Light-Related Photosynthetic Characteristics of Lotic Macroalgae

Photosynthetic characteristics in response to irradiance were analysed in 42 populations of 33 macroalgal species by two distinct techniques (chlorophyll fluorescence and oxygen evolution). Photosynthesis–irradiance (PI) curves based on the two techniques indicated adaptations to low irradiance reflected by low saturation values, high to moderate values of photosynthetic efficiency (α) and photoinhibition (β), for Bacillariophyta and Rhodophyta, which suggests they are typically shade-adapted algae. In contrast, most species of Chlorophyta were reported as sun adapted algae, characterized by high values of I _k and low of α, and lack of or low photoinhibition. Cyanophyta and Xanthophyta were intermediate groups in terms of light adaptations. Photoinhibition was observed in variable degrees in all algal groups, under field and laboratory conditions, which confirms that it is not artificially induced by experimental conditions, but is rather a common and natural phenomenon of the lotic macroalgae. Low values of compensation irradiance (I _c) were found, which indicate that these algae can keep an autotrophic metabolism even under very low irradiances. High ratios (>2) of photosynthesis/respiration were found in most algae, which indicates a considerable net gain. These two physiological characteristics suggest that macroalgae may be important primary producers in lotic ecosystems. Saturation parameters (I _k and I _s) occurred in a relatively narrow range of irradiances (100–400 μmol photons m^?2s^?1), with some exceptions (higher in some filamentous green algae or lower in red algae). These parameters were way below the irradiances measured at collecting sites for most algae, which means that most of the available light energy was not photochemically converted via photosynthesis. Acclimation to ambient PAR was observed, as revealed by lower values of I _k and I _cand higher values of α and quantum yield in algae from shaded streams, and vice versa. Forms living within the boundary layer (crusts) showed responses of shade-adapted species and had the highest values of P _max, α and quantum yield, whereas the opposite trend was observed in gelatinous forms (colonies and filaments). These results suggests adaptation to the light regime rather than functional attributes related to the growth form.     

Effect of nitrate concentration and UVR on photosynthesis,respiration, nitrate reductase activity,and phenolic compounds in <Emphasis Type="Italic">Ulva rigida</Emphasis> (Chlorophyta)

Seaweeds growing in the intertidal zone are exposed to fluctuating nitrate and ultraviolet radiation (UVR) levels. While it has been shown that elevated UVR levels and the decrease of nitrate concentration can reduce photosynthetic levels in seaweeds, less is known about the combined effect of nitrate levels and UVR on metabolism and photoprotection mechanisms of intertidal species. Consequently, the objective of this study was to evaluate the effect of nitrate concentration and UVR treatments on photosynthesis, respiration, nitrate reductase activity and phenolic compound levels of Ulva rigida (Chlorophyta). There was a two- to threefold increase in maximal gross photosynthesis (GP_max) and respiration rates, as nitrate increased from 0 to 50 μM NO₃⁻. Similarly, nitrate reductase activity increased linearly from low values in algae incubated at 0 μM NO₃ to high values in tissue incubated at 50 μM NO₃⁻. Phenolic compounds in the tissue of U. rigida increased approximately 60% under 50 μM NO₃⁻ relative to those incubated at 0 μM NO₃⁻. Algae exposed to UVR (8 h) showed a significant decrease in the effective quantum yield and respiration, however, no effect was observed in the phenolic compounds levels. Full recovery of effective quantum yield was observed after U. rigida was transferred for 48 h to low PAR. Nitrate reductase also decreased after an 8-h UVR exposure, but no differences were observed among the nitrate treatments. This study shows that high nitrate levels reduced the negative effect of UVR on the effective quantum yield and increased the recovery of key metabolic enzymes. It is possible that the increase of phenolic compounds in the thallus of U. rigida under high nitrate levels provide a photoprotective mechanism when exposed to high UV levels during low tides.     

Photosynthetic Fluorescence Induction and Oxygen Production in Corallinacean Algae Measured on Site

Photoinhibition of photosynthesis was measured in two Mediterranean Corallinaceae, Jania rubens and Corallina mediterranea, using pulse-amplitude modulation (PAM) fluorescence and oxygen production on site. Both algae were found to be adapted to low irradiances of solar radiation and easily inhibited by exposure to excessive radiation. Both algae were impaired even in their natural habitat under overhanging rocks which protected them from direct solar radiation, except for a few hours in the early morning. Recovery from photoinhibition of both the photosynthetic quantum yield, defined as F_v′/F_m′, and oxygen production took several hours and was not complete. Judging from both parameters indicated above, Jania seems to be even more sensitive than Corallina, even though the former alga was found in more exposed habitats.     

An in situ study of photosynthetic oxygen exchange and electron transport rate in the marine macroalga Ulva lactuca (Chlorophyta)

Direct comparisons between photosynthetic O₂ evolution rate and electron transport rate (ETR) were made in situ over 24 h using the benthic macroalga Ulva lactuca (Chlorophyta), growing and measured at a depth of 1.8 m, where the midday irradiance rose to 400–600 μmol photons m⁻² s⁻¹. O₂ exchange was measured with a 5-chamber data-logging apparatus and ETR with a submersible pulse amplitude modulated (PAM) fluorometer (Diving-PAM). Steady-state quantum yield ((F_m′−F_t)/F_m′) decreased from 0.7 during the morning to 0.45 at midday, followed by some recovery in the late afternoon. At low to medium irradiances (0–300 μmol photons m⁻² s⁻¹), there was a significant correlation between O₂ evolution and ETR, but at higher irradiances, ETR continued to increase steadily, while O₂ evolution tended towards an asymptote. However at high irradiance levels (600–1200 μmol photons m⁻² s⁻¹) ETR was significantly lowered. Two methods of measuring ETR, based on either diel ambient light levels and fluorescence yields or rapid light curves, gave similar results at low to moderate irradiance levels. Nutrient enrichment (increases in [NO₃ ⁻], [NH₄ ⁺] and [HPO₄ ^2-] of 5- to 15-fold over ambient concentrations) resulted in an increase, within hours, in photosynthetic rates measured by both ETR and O₂ evolution techniques. At low irradiances, approximately 6.5 to 8.2 electrons passed through PS II during the evolution of one molecule of O₂, i.e., up to twice the theoretical minimum number of four. However, in nutrient-enriched treatments this ratio dropped to 5.1. The results indicate that PAM fluorescence can be used as a good indication of the photosynthetic rate only at low to medium irradiances. This revised version was published online in June 2006 with corrections to the Cover Date.     

Chlorophyll fluorescence in micropropagated <Emphasis Type="Italic">Rhododendron ponticum</Emphasis> subsp. <Emphasis Type="Italic">baeticum</Emphasis> plants in response to different irradiances

The aim of this study was to investigate acclimation of micropropagated plants of Rhododendron ponticum subsp. baeticum to different irradiances and recovery after exposure to high irradiance. Plants grown under high (HL) or intermediate (IL) irradiances displayed higher values of maximum electron transport rate (ETR_max) and light saturation coefficient (E_k) than plants grown under low irradiance (LL). The capacity of tolerance to photoinhibition (as assessed by the response of photochemical quenching, q_p) varied as follows: HL > IL > LL. Thermal energy dissipation (q_N) was also affected by growth irradiance, with higher saturating values being observed in HL plants. Light-response curves suggested a gradual replacement of q_p by q_N with increasing irradiance. Following exposure to irradiance higher than 1500 μmol m⁻² s⁻¹, a prolonged reduction of the maximal photochemical efficiency of PS 2 (F_v/F_m) was observed in LL plants, indicating the occurrence of chronic photoinhibition. In contrary, the decrease in F_v/F_m was quickly reverted in HL plants, pointing to a reversible photoinhibition.     

Two components of onset and recovery during photoinhibition of Ulva rotundata

Short-term (up to 5 h) transfers of shade-adapted (100 mol · m^–2 · s^–1) clonal tissue of the marine macroalga Ulva rotundata Blid. (Chlorophyta) to higher irradiances (1700, 850, and 350 mol · m^–2 · s^–1) led to photoinhibition of room-temperature chlorophyll fluorescence and O₂ evolution. The ratio of variable to maximum (F_v/F_m) and variable (F_v) fluorescence, and quantum yield () declined with increasing irradiance and duration of exposure. This decline could be resolved into two components, consistent with the separation of photoinhibition into energy-dissipative processes (photoprotection) and damage to photosystem II (PSII) by excess excitation. The first component, a rapid decrease in F_v/F_m and in F_v, corresponds to an increase in initial (F_o) fluorescence and is highly sensitive to 1 mM chloramphenicol. This component is rapidly reversible under dim (40 mol · m^–2 · s^–1) light, but is less reversible with increasing duration of exposure, and may reflect damage to PSII. The second (after 1 h exposure) component, a slower decline in F_v/F_m and F_v with declining F_o, appears to be associated with the photoprotective interconversion of violaxanthin to zeaxanthin and is sensitive to dithiothreitol. The accumulation of zeaxanthin in U. rotundata is very slow, and may account for the predominance of increases in F_o at high irradiances.Abbreviations and Symbols CAP chloramphenicol - DTT dithiothreitol - F_o, F_m, F_v initial, maximum, and variable fluorescence - quantum yield - PFD photon flux density - PSII photosystem II To whom correspondence should be addressedWe are grateful to O. Björkman and S. Thayer, Carnegie Institution of Washington, Stanford, Cal., USA, for analysis of xanthophyll pigments reported here. This research was supported by National Science Foundation grant OCE-8812157 to C.B.O. and J.R. Support for G.L. was provided by a NSF-CNRS (Centre National de la Recherche Scientifique) exchange fellowship.     

Irradiance optimization of outdoor microalgal cultures using solar tracked photobioreactors

Photosynthetic activity and temperature regulation of microalgal cultures (Chlorella vulgaris and Scenedesmus obliquus) under different irradiances controlled by a solar tracker and different cell densities were studied in outdoor flat panel photobioreactors. An automated process control unit regulated light and temperature as well as pH value and nutrient concentration in the culture medium. CO₂ was supplied using flue gas from an attached combined block heat and power station. Photosynthetic activity was determined by pulse amplitude modulation fluorometry. Compared to the horizontal irradiance of 55 mol photons m^?2 d^?1 on a clear day, the solar tracked photobioreactors enabled a decrease and increase in the overall light absorption from 19 mol photons m^?2 d^?1 (by rotation out of direct irradiance) to 79 mol photons m^?2 d^?1 (following the position of the sun). At biomass concentrations below 1.1 g cell dry weight (CDW) L^?1, photoinhibition of about 35 % occurred at irradiances of ≥1,000 μmol photons m^?2 s^?1 photosynthetic active radiation (PAR). Using solar tracked photobioreactors, photoinhibition can be reduced and at optimum biomass concentration (≥2.3 g CDW L^?1), the culture was irradiated up to 2,000 μmol photons m^?2 s^?1 to overcome light limitation with biomass yields of 0.7 g CDW mol photons^?1 and high photosynthetic activities indicated by an effective quantum yield of 0.68 and a maximum quantum yield of 0.80 (F _v/F _m). Overheating due to high irradiance was avoided by turning the PBR out of the sun or using a cooling system, which maintained the temperature close to the species-specific temperature optima.     

On the significance of photoinhibition of photosynthesis in the field and its generality among species

Photoinhibition was examined in naturally exposed willow leaves in the field. In the afternoon on clear and warm days, the quantum yield of electron transport, derived from gas exchange data, was decreased by 28%. Besides this photoinhibition, decreases in the photosynthetic capacity and in the stomatal conductance were also observed. Of these three limitations of carbon assimilation, photoinhibition was the major one at limiting light only.To investigate the generality of photoinhibition, shade- and sun-acclimated leaves of fourteen different species were compared in a laboratory study. Photoinhibition was quantified by fluorescence measurements following exposure to moderate and high light for 30 min. The extent of photoinhibition was inversely related to the photochemical quenching, q_p, during exposure (the proportion of open PS II traps). This relationship was the same independent of the species, the light-acclimation state of the leaf and the light intensity. However, sun- and shade-acclimated leaves occupied opposite sides of the relationship: the sun-leaves showed lower photoinhibition and higher q_p. The sun-leaves were also more competent than shade-leaves by showing faster recovery from a given level of photoinhibition.Abbreviations F₀, F_V, F_M, F_S minimal, variable, maximal and steady-state fluorescence - q_N, q_i total and photoinhibitory non-photochemical quenching of variable fluorescence - q_p photochemical quenching     

The effects of temperature acclimation on the photoinhibitory responses of Ulva rotundata Blid.

The effect of acclimation to 25, 18, or 10° C on the relationship between photoprotection and photodamage was tested in low-light-grown (80 mol · m^–2 · s^–1) Ulva rotundata Blid. exposed to several higher irradiances at the acclimation temperature. Changes in chlorophyll fluorescence parameters (minimum fluorescence, F₀, and the ratio of variable to maximum fluorescence, F_v/F_m, measured after 5 min darkness) were monitored during 5 h transfers to 350, 850, and 1700 mol · m^–2 · s^–1, and during recovery after 1- or 5-h treatments. At all temperatures, rate of onset and final extent of photoinhibition, measured by a decrease in F_v/F_m, increased with increasing irradiance. At a given photoinhibitory irradiance, rate of onset was most rapid at 10 ° C, but the extent was temperature-independent. Recovery rates from mild light stress were similar at all temperatures, but recovery from the most extreme photoinhibitory treatment lagged 2 h at 10° C. De-epoxidation of xanthophyll-cycle components proceeded faster and to a lower epoxidation status at 25° C, but there was little difference in the pool size among the three growth conditions. Using chloramphenicol to inhibit chloroplast protein synthesis and dithiothreitol to inhibit violaxanthin de-epoxidation, it was shown that at the lowest light treatment given, the extent of photoinhibition could be attributed both to greater amounts of photodamage and to greater zeaxanthin-related photoprotection at 25 than at 10° C. While these two mechanisms for high-light-induced loss of photosynthetic efficiency were operating at 10° C, there was evidence for a relatively greater proportion of zeaxanthin-unrelated photoprotection at the low temperature. This photoprotective mechanism is related to a rapidly reversible increase in F₀ and is insentivite to both chloramphenicol and dithiothreitol.Abbreviations and Symbol CAP chloramphenicol - DTT dihiothreitol - F₀, F_m, F_v minimum, maximum, and variable fluorescence - quantum yield This research was conducted in partial fulfillment of the requirements for the Ph. D. degree in the Department of Botany, Duke University. The author wishes to thank E.-M. Aro, W.J. Henley, G. Levavasseur, C.B. Osmond, and J. Ramus for helpful discussions, and C. Lovelock for pigment standards. Funding was provided by Grants-in-Aid of Research from Sigma Xi and the Phycological Society of America, and a Lynde and Harry Bradley Foundation Fellowship to L.A.F., and National Science Foundation grant OCE-8812157 to C.B.O. and J.R.     

THE SHORT‐TERM EFFECT OF IRRADIANCE ON THE PHOTOSYNTHETIC PROPERTIES OF ANTARCTIC FAST‐ICE MICROALGAL COMMUNITIES1

Although sea‐ice represents a harsh physicochemical environment with steep gradients in temperature, light, and salinity, diverse microbial communities are present within the ice matrix. We describe here the photosynthetic responses of sea‐ice microalgae to varying irradiances. Rapid light curves (RLCs) were generated using pulse amplitude fluorometry and used to derive photosynthetic yield (Φ_PSII), photosynthetic efficiency (α), and the irradiance (E_k) at which relative electron transport rate (rETR) saturates. Surface brine algae from near the surface and bottom‐ice algae were exposed to a range of irradiances from 7 to 262 μmol photons · m^?2 · s^?1. In surface brine algae, Φ_PSII and α remained constant at all irradiances, and rETR_max peaked at 151 μmol photons · m^?2 · s^?1, indicating these algae are well acclimated to the irradiances to which they are normally exposed. In contrast, Φ_PSII, α, and rETR_max in bottom‐ice algae reduced when exposed to irradiances >26 μmol photons · m^?2 · s^?1, indicating a high degree of shade acclimation. In addition, the previous light history had no significant effect on the photosynthetic capacity of bottom‐ice algae whether cells were gradually exposed to target irradiances over a 12 h period or were exposed immediately (light shocked). These findings indicate that bottom‐ice algae are photoinhibited in a dose‐dependent manner, while surface brine algae tolerate higher irradiances. Our study shows that sea‐ice algae are able to adjust to changes in irradiance rapidly, and this ability to acclimate may facilitate survival and subsequent long‐term acclimation to the postmelt light regime of the Southern Ocean.     

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.     

Relationships between irradiance and photosynthesis for marine benthic green algae (Chlorophyta) of differing morphologies

Photosynthesis-irradiance relationships were determined in the field for five species of littoral and shallow sublittoral marine benthic green algae (Chlorophyta) of differing morphologies. Each species exhibited a linear increase in photosynthetic rate with increasing irradiance up to a maximum light-saturated value. Full sunlight (1405 to 1956 μE·m^?2·s^?1) inhibited photosynthesis of all species except the thick, optically dense, Codium fragile (Sur.) Har. Compensation irradiances ranged from 6.1 μE·m^?2·s^?1 for Enteromorpha intestinalis (L.) Link to 11.4 μE·m^?2·s^?1 for Ulva lobata (Kütz) S. & G. and did not reveal a consistent relationship to seaweed morphology. Saturation irradiances were determined statistically (I_k) and visually from graphical plots. with the latter technique resulting in values three to eight times higher and different comparative rankings of species than the former. I_k saturation irradiances were highest for Chaetomorpha linum (Müll.) Kütz. (81.9 μE·m^?2·s^?1) and lowest for Codium fragile (49.6 μE·m^?2·s^?1) and did not reveal a relationship with seaweed morphology. Regression equations describing light-limited photosynthetic rates and the relative magnitudes of the maximal net photosynthetic responses both strongly suggested a relationship with seaweed morphology. Highest net photosynthetic rates were obtained for the thin, sheet-like algae Ulva lobata (9.2 mg C·g dry wt^?1·h^?1), U. rigida C. Ag. (6.5 mg C·g dry wt^?1·h^?1) and the tubular form, Enteromorpha intestinalis (7.3 mg C·g dry wt^?1·h^?1), while lowest rates occurred for Codium fragile (0.9 mg C·g dry wt^?1·h^?1). Similarly, steepest light-limited slopes were found for the algae of simpler morphology, while the most gradual slope was determined for Codium fragile, the alga with greatest thallus complexity.     

Effects of light intensity and temperature on the photosynthetic irradiance response curves and chlorophyll fluorescence in three picocyanobacterial strains of Synechococcus

Chrococcoid cyanobacteria of the genus Synechococcus are the important component of marine and freshwater ecosystems. Picocyanobacteria comprise even 80% of total cyanobacterial biomass and contribute to 50% of total primary cyanobacterial bloom production. Chlorophyll (Chl) fluorescence and photosynthetic light response (P-I) curves are commonly used to characterize photoacclimation of Synechococcus strains. Three brackish, picocyanobacterial strains of Synechococcus (BA-132, BA-124, BA-120) were studied. They were grown under 4 irradiances [10, 55, 100, and 145 μmol(photon) m^?2 s^?1] and at 3 temperatures (15, 22.5, and 30°C). Photosynthetic rate was measured by Clark oxygen electrode, whereas the Chl fluorescence was measured using Pulse Amplitude Modulation fluorometer. Based on P-I, two mechanisms of photoacclimation were recognized in Synechococcus. The maximum value of maximum rate of photosynthesis (P _max) expressed per biomass unit at 10 μmol(photon) m^?2 s^?1 indicated a change in the number of photosynthetic units (PSU). The constant values of initial slope of photosynthetic light response curve (α) and the maximum value of P _max expressed per Chl unit at 145 μmol(photon) m^?2 s^?1 indicated another mechanism, i.e. a change in PSU size. These two mechanisms caused changes in photosynthetic rate and its parameters (compensation point, α, saturation irradiance, dark respiration, P _max) upon the influence of different irradiance and temperature. High irradiance had a negative effect on fluorescence parameters, such as the maximum quantum yield and effective quantum yield of PSII photochemistry (φ_PSII), but it was higher in case of φ_PSII.     

On the relationship between the quantum yield of Photosystem II electron transport,as determined by chlorophyll fluorescence and the quantum yield of CO2-dependent O2 evolution

We tested the two empirical models of the relationship between chlorophyll fluorescence and photosynthesis, previously published by Weis E and Berry JA 1987 (Biochim Biophys Acta 894: 198–208) and Genty B et al. 1989 (Biochim Biophys Acta 990: 87–92). These were applied to data from different species representing different states of light acclimation, to species with C₃ or C₄ photosynthesis, and to wild-type and a chlorophyll b-less chlorina mutant of barley. Photosynthesis measured as CO₂-saturated O₂ evolution and modulated fluorescence were simultaneously monitored over a range of photon flux densities. The quantum yields of O₂ evolution (ØO₂) were based on absorbed photons, and the fluorescence parameters for photochemical (q_p) and non-photochemical (q_N) quenching, as well as the ratio of variable fluorescence to maximum fluorescence during steady-state illumination (F'_v/F'_m), were determined. In accordance with the Weis and Berry model, most plants studied exhibited an approximately linear relationship between ØO₂/q_p (i.e., the yield of O₂ evolution by open Photosystem II reaction centres) and q_N, except for wild-type barley that showed a non-linear relationship. In contrast to the linear relationship reported by Genty et al. for q_p×F'_v/F'_m (i.e., the quantum yield of Photosystem II electron transport) and ØCO₂, we found a non-linear relationship between q_p×F'_v/F'_m and ØO₂ for all plants, except for the chlorina mutant of barley, which showed a largely linear relationship. The curvilinearity of wild-type barley deviated somewhat from that of other species tested. The non-linear part of the relationship was confined to low, limiting photon flux densities, whereas at higher light levels the relationship was linear. Photoinhibition did not change the overall shape of the relationship between q_p×F'_v/F'_m and ØO₂ except that the maximum values of the quantum yields of Photosystem II electron transport and photosynthetic O₂ evolution decreased in proportion to the degree of photoinhibition. This implies that the quantum yield of Photosystem II electron transport under high light conditions may be similar for photoinhibited and non-inhibited plants. Based on our experimental results and theoretical analyses of photochemical and non-photochemical fluoresce quenching processes, we conclude that both models, although not universal for all plants, provide useful means for the prediction of photosynthesis from fluorescence parameters. However, we also discuss that conditions which alter one or more of the rate constants that determine the various fluorescence parameters, as well as differential light penetration in assays for oxygen evolution and fluorescence emission, may have direct effect on the relationships of the two models.Abbreviations F₀ and F'₀ fluorescence when all Photosystem II reaction centres are open in dark- and light-acclimated leaves, respectively - F_m and F'_m fluorescence when all Photosystem II reaction centres are closed in dark and light, respectively - F_v variable fluorescence equal to F_m-F₀ - F_s steady state level of fluorescence in light - F'_v and F'_m variable (F'_m-F'₀) and maximum fluorescence under steady state light conditions - HEPES N-2-hydroxyethylpiperazine-N-2-ethane-sulphonic acid - Q_A the primary, stabile quinone acceptor of Photosystem II - q_N non-photochemical quenching of fluorescence - q_p photochemical quenching of fluorescence - ØO₂ quantum yield of CO₂-saturated O₂ evolution based on absorbed photons     

Differences in the photoacclimation and photoprotection exhibited by two species of the ciguatera causing dinoflagellate genus,Gambierdiscus

In culture, Gambierdiscus spp. have been shown to prefer irradiances that are relatively low (≤250 μmol photons m⁻² s⁻¹) versus those to which they are frequently exposed to in their natural environment (>500 μmol photons m⁻² s⁻¹). Although several behavioral strategies for coping with such irradiances have been suggested, it is unclear as to how these dinoflagellates do so on a physiological level. More specifically, how do long term exposures (30 days) affect cell size and cellular chlorophyll content, and what is the photosynthetic response to short term, high irradiance exposures (up to 1464 μmol photons m⁻² s⁻¹)? The results of this study reveal that cell size and chlorophyll content exhibited by G. carolinianus increased with acclimation to increasing photon flux density. Additionally, both G. carolinianus and G. silvae exhibited reduced photosynthetic efficiency when acclimated to increased photon flux density. Photosynthetic yield exhibited by G. silvae was greater than that for G. carolinianus across all acclimation irradiances. Although such differences were evident, both G. carolinianus and G. silvae appear to have adequate biochemical mechanisms to withstand exposure to irradiances exceeding 250 μmol photons m⁻² s⁻¹ for at least short periods of time following acclimation to irradiances of up to 150 μmol photons m⁻² s⁻¹.     

Influence of phytohormones on morphology and chlorophyll a fluorescence parameters in embryos of Fucus vesiculosus L. (Phaeophyceae)

While a variety of plant hormones from brown algae were described, there were few studies that examined the combined effects of these hormones on morphogenesis and photosynthetic physiology in developing fucoid embryos. We evaluated the effects of phytohormones to determine the extent, to which responses were similar to those of terrestrial plants. Kinetin, IAA, ABA, GA₃, and kinetin + IAA were added to seawater at a physiological concentration (1 mg/L), and embryos of Fucus vesiculosus L. were grown for 10 days. Photosynthetic activity of single embryos or embryo cells were characterized using the following fluorescence parameters: minimum fluorescence yield (F ₀), maximum quantum yield (F _v/F _m), relative maximum rate of electron transfer to photosystem II under saturation irradiances (rETR_max), photosynthetic efficiency under non-saturating irradiances (αETR) and saturation irradiance (E _k). In addition, embryo length and diameter and apical hair length and number were determined. Morphological changes associated with hormone treatments included an increase in the embryo length in the presence of IAA, an increase in the embryo diameter in the presence of IAA, kinetin, and kinetin + IAA, an increase in the maximum hair length and number in the presence of kinetin + IAA, and a decrease in the hair length and number in the presence of ABA. With respect to fluorescence parameters, significant effects of phytohormones included an increase in the F ₀ and F _v/F _m at kinetin treatment, a synergistic effect of kinetin + IAA on F _v/F _m, rETR_max, and αETR, a promotion of F _v/F _m by GA, and a decrease of the parameters by ABA. These results are consistent with the data on responses of land plants to the same hormones and suggest that brown algae have evolved regulatory mechanisms for morphogenesis and photosynthetic regulation similar to plants.     

Sex-specific light acclimation of Chara canescens (Charophyta)

Bisexual populations of the charophyte Chara canescens (Desv. et Loisel. in Loisel., 1810) containing male and female individuals are rarely found. Two experiments were carried out to study whether male and female algae from the same site exhibit different physiological capacities, especially with respect to light acclimation.Algae from two different shore levels and from laboratory cultures acclimated to six irradiance conditions (35–500 μmol photons m⁻² s⁻¹) were compared. Field measurements showed that both female and male algae of C. canescens are able to acclimate to daily changes in solar irradiance. The quantum yield of Photosystem II (PSII) decreased with increasing irradiance in the morning and increased with decreasing irradiance in the afternoon. Growth experiments showed increasing growth rates from 35 μmol photons m⁻² s⁻¹ (∼7 mg FW) up to 500 μmol photons m⁻² s⁻¹ (∼27 mg FW) in female and male C. canescens. The irradiance saturation point for photosynthesis (E_k) was about 140 μmol m⁻² s⁻¹ for both sexes within the whole range of acclimation irradiances. The maximum photosynthesis rate at saturating irradiances (P_max) of male algae was highest at E_k, whereas P_max of female algae was highest at 500 μmol photons m⁻² s⁻¹. The photosynthetic efficiency in the light-limited range (α) increased in female C. canescens and decreased in male C. canescens. The ratio of the non-photochemical quenching parameter (NPQ) to the relative electron transport rates rETR_(MT) increased in both sexes with irradiance, but showed a steeper increase in male than in female algae. Pigment analysis showed similar acclimation pattern for male and female C. canescens. Chl a/Chl b ratios of both sexes were constant over the whole range of E_g, whereas Chl a/carotenoid ratios in male and female C. canescens decreased from 70 μmol photons m⁻² s⁻¹ upwards. Pigment analysis pointed out that the carotenes α-, β- and γ-carotene were more prominent in male than in female algae.Our results indicate that female C. canescens are more efficient in light acclimation than male algae from the same site. Nevertheless, further investigations of bisexual C. canescens populations resolving CO₂-uptake mechanisms and/or genetic differences are needed.     

Photosynthetic acclimation in shade-developed leaves of Euterpe edulis Mart (Arecaceae) after long-term exposure to high light

To analyze acclimation of Euterpe edulis seedlings to changes in light availability, we transferred three-year-old seedlings cultivated for six months under natural shade understory [≈ 1.3 mol(photon) m^?2 d^?1] to a forest gap [≈ 25.0 mol(photon) m^?2 d^?1]. After the transfer, changes in chlorophyll fluorescence and leaf gas-exchange parameters, as well as in the light-response curves of photosynthesis and photosynthetic induction parameters, were analyzed during the following 110 days. Simultaneously measured photosynthetic characteristics in the shaded seedlings grown in understory served as the control. Despite the fact that the understory seedlings were under suboptimal conditions to achieve their light-saturated net photosynthetic rate (P _Nmax), light-response curves and photosynthetic induction parameters indicated that the species had the low respiration rate and a fast opening of stomata in response to the intermittent occurrence of sunflecks, which exerted a feed-forward stimulation on P _Nmax. Sudden exposure to high light induced photoinhibition during the first week after the transfer of seedlings to gap, as it was shown by the abrupt decline of the maximal quantum yield of PSII photochemistry (F_v/F_m). The photoinhibition showed the time-dependent dynamics, as the F_v/F_m of the seedlings transferred to the forest gap recovered completely after 110 days. Furthermore, the net photosynthetic rate increased 3.5-fold in relation to priorexposure values. In summary, these data indicated that more than 21 days was required for the shade-acclimated seedlings to recover from photoinhibition and to relax induction photosynthetic limitations following the sudden exposure to high light. Moreover, the species responded very quickly to light availability; it highlights the importance of sunflecks to understory seedlings.     

EFFECTS OF LONG TERM EXPOSURE TO LOW TEMPERATURE ON THE PHOTOSYNTHETIC APPARATUS OF DUNALIELLA TERTIOLECTA (CHLOROPHYCEAE)1

The effects of long term exposure to suboptimal growth temperature on the photosynthetic apparatus of Dunaliella tertiolecta Butcher were investigated using carbon fixation rate versus irradiance curves and the variable fluorescence induction method. Carbon fixation rates per unite chlorophyll a at saturating (p^B_m) and subsaturating (α^B) irradiances were 55% and 39% lower, respectively, at 12° C than at 20° C. Chlorophyll a quotas and the spectrally averaged in vivo absorption cross section normalized to chlorophyll a (a^*) were not significantly different at these two temperatures. Analysis of the fluorescence kinetics revealed 1) no significant variations of the amount of PSII photoactive reaction centers per unit chlorophyll a, 2) a 14% decrease of the PSII quantum yield(+) and 3) a 29% decrease of the energy transfer efficiency between the light harvesting chlorophyll a pigment bed and the PSII reaction centers. The decrease in energy transfer efficiency between the antennae and the PSII reaction centers at 12° C was interpreted as a mechanism to avoid photoinhibition.     

PHOTOSYNTHESIS AND CELL DIVISION BY ANTARCTIC MICROALGAE: COMPARISON OF BENTHIC,PLANKTONIC AND ICE ALGAE1

Irradiance-dependent rates of photosynthesis and cell division of six species of microalgae isolated from the benthos, plankton and sea ice microbial community in McMurdo Sound, Antarctica were compared. Microalgae isolated from different photic environments had distinct photosynthetic and growth characteristics. For benthic and ice algae, photosynthesis saturated at 6 to 20 μE.m^?2.s^?1 and was photoinhibited at 10 to 80 μE.m^?2.s^?1 while for the planktonic algae, saturation irradiances were up to 13 times higher and photoinhibition was not detected. The slope of the light-limited portion of the P-I relationship was up to 50 times greater for the benthic algae than for either the ice or planktonic algae suggesting that benthic algae used the low irradiances more efficiently for carbon uptake. Cell division was dependent on the incubation irradiance for all but one microalga examined. The dependence of division rates on irradiance was however much smaller than for carbon uptake, suggesting that cell division buffers the influence of short term variations of irradiance on cellular metabolism.