Diurnal and seasonal variation in photosynthesis of peach palms grown under subtropical conditions

The Amazonian peach palm (Bactris gasipaes Kunth) has been grown for heart-of-palm production under subtropical conditions. As we did not see any substantial study on its photosynthesis under Amazonian or subtropical conditions, we carried out an investigation on the diurnal and seasonal variations in photosynthesis of peach palms until the first heart-of-palm harvest, considering their relationship with key environmental factors. Spineless peach palms were grown in 80-L plastic pots, under irrigation. Gas exchange and chlorophyll fluorescence emission measurements were taken in late winter, mid spring, mid summer and early autumn, from 7:00 to 18:00 h, with an additional chlorophyll fluorescence measurement at 6:00 h. The highest net CO₂ assimilation (P _N), observed in mid summer, reached about 15 μmol m⁻² s⁻¹, which was about 20% higher than the maximum values found in autumn and spring, and 60% higher than that in winter The same pattern of diurnal course for P _N was observed in all seasons, showing higher values from 8:00 to 9:00 h and declining gradually from 11:00 h toward late afternoon. The diurnal course of stomatal conductance (g _s) followed the same pattern of P _N, with the highest value of 0.6 mol m⁻² s⁻¹ being observed in February and the lowest one (0.23 mol m⁻² s⁻¹) in September. The maximal quantum yield of photosystem II (F_v/F_m) was above 0.75 in the early morning in all the months. The reversible decrease was observed around midday in September and October, suggesting the occurrence of dynamic photoinhibition. A significant negative correlation between the leaf-air vapour pressure difference (VPD_leaf-air) and P _N and a positive correlation between P _N and g _s were observed. The photosynthesis of peach palm was likely modulated mainly by the stomatal control that was quite sensible to atmospheric environmental conditions. Under subtropical conditions, air temperature (T _air) and VPD_leaf-air impose more significant effects over P _N of peach palm than an excessive photosynthetic photon flux density (PPFD). The occurrence of dynamic photoinhibition indicates that under irrigation, peach palms appeared to be acclimated to the full-sunlight conditions under which they have been grown.     

The kinetics of photoinhibition of the photosynthetic apparatus in pea chloroplasts

Abstract The kinetics of a range of chlorophyll fluorescence parameters, non-cyclic electron transport and the capacity of the thylakoids to bind Atrazine were examined during photoinhibition treatment of intact pea chloroplasts. Parameters of fluorescence induction of chloroplasts in the presence and absence of 3-(3,4-dichlorophenyl)-1,1-dimethyl urea at 20 °C and at 77 K were determined. The contributions of photochemical and non-photochemical quenching processes to the loss of fluorescence during photoinhibitory treatment were assessed. Two distinct phases of photoinhibitory damage were observed. During the initial 5 min period of exposure to light the minimal fluorescence level (F_o) increased, whilst the maximal fluorescence level (F_P) decreased, both coupled and uncoupled non-cyclic electron transport to methyl viologen decreased and the ability to bind Atrazine to the thylakoids decreased. Fluorescence analyses demonstrated that during this period thylakoids were becoming increasingly less efficient at generating and maintaining a transmembrane proton electrochemical gradient. Photoinhibitory damage that occurred at later times between 5 and 20 min was of a very different nature. Both F_o and F_P declined, a loss of coupled and uncoupled non-cyclic electron transport was observed together with a loss of the capacity to photo-oxidize water. However, no further loss of Atrazine-binding was associated with such changes. A consistent decrease in the quantum yield of non-cyclic electron transport was also observed throughout photoinhibition treatment. The possibility of two distinct mechanisms of photoinhibitory damage to the photosynthetic apparatus is discussed.     

Genotypic variation within Zea mays for susceptibility to and rate of recovery from chill-induced photoinhibition of photosynthesis

Six genotypes of Zea mays L. were grown in pots inside a glasshouse at a mean temperature of 22±2°C and a minimum photosynthetic photon flux density (Q) during the daylight period of 400 μmol m^?2 s^?1. Chilling-dependent photoinhibition was induced by exposing plants to a temperature of 7°C and a Q of 1 000 μmol m^?2 s^?1 for 6 h. Recovery from photoinhibition was then followed at a temperature of 25°C and a Q of 200 μmol m^?2 s^?1. Leaf gas exchange and chlorophyll fluorescence were measured on attached leaves at room temperature prior to the photoinhibitory treatments and at 6 sampling intervals from 0 to 24 h during the recovery period. The relative water content (RWC) was also measured during the recovery period. The results showed a significant genotypic variation in the susceptibility to and rate of recovery from chilling-dependent photoinhibition of photosynthesis in Zea mays seedlings. The Highland Pool 1a from highland sites in Mexico was the least susceptible to chill-induced photoinhibition, but had the slowest rate of recovery. The hybrid variety LG11 showed the highest rate of recovery, whilst the inbred line ZPF307 was the most susceptible to chill-induced photoinhibition. Susceptibility to photoinhibition and subsequent recovery were at least partially independent, suggesting that selection for improved genotypes will require independent selection for both tolerance and capacity for recovery. Although chlorophyll fluorescence provided a more rapid method of assessing the occurrence of photoinhibition, it was not as effective as direct gas-exchange measurements of the maximum quantum yield of photosynthesis (φ) in separating genotypes with respect to their susceptibility to photoinhibition, especially in the most vulnerable genotypes such as ZPF307. Water stress induced by chilling and high Q treatments appeared to impair the recovery processes. Decreases in stomatal conductance (g_s) produce a significant decrease in intercellular CO₂ concentration (C_i), although this decrease was never so extreme that it limited photosynthetic rates at the light intensities used to determine φ. Nevertheless, closure of stomata in patches, producing local restriction of CO₂ supply, would explain the poor correlation between chlorophyll fluorescence and quantum yield measurements in some genotypes immediately after photoinhibitory treatments.     

Inhibition of photosynthesis in current-year needles of unfertilized and fertilized Norway spruce [Picea abies (L.) Karst.] during autumn and early winter

Inhibition of photosynthesis was followed during autumn and early winter in current-year sun and shade needles of unfertilized and fertilized Norway spruce [Picea abies (L.) Karst.] by simultaneous measurements of photosynthetic O₂ evolution and chlorophyll a fluorescence at 20 °C. The CO₂-saturated rate of O₂ evolution was generally higher in sun needles of fertilized trees than in those of unfertilized trees over a wide range of incident photon flux densities (PFDs). Furthermore, the maximum photo-chemical efficiency of photosystem (PS) II, as indicated by the ratio of variable to maximum fluorescence (F_V/F_M) was generally higher for sun needles of fertilized trees. The depression of f_v/f_m during frost periods was more pronounced in sun needles than in shade needles, indicating that winter inhibition in Norway spruce is strongly light-dependent. However, the inhibition of the rate of O₂ evolution at high PFDs in needles of fertilized trees during early winter was partly independent of the light regime experienced by those needles in the field, which appeared to result in a pronounced decrease in the proportion of oxidized PS II reaction centres in shade needles. A nearly identical linear relationship between the quantum yield of PS II electron transport determined by chlorophyll fluorescence and the quantum yield of O₂ evolution (gross rate of O₂ evolution/PFD) was obtained for the investigated types of needles during autumn and early winter. Except for shade needles of fertilized trees, this appeared to be largely achieved by adjustments in thermal energy dissipation within PS II.     

Photoinhibition in the Antarctic moss Grimmia antarctici Card when exposed to cycles of freezing and thawing

Freezing and thawing of the endemic moss species Grimmia antarctici Card, caused photoinhibition. When snow cover was removed from moss in the field, resulting in exposure to fluctuating temperatures and light conditions, photoinhibition, measured as a reduction in the ratio of variable to maximum chlorophyll a fluorescence (F_v/F_m), was observed. The extent of photoinhibition was highly variable and appeared to be reversible during periods of warmer temperatures. A series of controlled laboratory studies found that the light conditions that prevail between freezing and thawing events influenced the recovery from photoinhibition observed during freezing and thawing, with low light conditions facilitating the greatest rates of recovery. After four cycles of freezing and thawing, recovery from photoinhibition in hydrated moss was achieved within 12 h of transfer to 5°C and 15 μmol quanta m^?2 s^?1. These results favour the hypothesis that photoinhibition observed during freezing represents a protective process involving the down-regulation of photo-system II when photosynthetic carbon assimilation is limited by low temperatures.     

Photoinhibition as a control on photosynthesis and production of Sphagnum mosses

The effect of high light intensity on photosynthesis and growth of Sphagnum moss species from Alaskan arctic tundra was studied under field and laboratory conditions. Field experiments consisted of experimental shading of mosses at sites normally exposed to full ambient irradiance, and removal of the vascular plant canopy from above mosses in tundra water track habitats. Moss growth was then monitored in the experimental plots and in adjacent control areas for 50 days from late June to early August 1988. In shaded plots total moss growth was 2–3 times higher than that measured in control plots, while significant reductions in moss growth were found in canopy removal plots. The possibility that photoinhibition of photosynthesis might occur under high-light conditions and affect growth was studied under controlled laboratory conditions with mosses collected from the arctic study site, as well as from a temperate location in the Sierra Nevada, California. After 2 days of high-light treatment (800 mol photons m^–2 s^–1) in a controlled environmental chamber, moss photosynthetic capacity was significantly lowered in both arctic and temperate samples, and did not recover during the 14-day experimental period. The observed decrease in photosynthetic capacity was correlated (r ²=0.735, P<0.001) with a decrease in the ratio of variable to maximum chlorophyll fluorescence (F _v/F _m) in arctic and temperate mosses. This relationship indicates photoinhibition of photosynthesis in both arctic and temperate mosses at even moderately high light intensities. It is suggested that susceptibility to photoinhibition and failure to photoacclimate to higher light intensities in Sphagnum spp. may be related to low tissue nitrogen levels in these exclusively ombrotrophic plants. Photoinhibition of photosynthesis leading to lowered annual carbon gain in Sphagnum mosses may be an important factor affecting CO₂ flux at the ecosystem level, given the abundance of these plants in Alaskan tussock tundra.     

Photoinhibition at chilling temperature

The effects of moderate light at chilling temperature on the photosynthesis of unhardened (acclimated to +18° C) and hardened (cold-acclimated) spinach (Spinacea oleracea L.) leaves were studied by means of fluorescence-induction measurements at 20° C and 77K and by determination of quantum yield of O₂ evolution. Exposure to 550 mol photons·m^-2·s^-1 at +4° C induced a strong photoinhibition in the unhardened leaves within a few hours. Photoinhibition manifested by a decline in quantum yield was characterized by an increase in initial fluorescence (F _o) and a decrease in variable fluorescence (F _v) and in the ratio of variable to maximum fluorescence (F _V/F _M), both at 77K and 20° C. The decline in quantum yield was more closely related to the decrease in the F _V/F _M ratio measured at 20° C, as compared with F _V/F _M at 77K. Quenching of the variable fluorescence of photosystem II was accompanied by a decline in photosystem-I fluorescence at 77K, indicating increased thermal de-excitation of pigments as the main consequence of the light treatment. All these changes detected in fluorescence parameters as well as in the quantum yield of O₂ evolution were fully reversible within 1–3 h at a higher temperature in low light. The fast recovery led us to the view that this photoinhibition represents a regulatory mechanism protecting the photosynthetic apparatus from the adverse effects of excess light by increasing thermal energy dissipation. Long-term cold acclimation probably enforces other protective mechanisms, as the hardened leaves were insensitive to the same light treatment that induced strong inhibition of photosynthesis in unhardened leaves.Abbreviations F ₀ initial fluorescence - F _M maximum fluorescence - F _V variable fluorescence (F _M-F ₀ - PFD photon flux density - PS photosystem     

Down regulation of photosynthesis in Artabotrys hexapetatus by high light

Using variable to maximum fluorescence (F_v/F_m) as the criterion, the down regulation of photosynthesis by high light stress was characterized in the detached leaves of Artabotrys hexapetatus. The decrease in F_v/F_m was corelated with the decrease in oxygen evolution by thylakoids isolated from high light exposed leaves. The decrease in F_v/F_m was linear with increasing time of exposure to high light. A comparison of recovery measured as F_v/F_m, in low light versus dark, revealed that the recovery in darkness was as significant as in low light. Since the relaxation of fluorescence was a rapid response after exposure to high light and the fact that the recovery occurs in total darkness, it is concluded that photoinhibition and down regulation of photosynthesis by high light are independent events.Abbreviation Fpl- initial plateau - F_m- maximum fluorescence - F_o- prompt fluorescence - F_v- variable fluorescence - PFD- photon flux density - PS I (II)- Photosystem I (II)     

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.     

Photosystem 2 is more tolerant to high temperature in apple (<Emphasis Type="Italic">Malus domestica</Emphasis> Borkh.) leaves than in fruit peel

Tolerance of photosystem 2 (PS2) to high temperature in apple (Malus domestica Borkh. cv. Cortland) leaves and peel was investigated by chlorophyll a fluorescence (OJIP) transient after exposure to 25 (control), 40, 42, 44, and 46 °C in the dark for 30 min. The positive L-step was more pronounced in a peel than in leaves when exposed to 44 °C. Heat-induced K-step became less pronounced in leaves than in peel when exposed to 42 °C or higher temperature. Leaves had negative L-and K-steps relative to the peel. The decrease of oxygen-evolving complex (OEC) by heat stress was higher in the peel than in the leaves. OJIP transient from the 46 °C treated peel could not reach the maximum fluorescence (F_m). The striking thermoeffect was the big decrease in the relative variable fluorescence at 30 ms (V_I), especially in the leaves. Compared with the peel, the leaves had less decreased maximum PS2 quantum efficiency (F_v/F_m), photochemical rate constant (K_P), F_m and performance index (PI) on absorption basis (PI_abs) and less increased minimum fluorescence (F₀) and non-photochemical rate constant (K_N), but more increased reduction of end acceptors at PS1 electron acceptor side per cross section (RE₀/CS₀) and per reaction center (RE₀/RC₀), quantum yield of electron transport from Q_A ⁻ to the end acceptors (ϕ _R0) and total PI (PI_abs,total) when exposed to 44 °C. In conclusion, PS2 is more thermally labile than PS1. The reduction of PS2 activity by heat stress primarily results from an inactivation of OEC. PS2 was more tolerant to high temperature in the leaves than in the peel.     

Photoinhibition of photosynthesis in intact kiwifruit (Actinidia deliciosa) leaves: Recovery and its dependence on temperature

Recovery of photoinhibition in intact leaves of shade-grown kiwifruit was followed at temperatures between 10° and 35° C. Photoinhibition was initially induced by exposing the leaves for 240 min to a photon flux density (PFD) of 1 500 mol·m^-2·s^-1 at 20° C. In additional experiments to determine the effect of extent of photoinhibition on recovery, this period of exposure was varied between 90 and 400 min. The kinetics of recovery were followed by chlorophyll fluorescence at 77K. Recovery was rapid at temperatures of 25–35° and slow or negligible below 20° C. The results reinforce those from earlier studies that indicate chilling-sensitive species are particularly susceptible to photoinhibition at low temperatures because of the low rates of recovery. At all temperatures above 15° C, recovery followed pseudo first-order kinetics. The extent of photoinhibition affected the rate constant for recovery which declined in a linear fashion at all temperatures with increased photoinhibition. However, the extent of photoinhibition had little effect on the temperature-dependency of recovery. An analysis of the fluorescence characteristics indicated that a reduction in non-radiative energy dissipation and repair of damaged reaction centres contributed about equally to the apparent recovery though biochemical studies are needed to confirm this. From an interpretation of the kinetics of photoinhibition, we suggest that recovery occurring during photoinhibition is limited by factors different from those that affect post-photoinhibition recovery.Abbreviations and symbols 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, transfer to photosystem I - K(PI), k(R) rate constants for photoinhibition and recovery - PFD photon flux density - PSI, II photosystem I, II - _i photon yield of photosynthesis (incident light)     

Sensitivity of Photosystem 2 of Antarctic Lichens to High Irradiance Stress: Fluorometric Study of Fruticose (Usnea antarctica) and Foliose (Umbilicaria decussata) Species

Two lichen species collected in maritime Antarctica (King George Island) were exposed under laboratory conditions to excess irradiance to evaluate the response of photosystem 2 (PS2). The response was measured on fully hydrated lichen thalli at 5 °C by means of a modulated fluorometer using chlorophyll (Chl) fluorescence induction curve supplemented with analysis of quenching mechanisms. Chl fluorescence parameters [i.e. ratio of variable to maximum Chl fluorescence (F_V/F_M), quantum yield of PS2 photochemical reactions (₂), quenching coefficients] were evaluated before and several times after exposition to high irradiance in order to characterise the extent of photoinhibition, fast and slow phase of recovery. Strong irradiance (2 000 mol m^–2 s^–1) caused high degree of photoinhibition, particularly higher in fruticose (Usnea antarctica) than in foliose (Umbilicaria decussata) lichen species. Fast phase of recovery from photoinhibition, corresponding to regulatory mechanisms of PS2, was more apparent in U. decussata and ₂ than in U. antarctica and F_V/F_M and ₂ within 40 min after photoinhibitory treatment. It was followed by a slow phase lasting several hours, corresponding to repair and re-synthesis processes. After photoinhibitory treatment, recovery of non-photochemical quenching (NPQ) was faster and more pronounced in U. decussata than in U. antarctica. Significant differences were found between the two species in the rate of recovery in fast-(q_E) and slow-recovering (q_T+I) component of NPQ.     

Reversible photoinhibition of unhardened and cold-acclimated spinach leaves at chilling temperatures

The photoinhibition of photosynthesis at chilling temperatures was investigated in cold-acclimated and unhardened (acclimated to +18° C) spinach (Spinacia oleracea L.) leaves. In unhardened leaves, reversible photoinhibition caused by exposure to moderate light at +4° C was based on reduced activity of photosystem (PS) II. This is shown by determination of quantum yield and capacity of electron transport in thylakoids isolated subsequent to photoinhibition and recovery treatments. The activity of PSII declined to approximately the same extent as the quantum yield of photosynthesis of photoinhibited leaves whereas PSI activity was only marginally affected. Leaves from plants acclimated to cold either in the field or in a growth chamber (+1° C), were considerably less susceptible to the light treatment. Only relatively high light levels led to photoinhibition, characterized by quenching of variable chlorophyll a fluorescence (F_V) and slight inhibition of PSII-driven electron transport. Fluorescence data obtained at 77 K indicated that the photoinhibition of cold-acclimated leaves (like that of the unhardened ones) was related to increased thermal energy dissipation. But in contrast to the unhardened leaves, 77 K fluorescence of cold-acclimated leaves did not reveal a relative increase of PSI excitation. High-light-treated, cold-acclimated leaves showed increased rates of dark respiration and a higher light compensation point. The photoinhibitory fluorescence quenching was fully reversible in low light levels both at +18° C and +4° C; the recovery was much faster than in unhardened leaves. Reversible photoinhibition is discussed as a protective mechanism against excess light based on transformation of PSII reaction centers to fluorescence quenchers.Abbreviations F_O initial fluorescence - F_M maximal fluorescence - F_V devariable fluorescence (f_m-f_o) - PFD photon flux density - PS photosystem - SD standard deviation The authors thank the Deutsche Forschungsgemeinschaft and the Academy of Finland for financial support.     

Changes in photosynthesis and fluorescence in response to flooding in emerged and submerged leaves of Pouteria orinocoensis

In the seasonally flooded forest of the Mapire River, a tributary of the Orinoco, seedlings remain totally covered by flood water for over six months. In order to characterize the physiological response to flooding and submergence, seedlings of the tree Pouteria orinocoensis, an important component of the forest vegetation, were subjected experimentally to flooding. Flooding was imposed gradually, the maximum level of flood including submerged and emerged leaves. After 45 d a severe reduction of net photosynthetic rate (P _N) and stomatal conductance (g _s) was observed in emerged leaves, whereas leaf water potential remained constant. The decrease in P _N of emerged leaves was associated to an increase in both relative stomatal and non-stomatal limitations, and the maintenance of the internal/air CO₂ concentration (C _i/C _a) for at least 20 d of flooding. After this time, both P _N and g _s became almost zero. The decrease in photosynthetic capacity of emerged leaves with flooding was also evidenced by a decrease in carboxylation efficiency; photon-saturated photosynthetic rate, and apparent quantum yield of CO₂ fixation. Oxygen evolution rate of submerged leaves measured after three days of treatment was 7 % of the photosynthetic rate of emerged leaves. Submersion determined a chronic photoinhibition of leaves, viewed as a reduction in maximum quantum yield in dark-adapted leaves, whereas the chlorophyll fluorescence analysis of emerged leaves pointed out at the occurrence of dynamic, rather than chronic, photoinhibition. This was evidenced by the absence of photochemical damage, i.e. the maintenance of maximum quantum yield in dark-adapted leaves. Nevertheless, the observed lack of complementarity between photochemical and non-photochemical quenching after 12 d of flooding implies that the capacity for photochemical quenching decreased in a non-co-ordinate manner with the increase in non-photochemical quenching.     

Estimation of the effect of photoinhibition on the carbon gain in leaves of a willow canopy

The occurrence of photoinhibition of photosynthesis in leaves of a willow canopy was examined by measuring the chlorophyll-a fluorescence ratio of F _V/F _M (F_M is the maximum fluorescence level of the induction curve, and F_V is the variable fluorescence, F _V=F _M–F ₀, where F₀ is the minimal fluorescence). The majority of the leaves situated on the upper parts of peripheral shoots showed an afternoon inhibition of this ratio on clear days. This was the consequence of both a decrease in F _M and a rise in F _O. In the same leaves the diurnal variation in intercepted photosynthetic photon flux density (PPFD) was monitored using leaf-mounted sensors. Using the multivariate method, partial least squares in latent variables, it is shown that the dose of PPFD, integrated and linearly weighted over the last 6-h period, best predicts photoinhibition. Photoinhibition occurred even among leaves that did not intercept PPFDs above 1000 mol·m^–2·s^–1. Exposure of leaves to a standard photoinhibitory treatment demonstrated that the depression in the F _V/F _M ratio was paralleled by an equal depression in the maximal quantum yield of CO₂ uptake and a nearly equal depression in the rate of bending (convexity) of the light-response curve of CO₂ uptake. As a result, the rate of net photosynthesis is depressed over the whole natural range of PPFD. By simulating the daily course in the rate of net photosynthesis, it is estimated that in the order of one-tenth of the potential carbon gain of peripheral willow shoots is lost on clear days as a result of photoinhibition. This applies to conditions of optimal temperatures. Photoinhibition is even more pronounced at air temperatures below 23° C, as judged from measurements of the F_V/F_M ratio on clear days: the afternoon inhibition of this ratio increased in a curvilinear manner from 15% to 25% with a temperature decrease from 23° to 14° C.Abbreviations and Symbols F_O minimum fluorescence - F_V variable fluorescence - F_M maximum fluorescence - PLS partial least squares in latent variables - PPFD photosynthetic photon flux density - VPD water vapour-pressure deficit This study was supported by the Swedish Natural Science Research Council. We are indebted to Dr. Jerry Leverenz (Department of Plant Physiology, University of Umeå, Sweden) for guidance with the modelling of the photosynthesis data.     

Seasonal Effects on Photosynthetic Electron Transport and Fluorescence Properties in Isolated Chloroplasts of Pinus silvestris

Chloroplasts were isolated from primary needles of 1-year-old seedlings and from secondary needles of a 20-year-old pine tree in a natural stand. In autumn the electron transport capacities of PSII, PSI and PS (II + I) decreased and the electron transport between PSII and PSI became inhibited in October in the 20-year-old tree. This inhibition lasted until May the following year. The partial reactions of PSI and PSII still showed low but fairly constant rates during the whole winter seedlings. Seasonal changes in the electron transport properties of 1-year-old showed the same general trends as observed in the 20-year-old tree, but the changes were less pronounced. However, in snow-covered seedlings the PSI-mediated electron transport and the electron transport from H₂O to NADP increased during the late winter when the seedlings were still covered by snow. The total chlorophyll content of the needles decreased in autumn and winter. Low temperature fluorescence ratios of F692/F680 and F726/F680 indicated more severe destruction of the chlorophyll a antennae closely associated with the two photosystems than of the light harvesting chlorophyll a/b complex. In this case, too, the changes were more pronounced in the 20-year-old tree than in the 1-year-old seedlings. The chlorophyll/P700 ratios indicated a more marked reduction in the reaction centre molecules during autumn than in the antennae chlorophyll molecules. The changes in electron transport and low temperature fluorescence properties which occurred during autumn and winter were mainly reversed during spring.     

Performance of active Photosystem II centers in photoinhibited pea leaves

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

Development of photoautotrophy and photoinhibition of Gardenia jasminoides plantlets during micropropagation

This paper reports on the fast fluorescence responses of Gardenia jasminoides Ellis plantlets, at two successive stages (shoot multiplication and root induction) of culture in vitro. We test whether plantlets in vitro suffer photoinhibition during culture and whether the degree of photoautotrophy of these mixotrophic plantlets has any effect on the extent of photoinhibitory impairment. In this regard the effects of different sucrose levels in the medium and PPFD during growth on the development of photoautotrophy and the extent of photoinhibition were evaluated. Plantlets were grown under low, intermediate, and high (50, 100, and 300 mol m^-2 s^-1) PPFD, and at 3 different sucrose concentrations (0.5, 1.5, and 3.0%, w/v) in the medium, during shoot multiplication. During root induction the same growth conditions were assayed except for the high PPFD. The development of photoautotrophy was assessed via the difference between the stable carbon isotope composition of sucrose used as heterotrophic carbon source and that of leaflets grown in vitro. Plantlets from root induction showed more developed photoautotrophy than those from shoot multiplication. For both stages the low-sucrose medium stimulated the photoautotrophy of plantlets in vitro. In addition, intermediate PPFD induced photoautotrophy during shoot multiplication. For plantlets of both culture stages at the lowest PPFD no photoinhibition occurred irrespective of the sucrose concentration in media. However, during the shoot multiplication stage chlorophyll fluorescence measurements showed a decrease in F _v /F _m and in t _1/2 as growing PPFD increased, indicating photoinhibitory damage. The decline of F _v /F _m was caused mostly by an increase in F _o , indicating the inactivation of PSII reaction centers. However plantlets growing under low sucrose showed reduced susceptibility to photoinhibition. During root induction, only plantlets cultured with high sucrose showed a decrease in F _v /F _m as PPFD increased, although t _1/2 remained unchanged. In this case, the decline of F _v /F _m was mostly due to a decrease in F _m , which indicates increased photoprotection rather than occurrence of photodamage. Therefore, growth in low-sucrose media had a protective effect on the resistance of PSII to light stress. In addition, plantlets were more resistant to photoinhibition during root induction than during shoot multiplication. Results suggest that increased photoautotrophy of plantlets reduces susceptibility to photoinhibition during gardenia culture in vitro.Abbreviations AP apparent photosynthesis - Chl total chlorophyll content - Chl a/b chlorophyll a-to-b ratio - Chl/Car total chlorophyll-to-carotenoids ratio - ¹³C ratio of ¹³C/¹²C relative to PeeDee belemnite standard - F _m maximum chlorophyll fluorescence - F _o fluorescence emission when all reaction centres are open and the photochemical quenching is minimal - F _v variable chlorophyll fluorescence (F _m -F _o ) - F _v /F _m the ratio of variable to maximum chlorophyll fluorescence, indicator photochemical efficiency of PSII - MS medium Murashige and Skoog (1962) medium - PPFD photosynthetic photon flux density - Rd dark respiration, t _1/2 the half-time of the increase from F _o to F _m - IAA indole butyric acid     

Gap size effects on photoinhibition in understorey saplings in tropical rainforest

The sudden increase in irradiance after canopy disturbance in primary forest together with the accompanying increase in leaf temperatures is known to cause photoinhibition in shade acclimated foliage of understorey plants. We hypothesized that there is species specific variation among understorey saplings in the magnitude of photoinhibition in response to gap creation, which is related to their requirement for overstorey disturbance. Eleven more or less circular gaps were created varying in size from 60 up to 1459 m². Photoinhibition was assessed by determining predawn and midday F_v/F_m using chlorophyll fluorescence at two occasions during the first 3 weeks after creation of the gaps. The light environment was assessed using hemispherical photography. Five species that occurred in sufficient numbers in the understorey after gap creation were measured. They all showed an increase of photoinhibition with increasing gap size. Variation in exposure to direct sunlight within gaps contributed also to variation in photoinhibition. Dynamic photoinhibition, the overnight increase in F_v/F_m, was about 20% of total photoinhibition as measured at midday. The species responded quantitatively different. Oxandra asbeckii was most sensitive as evident from a decrease of predawn F_v/F_m from 0.79 in the understorey of undisturbed forest to 0.70 in the smallest and further to 0.41 in the largest gaps. Catostemma fragrans, the least sensitive species showed hardly any photoinhibition in the smallest gaps and less in the largest ones, whereas Lecythis concertiflora, Licania heteromorpha, and Chlorocardium rodiei had intermediate responses. Species rank order in sensitivity to photoinhibition was maintained across the whole range of gap sizes. The relationship between sensitivity to photoinhibition and species-specific gap size preference for regeneration is discussed.     

Photoinhibition of photosynthesis on a coral reef

Photoinhibition of macroalgae in the epilithic algal community (KAC) of coral reefs was studied using chlorophyll fluorescence techniques at One Tree Island, Great Barrier Reef, Australia. F_v/F_m (variable to maximum fluorescence, darkened samples) of shallow macroalgae declined by 50% on fine summer and winter days, recovering in late afternoon. Within a species, thalli from low-light habitats were more photoinhibited (2h at 1400μimol m^?2 s^?1) than those from high-light habitats. The sensitivity of Lobophora variegata (Phacophyta) and Chlorodesmis fastigiata (Chlorophyta) increased with depth (1 versus 20 m). However, shallow Halimeda tuna (Chlorophyta) plants growing between corals were more photoinhibited than those from deep, open areas. Photoinhibition and recovery were depth- and species-specific. Shallow Lobophora and Chlorodesmis maintained a greater degree of Q _A oxidation during photoinhibition. In deep thalli, reduced effective quantum yield of open photosystem II centres reflected lower proportions and excitation capture efficiencies of open centres. In Lobophora, zeaxanthin formation accompanied non-photochemical fluorescence quenching (NPQ), but in Chlorodesmis NPQ was limited and no zeaxanthin or antherxanthin formed. Higher photosynthetic efficiency in the lower storey of the EAC may compensate for photoinhibition in the upper storey, thereby reconciling photoinhibition of individual thalli with previous observations of no net inhibition of community productivity.