (4.0%; P<0.1), or the combination of frostless nights and shade (8.4%; P<0.001) in C. leptosepala. Corresponding increases in F_v/F_m were less in E. grandiflorum (1.8% following frostless nights, P<0.05; 3.7% with shade, P<0.05; and 5.1% with both, P<0.001). Plants of E. grandiflorum in the process of emerging through snowbanks had a 10% increase in F_v/F_m with experimental shading (P<0.05). In both species, depressions in F_v/F_m that resulted from natural frost and high sunlight exposure recovered fully by sunset the same day, and depressed F_v/F_m was associated with greater non‐photochemical quenching. Thus, only slight and reversible LTP was apparent, and both species appeared well‐adapted for maintaining carbon gain on days following frost and high sunlight exposure.