Photosynthetic characteristics of phycoerythrin-containing marine Synechococcus spp.

Marine Synechococcus spp. are sufficiently abundant to make a significant contribution to primary productivity in the ocean. They are characterized by containing high cellular levels of phycoerythrin which is highly fluorescent in vivo. We sought (Jan.–Apr., 1984) to determine the adaptive photosynthetic features of two clonal types of Synechococcus spp., and to provide a reliable physiological basis for interpreting remote sensing data in terms of the biomass and productivity of this group in natural assemblages. It was found that the two major clonal types optimize growth and photosynthesis at low photon flux densities by increasing the numbers of photosynthetic units per cell and by decreasing photosynthetic unit size. The cells of clone WH 7803 exhibited dramatic photoinhibition of photosynthesis and reduction in growth rate at high photon flux densities, accompanied by a large and significant increase in phycoerythrin fluorescence. Maximal photosynthesis of cells grown under 10–50 E m^-2 s^-1 was reduced by 20 to 30% when the cells were exposed to photon flux densities greater than 150 E m^-2 s^-1. However, steady-state levels of photosynthesis maintained for brief periods under these conditions were higher than those of cells grown continuously at high photon flux densities. No photoinhibition occurred in clone WH 8018 and rates of photosynthesis were greater than in WH 7803. Yields of in-vivo phycoerythrin fluorescence under all growth photon flux densities were lower in clone WH 8018 compared to clone WH 7803. Since significant inverse correlations were obtained between phycoerythrin fluorescence and P_max and for both clones grown in laboratory culture, it may be possible to provide a reliable means of assessing the physiological state, photosynthetic capacity and growth rate of Synechococcus spp. in natural assemblages by remote sensing of phycoerythrin fluorescence. Poor correlations between phycoerythrin fluorescene and pigment content indicate that phycoerythrin fluorescence may not accurately estimate Synechococcus spp. biomass based on pigment content alone.     

Photoadaptation of photosynthesis in Gonyaulax polyedra

Gonyaulax polyedra Stein exhibited a combination of photoadaptive strategies of photosynthesis when only a single environmental variable, the light intensity during growth, was altered. Which of several biochemical/physiological adjustments to the light environment were employed depended on the level of growth irradiance. The photoadaptive strategies employed over any small range of light levels appeared to be those best suited for optimizing photosynthetic performance and not photosynthetic capacity. (Photosynthetic performance, P _i, is defined as the rate of photosynthesis occurring at the level of growth irradiance.) Among all photosynthetic parameters examined, only photosynthetic performance showed a consistent correspondence to growth rates of G. polyedra. Above 3500 to 4000 W cm^-2, where photosynthetic performance was equal to photosynthetic capacity, cells were not considered light-limited in either photosynthesis or growth. At these higher light levels, photosynthetic perfomance, cell volume, growth rates and respiration rates remained maximal; photosynthetic pigment content varied only slightly, while the photosynthetic capacity of the cells declined. At intermediate light levels (3000 to 1500 W cm^-2), photosynthesis, not growth, was light-limited, and photoadaptive strategies were induced which enhance absorption capabilities and energy transfer efficiencies of chlorophyll a to the reaction centers of G. polyedra. Photosynthetic capacity remained constant at about 280 mol O₂ cm^-3 h^-1, while photosynthetic performance ranged from 100 to 130 mol O₂ cm^-3 h^-1. Major increases in photosynthetic pigments, especially peridinin-chlorophyll a-proteins and an unidentified chlorophyll c component, accompanied photoadaptation to low irradiances. Maximal growth rates of 0.3 divisions day^-1 were maintained, as were respiration rates of about-80 mol O₂ cm^-3 h^-1 and cell volumes of about 5.4×10^-8 cm^-3 cell^-1. Below about 1250 W cm^-2, photosynthesis in G. polyedra was so light-limited that photosynthetic performance was unable to support maximal growth rates. Under these conditions, G. polyedra displayed photostress responses rather than photoadaptive strategies. Photostress was manifested as reduced cell volumes, slower growth, and drastic reductions in pigmentation, photosynthetic capacity, and rates of dark respiration.     

High-temperature tolerance of photosynthesis in the red alga Chondrus crispus

Chondrus crispus (Stackhouse) is a perennial red seaweed, common in intertidal and shallow sublittoral communities throughout the North Atlantic Ocean. In the intertidal zone, C. crispus may experience rapid temperature changes of 10 to 20C° during a single immerison-emerision cycle, and may be exposed to temperatures that exceed the thermal limits for long-term survival. C. crispus collected year-round at Long Cove Point, Chamberlain, Maine, USA, during 1989 and 1990, underwent phenotypic acclimation to growth temperature in the laboratory. This phenotypic acclimation enhanced its ability to withstand brief exposure to extreme temperature. Plants grown at summer seawater temperature (20°C) were able to maintain constant rates of lightsaturated photosynthesis at 30°C for 9 h. In contrast, light-saturated photosynthetic rates of plants grown at winter seawater temperature (5°C) declined rapidly following exposure to 30°C, reached 20 to 25% of initial values within 10 min, and then remained constant at this level for 9 h. The degree of inhibition of photosynthesis at 30°C was also dependent upon light intensity. Inhibition was greatest in plants exposed to 30°C in darkness or high light (600 mol photons m^-2s^-1) than in plants maintained under moderate light levels (70 to 100 mol photons m^-2s^-1). Photosynthesis of 20°C-acclimated plants was inhibited by exposure to 30°C in darkness or high light, but the degree of inhibition was less than that exhibited by 5°C-grown plants. Not only was light-saturated photosynthesis of 20°C plants less severely inhibited by exposure to 30°C than that of 5°C plants, but the former also recovered faster when they were returned to growth conditions. The mechanistic basis of this acclimation to growth temperature is not clear. Our results indicate that there were no differences between 5 and 20°C-grown plants in the thermal stability of respiration, electron transport associated with Photosystems I or II, Rubisco or energy transfer between the phycobilisomes and Photosystem II. Overall, our results suggest that phenotypic acclimation to seawater temperature allows plants to tolerate higher temperatures, and may play an important role in the success of C. crispus in the intertidal environment.     

Ecotypic differentiation in the marine diatom Skeletonema costatum: influence of light intensity on the photosynthetic apparatus

Three genetically distinct clones of Skeletonema costatum (Grev.) Cleve were grown at 20°C under high (274 E m^-2 s^-1) and low (27 E m^-2 s^-1) light conditions and their photoadaptive photosynthetic responses compared. When all three clones were grown under low light, pigment analyses and fluorescence excitation spectra demonstrated that the accessory pigments, chlorophyll c and fucoxanthin, became more important in light-harvesting compared to chlorophyll a. Photosynthetic unit sizes increased for Photosystems I and II in low light, but photosynthesis vs irradiance characteristics were not reliable predictors of photosynthetic unit features. Fluorescence excitation spectra and photosynthesis vs irradiance (P-I) relationships indicated that changes in energy transfer occurred independent of changes in pigment content. Large increases in accessory pigment content were not accompanied by large increases in excitation from these pigments. Changes in energy transfer properties were as important as changes in PSU size in governing the photoadaptive responses of S. costatum. When the three clones were grown under identical conditions, each had a separate and distinct pattern of photoadaptation. Significant differences among clones were found for pigment ratios, photosynthetic unit sizes for Photosystems I and II and efficiency of energy transfer between pigments. These strikingly different photoadaptive strategies among clones may partially account for the great ecological success of the diatom species. This is the first quantitative investigation of the importance of both chlorophyll c and fucoxanthin to the adaptive responses of diatoms to light intensity, and represents the most complete characterization of the photoadaptive responses of a single species of marine phytoplankter to differences in light environment.     

Photoinhibition at low quantum flux densities in a marine dinoflagellate (Amphidinium carterae)

Growth and photosynthetic properties of the marine dinoflagellate Amphidinium carterae Hulbert were examined under continuous illumination in batch cultures at four different irradiances between 2 and 150 E m^-2 s^-1. The slope of both cell- and Chl a-based photosynthesis versus the irradiance curves was greatest for cells grown at 15 E m^-2 s^-1. The relative Chl a values cell^-1 were 1, 1.5 and 2 for cultures grown at 150, 80 and 15 E m^-2 s^-1, respectively. A low-temperature (-196°C) fluorescence technique was used to examine cells for photoinhibiton. Photoinhibition was greatest for cells grown at 150 E m^-2 s^-1. However, significant photoinhibition of this species was noted even at 80 E m^-2 s^-1. No significant difference in the fluorescence pattern was found between cells grown at 2 and 15 E m^-2 s^-1. Time course studies indicate that photoinhibition may occur within 2 h following exposure to 350 E m^-2 s^-1 in cells grown at 15 E m^-2 s^-1 and is reversible when light levels are lowered within 4 h. The ecological significance of phytoplankton unable to cope with excess photosynthetic excitation energy is discussed.     

Photoadaption in marine phytoplankton: Response of the photosynthetic unit

Some species of phytoplankton adapt to low light intensities by increasing the size of the photosynthetic unit (PSU), which is the ratio of light-harvesting pigments to P700 (reaction-center chlorophyll of Photosystem I). PSU size was determined for 7 species of marine phytoplankton grown at 2 light intensities: high (300 E m^-2 s^-1) and low (4 E m^-2 s^-1); PSU size was also determined for 3 species grown at only high light intensity. PSU size varied among species grown at high light from 380 for Dunaliella euchlora to 915 for Chaetoceros danicus. For most species grown at low light intensity, PSU size increased, while the percentage increase varied among species from 13 to 130%. No change in PSU size was observed for D. euchlora. Photosynthetic efficiency per chlorophyll a (determined from the initial slope of a curve relating photosynthetic rate to light intensity) varied inversely with PSU size. In contrast, photosynthetic efficiency per P700 was enhanced at larger PSU sizes. Therefore, phytoplankton species with intrinsically large PSU sizes probably respond more readily to the rapid fluctuations in light intensity that such organisms experience in the mixed layer.Contribution No. 1180 from the Department of Oceanography, University of Washington, Seattle, Washington, USA     

Photosynthesis and amelioration of desiccation in the intertidal saccate alga Colpomenia peregrina

Rates of gross photosynthesis for the intertidal saccate alga Colpomenia peregrina (Sauv.) Hamel were determined under submersed and emersed conditions. Maximal photosynthetic rates were lower than for most seaweeds but comparable with other saccate members of the genus. By fitting the data to a hyperbolic tangent function, maximal photosynthetic rates were estimated to be 5.29 mmol CO₂ m^-2 h^-1 under submersed conditions and 2.06 mmol CO₂ m^-2 h^-1 under emersed conditions. I_k for submersed thalli was 69.1 E m^-2 s^-1, wherea for emersed thalli it was 149.0 E m^-2 s^-1, or 2.2 times higher. At low tide in the field and under saturating irradiance, carbon from seawater retained within the thallus cavity was assimilated at 0.9 mmol CO₂ m^-2 h^-1. In the laboratory under emersed conditions, carbon from this source was taken up at 0.6 mmol CO₂ m^-2 h^-1 at 20°C and at 0.34 mmol CO₂ m^-2 h^-1. Retained seawater also greatly reduced drying under desiccating conditions. Experimental thalli from which seawater had been removed lost thallus water continuously throughout the drying period (120 min). On the other hand, control, thalli lost water for the first 15 min, after which no further water loss occurred. At the termination of the experiment, control thalli had lost 7.2% of their water, whereas experimental thalli had lost 39.2%. Desiccation affected the emersed photosynthetic rate of experimental and control thalli. Emersed photosynthetic rates for thalli dried for 15 min were higher than for fullyhydrated thalli. However, emersed photosynthesis of thalli dried for longer than 15 min was lower than fully-hydrated rates and was directly related to percent water loss. Utilizing data from this study, a model was constructed to determine total photosynthetic production of C. peregrina over a single daylight period. From these calculations it was determined that emersed photosynthesis can increase daily photosynthetic production of C. peregrina by 50%.     

Photosynthetic characteristics of Prochloron sp./ascidian symbioses

The prokaryotic green alga Prochloron sp. (Prochlorophyta) is found in symbiotic association with colonial didemnid ascidians that inhabit warm tropical waters in a broad range of light environments. We sought to determine the light-adaptation features of this alga in relation to the natural light environments in which the symbioses are found, and to characterize the temperature sensitivity of photosynthesis and respiration of Prochloron sp. in order to assess its physiological role in the productivity and distribution of the symbiosis. Colonies of the host ascidian Lissoclinum patella were collected from exposed and shaded habitats in a shallow lagoon in Palau, West Caroline Islands, during February and March, 1983. Some colonies from the two light habitats were maintained under conditions of high light (2 200 E m^–2 s^–1) and low light (400 E m^–2 s^–1) in running seawater tanks. The environments were characterized in terms of daily light quantum fluxes, daily periods of light-saturated photosynthesis (H_sat), and photon flux density levels. Prochloron sp. cells were isolated from the hosts and examined for their photosynthesis vs irradiance relationships, respiration, pigment content and photosynthetic unit features. In addition, daily P:R ratios, photosynthetic quotients, carbon balances and photosynthetic carbon release were also characterized. It was found that Prochloron sp. cells from low-light colonies possessed lower chlorophyll a/b ratios, larger photosynthetic units sizes based on both reaction I and reaction II, similar numbers of reaction center I and reaction center II per cell, lower respiration levels, and lower P_max values than cells from high-light colonies. Cells isolated from low-light colonies showed photoinhibition of P_max at photon flux densities above 800 E m^–2 s^–1. However, because the host tissue attenuates about 60 to 80% of the incident irradiance, it is unlikely that these cells are normally photoinhibited in hospite. Collectively, the light-adaptation features of Prochloron sp. were more similar to those of eukaryotic algae and vascular plant chloroplasts than to those of cyanobacteria, and the responses were more sensitive to the daily flux of photosynthetic quantum than to photon flux density per se. Calculation of daily minimum carbon balances indicated that, though high-light cells had daily P:R ratios of 1.0 compared to 4.6 for low-light cells, the cells from the two different light environments showed nearly identical daily carbon gains. Cells isolated from high-light colonies released between 15 and 20% of their photosynthetically-fixed carbon, levels sufficient to be important in the nutrition of the host. Q₁₀ responses of photosynthesis and respiration in Prochloron sp. cells exposed briefly (15–45 min) to temperatures between 15° and 45°C revealed a discontinuity in the photosynthetic response at the ambient growth temperatures. The photosynthetic rates were found to be more than twice as sensitive to temperatures below ambient (Q₁₀=3.47) than to temperatures above ambient (Q₁₀=1.47). The Q₁₀ for respiration was constant (Q₁₀=1.66) over the temperature range examined. It appears that the photosynthetic temperature sensitivity of Prochloron sp. may restrict its distribution to warmer tropical waters. The ecological implications of these findings are discussed in relation to published data on other symbiotic systems and free-living algae.     

Diurnal patterns in photosynthetic capacity and depth-dependent photosynthesis-irradiance relationships inSynechococcus spp. and larger phytoplankton in three water masses in the Northwest Atlantic Ocean

The photosynthetic characteristics of prokaryotic phycoerythrin-rich populations of cyanobacteriaSynechococcus spp. and larger eukaryotic algae were compared at a neritic frontal station (Pl), in a warm-core eddy (P2), and at Wilkinson's Basin (P3) during a cruise in the Northwest Atlantic Ocean in the summer of 1984.Synechococcus spp. numerically dominated the 0.6 to 1 m fraction, and to a lesser extent the 1 to 5 m size fractions, at most depths at all stations. At P2 and P3, all three size categories of phytoplankton (0.6 to 1 m, 1 to 5 m, and >5 m) exhibited similar depth-dependent chages in both the timing and amplitude of diurnal periodicities of chlorophyllbased and cell-based photosynthetic capacity. Midday maxima in photosynthesis were observed in the upper watercolumn which damped-out in all size fractions sampled just below the thermocline. For all size fractions sampled near the bottom of the euphotic zone, the highest photosynthetic capacity was observed at dawn. At all depths, theSynechococcus spp.-dominated size fractions had lower assimilation rates than larger phytoplankton size fractions. This observation takes exception with the view that there is an inverse size-dependency in algal photosynthesis. Results also indicated that the size-specific contribution to potential primary production in surface waters did not vary appreciably over the day. However, estimates of the percent contribution ofSynechococcus spp. to total primary productivity in surface waters at the neritic front were significantly higher when derived from short-term incubator measurements of photosynthetic capacity rather than from dawn-to-duskin situ measurements of carbon fixation. The discrepancy was not due to photoinhibitory effects on photosynthesis, but appeared to reflect increased selective grazing pressure onSynechococcus spp. in dawn-to-dusk samples. Low-light photoadaptation was evident in analyses of the depth-dependency ofP-I parameters (photosynthetic capacity,P _max; light-limited slope, alpha;P _max alpha,I _k ; light-intensity beyond which photoinhibition occurs,I _b ) of the > 0.6 m communities at all three stations and was attributable to stratification of the water column. There was a decrease in assimilation rates andI _k with depth that was associated with increases in light-limited rates of photosynthesis. No midday photoinhibition ofP _max orI _b was observed in any surface station. Marked photoinhibition was detected only in the chlorophyll maximum at the neritic front and below the surface mixed-layer at Wilkinson's Basin, where susceptibility to photoinhibition increased with the depth of the collected sample. The 0.6 to 1 m fraction always had lower light requirements for light-saturated photosynthesis than the > 5 m size fraction within the same sample. Saturation intensities for the 1 to 5 m and 0.6 to 1 m size fractions were more similar whenSynechococcus spp. abundances were high in the 1 to 5 m fraction. The > 5 m fraction appeared to be the prime contributor to photoinhibitory features displayed in mixed samples (> 0.6 m) taken from the chlorophyll maxima. InSynechococcus spp.-dominated 0.6 to 1 and 1 to 5 m size fractions, cellular chlorophylla content increased 50- to 100-fold with depth and could be related to increases in maximum daytime rates of cellularP _max at the base of the euphotic zone. Furthermore, the 0.6 to 1 m and > 5 m fractions sampled at the chlorophyll maximum in the warm-core eddy had lower light requirements for photosynthesis than comparable surface samples from the same station. Results suggest that photoadaptation in natural populations ofSynechococcus spp. is accomplished primarily by changing photosynthetic unit number, occuring in conjuction with other accommodations in the efficiency of photosynthetic light reactions.     

Photosystem II photoinhibition and altered kinetics of photosynthesis during nutrient-dependent high-light photoadaptation in Gonyaulax polyedra

Gonyaulax poledra Stein was transferred at different cell densities from increasingly nutrient-limited low-light (LL, 80 E m^-2 s^-1) batch-cultures to high-light (HL, 330 E m^-2 s^-1) growth conditions. Several age-dependent differences in HL-adaptation strategies were apparent. Short-term (3h) susceptibility to photosynthetic photoinhibition increased with culture age, with light-limited rates of photosynthesis exhibiting greater photosuppression than light-saturated rates at all stages of growth. These shortterm changes were not accompanied by photobleaching of chlorophyll but were directly related to age-dependent photoinactivation of Photosystem II electron-transport rates. The capacity of electron transport by Photosystem I was only slightly affected. Prolonged exposure of LL log-phase cells to HL conditions did induce photobleaching of chlorophyll associated with increased cell volume, a transient decrease of organic carbon and nitrogen content, enhanced cellular-, carbon-and chlorophyll-based rates of light-saturated photosynthesis (P _max) and suppressed cellular rates of light-limited photosynthesis. As a result, the density of LL log-phase cells doubled and their cellular photosynthetic performance nearly tripled within 1 d of HL exposure while cellular respiratory demands remained unchanged. By contrast, prolonged HL incubation of LL stationary populations induced a transitory burst in cell division and a large reduction in cell volume, leading to a short-term increase in volume-based organic carbon and nitrogen content. Despite reduced cell volume and lowered carbon demand, the cellular-, carbon-and chlorophyll-based rates of P _max in nondividing populations fell by 64, 48 and 27%, respectively, over a 4 d exposure to HL, while light-limited rates were almost fully suppressed within 1 d and chlorophyll a content was reduced by 56%. As a result, the photosynthetic performance of LL-aged cells declined immediately under HL conditions. Addition of inorganic nutrients to LL stationary cultures at the time of HL transfer led to immediate and complete suppression of photosynthesis and cell lysis within 1 d. Addition of nutrients following transfer to HL induced cell responses intermediate to those described for LL log and aged cells exposed to HL. Results support the view that declining nutrient-status impairs HL photoadaptive responses in phytoplankton populations and that the rate and pattern of photoadaptive responses may be used as physiological growth indicators in field studies. The study was conducted from March 1981 to May 1983.     

Photoinhibition and recovery of the kelp Laminaria saccharina at optimal and superoptimal temperatures

Effects of high irradiance on photosynthetic characteristics were examined in sporophytes of the kelp Laminaria saccharina Lamour. from 1992 to 1994. Exposure to high irradiance (700 mol photons m^-2s^-1) for 1 h at optimal temperature (12°C) caused a 40 to 60% decline in photosynthetic efficiency (alpha), quantum yield, and the ratio of variable to maximum chlorophyll fluorescence (F_v/F_m), an indicator of Photosystem II efficiency. Although the photoinhibition effects were partly attributable to protective mechanisms, a concurrent increase in minimal fluorescence (F_o) indicated damage to Photosystem II reaction centers. The magnitude of photoinhibition was proportional to irradiance and duration; however, F_v/F_m was significantly reduced after exposure to irradiances as low as 40 to 50 mol photons m^-2s^-1 for 1 h, or to 700 mol photons m^-2s^-1 for only 5 min. In contrast, photosynthetic capacity (P_max) was affected only at much higher irradiance. Superoptimal temperatures up to 24°C did not exacerbate high-light effects. At 25°C, however, alpha and P_max were more susceptible to photoinhibition than at lower temperatures. Recovery from photoinhibition was examined by following F_v/F_m and F_o for 24 h after exposure to high light. Recovery of F_v/F_m was fastest during the first 1 to 3 h, and slowed or ceased after 6 to 12 h, while recovery of F_o was relatively constant over 12 h. Dithiothreitol, which blocks formation of energy-dissipating xanthophylls, reduced both the initial rate and extent of recovery. Chloramphenicol, which blocks chloroplast-encoded protein synthesis, had little effect on initial rates of recovery, but stopped recovery after 3 h. Thus, L. saccharina appears to rely on the xanthophyll cycle to protect the photosynthetic apparatus, and reversal of this protective mechanism causes the rapid initial recovery in F_v/F_m. Longterm recovery depends on repair of damaged reaction centers. Both the rate and extent of recovery were temperature-dependent. The initial rate was higher at 18 to 22°C than at 12°C, but the extent of recovery over 24 h declined with increasing temperature. High temperatures, therefore, appear to enhance protective mechanisms, but disrupt repair processes. L. saccharina from Long Island Sound, an ecotype adapted to low light and high temperature, showed slightly but consistently greater effects of photoinhibition than plants from the Atlantic coast of Maine, but exhibited faster recovery at superoptimal temperatures.     

Time-course of photoadaptation in the photosynthesis-irradiance relationship of a dinoflagellate exhibiting photosynthetic periodicity

Cultures of the marine dinoflagellate Glenodinium sp. were light-shifted and rates of photoadaptation determined by monitoring changes in cell volume, growth rate, pigmentation, parameters of the photosynthesisirradiance (P-I) curves and respiration. To approximate physiological conditions of field populations, cells were cultured on an alternating light-dark cycle of 12hL:12hD, which introduced a daily periodicity of photosynthesis. One result of the present study was to demonstrate how specific parameters of the P-I relationship influenced by periodicity of the light: dark cycle are distinguished from photosynthetic parameters influenced by changes in light level. Under steady-state conditions, rates of both light-saturated (P_max) and light-limited photosynthesis changed in unison over the day; these changes were not related to pigmentation, and displayed their maxima midday. This close relationship between P_max and the slope (a) of the cellular P-I curves in steadystate conditions was quickly adjusted when growth illumination was altered. Rates of light-limited photosynthesis were increased under low light conditions and the periodicity of cellular photosynthesis was maintained. The short-term responses of the P-I relationship to changing light level was different, depending on (1) whether the light shift was from high to low light or vice versa, and (2) whether the high light levels were sufficient to promote maximal photosynthesis rates. Major increases in the photosynthetic carotenoid peridinin, associated with a single type of light-harvesting chromo protein in the chloroplast, was observed immediately upon shifting high light cultures to low light conditions. Following pigment synthesis, significant increases in rates of light-limited photosynthesis were observed in about one-tenth the generation time, while cellular photosynthetic potential was unaffected. it is suggested that general results were consistent with suggested that general results were consistent with earlier reports that the major photoadaptive strategy of Glenodinium sp. is to alter photosynthetic unit (PSU) size. Photoadaptive response times to high light were light-dependent, but appeared to be shower than photoadaptive responses to low light. If light intensities were bright enough to maximize growth rates, photosynthetic response times were on the order of a generation period and pigmentation fell quickly as cells divided at a faster rate. If light-intensities were not sufficient to maximize growth rates, then pigment content did not decline, while rates of light-limited photosynthesis declined quickly. In all cases, photoadaptation was followed best by monitoring fast changes in half saturation constants for photosynthesis, rather than fluctuating changes in pigmentation. Results compared well with time-course phenomena reported for other groups of phytoplankton. Overall, results suggest phytoplankton can bring about photo-induced changes in photosynthesis very quickly and thus accommodate widely fluctuating light regimes over short periods of time.     

Effect of freezing on photosynthesis of intertidal macroalgae: relative tolerance of Chondrus crispus and Mastocarpus stellatus (Rhodophyta)

The effect of freezing on photosynthetic metabolism was studied in the red algae, Chondrus crispus and Mastocarpus stellatus. Plants of both species were collected from the intertidal at Chamberlain or Kresge Point, Maine, USA (43°56N, 69°54W) between February and March 1987. Photosynthetic rates were measured immediately after freezing at-20°C and following recovery periods in seawater. Photosynthesis in C. crispus declined rapidly following freezing, falling to 70% of control values within 1 h and 30% after 3 h exposure. Minimum photosynthetic rates (7 to 9% of controls) occurred following freezing exposures of 12 h or more. Full photosynthetic recovery in C. crispus after 3 h at-20°C required 48 h. Photosynthesis in C. crispus did not fully recover in plants frozen for 6 h or more. In contrast, photosynthesis in M. stellatus was relatively unaffected by freezing exposures of <12 h. Twelve hours or more at-20°C reduced photosynthesis to 55% of controls. Photosynthesis in M. stellatus fully recovered from 24 h at-20°C within 24 h. In both species the reduction of photosynthesis by freezing was associated with damage to the plasma membrane and reduced efficiency of energy transfer from phycobilisomes to chlorophyll a, but did not appear to involve ribulose-1,5-bisphosphate carboxylase oxygenase activity. The freezing tolerance of C. crispus and M. stellatus positively correlates with their respective intertidal distributions, suggesting that freezing may be involved in controlling the distributions of these species on the shore.     

Photosynthetic characteristics of giant kelp (Macrocystis pyrifera) determined in situ

Rates of net photosynthesis and nocturnal respiration by individual blades of the giant kelp Macrocystis pyrifera (L.) C. Agardh in southern California, were determined in situ by measuring oxygen production in polyethylene bags during spring/summer of 1983. Mature blades from different depths in the water column exhibited different photosynthetic characteristics. Blades from the surface canopy (0 to 1 m depth) exhibited higher photosynthetic capacity under saturating irradiance and higher photosynthetic efficiency at low irradiances than blades from 3 to 5 or 7 to 9 m depths. Saturating irradiance was lower for canopy blades than for deeper blades. Canopy blades showed no short-term photoinhibition, but photosynthetic rates of deeper blades were significantly reduced during 1 to 2 h incubations at high irradiances. Results of 1 to 2 wk acclimation experiments indicated that differences between photosynthetic characteristics of blades from different depths were primarily attributable to acclimation light conditions. Vertical displacement of blades within the kelp canopy occurred on a time-scale of 1 min to 1 h. Blades continually moved between the unshaded surface layer and deeper, shaded layers. Vertical movement did not maximize photosynthesis by individual blades; only a small proportion of blades making up a dense surface canopy maintained light-saturated photosynthetic rates during midday incubations. The relatively high photosynthetic rates exhibited by canopy blades over the entire range of light conditions probably resulted from acclimation to intermittent high and low irradiances, a consequence of vertical displacement. Vertical displacement also reduced the afternoon depression in photosynthesis of individual canopy blades. The overall effect of vertical displacement was optimization of total net photosynthesis by the kelp canopy and, therefore, optimization of whole-plant production.     

Polychlorinated biphenyl (PCB) effects on marine phytoplankton photosynthesis and cell division

Inhibition of photosynthesis and cell division by polychlorinated biphenyls (PCBs) was studied using 7 marine phytoplankton species representing 4 algal classes. PCB concentrations as low as 1.0 g l^-1 reduced cell division of Thalassiosira pseudonana 3H and Isochrysis galbana. Both photosynthesis and cell division of T. pseudonana 3H, Chaetoceros socialis, Skeletonema costatum, T. pseudonana 13-1, Monochrysis, lutheri and I. galbana were inhibited at a PCB concentration of 10.0 g l^-1. The effects on photosynthesis were immediate and probably resulted in reduced rates of cell division. Interspecific differences in susceptibility were observed. These differences have significance with respect to primary production and the species composition of phytoplankton communities. The initial slopes of photosynthesis-irradiance (P-I) curves for the diatoms S. costatum and T. pseudonana 3H were reduced in the presence of PCBs. These results suggest that PCBs affect the photosynthetic light reactions.     

Photosynthetic studies of Chondrus crispus

The net photosynthesis of intertidal, subtidal, carposporic, tetrasporic, and winter versus summer acclimatized plants of Chondrus crispus Stackhouse were evaluated under different temperatures and quantities of light. The optimum temperature and light conditions for net photosynthesis of C. crispus are seasonally and spatially variable, and there is an adaptive shift in the photosynthetic capacity at different seasons and positions on the shore. Plants collected during the fall and winter had lower light optima (465 to 747 ft-c) for net photosynthesis than spring and summer specimens (about 1000 ft-c). Intertidal populations exhibited a higher rate of net photosynthesis between 250 and 2819 ft-c than subtidal plants. Summer materials have a greater tolerance to high temperatures and a higher temperature optimum than winter materials. Shallow subtidal populations (-6m) exhibited a higher temperature optimum than deep subtidal plants (-12m). Tetrasporic plants (diploid) showed a higher rate of net photosynthesis than carposporic plants (haploid). It is suggested that the diploid plants of C. crispus may extend deeper in the subtidal zone, because they have a higher rate of net photosynthesis than carposporic plants. The results of the present studies are compared with previous physiological studies of C. crispus.Published with the approval of the Director of the New Hampshire Agriculture Experiment Station as Scientific Contribution Number 742.     

Photosynthetic activity of the non-dormant <Emphasis Type="Italic">Posidonia oceanica</Emphasis> seed

The photosynthetic adaptive features of non-dormant seeds in Posidonia oceanica were studied in order to evaluate the effects of light on germination success. Transmission electron micrographs showed the presence of chloroplasts in the epidermal cells, close to the nucleus at the periphery of the cytoplasm. The well-developed thylakoid membranes and the presence of starch granules indicated that the chloroplasts were photosynthetically active. The relationship between photosynthesis versus irradiance in P. oceanica seeds incubated at 15 and 21°C was analysed. The net photosynthesis in the non-dormant seed of P. oceanica was positive and compensated its respiration demand (90 μmol quanta m⁻² s⁻¹) at both temperatures. Net photosynthesis was negative at the other irradiance values. To test the effects of light on germination success, seeds were placed both in dark and light conditions. Germination success was significantly higher in light rather than in dark condition. The characteristics observed in the photosynthesis in P. oceanica seed could be a mechanism to guarantee seedling survival in temperate waters, demonstrating though the specialized nature of this species.     

Photosynthesis-irradiance responses and photosynthetic periodicity in the sea anemone Aiptasia pulchella and its zooxanthellae

Sea anemones (Aiptasia pulchella) containing zooxanthellae (Symbiodinium microadriaticum) were maintained in a long-term laboratory culture on a 12 h light (100 E m^-2 s^-1):12 h dark cycle. Photosynthetic oxygen production was measured for the symbiotic association and for freshlyisolated zooxanthellae. Light utilization efficiencies () were similar for both sets of zooxanthellae, suggesting negligible shading of zooxanthellae by animal tissue in this association. Whereas freshly-isolated zooxanthellae were photoinhibited at high irradiances (800 to 1 800 E m^-2 s^-1), zooxanthellae in the host continued to function at photosynthetic capacity. Time of day may influence photosynthetic measurements in symbiotic organisms, as it was found that photosynthesis in A. pulchella followed a diel periodicity at both light-saturating (1 200 E m^-2 s^-1) and subsaturating (150 E m^-2 s^-1) irradiances. There was a peak period of photosynthesis between 12.00 and 14.00 hrs. Light stimulated dark respiration rates of A. pulchella. Dark respiration of sea anemones increased somewhat towards the end of the light cycle and was always greater after exposure to high irradiances.     

Gene flow among population of a teleost (painted greenling,Oxylebius pictus) from Puget Sound to southern California

Net photosynthetic oxygen evolution in Amphiroa anceps (Lamarck) Decaisne is inhibited at high oxygen concentrations. Photosynthesis is highest between pH 6.5 and 7.5. At pH 9 to 10 there is still a significant photosynthetic rate, suggesting that this alga can use HCO _- ³ as a substrate for photosynthesis. At pH 7.0 to 8.5, the photosynthetic rate saturates at a total inorganic carbon concentration (C_i) greater than 3 mM. At pH 8.5 and 8.8, calcification rate continues to increase with increasing concentration of C_i. Between pH 7 and 9, the calcification rate in the light in A. foliacea Lamouroux is proportional to the photosynthetic rate, whereas at higher pH where the photosynthetic rate is very low, the calcification rate is stimulated by the higher concentration of CO _2- ³ ion. At all pH values examined, the calcification rate of living plants in the dark and of dead plants is directly proportional to the CO _2- ³ ion concentration, suggesting little metabolic involvement in calcification processes in the dark, whereas calcification in live A. foliacea in the light is influenced both by the photosynthetic rate and the CO _2- ³ ion concentration in the medium.     

Photosynthetic and optical properties of the marine chlorophyte Dunaliella tertiolecta grown under fluctuating light caused by surface-wave focusing

Photosynthetic and optical properties of the marine chlorophyte Dunaliella tertiolecta Butcher were studied in response to irradiance fluctuations caused by surface-wave focusing. The experimental conditions simulated the prominent features of the light field (high average irradiance, spectral composition and statistical properties) in the uppermost few meters of the water column under sunny surface conditions. The properties of algae grown under high-frequency fluctuations were compared with control cells grown under constant light at the same average irradiance (800 mol quantam^-2s^-1). No significant differences were found for a number of parameters, including growth rate, cellular chlorophyll a and pigment ratios, photosynthetic unit size and density of Photosystem I reaction centers, the rate of photosynthesis at the growth irradiance, dark respiration, and in vivo fluorescence of chlorophyll a per cell. Photosynthetic parameters were not affected by whether the incident light for oxygen exchange measurements was fluctuating or constant. This was the case whether the cells had been previously acclimated to either fluctuating or constant irradiance. Such a photosynthetic response indicates that cells are accomplishing a time integration of the fluctuating light. In addition, although D. tertiolecta is capable of dramatically changing its optical properties in response to low or high growth irradiance levels, the refractive index of the cells, the efficiency factors for light absorption and scattering by individual cells, and chlorophyll-specific absorption and scattering coefficients of cell suspensions, were all very similar under high irradiance, whether or not wave focusing was present.Contribution to the program of GIROQ (Groupe Interuniversitaire de Recherches Océanographiques du Québec)