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.     

Sun/shade adaptations of the photosynthetic apparatus of <Emphasis Type="Italic">Hoya carnosa</Emphasis>, an epiphytic CAM vine,in a subtropical rain forest in northeastern Taiwan

Most past work on the ecophysiology of the Crassulacean acid metabolism (CAM) plant, Hoya carnosa, in the lab and in situ in Australia indicates that this epiphytic vine is better adapted to shaded, not exposed, locations, although a recent study of this species in Taiwan presents findings that run counter to this conclusion. Thus, photosynthetic characteristics of shaded and exposed individuals of H. carnosa were compared in situ in a subtropical rain forest in northeastern Taiwan in order to determine whether this CAM epiphyte is better adapted to the shade or the sun. Although leaves of shade plants had much greater chlorophyll concentrations than did those of sun plants, chlorophyll a/b ratios did not differ between the two groups of plants. Fluorescence measurements revealed some ability of leaves to acclimate to both shade and sun, although some evidence for photoinhibition (photoprotection) was observed in more exposed plants. Despite the latter, both exposed and shaded plants exhibited CAM, measured as diel fluctuations in leaf acidity, and CAM was more consistently found in the exposed plants. Furthermore, some evidence for more CAM at higher light availabilities was found. Overall, the results of this investigation reveal that H. carnosa in this subtropical rain forest in Taiwan exhibits adaptations to both high and low light levels, which should prove adaptive for an epiphytic vine with leaves on the same individual exposed to a wide range of exposure and shade in the host tree canopy.     

Photoinhibition and recovery in tropical plant species: response to disturbance

Disturbance or rainforest is often followed by mass mortality of understorey seedlings. Transitions of shade grown plants to full sunlight can cause reductions in the efficiency with which light is used in photosynthesis, called photoinhibition. In order to assess the influence of photoinhibition on mortality and growth after rainforest disturbance this study examined photoinhibition in both simulated and real forest disturbances in northern Papua New Guinea. In an experiment simulating rainforest disturbance, exposure of shade-grown plants to full sunlight resulted in abrupt decreases in the chlorophyll fluorescence parameter F _v/F _m that is characteristic of photoinhibition. However, in the well-watered plants used in these experiments there were no fatalities during 3 weeks after exposure to full sunlight. Thus, it is unlikely that photoinhibition, alone, is responsible for seedling fatalities after rainforest disturbances, but more likely that fatalities are due to photoinhibition in conjunction with other environmental stress. There were differences between the response of species to the simulated disturbance that concurred with their preferred habitats. For example, species form the genus Barringtonia, which is commonly found in shaded understorey environments, underwent greater reductions in F _v/F _m and were slower to recover than species that usually inhabit high solar radiation environments. The extent of photoinhibition and the rate of recovery appeared to be dependent on avoidance of direct solar radiation by altering leaf angles and on increasing maximum photosynthetic rates. A field survey of photoinhibition in man-made rainforest gaps corroborated the findings of the simulated disturbance experiment showing that plant species commonly found in shaded environments showed a greater degree of photoinhibition in forest gaps at midday than those species which are classified as species that benefit from gaps or specialist gap inhabitors.     

Diurnal changes in photochemical efficiency,the reduction state of Q,radiationless energy dissipation,and non-photochemical fluorescence quenching in cacti exposed to natural sunlight in northern Venezuela

Summary Diurnal measurements of chlorophyll a fluorescence from cacti (Nopalea cochenillifera, Opuntia ficus-indica, and Opuntia wentiana) growing in northern Venezuela were used to determine photochemical fluorescence quenching related to the reduction state of the primary electron acceptor of PS II as well as non-photochemical fluorescence quenching which reflects the fraction of energy going primarily into radiationless deexcitation. The cladodes used in this study were oriented such that one surface received direct sunlight in the morning and the other one during the afternoon. Both surfaces exhibited large increases in radiationless energy dissipation from the photochemical system accompanied by decreases in PS II photochemical efficiency during direct exposure to natural sunlight. During exposure to sunlight in the morning, dissipation of absorbed light energy through photosynthesis and radiationless energy dissipation was sufficient to maintain Q, the primary electron acceptor for PS II, in a low reduction state. During exposure to sunlight in the afternoon, however, the reduction state of Q rose to levels greater than 50%, presumably due to a decrease in photosynthetic electron transport as the decarboxylation of the nocturnally accumulated malic acid was completed. Exposure to direct sunlight in the afternoon also led to more sustained increases in radiationless energy dissipation. Furthermore, the increases in radiationless energy dissipation during exposure of a water-stressed cladode of O. wentiana to direct sunlight were much greater than those from other well-watered cacti, presumably due to sustained stomatal closure and decreased rates of photosynthetic electron transport. These results indicate that the radiationless dissipation of absorbed light is an important process in these CAM plants under natural conditions, and may reflect a protective mechanism against the potentially damaging effects of the accumulation of excessive energy, particularly under conditions where CO₂ availability is restricted.Abbreviations CAM crassulacean acid metabolism - F _o instantaneous fluorescence emission - F _M maximum fluorescence emission - F _v variable fluorescence emission - K _D rate constant for radiationless energy dissipation in the antenna chlorophyll - PFD photon flux density - PS I photosystem I - PS II photosystem II - Q primary electron acceptor of photosystem II - q _NP non-photochemical fluorescence quenching - q _P photochemical fluorescence quenching - T _C cladode temperature     

Comparative measurements of chlorophyll a fluorescence, acid accumulation and gas exchange in exposed and shaded plants of Clusia minor L. and Clusia multiflora H. B. K. in the field

 Changes in chlorophyll a fluorescence during the day and diurnal-changes of net CO₂-exchange and organic acid contents were determined in two species of the genus Clusia during the dry season in Venezuela. The investigations included plants of the C₃/CAM intermediate species Clusia minor and the C₃ species C. multiflora growing at exposed and shaded sites. Both species showed a C₃ pattern of net CO₂-exchange at the exposed site. In the shade under extreme drought stress C. minor showed a weak expression of CAM without CO₂-uptake during the afternoon (phase IV of CAM). C. multiflora growing in the shade exhibited a C₃-pattern of net CO₂-exchange and a small but significant nocturnal accumulation of citrate. Shaded plants of C. minor were able to double their light utilisation for electron transport and to reduce non-photochemical quenching during phase III compared to phase II of CAM. Furthermore, increase of electron transport rate through photosystem II in phase III of CAM is correlated to decarboxylation of malate. At the exposed site C. multiflora was less negatively affected by high PPFD than C. minor. This was shown by a lower reduction of potential electron quantum yield (F_v/F_m) and higher light utilisation of electron transport of C. multiflora compared to C. minor. At the exposed site C. minor did not make use of the CAM option to increase light utilisation of electron transport and to reduce non-photochemical quenching as did the plants growing in the shade. Received: 20 March 1996 / Accepted: 24 June 1996     

Photosynthetic characteristics of ornamental passion flowers grown under different light intensities

Responses of leaf gas exchange, fluorescence emission, chlorophyll concentration, and morpho-anatomical features to changes in photosynthetic photon flux density (PPFD) were studied in three wild ornamental species of Passiflora L. to select sun and shade species for landscaping projects. Artificial shade was obtained with different shading nylon nets, under field conditions, which allowed the reduction of 25, 50, and 75% of global radiation, along with a control treatment under full sunlight. For Passiflora morifolia the highest mean values of light-saturated net photosynthetic rate (P _Nmax) and light compensation point (LCP) were observed at 50 and 25% shade, respectively, while the highest values of dark respiration rate (R _D) and apparent quantum yield (α) were observed at 75% shade. For Passiflora suberosa litoralis the highest value of P _max was observed at full sunlight. The highest mean values for P _max, R _D, and LCP for Passiflora palmeri var. sublanceolata were obtained at 25% shade. The highest values of net photosynthetic rate (P _N) for P. morifolia, P. palmeri var. sublanceolata, and P. suberosa litoralis were 21.09, 16.15, and 12.36 μmol(CO₂) m⁻² s⁻¹, observed at 50 and 75% shade and full sunlight, respectively. The values of the minimal chlorophyll fluorescence (F₀) were significantly different in P. suberosa litoralis and P. palmeri var. sublanceolata, increasing with the increase of the irradiance. In contrast, the values of maximum photochemical efficiency of PSII (F_v/F_m) were significantly different only in P. suberosa litoralis, being higher at 75%, progressively reducing with the increase of PPFD levels. The total concentration of chlorophyll (Chl) was higher in shaded plants than in the ones cultivated in full sunlight. On the other hand, the values of Chl a/b ratio were reduced in shaded plants. A significant effect of shade levels on leaf area (LA) and specific leaf area (SLA) was found for the three species, whose highest mean values were observed at 75% shade. The thickness of foliar tissues was significantly higher for the three species at full sunlight and 25% shade. These results suggested that P. morifolia and P. palmeri var. sublanceolata appeared to be adapted to moderate shade conditions. P. suberosa litoralis presented higher plasticity to greater variation of the irradiance levels, while the photoinhibition was one of the limiting factors for this species at full sunlight.     

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

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

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.     

Photosynthetic regulation of C4 desert plant Haloxylon ammodendron under drought stress

About 20-year-old desert plants of C₄ species, Haloxylon ammodendron, growing at the southern edge of the Badain Jaran Desert in China, were selected to study the photosynthetic characteristics and changes in chlorophyll fluorescence when plants were subject to a normal arid environment (AE), moist atmospheric conditions during post-rain (PR), and the artificial supplement of soil water (SW). Results showed that under high radiation, in the AE, the species down-regulated its net assimilation rate (A) and maximum photochemical efficiency of PS II (Fv/Fm), indicating photoinhibition. However, under the PR and SW environments, A was up-regulated, with a unimodal diurnal course of A and a small diurnal change in Fv/Fm, suggesting no photoinhibition. When the air humidity or SW content was increased, the light compensation points were reduced; light saturation points were enhanced; while light saturated rate of CO₂ assimilation (A _max) and apparent quantum yield of CO₂ assimilation (Φ_C) increased. Φ_C was higher while the A _max was reduced under PR relative to the SW treatment. It was concluded that under high-radiation conditions drought stress causes photoinhibition of H. ammodendron. Increasing air humidity or soil moisture content can reduce photoinhibition and increase the efficiency of solar energy use.     

Chilling stress and chilling tolerance of sweet potato as sensed by chlorophyll fluorescence

We studied changes in the chlorophyll (Chl) fluorescence components in chilling-stressed sweet potato (Ipomoea batatas L. Lam) cv. Tainung 57 (TN57, chilling-tolerant) and cv. Tainung 66 (TN66, chilling-susceptible). Plants under 12-h photoperiod and 400 μmol m⁻² s⁻¹ irradiance at 24/20 °C (day/night) were treated by a 5-d chilling period at 7/7 °C. Compared to TN66, TN57 exhibited a significantly greater basic Chl fluorescence (F₀), maximum fluorescence (F_m), maximum fluorescence yield during actinic irradiation (F_m′ ), and the quantum efficiency of electron transport through photosystem 2, PS2 (Φ_PS2). Chilling stress resulted in decrease in the potential efficiency of PS2 (F_v/F_m), Φ_PS2, non-photochemical fluorescence quenching (NPQ), non-photochemical quenching (q_N), and the occurrence of chilling injury in TN66. Chilling increased the likelihood of photoinhibition, characterized by a decline in the Chl fluorescence of both cultivars, and photoinhibition during low temperature stress generally occurred more rapidly in TN66.     

Crassulacean acid metabolism in the shade. Studies on an epiphytic fern,Pyrrosia longifolia,and other rainforest species from Australia

Summary Crassulacean acid metabolism (CAM) was studied in a tropical epiphytic fern, Pyrrosia longifolia, from a fully sun-exposed and from a very shaded site in Northern Queensland, Australia. Measurements of instantaneous net CO₂ exchange showed carbon gain via CO₂ dark fixation with some net CO₂ uptake also occuring during late afternoon, in both sun and shade fronds. Maximum rates of net CO₂ uptake and the nocturnal increase in titratable acidity were lower in shade than in sun fronds. ¹³C values of sun and shade fronds were not significantly different, and ranged between-14 and-15 suggesting that, in the long term, carbon gain was mainly via CO₂ dark fixation. Sun fronds had a higher light compensation point of photosynthesis than shade fronds but the same quantum yield. Yet there was no acclimation of photosynthetic O₂ evolution, (measured at 5% CO₂) in sun and shade fronds and photosynthesis saturated at between 200 and 400 mol quanta m^-2 s^-1. Use of higher light intensities for photosynthesis of sun fronds was probably precluded by low nutrient availability. Total nitrogen was less than 1% of dry weight in fully expanded sun and shade fronds. Exposure of shade fronds to full sunlight for 6 h led to a 60% decline in the quantum yield of photosynthesis and to a decline in variable fluorescence measured at room temperature. Photoinhibition by high light was also observed in Hoya nicholsoniae, a rainforest climber growing in deep shade. This species also exhibited CAM as demonstrated by nocturnal net CO₂ uptake, nocturnal acidification and a ¹³C value of-14. Photosynthetic O₂ evolution in this species was saturated at 2.5% of full sunlight. Two species of Dendrobium (Orchidaceae) from sun-exposed sites, one species exhibiting CAM and the other one exhibiting net CO₂ uptake exclusively during daytime via conventional C₃ photosynthesis, showed similar light response curves and the same quantum yield for photosynthetic O₂ evolution.     

Susceptibility of green leaves and green flower petals of CAM orchid <Emphasis Type="Italic">Dendrobium</Emphasis> cv. Burana Jade to high irradiance under natural tropical conditions

Photosynthetic rates of green leaves (GL) and green flower petals (GFP) of the CAM plant Dendrobium cv. Burana Jade and their sensitivities to different growth irradiances were studied in shade-grown plants over a period of 4 weeks. Maximal photosynthetic O₂ evolution rates and CAM acidities [dawn/dusk fluctuations in titratable acidity] were higher in leaves exposed to intermediate sunlight [a maximal photosynthetic photon flux density (PPFD) of 500–600 μmol m⁻² s⁻¹] than in leaves grown under full sunlight (a maximal PPFD of 1 000–1 200 μmol m⁻² s⁻¹) and shade (a maximal PPFD of 200–250 μmol m⁻² s⁻¹). However, these two parameters of GFP were highest in plants grown under the shade and lowest in full sun-grown plants. Both GL and GFP of plants exposed to full sunlight had lower predawn F_v/F_m [dark adapted ratio of variable to maximal fluorescence (the maximal photosystem 2 yield without actinic irradiation)] than those of shade-grown plants. When exposed to intermediate sunlight, however, there were no significant changes in predawn F_v/F_m in GL whereas a significant decrease in predawn F_v/F_m was found in GFP of the same plant. GFP exposed to full sunlight exhibited a greater decrease in predawn F_v/F_m compared to those exposed to intermediate sunlight. The patterns of changes in total chlorophyll (Chl) content of GL and GFP were similar to those of F_v/F_m. Although midday F_v/F_m fluctuated with prevailing irradiance, changes of midday F_v/F_m after exposure to different growth irradiances were similar to those of predawn F_v/F_m in both GL and GFP. The decreases in predawn and midday F_v/F_m were much more pronounced in GFP than in GL under full sunlight, indicating greater sensitivity in GFP to high irradiance (HI). In the laboratory, electron transport rate and photochemical and non-photochemical quenching of Chl fluorescence were also determined under different irradiances. All results indicated that GFP are more susceptible to HI than GL. Although the GFP of Dendrobium cv. Burana Jade require a lower amount of radiant energy for photosynthesis and this plant is usually grown in the shade, is not necessarily a shade plant.     

Susceptibility to photoinhibition of three deciduous broadleaf tree species with different successional traits raised under various light regimes

The susceptibility to photoinhibition of tree species from three different successional stages were examined using chlorophyll fluorescence and gas exchange techniques. The three deciduous broadleaf tree species were Betula platyphylla var. japonica, pioneer and early successional, Quercus mongolica, intermediate shade‐tolerant and mid‐successional, and Acer mono, shade‐tolerant and late successional. Tree seedlings were raised under three light regimes: full sunlight (open), 10% full sun, and 5% full sun. Susceptibility to photoinhibition was assessed on the basis of the recovery kinetics of the ratio of vaviable to maximum fluorescence (F_v/F_m) of detached leaf discs exposed to about 2000 μmol m^?1 s^?1 photon flux density (PFD) for 2 h under controlled conditions (25 to 28 °C, fully hydrated). Differences in susceptibility to photodamage among species were not significant in the open and 10% full sun treatments. But in 5% full sun, B. platyphylla sustained a significantly greater photodamage than other species, probably associated with having the lowest photosynthetic capacity indicated by light‐saturated photosynthetic rate (B. platyphylla, 9·87, 5·85 and 2·82; Q. mongolica, 8·05, 6·28 and 4·41; A. mono, 7·93, 6·11 and 5·08 μmol CO₂ m^?1 s^?1for open, 10% and 5% full sun, respectively). To simulate a gap formation and assess its complex effects including high temperature and water stress in addition to strong light on the susceptibility to photoinhibition, we examined photoinhibition in the field by means of monitoring ΔF/F′_m on the first day of transfer to natural daylight. Compared with ΔF/F′_m in AM, the lower ΔF/F′_m in PM responding to lower PFD following high PFD around noon indicated that photoinhibition occurred in plants grown in 10 and 5% full sun. The diurnal changes of ΔF/F′_m showed that Q. mongolica grown in 5% full sun was less susceptible to photoinhibition than A. mono although they showed little differences both in photosynthetic capacity in intact leaves and susceptibility to photoinhibition based on leaf disc measurements. These results suggest that shade‐grown Q. mongolica had a higher tolerance for additional stresses such as high temperature and water stress in the field, possibly due to their lower plasticity in leaf anatomy to low light environment.     

'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.     

Photoadaptation, photoinhibition and productivity in the blue-green alga, Spirulina platensis grown outdoors

Two Spirulina platensis strains, SP-G and SP-RB, resistant and sensitive to photoinhibition of photosynthesis, respectively, were grown outdoors in dense cultures and under different photon fluxes provided by shading. Cultures of both strains grown under full sunlight were more resistant to photoinhibition than those grown under nets with 15–50% decreases in the incident photon flux. Cultures grown outdoors were more resistant to photoinhibition than the laboratory ones. At noon, the photosynthetic activity, as expressed by O₂ evolution, was higher for cultures grown under 50% shade, as compared with unshaded cultures. Productivity of the shaded cultures, in terms of biomass produced per day, was always higher when the cultures were protected from photoinhibition.     

The photosynthetic apparatus of C3 and CAM-induced Mesembryanthemum crystallinum L.

Accompanying the CAM induction of Mesembryanthemum crystallinum L. grown in high salinity there are changes in the enzymes of carbon metabolism. However, there are no changes in the electron transport activities, Chla/b ratios or in the distribution of chlorophyll amongst the various pigment-protein complexes of isolated thylakoids. Hence with CAM induction there are no changes in the photochemical apparatus of M. crystallinum thylakoids. Despite comparable amounts of chlorophylla/b-proteins of photosystem II to those found in typical C₃ sun plants, both the C₃ and CAM M. crystallinum chloroplasts have relatively more photosystem II, and, concommitantly, less photosystem I complex. This is consistent with greater fluorescence emission at 685 and 695 nm, and lower emission at 735 nm (measured at 77 K) than typically found for C₃ plants, whether sun or shade species. Photoinhibition of isolated C₃ and CAM thylakoids by white light led to comparable decreases in electron transport capacities and fluorescence emission at 77 K with photosystem II being more affected than PSI. We suggest however, that the presence of more core PSII complexes relative to PSI complexes in this CAM-inducible plant, may provide an additional strategy to mitigate photoinhibition in the short-term.     

Differential responses of antioxidant enzymes in pioneer and late-successional tropical tree species grown under sun and shade conditions

To investigate the ability of pioneer and late-successional species to adapt to a strong light environment in a reforestation area, we examined the activities of antioxidant enzymes in relation to photosystem II, chlorophyll a fluorescence and photosynthetic pigment concentration for eight tropical tree species grown under 100% (sun) and 10% (shade) sunlight irradiation. The pioneer (early-succession) species (PS) were Cecropia pachystachya, Croton urucurana, Croton floribundus and Schinus terebinthifolius. The non-pioneer (late succession) species (LS) were Hymenaea courbaril L. var. stilbocarpa, Esenbeckia leiocarpa, Cariniana legalis and Tabebuia roseo-alba. We observed a greater decline in the ratio of variable to maximum chlorophyll a fluorescence (F_v/F_m) under full sunlight irradiation in the late-successional species than in the pioneer species. The LS species most sensitive to high irradiance were C. legalis and H. courbaril. In LS species, chlorophyll a, chlorophyll b and total chlorophyll concentrations were higher in the shade-grown plants than in plants that developed under full sunlight, but in the PS species C. floribundus and C. pachystachya, we did not observe significant changes in chlorophyll content when grown in the two contrasting environments. The carotenoids/total chlorophyll ratio increased significantly when plants developed under high-sunlight irradiation, but this response was not observed in the PS species S.terebinthifolius and C. pachystachya. The improved performance of the pioneer species in high sunlight was accompanied by an increase in superoxide dismutase (SOD, EC 1.15.1.1) activity, though no light-dependent increase in the activity of ascorbate peroxidase (APX, EC 1.11.1.11) was observed. The activity of catalase (CAT, EC 1.11.1.6) was reduced by high irradiation in both pioneer and late-successional species. Our results show that pioneer species perform better under high-sunlight irradiation than late-successional species, as indicated by increased SOD activity and a higher F_v/F_m ratio. C. legalis was the LS species most susceptible to photoinhibition under full sunlight conditions. These results suggest that pioneer plants have more potential tolerance to photo-oxidative damage than late-successional species associated with the higher SOD activity found in pioneer species. Reduced photoinhibition in pioneer species probably results from their higher photosynthetic capacities, as has been observed in a previous survey carried out by our group.     

Effect of high-light treatments in inducing photoinhibition of photosynthesis in intact leaves of low-light grown Phaseolus vulgaris and Lastreopsis microsora

Photoinhibition studies, using gas-exchange techniques, were conducted with leaflets of Phaseolus vulgaris L. plants that were grown under low photonfluence rates. Comparative measurements were made on attached, intact leaflets and in subsequently isolated chloroplasts. Photoinhibition studies were also conducted with attached fronds of the deep-shade fern Lastreopsis microsora (Endl.) Tindale. Leaflets of lowlight-grown Phaseolus vulgaris and fronds of the shade fern were found to be subject to similar photoinhibition when exposed to photon-fluence rates in excess of those at which they were grown. Photoinhibition following exposure to a photon fluence-rate approximating full sunlight is manifested as a reduction in the capacity for both light-saturated and light-limited carbon uptake and is reflected at the chloroplast level as substantial inhibition of electron flow through photosystem (PS) II, with little effect on PS I. The extent of photoinhibition is markedly dependent on the length of exposure to a high-light regime and on the actual photon-fluence rate maintained during treatment. A greater degree of photoinhibition is evident if carbon metabolism is prevented by the removal of CO₂ than when maximum rates of CO₂ uptake prevail throughout the exposure to a high photonfluence rate. Apparently a certain level of CO₂ turnover is beneficial in providing a sink for photochemically generated energy. When leaf material is exposed to photon-fluence rates well in excess of the rate present during growth apparently the potentials of the various biophysical and photochemical means of dissipating excitation energy are exceeded and photoinhibition of photosynthesis results.Abbreviation PFR photon fluence rate     

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.     

Photosynthetic responses to light in seedlings of selected Amazonian and Australian rainforest tree species

Summary Seedlings of the Caesalpinoids Hymenaea courbaril, H. parvifolia and Copaifera venezuelana, emergent trees of Amazonian rainforest canopies, and of the Araucarian conifers Agathis microstachya and A. robusta, important elements in tropical Australian rainforests, were grown at 6% (shade) and 100% full sunlight (sun) in glasshouses. All species produced more leaves in full sunlight than in shade and leaves of sun plants contained more nitrogen and less chlorophyll per unit leaf area, and had a higher specific leaf weight than leaves of shade plants. The photosynthetic response curves as a function of photon flux density for leaves of shade-grown seedlings showed lower compensation points, higher quantum yields and lower respiration rates per unit leaf area than those of sun-grown seedlings. However, except for A. robusta, photosynthetic acclimation between sun and shade was not observed; the light saturated rates of assimilation were not significantly different. Intercellular CO₂ partial pressure was similar in leaves of sun and shade-grown plants, and assimilation was limited more by intrinsic mesophyll factors than by stomata. Comparison of assimilation as a function of intercellular CO₂ partial pressure in sun- and shade-grown Agathis spp. showed a higher initial slope in leaves of sun plants, which was correlated with higher leaf nitrogen content. Assimilation was reduced at high transpiration rates and substantial photoinhibition was observed when seedlings were transferred from shade to sun. However, after transfer, newly formed leaves in A. robusta showed the same light responses as leaves of sun-grown seedlings. These observations on the limited potential for acclimation to high light in leaves of seedlings of rainforest trees are discussed in relation to regeneration following formation of gaps in the canopy.