A mechanistic model of photoinhibition

A mechanistic model was developed, to simulate the main facets of photoinhibition in phytoplankton. Photoinhibition is modelled as a time dependent decrease in the initial slope of a photosynthesis versus irradiance curve, related to D1 (photosystem II reaction centre protein) damage and non-photochemical quenching. The photoinhibition model was incorporated into an existing ammonium-nitrate nutrition interaction model capable of simulating photoacclimation and aspects of nitrogen uptake and utilization. Hence the current model can simulate the effects of irradiance on photosynthesis from sub-saturating to inhibitory photon flux densities, during growth on different nitrogen sources and under nutrient stress. Model output conforms well to experimental data, allowing the extent of photoinhibition to be predicted under a range of nutrient and light regimes. The ability of the model to recreate the afternoon depression of photosynthesis and the enhancement of photosynthesis during fluctuating light suggests that these two processes are related to photoinhibition. The model may be used to predict changes in biomass and/or carbon fixation under a wide range of oceanographic situations, and it may also help to explain the progression to dominance of certain algal species, and bloom formation under defined irradiance and nutrient conditions.     

喜光榕树和耐荫榕树光适应机制的差异

100%和36%光强下生长的喜光的斜叶榕的光合能力高于耐荫的假斜叶榕,而热耗散能力与之相似,说明强光下斜叶榕主要通过光合作用利用光能和热耗散、假斜叶榕主要通过热耗散防御光破坏.100%光强下生长的两种榕树的日间光抑制程度相似,但叶表光强相同情况下各光强下生长的假斜叶榕的光抑制均比斜叶榕严重.100%光强下假斜叶榕叶片悬挂角大于斜叶榕,导致日间叶表光强低于斜叶榕,这可能是两种榕树日间光抑制程度相似的原因,表明叶片悬挂角的适应变化对假斜叶榕有重要的意义.     

CARBON UPTAKE IN A MARINE DIATOM DURING ACUTE EXPOSURE TO ULTRAVIOLET B RADIATION: RELATIVE IMPORTANCE OF DAMAGE AND REPAIR1

Experiments on a marine diatom, Thalassiosira pseudonana (Hustedt) clone 3H, demonstrate that under moderate photon flux densities (75 μmol quanta·m^?2·s^?1) of visible light the inhibition of photosynthesis by supplemental ultraviolet (UV) radiation (UV-B: 280–320 nm) is well described as a hyperbolic function of UV-B irradiance for time scales of 0.5–4 h. Results are consistent with predictions of a recently developed model of photosynthesis under the influence of UV and visible irradiance. Although net destruction of chlorophyll occurs during a 4-h exposure to UV-B, and the effect is a function of exposure, the principal effect of UV-B is a decrease in chlorophyll-specific photosynthetic rate. The dependence of photoinhibition on dosage rate, rather than cumulative dose, and the hyperbolic shape of the relationship are consistent with net photoinhibition being an equilibrium between damage and repair. The ratio of damage to repair is estimated by a mathematical analysis of the inhibition of photosynthesis during exposures to UV-B. A nitrate-limited culture was much more sensitive to UV-B than were the nutrient-replete cultures, but the kinetics of photoinhibition were similar. The analysis suggests that the nutrient-limited culture was more sensitive than the nutrient-replete cultures because repair or turnover of critical proteins associated with photosynthesis is inhibited. An inhibitor of chloroplast protein synthesis was used to suppress repair processes. Photoinhibition by UV-B was enhanced, and inhibition was a function of cumulative dose, as would be expected if damage were not countered by repair. The fundamental importance of repair processes should be considered in the design of field experiments and models of UV-B effects in the environment, especially in the context of vertical mixing. Repair processes must also be considered whenever biological weighting functions are developed.     

Dynamic discrete model of flashing light effect in photosynthesis of microalgae

For a photobioreactor for mass-culturing microalgae, it is known that flashing light effect enhances the efficiency of photosynthesis. A dynamic model for photosynthesis was developed to elucidate this effect. A particular feature of the model is that discrete RuBP particles circulate in the Calvin cycle and their speeds in the cycle are determined by the amount of ATP generated in the photon reception process. This can realise the light saturation under continuous light and the flashing light effect under fluctuating illumination. Laboratory experiments were conducted to obtain model parameters by curve-fitting for Chaetoceros calcitrans. The present model demonstrates the light flashing effect moderately well and elucidates its mechanism reasonably.     

SEASONAL VARIATIONS IN PHOTOSYNTHESIS-IRRADIANCE RESPONSES BY BIOFILMS IN MEDITERRANEAN STREAMS

The relationships between primary production and irradiance were analyzed over an annual cycle in natural biofilms of two undisturbed streams: La Solana (LS), an open calcareous stream, and Riera Major (RM), a shaded siliceous stream. In LS, low photosynthetic efficiency (α^chl and α^area) and high values of both the light saturation parameter (I_k) and the carotenoid / chlorophyll ratio indicated adaptation to high light regimes. On the other hand, higher photosynthetic efficiency and lower I_k as well as photoinhibition at high irradiance found in the biofilms of RM indicated shade adaptation. However, the lack of correlation between light availability in nature and the photosynthetic parameters studied in the laboratory suggested that light was not the most important factor in determining seasonal changes in the photosynthetic behavior in this stream. Both in the open and shaded streams, algal patches collected simultaneously exhibited different photosynthesis-irradiance (P-I) curues, showing that community composition influenced the P-I parameters. In the open stream (LS), however, significant negative correlations between α^area and chlorophyll a and between P _max^chl and chlorophyll a ( r = -0.994 , P < 0.001, and r = -0.929 , P < 0.05, respectively) suggested that photosynthesis was affected by self-shading. Due to the absence of photoinhibition in the biofilms of LS, high photosynthetic rates were maintained at the ambient high light environment, thus compensating for low photosynthesis at low irradiance. In the shaded stream (RM), because photosynthesis was saturated at low irradiances, primary production was relatively high given the low light conditions .     

Activation of photosynthesis and resistance to photoinhibition in cyanobacteria within biological desert crust

Filamentous cyanobacteria are the main primary producers in biological desert sand crusts. The cells are exposed to extreme environmental conditions including temperature, light, and diurnal desiccation/rehydration cycles. We have studied the kinetics of activation of photosynthesis during rehydration of the cyanobacteria, primarily Microcoleus sp., within crust samples collected in the Negev desert, Israel. We also investigated their susceptibility to photoinhibition. Activation of the photosynthetic apparatus, measured by fluorescence kinetics, thermoluminescence, and low temperature fluorescence emission spectra, did not require de novo protein synthesis. Over 50% of the photosystem II (PSII) activity, assembled phycobilisomes, and photosystem I (PSI) antennae were detected within less than 5 min of rehydration. Energy transfer to PSII and PSI by the respective antennae was fully established within 10 to 20 min of rehydration. The activation of a fraction of PSII population (about 20%-30%) was light and temperature-dependent but did not require electron flow to plastoquinone [was not inhibited by 3-(3,4-dichlorophenyl)-1,1-dimethylurea]. The cyanobacteria within the crusts are remarkably resistant to photoinhibition even in the absence of protein synthesis. The rate of PSII repair increased with light intensity and with time of exposure. Consequently, the extent of photoinhibition in high-light-exposed crusts reached a constant, relatively low, level. This is in contrast to model organisms such as Synechocystis sp. strain PCC 6803 where PSII activity declined continuously over the entire exposure to high illumination. Ability of the crust's organisms to rapidly activate photosynthesis upon rehydration and withstand photoinhibition under high light intensity may partly explain their ability to survive in this ecosystem.     

A cell-based model for the photoacclimation and CO(2)-acclimation of the photosynthetic apparatus

We have developed a mathematical model based on the underlying mechanisms concerning the responses of the photosynthetic apparatus of a microalga cell which grows under constant incident light intensity and ambient CO(2) concentration. Photosynthesis involves light and carbon-fixation reactions which are mutually dependent and affect each other, but existing models for photosynthesis don't account for both reactions at once. Our modeling approach allows us to derive distinct equations for the rates of oxygen production, NADPH production, carbon dioxide fixation, carbohydrate production, and rejected energy, which are generally different. The production rates of the photosynthesis products are hyperbolic functions of light and CO(2) concentration. The model predicts that in the absence of photoinhibition, CO(2)-inhibition, photorespiration, and chlororespiration, a cell acclimated to high light and/or CO(2) concentration has higher photosynthetic capacity and lower photosynthetic efficiency than does a cell acclimated to low conditions. This results in crossing between the two curves which represent the oxygen production rates and carbon fixation rates in low and high conditions. Finally, in the absence of photoinhibition and CO(2)-inhibition, the model predicts the carbohydrate production rate in terms of both light intensity and CO(2) concentration.     

Kinetics of photoacclimation in corals

Traditional models describing the relationship between photosynthesis (P) and irradiance (I) do not account for photoacclimation to short-term variation in irradiance. Here we develop and test a model that predicts the rate of photosynthesis under fluctuating irradiances at the scale of days to weeks. Using oxygen respirometry, we measured the rates of change in the P-I model parameters P(max) (maximum rate of gross photosynthesis) and I(k) (sub-saturation irradiance) of the photo-symbiotic coral Turbinaria mesenterina (Lamarck) following large and small increases and decreases in growth irradiance. We analyse the behaviour of the dynamic P-I model in turbid-water conditions using a dataset of 3-month continuous irradiance as the input variable. In response to upward or downward changes in experimental growth irradiance, I(k) values decreased or increased exponentially, reaching new and stable levels within 5-10 days. I(k) responded 4 times stronger than P(max) to changes in growth irradiance. The kinetics of I(k) did not show hysteresis, and changed in similar ways when irradiance was increased or decreased in small or large amounts. This suggests that mechanisms associated with photo-protection during increases in irradiance, and the maximisation of photosynthetic efficiency during decreases in irradiance, are equally potent. On the scale of months, the dynamic P-I model did not predict higher rates of photosynthesis than the static P-I model, but buffered the variation in photosynthesis during periods of reduced irradiance. Fourier analysis indicated that the kinetics of I(k) closely matches the main periodicities in daily irradiance (1-2 weeks). The recorded kinetics of photoacclimation in the Turbinaria-zooxanthella symbiosis is comparable to that of free-living phytoplankton and faster than that of higher plants.     

Characterization of Photosynthetic Rhythms in Marine Dinoflagellates: II. Photosynthesis-Irradiance Curves and in Vivo Chlorophyll a Fluorescence

Using data from light-dark cultures of Gonyaulax polyedra entrained to a 24-hour cycle, whole cell absorption curves and photosynthesis-irradiance curves were constructed for various circadian times. While whole cell absorbance and half-saturation constants of photosynthesis showed no statistical difference that could be directly related to the photosynthetic rhythm, the initial slope of the photosynthesis-irradiance curve was a time-dependent parameter which altered in direct proportion to the change in photosynthetic capacity. The results indicated a temporal change in the relative quantum yield of photosynthesis, and the circadian rhythmicity of light-limited photosynthesis was established under constant conditions. Circadian rhythmicity was detected in room temperature chlorophyll fluorescence yield. Low temperature fluorescence kinetics also showed fluctuations. The results suggest that regulation of photosynthesis by the biological clock of Gonyaulax may be mediated through the membrane-bound light reactions and a partial explanation of the underlying mechanism is proposed.     

Consequences of photosystem‐I damage and repair on photosynthesis and carbon use in Arabidopsis thaliana

Natural growth environments commonly include fluctuating conditions that can disrupt the photosynthetic energy balance and induce photoinhibition through inactivation of the photosynthetic apparatus. Photosystem II (PSII) photoinhibition is efficiently reversed by the PSII repair cycle, whereas photoinhibited photosystem I (PSI) recovers much more slowly. In the current study, treatment of the Arabidopsis thaliana mutant proton gradient regulation 5 (pgr5) with excess light was used to compromise PSI functionality in order to investigate the impact of photoinhibition and subsequent recovery on photosynthesis and carbon metabolism. The negative impact of PSI photoinhibition on CO₂ fixation was especially deleterious under low irradiance. Impaired starch accumulation after PSI photoinhibition was reflected in reduced respiration in the dark, but this was not attributed to impaired sugar synthesis. Normal chloroplast and mitochondrial metabolisms were shown to recover despite the persistence of substantial PSI photoinhibition for several days. The results of this study indicate that the recovery of PSI function involves the reorganization of the light‐harvesting antennae, and suggest a pool of surplus PSI that can be recruited to support photosynthesis under demanding conditions.     

SEASONAL PATTERN OF DIEL PERIODICITY IN PHOTOSYNTHESIS BY POLAR PHYTOPLANKTON: SPECIES-SPECIFIC RESPONSES1

The diel patterns of light-saturated and light-limited photosynthesis were measured for three diatom species in McMurdo Sound, Antarctica during the transition from late austral winter to summer. Maximum photosynthetic capacity occurred around mid-day during September, when there was a well defined light/dark cycle, and progressively shifted to about midnight by late october when irradiance was continuous. There was a concomitant shift in minimum photosynthetic capacity from midnight to midday. Rates of light-saturated and -limited photosynthesis covaried, and the magnitude of seasonal and diel changes in photosynthetic characteristics were similar. The linear relationship between light-saturated and -limited photosynthesis suggests that the shapes of the photosynthesis-irradiance curves remained relatively constant over the day and througout the season. The unique diel patterns of photosynthesis of these polar phytoplankton appear to be a response to the persistently low, yet continuous irradiance of the polar summer.     

Effects of environmental parameters on net photosynthesis of a free-living brown seaweed,Cystoseira barbata formarepens: determination of optimal photosynthetic culture conditions

The net photosynthesis of the Mediterranean brown seaweedCystoseira barbata f.repens is measured according to irradiance, temperature and salinity. There is not only, a good utilization of low light intensities (light-shade adaptation), but also a specific ability to use a broad range of irradiance, which corresponds in the photosynthesis-irradiance curves to a high initial slope and an extended light saturation level from 300 to 1500 mol photon m^–2 s^–1; only very high irradiances induce photoinhibition. Maximum net photosynthesis occurred at temperatures ranging from 20 °C to 30 °C. The alga tolerates not only a low level of salinity, but also a slight increase in salinity; however, at more than 47.5 g 1^–1 NaCl, oxygen exchange is significantly reduced.Light, temperature and salinity requirements are discussed, taking into account ecological considerations. Yields and quality of alginic acid are presented according to the irradiance and yearly evolutionin situ in order to aid future cultivation of this species.     

A cell-based model for the photoacclimation and CO2-acclimation of the photosynthetic apparatus

We have developed a mathematical model based on the underlying mechanisms concerning the responses of the photosynthetic apparatus of a microalga cell which grows under constant incident light intensity and ambient CO₂ concentration. Photosynthesis involves light and carbon-fixation reactions which are mutually dependent and affect each other, but existing models for photosynthesis don't account for both reactions at once. Our modeling approach allows us to derive distinct equations for the rates of oxygen production, NADPH production, carbon dioxide fixation, carbohydrate production, and rejected energy, which are generally different. The production rates of the photosynthesis products are hyperbolic functions of light and CO₂ concentration. The model predicts that in the absence of photoinhibition, CO₂-inhibition, photorespiration, and chlororespiration, a cell acclimated to high light and/or CO₂ concentration has higher photosynthetic capacity and lower photosynthetic efficiency than does a cell acclimated to low conditions. This results in crossing between the two curves which represent the oxygen production rates and carbon fixation rates in low and high conditions. Finally, in the absence of photoinhibition and CO₂-inhibition, the model predicts the carbohydrate production rate in terms of both light intensity and CO₂ concentration.     

Dynamics of Phytoplankton Communities Under Photoinhibition

We analyzed a model of phytoplankton competition for light in a well-mixed water column. The model, proposed by Gerla et al. (Oikos 120:519–527, 2011), assumed inhibition of photosynthesis at high irradiance (photoinhibition). We described the global behavior through mathematical analyses, providing a general solution to the multi-species competition for light with photoinhibition. We classified outcomes of 2- and 3-species competitions as examples, and evaluated feasibility of the theoretical predictions using empirical relationships between photosynthetic production and irradiance. Numerical simulations with published p–I curves indicate that photoinhibition may often lead to strong Allee effects and competitive facilitation among species. Hence, our results suggest that photoinhibition may play a major role in organizing phytoplankton communities.     

Light distribution in leaf chambers and its consequences for photosynthesis measurements

The impact of a heterogeneous distribution of actinic light within a leaf chamber for photosynthetic measurements by gas exchange on the photosynthesis-irradiance relationship was investigated. High-resolution light distributions were measured over the area of a commercially available clamp-on leaf chamber equipped with build-in red and blue LEDs, as well as over the area of a custom-made leaf chamber with external light source, using a low-cost digital camera and freely available software. The impact of the measured heterogeneity on the photosynthesis-irradiance response curve was calculated for two realistic scenarios. When the average light intensity over the leaf chamber area was estimated accurately, heterogeneity had minor effects on the photosynthesis-irradiance response curve. However, when the irradiance was measured in the chamber centre, which is common practice, and assumed to be homogeneous, for both leaf chambers the photosynthesis-irradiance response curve was subject to considerable error and led to serious underestimation of the light-limited quantum yield of photosynthesis. Additionally, mixed light sources with different heterogeneity patterns per light source, such as in the clamp-on leaf chamber, potentially increase errors due to heterogeneous physiological responses to light spectrum. High-resolution quantification of the leaf-chamber light distribution enables calculation of the correct average light intensity and already resolves the most pressing problems associated with heterogeneity. To exclude any light-distribution related errors in gas-exchange measurements a leaf chamber and actinic irradiance source design with a homogeneous light distribution is an absolute requirement.     

Mathematical modelling of the light response curve of photoinhibition of Photosystem II

The light response curves of the acceptor and donor side mechanisms of photoinhibition of Photosystem II were calculated, using Arabidopsis as a model organism. Acceptor-side photoinhibition was modelled as double reduction of Q_A, noting that non-photochemical quenching has the same effect on the quantum yield of Q_A double reduction in closed PSII centres as it has on the quantum yield of electron transport in open centres. The light response curve of acceptor-side photoinhibition in Arabidopsis shows very low efficiency under low intensity light and a relatively constant quantum yield above light saturation of photosynthesis. To calculate the light response curve of donor-side photoinhibition, we built a model describing the concentration of the oxidized primary donor P₆₈₀⁺ during steady-state photosynthesis. The model is based on literature values of rate constants of electron transfer reactions of PSII, and it was fitted with fluorescence parameters measured in the steady state. The modelling analysis showed that the quantum yield of donor-side photoinhibition peaks under moderate light. The deviation of the acceptor and donor side mechanisms from the direct proportionality between photoinhibition and photon flux density suggests that these mechanisms cannot solely account for photoinhibition in vivo, but contribution of a reaction whose quantum yield is independent of light intensity is needed. Furthermore, a simple kinetic calculation suggests that the acceptor-side mechanism may not explain singlet oxygen production by photoinhibited leaves. The theoretical framework described here can be used to estimate the yields of different photoinhibition mechanisms under different wavelengths or light intensities.     

Photosynthetic acclimation to solar UV radiation of marine red algae from the warm-temperate coast of southern Spain: A review

A review is presented of the physiological mechanisms developed bybenthic macroalgae to cope with deleterious wavelengths, particularly UVradiation. Photoinhibition of photosynthesis, is a photoprotectivemechanism in various species studied in southern Spain. Incubations inoutdoor systems and transplantation experiments under natural radiationallowed to led to understanding of some the photoprotective strategies usedby red algae. Under conditions of enhanced UV-B radiation, algae inshallow sites show marked photoinhibition and rapid recovery ofphotosynthesis (dynamic photoinhibition), whereas algae from deeperlocations can suffer photodamage (chronic photoinhibition). Theexpression of this photoprotective strategy by intertidal species representsan efficient physiological adaptation to tolerate deleterious irradiance, whenlow tides coincide with the onset of solar radiation. Subtidal species canbe also exposed to high doses of UV radiation. This is particularly evidentin clear, Mediterranean waters, where light (including UV-B) can reach to10 m depth. The implications of photoacclimation processes formacroalgal ecology in warm-temperate littorals and the possibleconsequences for outdoor cultivation are outlined in terms of environmentalUV variability.     

水淹导致皇冠草光合机构发生变化并加剧其出水后光抑制

通过气体交换和叶绿素荧光等方法研究了水淹及胁迫解除后皇冠草不同功能叶的光合特性及光抑制的变化.结果表明:与对照相比,气生叶(全淹组淹水前形成的功能叶)在水淹条件下叶片大小和气孔没有明显变化,但沉水叶(全淹组淹水后新生的功能叶)的叶面积增加,气孔变小,上表皮气孔密度增加.水淹导致气生叶碳同化能力、光化学效率和叶绿素含量下降.沉水叶在发育过程中碳同化能力、光化学效率和叶绿素逐渐升高.气生叶和沉水叶出水后其活体叶片在强光下的相对含水量急剧下降,发生明显的光抑制;而弱光下无明显光抑制发生.出水后离体叶片强光照射下6h后两种功能叶均发生严重光抑制,且弱光下不能恢复.因此,可以认为淹水条件下,沉水叶上表皮气孔密度的增加使其蒸腾速率提高;沉水叶较强的碳同化能力和增加的叶面积是确保其植株水下生存的重要因素;强光使气生叶和沉水叶出水后均发生严重光抑制,导度和蒸腾速率提高导致的叶片失水则加剧了这一过程,两者共同作用导致自然条件下两种功能叶的出水死亡.     

Outdoor continuous culture of Porphyridium cruentum in a tubular photobioreactor: quantitative analysis of the daily cyclic variation of culture parameters

The present work reports on the daily cyclic variation of oxygen generation rates, carbon consumption rates, photosynthetic activities, growth rates and biochemical composition of the biomass in a pilot plant continuous outdoor culture of the microalgae Porphyridium cruentum. A linear relationship between the external irradiance and the average irradiance inside the culture was found. In addition, the oxygen generation and carbon consumption rates were found to be a function of the average irradiance inside the culture. A reduction in photosynthetic activity of the cells at noon and recovery in the afternoon was also observed. Therefore, the cells showed a short-term response of parameters such as oxygen generation rate as well as carbon consumption rate with external and average irradiance; a model of photosynthesis rate considering photoinhibition is proposed. This model is a useful tool for the operation and scaleup of tubular photobioreactors, and can be used for determining CO₂ requirements of the system. The higher the photosynthesis rates, the lower the carbon losses, ranging from 25% at noon to 100% during the night. The growth rate showed a linear relationship with the daily mean average irradiance inside the culture with a long-term response. Likewise, a linear relationship among the oxygen generation rate and the growth rate was obtained. With respect to the biochemical composition of the biomass, the cells showed a long-term response of metabolic routes to mean daily culture conditions. During the illuminated period, energy was stored as carbohydrates and synthesis of proteins was low. During the night, the stored carbohydrates were consumed. The fatty acid dry weight (DW) content decreased during the daylight period, whereas the fatty acid profile, as total fatty acids, was a function of growth rate. A short-term variation of exopolysaccharides synthesis with solar irradiance was also observed, i.e. the higher the external irradiance the higher the excretion of exopolysaccharides as a protection against adverse culture conditions.     

Mechanisms of photoinhibition induced by high light in Hosta grown outdoors

Aims It has long been recognized that photoinhibition of photosynthesis is induced by high light. However, our recent studies are not consistent with this traditional view. Therefore, the objective of this study is to explore the induction of photoinhibition and its mechanisms under full sunlight outdoors.
Methods Changes of leaf morphology, gas exchange, and chlorophyll a fluorescence were measured to investigate the induction and mechanisms of photoinhibition under high light in Hosta, which is a typical shade-tolerant plant.
Important findings Hosta plants grown under full sunlight (HT) and low light (LT) developed sun- and shade-type leaf morphological characteristics, respectively. Under a full sunlight, Hosta plants had lower photosynthetic rate and chlorophyll content than under the LT; whereas, there were only slight difference in the maximum quantum yield of photosystem II (F_v/F_m) between the two treatments, suggesting that Hosta plants could grow normally under full sunlight without severe photoinhibition. After transition from the low to a high light (LHT), the photosynthetic rate and maximum quantum yield of photosystem II decreased sharply, reflecting that the LHT treatment led to irreversibly inactivation of photosystem II. Additionally, the shape of chlorophyll a fluorescence transients also changed significantly; the relative fluorescence yield of the K and J steps were reduced by 24.3% and 34.2%, respectively, indicating that the acceptor side of photosystem II was damaged more severely than the donor side. Consequently, we postulate that photoinhibition in Hosta leaves is mainly induced by the sudden enhancement of light intensity outdoors. Hosta can acclimate to high irradiance through leaf development outdoors. Our finding is of great significance in understanding the acclimation of plants to high light and cultivation of shade-tolerant plants in field.