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.     

Intraspecific competition in<Emphasis Type="Italic"> Fucus serratus</Emphasis> and<Emphasis Type="Italic"> F</Emphasis>.<Emphasis Type="Italic"> evanescens</Emphasis> (Phaeophyceae: Fucales) germlings: effects of settlement density,nutrient concentration,and temperature

Survival and growth of early post-settlement stages are critical for the development of seaweed populations. Fucoid germlings commonly settle in dense monospecific aggregates, where intraspecific competition and environmental variables (e.g. nutrient concentration and temperature) may affect survival and growth. Using factorial experiments, we determined the effects of settlement density (~10, ~50 and ~250 germlings cm^–2), nutrient enrichment (from ~10 to ~40 µM N and from ~0.5 to ~2.5 µM P), and temperature (7°C and 17°C) on Fucus serratus and F. evanescens germlings in laboratory cultures over 3 months. Settlement density, nutrient concentration and temperature interactively affected growth of germlings, and the magnitude of this interaction varied between the two species. This represents the first record of such factorial interactions in Fucus spp. germlings. Intraspecific competition, estimated as the relative reduction in germling growth and survival from low to high densities, increased with decreasing nutrient concentration and increasing temperature in both species. While temperature and nutrient concentration had little effect on germling size distributions, size inequality and skewness generally increased with germling density, indicating that a few large individuals gained dominance and suppressed many smaller ones at high density. Self-thinning increased with settlement density and depended on nutrient concentration and species at high density. At high density, self-thinning increased with decreasing nutrient levels in F. evanescens, but not in F. serratus. At low density, nutrient enrichment increased germling growth in F. evanescens, but not in F. serratus, whereas growth in both species was stimulated by nutrient enrichment at higher densities. These results suggest that germling growth and self-thinning are more sensitive to variation in nutrient concentration in F. evanescens than in F. serratus. The potential implications of our findings for the understanding of eutrophication-related abundance changes in both species in southern Norway are discussed.Communicated by L. Hagerman, Helsingør     

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.     

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.     

Response of zooxanthellae in symbiosis with the Mediterranean corals Cladocora caespitosa and Oculina patagonica to elevated temperatures

Scleractinian symbiotic corals living in the Ligurian Sea (NW Mediterranean Sea) have experienced warm summers during the last decade, with temperatures rapidly increasing, within a few days, to 3–4°C above the mean value of 24°C. The effect of elevated temperatures on the photosynthetic efficiency of zooxanthellae in symbiosis with temperate corals has not been well investigated. In this study, the corals, Cladocora caespitosa and Oculina patagonica were collected in the Ligurian Sea (44°N, 9°E), maintained during 2 weeks at the mean summer temperature of 24°C and then exposed during 48 h to temperatures of 24 (control), 27, 29 and 32°C. Chlorophyll (chl) fluorescence parameters [F _v/F _m, electron transport rate (ETR), non-photochemical quenching (NPQ)] were measured using pulse amplitude modulated (PAM) fluorimetry before, during the thermal increase, and after 1 and 7 days of recovery (corals maintained at 24°C). Zooxanthellae showed a broad tolerance to temperature increase, since their density remained unchanged and there was no significant reduction in their maximum quantum yield (F _v/F _m) or ETR up to 29°C. This temperature corresponded to a 5°C increase compared to the mean summer temperature (24°C) in the Ligurian Sea. At 32°C, there was a significant decrease in chl contents for both corals. This decrease was due to a reduction in the chl/zooxanthellae content. For C. caespitosa, there was also a decrease in ETR_max, not associated with a change in F _v/F _m or in the non-photochemical quenching (NPQ); for O. patagonica, both ETR_max and F _v/F _m significantly decreased, and NPQ_max showed a significant increase. Damages to the photosystem II appeared to be reversible in both corals, since F _v/F _m values returned to normal after 1 day at 24°C. Zooxanthellae in symbiosis with the Mediterranean corals investigated can therefore be considered as resistant to short-term increases in temperature, even well above the maximum temperatures experienced by these corals in summer.     

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.     

Age-specific responses to elevated salinity in the coastal marsh plant black needlerush (Juncus roemerianus Scheele) as determined through polyphasic chlorophyll a fluorescence transients (OJIP)

This study employed polyphasic chlorophyll a fluorescence transients (OJIP), a non-invasive marker of environmental stress in plants, to evaluate salt tolerance in three different Juncus roemerianus age classifications (6-, 24-, and 60-months). Following exposure to elevated salts (30 psu), the younger plants sustained growth, which was comparable to freshwater controls. While older (60-month) plants receiving only freshwater also grew over the 8-week study, the older salt-treated plants did not increase in size. Similarly, there were significant declines in variable chlorophyll a fluorescence parameters (F _v/F _m and F _v/F _o), electron transport flux per reaction center (ET_o/RC), and photosystem II performance index (PI_ABS) for 60-month J. roemerianus following salt treatment. These responses were not evident in the two younger salt-treated age classifications. Our results suggest that older J. roemerianus are less tolerant to rapid and sudden increases in salinity relative to younger plants and that this age-specific response may help explain observed discrepancies in salt tolerance in J. roemerianus.     

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.     

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.     

Is acclimation beneficial to scleractinian corals,Porites spp.?

With coral reefs impacted by climate change, attention is turning to the extent to which scleractinian corals can acclimatize to new physical conditions. The implicit assumption that acclimatization is beneficial has not been tested for scleractinians, although it has been investigated in other systems and has been referred to as the beneficial acclimation hypothesis (BAH). This study tests this hypothesis for scleractinians in experiments on massive Porites spp. from Moorea, French Polynesia (17° 28.564S, 149° 49.018 W). Corals were acclimated for 15–21 days to three temperatures within the range experienced in the collection habitat and then transferred to each of the same temperatures for a treatment period of 14–15 days. The response of the holobiont was measured as growth, the response of the Symbiodinium populations as maximum photochemical efficiency of open reaction centers II (F _v/F _m). An ANOVA with polynomial contrasts was used to distinguish among the BAH, three alternative hypotheses and a null hypothesis describing the consequences of acclimation. In the first experiment (2009), massive Porites spp. were unresponsive to temperature. In the second experiment (2013), the BAH was not supported, but growth of the holobiont conformed to the “hotter is better” (HIB) hypothesis; the response of Symbiodinium populations conformed to developmental buffering. These results suggest that acclimation by massive Porites spp. to temperatures experienced routinely in the natural environment does not have clear beneficial value for growth or photochemical efficiency (i.e., BAH was not supported), but they reveal that acclimation to increased temperature can have value in responding to a variety of subsequent temperatures (i.e., support for HIB).     

Ocean acidification and warming alter photosynthesis and calcification of the symbiont-bearing foraminifera Marginopora vertebralis

The impact of elevated CO₂ and temperature on photosynthesis and calcification in the symbiont-bearing benthic foraminifer Marginopora vertebralis was studied. Individual specimens of M. vertebralis were collected from Heron Island on the southern Great Barrier Reef (Australia). They were maintained for 5 weeks at different temperatures (28, 32 °C) and pCO₂ (400, 1,000 µatm) levels spanning a range of current and future climate-change scenarios. The photosynthetic capacity of M. vertebralis was measured with O₂ microsensors and a pulse-amplitude-modulated chlorophyll (Chl) fluorometer, in combination with estimates of Chl a and Chl c ₂ concentrations and calcification rates. After 5 weeks, control specimens remained unaltered for all parameters. Chlorophyll a concentrations significantly decreased in the specimens at 1,000 µatm CO₂ for both temperatures, while no change in Chl c ₂ concentration was observed. Photoinhibition was observed under elevated CO₂ and temperature, with a 70–80 % decrease in the maximum quantum yield of PSII. There was no net O₂ production at elevated temperatures in both CO₂ treatments as compared to the control temperature, supporting that temperature has more impact on photosynthesis and O₂ flux than changes in ambient CO₂. Photosynthetic pigment loss and a decrease in photochemical efficiency are thus likely to occur with increased temperature. The elevated CO₂ and high temperature treatment also lead to a reduction in calcification rate (from +0.1 to >?0.1 % day^?1). Thus, both calcification and photosynthesis of the major sediment-producing foraminifer M. vertebralis appears highly vulnerable to elevated temperature and ocean acidification scenarios predicted in climate-change models.     

Seasonal variation in ecophysiological patterns in macroalgae from an Arctic fjord. I. Sensitivity of photosynthesis to ultraviolet radiation

Marine macroalgae inhabiting Arctic coastal ecosystems are exposed to pronounced seasonal variations in the radiation regime, including harmful UVB radiation. This study presents the first data on the seasonal changes in the sensitivity of macroalgal photosynthesis towards UV exposure by comparing under-ice, clear-water and turbid-water conditions characteristic for late winter, spring and summer. Various brown (Laminaria saccharina, L. digitata, L. solidungula, Saccorhiza dermatodea, Desmarestia aculeata), red (Palmaria palmata, Devaleraea ramentacea) and one green macroalgal species (Monostroma aff. arcticum) were collected at the same water depth throughout the seasons in the Kongsfjord (Spitsbergen, Svalbard, Norway). Maximum quantum yield (F_v/F_m) and maximum photosynthetic electron transport rates (ETR_max) were determined immediately after collection, after 2 h exposure to artificial UV radiation and after 18 h recovery in dim white light. Photosynthesis of the studied species showed different responses depending on their morpho-functional and physiological characteristics, their life strategies, phenology and depth distribution. Within the genus Laminaria, maximum quantum yield of adult specimens of the deep-water species L. solidungula was most strongly UV sensitive. Adult L. saccharina exhibited a lower UV sensitivity than a 6-month-old specimen. Inhibition of photosynthesis after UV exposure remained at the same level throughout the study period, both in adult L. saccharina and S. dermatodea. However, adult specimens of L. saccharina collected in May showed partial recovery only, whereas photosynthesis of specimens from both species collected later recovered fully. D. aculeata exhibited a remarkable decrease of UV sensitivity during the study period. Photosynthesis of specimens collected under the ice was strongly inhibited by UV, but the degree of inhibition decreased during spring and summer. Concomitantly ETR_max values were low after UV exposure in specimens collected in June, but increased later in the season. P. palmata exhibited a relatively flexible response. Photosynthesis in specimens collected under the ice in June or in turbid water in July/August was relatively strongly inhibited; specimens collected during sunny periods and in clear water in spring showed a much lower degree of photoinhibition after UV exposure. The seasonal pattern of low/high ETR_max values in spring/summer is probably a characteristic of the life strategy of this species. The UV sensitivity of D. ramentacea exhibited a similar seasonal pattern. In M. aff. arcticum, UV sensitivity increased and ETR_max values decreased during the study period, reflecting the life strategy of this annual late winter/spring species. The physiological basis for the seasonal changes in UV sensitivity of photosynthesis is presented in a companion paper (this issue).     

Effect of light and temperature on the population dynamics of two toxic bloom forming Cyanobacteria – Microcystis ichthyoblabe and Anabaena aphanizomenoides

The effect of light and temperature on the growth of Microcystis ichthyoblabe and Anabaena aphanizomenoides, isolated from the subtropical Oued Mellah lake, Morocco (33°30′N–07°20′W), were investigated in batch culture. Growth rates at 66 light–temperature combinations were determined and fitted with different mathematical models. The results show that the two Cyanobacteria grow at all light intensities and temperatures, except at 10 °C for A. aphanizomenoides, where the growth was strongly limited. The μ_max of M. ichthyoblabe increased with temperature from 0.56 d^?1 at 10 °C to 1.32 d^?1 at 35 °C. At all tested temperatures, a relative photoinhibition within the studied range of irradiance was observed and the photosensitivity was thermodependent. For Anabaena, the obtained μ_max ranged between 0.07 d^?1 at 10 °C and 1.46 d^?1 at 35 °C, and a weak photoinhibition was observed at 15 °C. The positive correlation between μ_max and I_opt (r²≥0.93) indicates a close interaction between light and temperature on the cyanobacteria growth. The results obtained in this work suggest that the growth of these two species is possible under low light and low temperature.     

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     

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.     

Mechanisms of range expansion in the intertidal brown alga <Emphasis Type="Italic">Fucus</Emphasis><Emphasis Type="Italic">serratus</Emphasis> in northern Spain

The intertidal brown alga Fucus serratus is naturally expanding its distributional range along the north shore of Spain. Shores being colonised by F. serratus harbour saturated algal communities, with almost no free space. Processes responsible for the successful expansion were investigated in a locality where the alga was experimentally introduced. After a short period, the number of F. serratus plants increased exponentially at a monthly intrinsic rate of 0.111. At early and intermediate stages of invasion, colonisation occurred by the growth of patches of the invader around the transplant points. F. serratus is a bad disperser, with most of propagules settling in near proximity to parent plants. Colonisation of manipulated and control areas was influenced by the proximity of propagule sources. Experimental removals of canopies, irrespective of whether the turf was removed or left intact, accelerated the expansion, as this component of the community had a negative influence on recruitment of both F. serratus and F. vesiculosus. The unmanipulated algal beds were, however, invasible, as some control quadrats close (<10 m) to F. serratus patches had a 100% cover of F. serratus at the end of the study. Prior or concurrent to increases of F. serratus, a decrease in cover of F. vesiculosus and other resident macroalgae was evident. No formal competition experiment was done but results suggest that F. serratus outcompetes F. vesiculosus and Bifurcaria bifurcata at mid-tidal levels. In scraped quadrats, in which both Fucus species recruited at large densities, F. serratus grew at a higher rate than F. vesiculosus and eventually monopolised the experimental surfaces.     

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     

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.     

Hitchhikers reveal cryptic host behavior: new insights from the association between Planes major and sea turtles in the Pacific Ocean

Studies that incorporate information from habitat-specific ecological interactions (e.g., epibiotic associations) can reveal valuable insights into the cryptic habitat-use patterns and behavior of marine vertebrates. Sea turtles, like other large, highly mobile marine vertebrates, are inherently difficult to study, and such information can inform the implementation of conservation measures. The presence of epipelagic epibionts, such as the flotsam crab Planes major, on sea turtles strongly suggests that neritic turtles have recently occupied epipelagic habitats (upper 200 m in areas with >200 m depth) and that epipelagic turtles spend time at or near the surface. We quantified the effects of turtle species, turtle size, and habitat (neritic or epipelagic) on the frequency of epibiosis (F ₀) by P. major on sea turtles in the Pacific Ocean. In neritic habitats, we found that loggerhead (F ₀ = 27.6 %) and olive ridley turtles (F ₀ = 26.2 %) host crabs frequently across a wide range of body sizes, and green turtles almost never host crabs (F ₀ = 0.7 %). These results suggest that loggerheads and olive ridleys display variable/flexible epipelagic-neritic transitions, while green turtles tend to transition unidirectionally at small body sizes. In epipelagic habitats, we found that loggerheads host crabs (F ₀ = 92.9 %) more frequently than olive ridleys (F ₀ = 50 %) and green turtles (F ₀ = 38.5 %). These results suggest that epipelagic loggerheads tend to spend more time at or near the surface than epipelagic olive ridleys and green turtles. Results of this study reveal new insights into habitat-use patterns and behavior of sea turtles and display how epibiont data can supplement data from more advanced technologies to gain a better understanding of the ecology of marine vertebrates during cryptic life stages.     

The theoretical basis for estimating phytoplankton production and specific growth rate from chlorophyll, light and temperature data

Phytoplankton maximum specific growth rate, μ_max, and maximum photosynthetic quantum yield, Φ_max, can be related mathematically via the photosynthetic light curve, P(I). A model is presented in which maximum quantum yield defines the initial slope of the light curve and is assumed to be a known constant, while maximum specific growth rate defines the light-saturated region of the curve and is assumed to be a known function of temperature. The effect of introducing μ_max(T) into P(I, Φ_max) is to replace the unknown, temperature-dependent light saturation parameter with a term involving the ratio μ_max/Φ_max. The advantage of writing P(I) in terms of both μ_max and Φ_max is that those parameters are particularly well documented in the literature. Consequently, estimates of nutrient-unlimited phytoplankton growth and production rates can be based solely on the constants μ_max, Φ_max and k_c (light absorption per unit of chlorophyll) and the free variables light, temperature and chlorophyll concentration. Rate estimates appear to be accurate to within a factor of two for an extremely wide range of conditions.One particularly significant result of introducing μ_max into P(I, Φ_max) is that the carbon : chlorophyll ration, θ, appears explicitly. It is possible to derive an expression for optimum θ based on the assumption that adaptive changes in carbon/chlorophyll occur so as to maximize the specific growth rate for given conditions of light and temperature. Laboratory and field data are compiled from the literature to test the formulae presented here.