Effect of sub-lethal damage to juvenile colonies of massive Porites spp. under contrasting regimes of temperature and water flow

In this study, juvenile colonies of massive Porites spp. (a combination of P. lutea and P. lobata) from the lagoon of Moorea (W 149°50′, S 17°30′) were damaged and exposed to contrasting conditions of temperature and flow to evaluate how damage and abiotic conditions interact to affect growth, physiological performance, and recovery. The experiment was conducted in April and May 2008 and consisted of two treatments in which corals were either undamaged (controls) or damaged through gouging of tissue and skeleton in a discrete spot mimicking the effects of corallivorous fishes that utilize an excavating feeding mode. The two groups of corals were incubated for 10 days in microcosms that crossed levels of temperature (26.7 and 29.6°C) and flow (6 and 21 cm s⁻¹), and the response assessed as overall colony growth (change in weight), dark-adapted quantum yield of PSII (F _v/F _m), and healing of the gouged areas. The influence of damage on growth was affected by temperature, but not by flow. When averaged across flow treatments, damage promoted growth by 25% at 26.7°C, but caused a 25% inhibition at 29.6°C. The damage also affected F _v/F _m in a pattern that differed between flow speeds, with a 10% reduction at 6 cm s⁻¹, but a 4% increase at 21 cm s⁻¹. Regardless of damage, F _v/F _m at 21 cm s⁻¹ was 11% lower at 26.7°C than at 29.6°C, but was unaffected by temperature at 6 cm s⁻¹. The lesions declined in area at similar rates (4–5% day⁻¹) under all conditions, although the tissue within them regained a normal appearance most rapidly at 26.7°C and 6 cm s⁻¹. These findings show that the response of poritid corals to sub-lethal damage is dependent partly on abiotic conditions, and they are consistent with the hypothesis that following damage, calcification and photosynthesis can compete for metabolites necessary for repair, with the outcome affected by flow-mediated mass transfer. These results may shed light upon the ways in which poritid corals respond to biting by certain corallivorous fishes.     

Juvenile life stages of the brown alga Fucus serratus L. are more sensitive to combined stress from high copper concentration and temperature than adults

The combined effects of exposure to copper and temperature were investigated in adult specimens and germlings of the canopy-forming brown alga Fucus serratus. A matrix of four temperatures, 6, 12, 17 and 22 °C, and three concentrations of copper, 0, 100 and 1,000 nM total copper were used. Measured endpoints were growth rate, chlorophyll fluorescence parameters and for germlings also survival. The growth rate of adult specimens of F. serratus changed with increasing temperature. Growth tended to be negatively affected by high concentrations of copper when exposed to heat (22 °C) though not significantly so. The photosynthetic performance (i.e., chlorophyll fluorescence parameters: F _v/F _m, maximum electron transport rate (ETR_max) and maximum non-photosynthetic quenching (NPQ_max) of adults was largely unaffected by both copper and temperature. Germling survival, growth rate and chlorophyll fluorescence parameters were affected by the combination of copper concentration and temperature. Increasing temperature led to reduced survival, increased rhizoid growth and higher F _v/F _m and ETR_max, whereas high copper concentration had a negative effect on the latter three endpoints. The negative effect of high copper concentration was amplified by high temperature. We conclude that juveniles of F. serratus are more susceptible to environmental stressors than adult specimens and recommend therefore including early life stages when assessing the risk of exposure to toxic compounds. Considering the response of adult specimens only may lead to false conclusions regarding the ecological impact of environmental stress.     

Effect of acclimatization to low temperature and reduced light on the response of reef corals to elevated temperature

This study tested the effects of acclimatization on the response of corals to elevated temperature, using juvenile massive Porites spp. and branching P. irregularis from Moorea (W149°50′, S17°30′). During April and May 2006, corals were acclimatized for 15 days to cool (25.7°C) or ambient (27.7°C) temperature, under shaded (352 μmol photons m⁻² s⁻¹) or ambient (554 μmol photons m⁻² s⁻¹) natural light, and then incubated for 7 days at ambient or high temperature (31.1°C), under ambient light (659 μmol photons m⁻² s⁻¹). The response to acclimatization was assessed as biomass, maximum dark-adapted quantum yield of PSII (F _v/F _m), and growth, and the effect of the subsequent treatment was assessed as F _v/F _m and growth. Relative to the controls (i.e., ambient temperature/ambient light), massive Porites spp. responded to acclimatization through increases in biomass under ambient temperature/shade, and low temperature/ambient light, whereas P. irregularis responded through reduced growth under ambient temperature/shade, and low temperature/ambient light. Acclimatization affected the response to thermal stress for massive Porites spp. (but not P. irregularis), with an interaction between the acclimatization and subsequent treatments for growth. This interaction resulted from a lessening of the negative effects of high temperature after acclimatizing to ambient temperature/shade, but an accentuation of the effect after acclimatizing to low temperature/shade. It is possible that changes in biomass for massive Porites spp. are important in modulating the response to high temperature, with the taxonomic variation in this effect potentially resulting from differences in morphology. These results demonstrate that corals can acclimatize during short exposures to downward excursions in temperature and light, which subsequently affects their response to thermal stress. Moreover, even con-generic taxa differ in this capacity, which could affect coral community structure. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Interactive effects of UV radiation and enhanced temperature on photosynthesis, phlorotannin induction and antioxidant activities of two sub-Antarctic brown algae

Lessonia nigrescens and Durvillaea antarctica, two large sub-Antarctic brown algae from the southern Chilean coast, were exposed to solar UV radiation in an outdoor system during a summer day (for 11 h) as well as to artificial UV radiation under controlled laboratory conditions at two temperatures (15 and 20 °C) for 72 h. Chlorophyll a fluorescence–based photoinhibition of photosynthesis was measured during the outdoor exposure, while electron transport rates, lipid peroxidation, antioxidant activity and content of phlorotannins were determined at different time intervals during the laboratory exposure. Under natural solar irradiances in summer, both species displayed well-developed dynamic photoinhibition: F _v/F _m values decreased by 70 % at noon coinciding with the levels of PAR >1,500 μmol m^?2 s^?1 and UV-B radiation >1 W m^?2 and recovered substantially in the afternoon. In treatments including UV radiation, recovery in D. antarctica started already during the highest irradiances at noon. The results from laboratory exposures revealed that (a) elevated temperature of 20 °C exacerbated the detrimental effects of UV radiation on photochemical parameters (F _v/F _m and ETR); (b) peroxidative damage measured as MDA formation occurred rapidly and was strongly correlated with the decrease in F _v/F _m, especially at elevated temperature of 20 °C; (c) the antioxidant activity and increases in soluble phlorotannins were positively correlated mainly in response to UV radiation; (d) phlorotannins were rapidly induced but strongly impaired at 20 °C. In general, short-term (2–6 h) exposures to enhanced UV radiation and temperature were effective to activate the photochemical and biochemical defenses against oxidative stress, and they continued operative during 72 h, a time span clearly exceeding the tidal or diurnal period. Furthermore, when algae were exposed to dim light and control temperature of 15 °C for 6 h, F _v/F _m increased and lipid peroxidation decreased, indicating consistently that algae retained their ability for recovery. D. antarctica was the most sensitive species to elevated temperature for prolonged periods in the laboratory. Although no conclusive evidence for the effect of the buoyancy of fronds was found, the interspecific discrepancies in thermo-sensitivity in the UV responses found in this study are consistent with various ecological and biogeographical differences described for these species.     

Metabolic properties of Northern krill, Meganyctiphanes norvegica, from different climatic zones. I. Respiration and excretion

Adaptive processes linked to overall metabolism were studied in terms of oxygen consumption and ammonia excretion in each of three self-contained krill populations along a climatic gradient. In the Danish Kattegat, krill were exposed to temperatures which ranged from 4°C to 16°C between seasons and a vertical temperature gradient of up to 10°C during summer. In the Scottish Clyde Sea, water temperatures varied less between seasons and the vertical temperature gradient in summer was only 3°C. Temperatures in the Ligurian Sea, off Nice, were relatively constant around 12-13°C throughout the year, with a thin surface layer (20-30 m) of warm water developing during summer. The trophic conditions were rich in the Kattegat and, particularly, in the Clyde, but comparatively poor in the Ligurian Sea. Oxygen consumption increased exponentially with increasing experimental temperature, which ranged from 4°C to 16°C. Overall respiration rates were between 19.9 and 89.9 µmol O₂ g^-1 dry wt h^-1. Krill from the Kattegat, the Clyde Sea, and the Ligurian Sea all exhibited approximately the same level of oxygen consumption (30-35 µmol O₂ g^-1 dry wt h^-1) when incubated at the ambient temperatures found in their respective environments (9°C, 5°C, and 12°C). This indicates that krill adjust their overall metabolic rates to the prevailing thermal conditions. The exception to this were the respiration rates of Ligurian krill from winter/spring, which were about twice as high as the rates from summer krill despite the fact that the thermal conditions were the same. This effect appears to result from enhanced somatic activity during a short period of increased food availability and reproduction. Accordingly, krill appears to be capable of adapting to both changing thermal and trophic conditions, especially when nutrition is a limiting factor in physiological processes.     

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.     

Short-term variability in the photosynthetic activity of microphytobenthos as detected by measuring rapid light curves using variable fluorescence

Short-term variability in the photosynthetic activity of microphytobenthos assemblages was studied by measuring chlorophyll fluorescence rapid light curves (RLC), using pulse amplitude modulated (PAM) fluorometry. Measurements carried out on undisturbed samples under dark–light cycles revealed large diel oscillations in both the initial slope of the RLC () and in the maximum relative electron transport rate (ETR_m). Short-term variations in RLC parameters were also observed, closely following changes in incident photon irradiance (E). Increases in irradiance were followed by decreases in and increases in ETR_m, resulting in significant correlations between the light-saturation parameter E_k and E. These results were interpreted as resulting from the onset of reversible energy-dissipating, non-photochemical quenching mechanisms and of compensatory high light-induced activation of carbon metabolism activity. Short-term RLC variability was shown to result mainly from physiological causes and to be detectable only by using short (10–20 s) light steps during RLC construction. Dark-adapted samples kept under constant conditions exhibited apparently endogenous rhythms in RLC parameters and in the maximum quantum yield, F_v/F_m, coincident with vertical migratory movements occurring during subjective photoperiods. These fluctuations appeared to result from the interaction between migratory rhythms and the physiological responses, and from the endogenous activation of processes affecting both the efficiency of energy transfer from light-harvesting antennae to the photosystem II (PSII) reaction centres or from non-radiative pathways (F_v/F_m, ) and the reactions downstream of PSII (ETR_m).Communicated by M. Kühl, Helsingør     

Metabolic properties of Northern krill, Meganyctiphanes norvegica, from different climatic zones. II. Enzyme characteristics and activities

Activities and characteristics of two metabolic key enzymes, citrate synthase (CS) and pyruvate kinase (PK), were studied in the Northern krill, Meganyctiphanes norvegica, with respect to adaptive properties under different thermal conditions. Krill were sampled during late winter/spring and summer from the constantly warm Ligurian Sea (12-13°C below the thermocline), the colder but also comparatively constant Clyde Sea (7-8°C), and the variable Kattegat (2-16°C). Both enzymes showed distinct tissue- and organ-specific activities, which were highest in the pleopods - the principal locomotive organs. The fourth and fifth abdominal segments, however, were used for routine investigation due to lowest variability. Specific activity of CS and PK did not differ between seasons in krill from the Kattegat or the Clyde Sea. In the Ligurian Sea, in contrast, specific CS activities were significantly lower during summer. Analysis of individual data illustrated a decrease of CS activity with size and an increase of PK activity with size. Taking these allometric effects into account, as emphasized by calculating the ratio between both enzymes, variation of CS and PK activities turned out to be solely dependent on body size, which differed between locations and seasons. Ligurian krill from the summer, however, were unique in that they showed a lower CS/PK ratio than would be predicted by the scaling effect. Thermal characteristics of each enzyme were similar between locations and seasons. During the winter, in Kattegat and Clyde Sea krill, K_m values (Michaelis-Menten constant) of CS towards acetyl-coenzyme A exhibited an almost constant level over the experimental temperature range of 4-16°C. During summer, however, K_m values were lower at 8°C in the Clyde Sea and at 12°C in the Kattegat. In Ligurian krill from the summer, K_m values were consistently lower than those of winter krill over the entire experimental temperature range. In conclusion, Kattegat and Clyde Sea krill show only minor adaptations to their respective thermal environments in terms of CS and PK characteristics. Ligurian krill, in contrast, exhibited decreased specific CS activity during summer, which might be compensated by elevated enzyme-substrate affinity as indicated by lower K_m values. Since temperature was constant during both seasons, this effect cannot be explained as a reaction to thermal conditions. Consequently, oligotrophic conditions in the Ligurian Sea during summer may entail a reduction in the somatic performance of krill, which is reflected by lower CS activity.     

Short-term variability and control of phytoplankton photosynthetic activity in a macrotidal ecosystem (the Strait of Dover, eastern English Channel)

Short-term changes in phytoplankton photosynthetic activity were studied during different periods of the years 2009 and 2010 in the coastal waters of a macrotidal ecosystem (the Strait of Dover, eastern English Channel). During each sampling period, samples were taken every 1.45 h., from sunrise to sunset, during at least 5 days distributed along a complete spring–neap tide cycle. The photosynthetic parameters were obtained by measuring rapid light curves using pulse amplitude modulated fluorometry and were related to environmental conditions and phytoplankton taxonomic composition. The maximum quantum yield (F _v/F _m) showed clear light-dependent changes and could vary from physiological maxima (0.68–0.60) to values close to 0.30 during the course of 1 day, suggesting the operation of photoprotective mechanisms. The maximum electron transport rate (ETR_m) and maximal light utilization efficiency (α) were generally positively correlated and showed large diel variability. These parameters fluctuated significantly from hour to hour within each day and the intraday pattern of variation changed significantly among days of each sampling period. Stepwise multiple linear regressions analyses indicated that light fluctuations explained a part of this variability but a great part of variability stayed unexplained. F _v/F _m, ETR_m and α were not only dependent on the light conditions of the sampling day but also on those of the previous days. A time lag of 3 days in the effect of light on ETR_m and α variation was highlighted. At these time scales, changes in phytoplankton community structure seemed to have a low importance in the variability in photosynthetic parameters. The photoacclimation index E _k showed a lower variability and was generally different from the incident irradiance, indicating a limited acclimation capacity with a poor optimization of light harvesting during the day. However, in well-mixed systems such as the Strait of Dover, the short-term photoacclimation is disrupted by the high level of variability in environmental conditions. Also, the variability observed in the present study can be associated with a particular kind of photosynthetic response: the “E _k-independent” variability. The physiological basis of this photosynthetic response is largely unresolved and further researches on this subject are still required to better explain the dynamics of phytoplankton activity in the Strait of Dover.     

Diurnal measurements of F v/F m used to improve productivity estimates in macroalgae

The ratio of variable fluorescence to maximal fluorescence (F _v/F _m) was measured during a night and day cycle in five different macroalgae growing in the littoral zone at the Swedish west coast; the green algae Ulva lactuca, Cladophora sp. and Enteromorpha flexousa, the red alga Ceramium nodulosum and the brown alga Fucus vesiculosus. All the green algae and C. nodulosum showed both diurnal fluctuations in F _v/F _m during days of high light intensities and a significant negative correlation between F _v/F _m and photon irradiance (PI). An attempt has been made to improve calculations of macroalgal net growth based on carbon fixation (α_net) considering this diurnal change in F _v/F _m. By assuming that the negative linear correlation between F _v/F _m and the maximum quantum yield for photosynthesis (Φ) is proportional to that between F _v/F _m and α_net, it was possible to include the daily variation of α_net due to photoinhibition. To compensate for the variation of F _v/F _m, a representative value for each day was obtained by weighting all values of F _v/F _m over the diurnal period in relation to total PI. For all algal species there was a fairly good agreement between this representative value and the F _v/F _m value measured around noon. As the daily representative F _v/F _m value showed a negative linear correlation with the daily mean PI, it was possible to correct α_net for differences of daily mean PI. Received: 10 March 1997 / Accepted: 25 August 1997     

Corallina and Ellisolandia (Corallinales,Rhodophyta) photophysiology over daylight tidal emersion: interactions with irradiance,temperature and carbonate chemistry

The photophysiology of three geniculate coralline algal species (Corallina officinalis, C. caespitosa and Ellisolandia elongata) was determined in intertidal rock pools in the south-west UK at Combe Martin (51°12′31N 4°2′19W) and Heybrook Bay (50°31′66N 4°11′41W), at the start, middle and end of summer (September 1 and 2) and winter (February 9 and 10) daylight tidal emersion periods, in relation to prevailing irradiance, temperature and carbonate chemistry conditions. Algal photophysiology was assessed from rapid light curves performed using pulse amplitude modulation fluorometry. Corallina and Ellisolandia experienced significant fluctuations in irradiance, temperature and carbonate chemistry over seasonal and tidal cycles. Rock pool carbonate chemistry was predictable (R ² = 0.82, P < 0.0001) by photodose (summed irradiance) plus water temperature, but not significantly related to photophysiology. In contrast, Corallina and Ellisolandia relative maximum electron transfer rate showed a significant negative relationship (R ² = 0.65, P < 0.0001) with irradiance plus water temperature. At a seasonal resolution, photoacclimation to maximize both light harvesting during winter months and photoprotection during summer months was observed for all species. Dynamic photoinhibition was apparent over both summer and winter tidal emersion, in relation to irradiance fluctuations. More effective photoinhibition was apparent during summer months, with greater sensitivity to irradiance and slower recovery in F _v/F _m, observed during winter. With sustained high irradiance over tidal emersion, the establishment of high pH/low inorganic carbon conditions may impact photochemistry. This study represents the first assessment of C. officinalis, C. caespitosa and E. elongata photophysiology underpinned by clear species concepts and highlights their ability to adapt to the dramatically fluctuating conditions experienced in intertidal rock pools.     

Size-dependent photosynthetic performance in the giant clam <Emphasis Type="Italic">Tridacna maxima</Emphasis>, a mixotrophic marine bivalve

Giant clams form a symbiosis with photosynthetic algae of the genus Symbiodinium that reside in clam mantle tissue. The allometry of symbiont photosynthetic performance was investigated as a mechanism for the increasing percentage of giant clam carbon respiratory requirements provided by symbionts as clam size increases. Chlorophyll fluorescence measurements of symbionts of the giant clam Tridacna maxima were measured during experiments conducted in September of 2009 using specimens 0.5–200 g tissue wet weight (3–25 cm long), collected from waters around southern Taiwan (N 21°36′, E 120°47′) from July to August of 2009. Light-dependent decreases in effective quantum yield (∆F/F _m′) calculated as the noontime maximum excitation pressure over PSII (Q _m), relative electron transport rates (rETR), and dark-adapted maximum quantum yield (F _v/F _m) all varied as a quadratic function of clam size. Both Q _m and rETR increased as clam size increased up to ~10–50 g then decreased as clam size increased. F _v/F _m decreased as clam size increased up to ~5–50 g then increased as clam size increased. Chlorophyll fluorescence measurements of rETR were positively correlated with gross primary production measured during chamber incubations. Overall, symbionts of mid-sized clams ~5–50 g exhibited the highest light-dependent decreases in effective photosynthetic efficiencies, the highest relative electron transport rates, and the lowest maximum photosynthetic efficiencies, and symbiont photosynthetic performance is allometric with respect to host clam size.     

Impact of temperature and species interaction on filamentous cyanobacteria may be more important than salinity and increased pCO2 levels

A future business-as-usual scenario (A1FI) was tested on two bloom-forming cyanobacteria of the Baltic Proper, Nodularia spumigena and Aphanizomenon sp., growing separately and together. The projected scenario was tested in two laboratory experiments where (a) interactive effects of increased temperature and decreased salinity and (b) interactive effects of increased temperature and elevated levels of pCO₂ were tested. Increased temperature, from 12 to 16 °C, had a positive effect on the biovolume and photosynthetic activity (F _v/F _m) of both species. Compared when growing separately, the biovolume of each species was lower when grown together. Decreased salinity, from 7 to 4, and elevated levels of pCO₂, from 380 to 960 ppm, had no effect on the biovolume, but on F _v/F _m of N. spumigena with higher F _v/F _m in salinity 7. Our results suggest that the projected A1FI scenario might be beneficial for the two species dominating the extensive summer blooms in the Baltic Proper. However, our results further stress the importance of studying interactions between species.     

Influence of ultraviolet-radiation on chlorophyll fluorescence and growth in different life-history stages of three species of Laminaria (phaeophyta)

Zoospores, gametophytes, young sporophytes and discs cut from mature sporophytes of Laminaria digitata, L. hyperborea and L. saccharina were exposed in the laboratory to UV-radiation, with a spectral composition and irradiance similar to natural sunlight, for periods ranging from 15 min to 8 d, and were then returned to white light. Germination of zoospores and the growth of gametophytes were reduced after exposures to UV longer than 1 h, whereas UV had little effect on the growth of young or mature sporophytes unless exposure continued for more than 48 h. The variable fluorescence (F _v:F_m) of all stages was strongly reduced immediately after short exposures to UV, but recovered almost completely within 24 h. However, exposure of gametophytes to UV for >4 h resulted in little or no recovery of F _v:F_m, whereas >16 h of UV were required to produce this result in young sporophytes, and >48 h in mature sporophytes. Thus, sensitivity to UV-radiation decreased from gametophytes to sporophytes, and with increasing age of sporophytes, but, in gametophytes, growth appeared to be a more sensitive indicator of UV-damage than F _v:F_m after 24 h recovery. The responses to UV of the zoospores and gametophytes of all three species were similar, but both growth and fluorescence measurements suggested that the sporophytes of L. saccharina were more sensitive to UV than those of the other two species.     

Brooded coral larvae differ in their response to high temperature and elevated pCO2 depending on the day of release

To evaluate the effects of temperature and pCO₂ on coral larvae, brooded larvae of Pocillopora damicornis from Nanwan Bay, Taiwan (21°56.179′N, 120°44.85′E), were exposed to ambient (419–470 μatm) and high (604–742 μatm) pCO₂ at ~25 and ~29 °C in two experiments conducted in March 2010 and March 2012. Larvae were sampled from four consecutive lunar days (LD) synchronized with spawning following the new moon, incubated in treatments for 24 h, and measured for respiration, maximum photochemical efficiency of PSII (F _v/F _m), and mortality. The most striking outcome was a strong effect of time (i.e., LD) on larvae performance: respiration was affected by an LD × temperature interaction in 2010 and 2012, as well as an LD × pCO₂ × temperature interaction in 2012; F _v/F _m was affected by LD in 2010 (but not 2012); and mortality was affected by an LD × pCO₂ interaction in 2010, and an LD × temperature interaction in 2012. There were no main effects of pCO₂ in 2010, but in 2012, high pCO₂ depressed metabolic rate and reduced mortality. Therefore, differences in larval performance depended on day of release and resulted in varying susceptibility to future predicted environmental conditions. These results underscore the importance of considering larval brood variation across days when designing experiments. Subtle differences in experimental outcomes between years suggest that transgenerational plasticity in combination with unique histories of exposure to physical conditions can modulate the response of brooded coral larvae to climate change and ocean acidification.     

Light-shade adaptation by the oceanic dinoflagellates Pyrocystis noctiluca and P. fusiformis

Species-specific rates of photosynthetic carbon uptake (P), chlorophyll a content and P versus irradiance (P-I), have been measured for cells of Pyrocystis noctiluca and P. fusiformis isolated from natural populations collected in the euphotic zone within and below the surface mixed layer in the Sargasso Sea. These same measurements and the assay for ribulose bis-phosphate carboxylase (RuBP-Case), have been made for cultures of P. noctiluca in a 12 h L: 12 h D photoperiod at 9 different constant or at changing light intensities. In nature chl a cell^-1 was constant throughout the euphotic zone. The photosynthetic capacity (P_max), of cells captured below the surface mixed layer was lower by a factor of 10 compared with cells collected from the surface mixed layer. The P_max for P. noctiluca collected and incubated within the surface mixed layer was the same as for cell cultures grown under high light, non nutrient-limiting conditions, suggesting that photosynthesis in the natural system was not nutrient limited. In laboratory cultures under constant low light intensities, chl a cell^-1 increased by a factor of 5 while both P_max and RuBPCase activity decreased by a factor of ca 4 compared with high light intensities. In changing light intensities both P_max and RuBPCase activities were decreased by factors of 4 during low light intervals while chl a cell^-1 approached a constant intermediate value. The change in chl a cell^-1 in response to prolonged exposure to constant low light intensities was first order with a rate constant of 0.33 d^-1. For all irradiance conditions in culture, the P-I dependence could be described by the simple Michaelis-Menten formula. The ratio of P_max to K_I, (the light intensity where P=P_max/2) was a constant with a Coefficient of Variation of 12%: The constancy of this ratio, the parallel changes in RuBPCase activity with P_max and the constant chl a cell^-1 in the Sargasso Sea imply that for P. noctiluca and presumably P. fusiformis in nature, a dark enzymatic step rather than changes in photosynthetic pigment concentrations may regulate the photosynthetic capacity in the changing photic environment.Contribution no. 1141 from McCollum-Pratt Institute and Department of Biology, The Johns Hopkins University. Supported by DOE contract no. EY 76S20 3278, NSF no. OCE 76-02571 and ONR no. N300014-81-C-0062     

Diurnal variation in chlorophyll fluorescence of Thalassia testudinum seedlings in response to controlled salinity and light conditions

Diurnal variability in chlorophyll fluorescence caused by dynamic irradiance conditions is an important issue when using pulse amplitude modulation fluorometry to measure physiological conditions of plants at the landscape scale. We examined the use of slopes and y-intercepts of diurnal effective photochemical efficiency of photosystem II (PSII) (ΔF/F _m′) versus photosynthetically active radiation (PAR) regressions in addition to direct measurements of maximum photochemical efficiencies of PSII (F _v/F _m) values to assess physiological status of Thalassia testudinum seedlings in a controlled mesocosm study. Seedlings were exposed to two light treatments (full sun and 50–70 % light reduction) and three salinity treatments (20, 35, and 50). Measurements were taken at 0600, 0900, 1200, 1500, 1800, and 2100 hours in order to assess the diurnal variation in photochemical efficiency of PSII and PAR, with measurements at 2100 providing F _v/F _m. Results indicated significant effects of light and salinity on regression y-intercepts and measured F _v/F _m values. Shaded seedlings had higher values for both parameters, suggesting low-light acclimation. The highest salinity treatment resulted in significant reductions for both parameters, suggesting stress. Stress was also indicated by significant reductions in both seedling leaf growth and mean differences between seedling leaves and media osmolalities in the hypersaline treatments (152.0 ± 26.4 vs. 630 ± 40.2 mmol kg^?1 for the control treatments). Slopes of ΔF/F _m′ versus PAR significantly differed with varying light treatments, with full sun seedlings exhibiting shallower slopes than shaded seedlings, indicating higher efficiency of dissipation of excess energy (photoprotection). These experimental results confirm field data suggesting that diurnal ΔF/F _m′ versus PAR regressions are responsive to changes in the physiological status of T. testudinum and that the y-intercepts of diurnal regressions may be used as a proxy for F _v/F _m.     

Microbial consortia increase thermal tolerance of corals

This study examined the response of a coral holobiont to thermal stress when the bacterial community was treated with antibiotics. Colonies of Pocillopora damicornis were exposed to broad and narrow-spectrum antibiotics targeting coral-associated α and γ-Proteobacteria. Corals were gradually heated from the control temperature of 26 to 31 °C, and measurements were made of host, zooxanthellar and microbial condition. Antibiotics artificially reduced the abundance and activity of bacteria, but had minimal effect on zooxanthellae photosynthetic efficiency or host tissue protein content. Heated corals without antibiotics showed significant declines in F _V /F _M , typical of thermal stress. However, heated corals treated with antibiotics showed severe tissue loss in addition to a decline in F _V /F _M . This study demonstrated that a disruption to the microbial consortium diminished the resilience of the holobiont. Corals exposed to antibiotics under control temperature did not bleach, suggesting that temperature may be an important factor influencing the activity, diversity and ecological function of the holobiont bacterial community.     

Adaptation of solitary corals and their zooxanthellae to low light and UV radiation

Adaptation of solitary corals, Fungia repanda and F. echinata, and their zooxanthellae to low light and ultraviolet light B (UV-B) was studied with respect to changes in their protein contents, photosynthetic pigment contents and the photosynthesis-irradiance (P-I) curves. The corals were collected from 1 to 50 m depths in the Republic of Belau (Paulau) in 1990 and 1991. The chlorophyll a content in a unit surface area of the coral did not change significantly with the depth of the habitat, whereas cellular chlorophyll a in the algae increased with the depth. Zooxanthellae density and protein content in a unit surface area of Fungia spp. decreased with the depth. Photosynthetic parameters normalized by a unit surface area of the Fungia spp., maximum gross photosynthetic rate (P _gmax area^-1) and dark respiration rate (R area^-1), were negatively correlated with the depth, while initial slope of the P-I curve () did not show significant correlation with the depth. Compensation light intensity (Ic) decreased with the depth. In isolated zooxanthellae, P _max chl a ^-1, and R chl a ^-1 decreased with the depth, while _{chl a} was constant. P _gmax cell^-1 and R cell^-1 did not change significantly but _cell increased with the depth. Ic decreased with the depth as in the intact corals. Reduction of protein content in a unit area of the coral from deeper habitat implies decrease of host animal tissues. Reduction of Ic can be explained by decrease of R area^-1, which may be due to the diminution of animal tissues. The photoadaptational response to low light intensity of intact Fungia spp. was found to be a combination of the photoadaptation of symbiotic algae and the decrease of host animal tissue. In order to study their adaptation to ultraviolet (UV) radiation, P-I curves of Fungia spp. and isolated zooxanthellae were analyzed before and after UV-B irradiation. 1 h UV-B irradiation showed no effect on the photosynthetic rate of the shallow water (1 m) corals, while it inhibited the photosynthesis of the deep water (30 m) corals and zooxanthellae isolated from both shallow and deep water corals. These results indicate that the host, Fungia spp., in shallow water have protective mechanism for intense UV-B in their habitat. These photoadaptational mechanisms seem to allow the Fungia spp. to have wide vertical distribution where light intensity spans more than two orders of magnitude.     

Evaluation of chlorophyll fluorescence, in vivo spectrophotometric pigment absorption and ion leakage as biomarkers of UV-B exposure in marine macroalgae

The photosynthetic fluorescence ratio F_v:F_m, in vivo absorption spectra and ion leakage were evaluated as biomarkers of ambient and elevated UV-B (280 to 320 nm) exposure of the intertidal alga Enteromorpha intestinalis (Chlorophyta) and the sublittoral alga Palmaria palmata (Rhodophyta). Measurements of thallus growth were also used to assess adverse biological effects. Ambient and elevated UV-B significantly inhibited photosynthesis in both species. It was shown that the F_v:F_m ratio is a sensitive, non-specific general biomarker of UV-B exposure in both species. Moreover, the in vivo absorption of what was tentatively identified as chlorophylls a and b as well as phycoerythrin and/or carotenoids, phycoerythrobilin and phycocyanin decreased in a dose-response dependent manner and was associated with a decrease in growth rate in P. palmata. The intertidal alga E. intestinalis showed a greater degree of tolerance to UV-B exposure. These results indicate that changes in the F_v:F_m ratio together with reductions in in vivo pigment absorption could provide an early quantitative warning of the detrimental effects of UV-B in marine macroalgae. Received: 16 May 1997 / Accepted: 16 July 1997