Sexuality and cyst formation of the spring-bloom dinoflagellate Scrippsiella hangoei in the coastal northern Baltic Sea

The temporal sequence and the magnitude of the sexual reproduction and subsequent cyst deposition of the common spring-bloom dinoflagellate Scrippsiella hangoei (Schiller) Larsen was studied during spring 1996 on the SW coast of Finland, Baltic Sea. The abundances of the different size of fractions of S. hangoei (14 to 18 μm, 18 to 22 μm and >22 μm) were monitored in the water column, and the deposition of resting cysts was measured using moored sediment traps. Cyst sedimentation rates were measured throughout the seasonal cycle in order to estimate cyst resuspension rates for the quantitative assessment of the fraction of population undergoing encystment. The onset of sexual reproduction, indicated by a significant increase of the small cells (14 to 18 μm) representing gametes, occurred in a nutrient replete environment well before the exponential growth phase and peak abundances of vegetative cells. Gamete formation was followed by high abundances of large cells (>22 μm) representing planozygotes, and subsequent sedimentation of resting cysts. Approximately 60% of the asexually growing bloom population was estimated to form planozygotes, suggesting that encystment was an important factor in bloom termination and possibly plays a role in the regulation of the magnitude of the bloom. Finally encystment accounted for 40% of the entire S. hangoei population, resulting in a considerable loss of the bloom population and an input of the vernal phytoplankton biomass to the benthos. Received: 11 December 1998 / Accepted: 8 April 1999     

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.     

Effects of light intensity on photosynthesis and dark respiration in six species of marine phytoplankton

Using an oxygen polarographic electrode, the shapes of photosynthetic curves and the effects of light on dark respiration in 6 species of marine phytoplankton wer examined. The species used were Skeletonema costatum, Ditylum brightwellii, Cyclotella nana (Thalassiosira pseudonana) (all Bacillariophyceae), Dunaliella tertiolecta (Chlorophyceae), Isochrysis galbana (Haptophyceae), and Gonyaulax tamarensis (Dinophyceae). A hysteresis was observed in all species examined with respect to increasing and decreasing light. Compensation light intensities varied by over 4 orders of magnitude, suggesting that the 1% light depth is an ambiguous measure of the euphotic zone. The data suggest that dark respiration accounts for ca. 25% of gross photosynthesis, but is species-dependent. In addition, respiration versus cell size does not describe an inverse exponential function over the size scales examined.This research was performed under the auspices of the United States Energy Research and Development Administration under Contract No. EY-76-C-02-0016.     

Effects of salinity and possible interactions with temperature and pH on growth and photosynthesis of <Emphasis Type="Italic">Halophila johnsonii</Emphasis> Eiseman

The effects of salinity, temperature, and pH variations on growth, survival, and photosynthetic rates of the seagrass Halophila johnsonii Eiseman were examined. Growth and survival responses to salinity were characterized by aquarium experiments in which plants were exposed to seven different salinity treatments (0, 10, 20, 30, 40, 50, and 60 psu) during 15 days. Photosynthetic behavior was assessed for short-term salinity exposures (1 or 20 h) by incubation experiments in biological oxygen demand (BOD) bottles and by measuring photosynthesis versus irradiance (PI) responses in an oxygen electrode chamber. In the bottle experiments the possible effects of interactions between salinity and temperature (15, 25, and 35°C) or pH (5, 6, 7, and 8.2) were also examined. Growth and survival of H. johnsonii were significantly affected by salinity, with maximum rates obtained at 30 psu. Salinity also altered the parameters of the PI curves. Light-saturated photosynthesis (P _max) and the photosynthetic efficiency at subsaturating light (α) increased significantly up to an optimum of 40 psu, decreasing again at the highest salinities. Dark respiration rates and compensating irradiance (I _c) showed minimum values at 40 and 50 psu, while light-saturation point (I _k) was maximum at 30–50 psu. An interaction between salinity and temperature was not found although an increase of temperature alone produced an increase in α, P _max, respiration rates, and I _k. An interaction between salinity and pH was only found in the P _max response: P _max increased with pH=5 at 30 psu. In addition, reducing the pH increased α significantly. In the BOD bottles experiment a significant reduction in the dark respiration with decreasing pH was observed, but the opposite trend was observed in the photosynthetic rate. These results suggest that the endemic seagrass H. johnsonii could be negatively affected by hypo- or hypersalinity conditions, although salinity changes did not seem to alter the tolerance of this species to other environmental factors, such as temperature or pH.     

Symbiont photosynthesis increases both respiration and photosynthesis in the symbiotic sea anemone Anemonia viridis

Dark respiration of the symbiotic sea anemone Anemonia viridis (Forskäl) was observed to increase by 34% when anemones were exposed to hyperoxic sea water (150% oxygen saturation) overnight, and by 39% after exposure to 6 h in the light at a saturating irradiance of 300 E m^-2 s^-1 at normoxia (100% oxygen saturation). No increase due to light stimulation was observed in aposymbiotic control anemones. In darkness, the oxygen concentration of the coelenteric fluid was hypoxic. However, within 10 min of anemones being illuminated, coelenteric fluid was hyperoxic, and it remained elevated throughout a 12 h light period. When measured over a 24 h period (12 h light: 12 h dark), the dark respiration rate increased gradually over the first 6 h of the light period until it was 35% above the dark night-time resting rate. It remained elevated throughout the remaining light period and for 2 h into the following dark period, after which it fell back to the resting rate. Gross photosynthesis (P ^gross) increased significantly when anemones were exposed to either hyperoxia (150% oxygen saturation) or 300 E m^-2 s^-1 at normoxia. This increase was not observed when symbiotic anemones were illuminated at a low-light intensity of 100 E m^-2 s^-1. The results of this study suggest that respiration in the dark is limited by oxygen diffusion and that normal respiration is restored in the daytime by utilisation of the oxygen released by photosynthesis. Furthermore, it appears that the increased respiration following exposure to high-light intensities provides a CO₂-rich intracellular environment which further enhances the photosynthetic rate of the zooxanthellae.     

Implications of salinity fluctuation for growth and nitrogen metabolism of the marine diatom Ditylum brightwellii in comparison with Skeletonema costatum

In 1987 effects of salinity fluctuations on growth of Ditylum brightwellii (West) Grunow, isolated from the Eastern Scheldt estuary (SW Netherlands) in 1981, were studied. D. brightwellii was grown in a 12 h light: dark cycle at constant salinity in brackish media. Ammonium-limited cultures were subjected to a salinity fluctuation. By decreasing the salinity to 4.8 photosynthesis and cell division were inhibited; cells were deformed. Protein and carbohydrate contents increased slightly, dark respiration was stimulated and cellular levels of glucose decreased at low salinity; this indicated a possible role of sugars in osmoregulation. Ammonium was accumulated in cultures, amino acids may have been stored; the role of the vacuole as a storage compartment was discussed. Both the ammonium uptake capacity and the affinity for ammonium decreased. Nitrogen limitation was relieved in the transient state. [With the activity of the nitrogen assimilation enzymes glutamine synthetase (GS) and glutamate synthase (GOGAT) being uninhibited by lower salinity.] Recovery from hypo-osmotic stress during a salinity increase was initiated by stimulated photosynthesis; chlorophyll a increased, but persistant contractions of cytoplasm (with chloroplasts) may have delayed cell growth. The glutamate dehydrogenase (GDH) activity decreased further whereas the cellular level of alanine increased in the presence of large ammonium pools; this may indicate a temporary activity of ADH (alanine dehydrogenase). Skeletonema costatum (Greville) Cleve, recovered faster from hypoosmotic stress than did D. brightwellii. Due to an osmotic shock from 13.6 to 7.1 S both species excreted amino acids and glucose; S. costatum accumulated more glucose, D. brightwellii accumulated more amino acids. S. costatum may with the competition for nitrogen in waters with an unstable salinity; it will replace D. brightwellii.Contribution no. 427 Delta Institute for Hydrobiological Research, Yerseke, The Netherlands     

The mechanism of calcification and its relation to photosynthesis and respiration in the scleractinian coral <Emphasis Type="Italic"> Galaxea fascicularis</Emphasis>

The mechanism of calcification and its relation to photosynthesis and respiration were studied with Ca²⁺, pH and O₂ microsensors using the scleractinian coral Galaxea fascicularis. Gross photosynthesis (Pg), net photosynthesis (Pn) and dark respiration (DR) were measured on the surface of the coral. Light respiration (LR) was calculated from the difference between Pg and Pn. Pg was about seven times higher than Pn; thus, respiration consumes most of the O₂ produced by the algal symbiont's photosynthesis. The respiration rate in light was ca. 12 times higher than in the dark. The coupled Pg and LR caused an intense internal carbon and O₂ cycling. The resultant product of this cycle is metabolic energy (ATP). The measured ATP content was about 35% higher in light-incubated colonies than in dark-incubated ones. Direct measurements of Ca²⁺ and pH were made on the outer surface of the polyp, inside its coelenteron and under the calicoblastic layer. The effects on Ca²⁺ and pH dynamics of switching on and off the light were followed in these three compartments. Ca²⁺ concentrations decreased in light on the surface of the polyp and in the coelenteron. They increased when the light was switched off. The opposite effect was observed under the calicoblastic layer. In light, the level of Ca²⁺ was lower on the polyp surface than in the surrounding seawater, and even lower inside the coelenteron. The concentration of calcium under the calicoblastic layer was about 0.6 mM higher than in the surrounding seawater. Thus Ca²⁺ can diffuse from seawater to the coelenteron, but metabolic energy is needed for its transport across the calicoblastic layer to the skeleton. The pH under the calicoblastic layer was more alkaline compared with the polyp surface and inside the coelenteron. This rise in pH increased the supersaturation of aragonite from 3.2 in the dark to 25 in the light, and brought about more rapid precipitation of CaCO₃. When ruthenium red was added, Ca²⁺ and pH dynamics were inhibited under the calicoblastic layer. Ruthenium red is a specific inhibitor of Ca-ATPase. The results indicated that Ca-ATPase transports Ca²⁺ across the calicoblastic layer to the skeleton in exchange for H⁺. Addition of dichlorophenyldimethylurea completely inhibited photosynthesis. The calcium dynamics under the calicoblastic layer continued; however, the process was less regular. Initial rates were maintained. We conclude that light and not energy generation triggers calcium uptake; however, energy is also needed.     

Effects of ambient levels of solar ultraviolet radiation on zooxanthellae and photosynthesis of the reef coral Montipora verrucosa

Paired flat plates of the hermatypic coral Montipora verrucosa from Kaneohe Bay, Oahu, Hawaii, were acclimated to photosynthetically active radiation (PAR) only and to full sunlight (PAR+UV) for several weeks in the summer of 1990. After the acclimation period, photosynthesis, both in PAR-only and PAR+UV as well as dark respiration were measured. Levels of the UV-absorbing compounds, S320, density of zooxanthellae, and chlorophyll a concentration were determined. Corals acclimated in PAR+UV had higher levels of the UV-protective compounds and lower areal zooxanthellae densities than corals acclimated in PAR-only. Chlorophyll a per unit volume of coral host and per algal cell did not differ between corals from the two acclimation treatments. Corals acclimated to PAR+UV displayed higher photosynthesis in full sunlight than corals acclimated to PAR-only, but when photosynthesis was measured in the light regime to which the corals had been acclimated, there were no differences in photosynthesis. Dark respiration was the same for corals from the two acclimation treatments regardless of the light quality immediately preceding the dark period.Contribution No. 902 HIMB     

Cyst formation of the noxious red tide flagellate Chattonella marina (Raphidophyceae) in culture

Cyst formation of the noxious red tide flagellate Chattonella marina (Raphidophyceae), cultured from a cyst in sediments of Suo-nada, Japan in 1986, was investigated by culture experiments. Nitrogen limitation was effective in inducing cyst formation. After incubation in N-limited medium at 25°C with 10000 lux on a 14 h light, 10 h dark photo-cycle, cultures were subjected to six different light intensities (0 10000 lux) in order to determine effects of light intensity on cyst formation. Cysts were formed under light intensities of 1000 lux or below, but not at 10000 lux. Cysts formed in culture displayed morphological characteristics quite similar to those natural cysts observed in sediments collected from the Inland Sea of Japan. Observations on nucleus, using diaminidinophenylindole (DAPI)-staining and epifluorescence microscopy, indicated that the majority of cysts formed in culture were uninucleate, a few were binucleate. Germination of cysts was observed under adequate conditions for germination (22°C, 3500 lux with a 14 h light, 10 h dark photo-cycle) even after storage at 11°C in the dark for more than 4 mo.     

Carbon and nitrogen metabolism by Monochrysis lutheri: Measurement of growth-rate-dependent respiration rates

Dark respiration rates were measured and carbon-excretion rates calculated for a nitrate-limited population of the marine chrysophyte Monochrysis lutheri grown in continuous culture at 20°C on a 12 h light-12 h dark cycle of illumination and over a series of 4 growth rates. A significant (P<0.05) positive correlation was found between dark respiration rate and growth rate. From a simple linear fit to the data, the respiration rate at maximum growth rate was estimated to be roughly 10.5% of the maximum gross-carbon-production rate, and more than three times higher than the extrapolated respiration rate at zero net-growth rate. Carbon-excretion rates showed no significant correlation with growth rate, and averaged less than 5% of the maximum gross-carbon-production rate. Mean cell nitrogen to carbon ratios were correlated in a virtually linear manner (r=0.994) with growth rate, and at a given growth rate were consistently higher than nitrogen to carbon ratios for the same species grown on continuous light. A comparison of carbon and nitrogen quotas as a function of growth rate for M. lutheri and other species suggests that the increase of cellular nitrogen at high growth rates under nitrate-limited growth conditions may be associated with the storage of cellular protein or amino acids rather than the presence of an inorganic nitrogen reservoir. The maximum nitrate uptake rate per cell during the day changed very little over the range of growth rates studied, and was comparable to the maximum uptake rate found for cells grown on continuous light. However, the cell nitrogen quota increased steadily with growth rate, causing a reduction in the maximum specific-uptake rate of nitrate during the day at high growth rates. The dark nitrate-uptake capacity of the population was clearly exceeded by the supply rate at the two higher growth rates, leading to a buildup of nitrate during the night which amounted to as much as 21% of the particulate nitrogen in the growth chamber by morning.Hawaii Institute of Marine Biology Contribution No. 478.     

Effects of high-frequency light fluctuations on growth and photoacclimation of the red algaChondrus crispus

High-frequency fluctuations due to wavefocusing are prominent characteristics of light in shallow marine environments. Effects of high-frequency (0.01 to 1 Hz) fluctuating light on growth rates of the red algaChondrus crispus Stackh., collected from Crane Neck Point, New York, USA, during July 1988, were determined by comparing plants grown under fluctuating and constant light regimes with similar daily irradiances. At high daily irradiance, growth rates were higher under fluctuating light than under constant light. Fluctuating light effects were frequency-dependent; growth was enhanced by fluctuations at 0.1 and 1 Hz, but not by fluctuations at 0.01 Hz. At low daily irradiance, growth rates were not affected by fluctuating light at any frequency tested. Enhancement of growth was not due to effects of high-frequency light fluctuations on photoacclimation responses ofC. crispus. Plants grown under fluctuating light at high daily irradiance actually exhibited lower photosynthetic capacity and efficiency (determined under constant light) than plants grown under constant light. These differences were attributable to variation in the density of Photosystem II reaction centers, which was low in plants grown under fluctuating light. Maximum turnover rate of whole-chain electron transport and activity of ribulose-1,5-bisphosphate carboxylase were affected by total daily irradiance, but not by high-frequency light fluctuations. Enhancement of growth under fluctuating light was partly attributable to reduced rates of dark respiration compared to rates of plants grown under constant light. The results also provided indirect evidence that high-frequency light fluctuations may enhance instantaneous photosynthetic rates. This effect could increase daily carbon gain and, therefore, stimulate growth ofC. crispus under high-frequency light fluctuations.     

Respiration,photosynthesis and carbon metabolism in planktonic ciliates

Release of¹⁴C-labelled carbon dioxide from uniformly labelled cells was used to measure respiration by individual ciliates in 2-h incubations in 1989 and 1990. In a strictly heterotrophic ciliate,Strobilidium spiralis (Leegaard, 1915), release of labelled carbon dioxide was equivalent to ca. 2.8% of cell C h^–1 at 20°C, and there was no difference between rates in the dark and light. In the chloroplast-retaining ciliatesLaboea strobila Lohmann, 1908,Strombidium conicum (Lohmann, 1908) Wulff, 1919 andStrombidium capitatum (Leegaard, 1915) Kahl, 1932, release of labelled carbon dioxide was less in the light than in the dark in experiments done at 15°C. InL. strobila release of radiolabel as carbon dioxide was equivalent to ca. 2.4% of cell C h^–1 in the dark but ca. 1% at 50µE m^–2 s^–1, an irradiance limiting to photosynthesis. InS. conicum release of radiolabel as carbon dioxide was equivalent to ca. 4.4% of cell C h^–1 in the dark, but at an irradiance saturating to photosynthesis (250 to 300µE m^–2 s^–1) there was no detectable release of labelled carbon dioxide. InS. capitatum release of radiolabel as carbon dioxide was equivalent to ca. 4.3% of cell C h^–1 in the dark but at an irradiance saturating to photosynthesis was ca. 2.4% of cell C h^–1. These data, combined with data from photosynthetic uptake experiments, indicate that¹⁴C uptake underestimates the total benefit of photosynthesis by 50% or more in chloroplastretaining ciliates.Contribution no. 7510 from the Woods Hole Oceanographic Institution     

Effects of light quality on initiation and development of meroplanktonic diatom blooms in a eutrophic shallow sea

We investigated the effects of light quality on resting stage cell germination and vegetative cell growth of meroplanktonic diatoms in a small port in Hakata Bay, Japan and in the laboratory. During the investigation over the year of 2006, the meroplanktonic diatom bloom first occurred in the end of May and then repeated wane and wax until October in the small port. From late April to middle May, light penetrating the water column was often strong and attenuations of all spectral lights were low. During this period, Skeletonema costatum, Thalassiosira minima, and Chaetoceros sp. appeared frequently, followed by the blooms of S. costatum and Chaetoceros sp. in late May. Thereafter, S. costatum and Chaetoceros sp. bloomed in late June but not in middle June, when pigmented flagellates bloom appeared. The attenuation of short-wavelength light such as violet and blue lights was markedly high during these diatom and flagellate blooms; all blooms disappeared within several days. Vegetative cell strains of the three diatoms under light emitting diodes (LEDs) with six different spectra (violet, blue, green, orange, red, and near-infrared) grew at a higher rate under short-wavelength light, violet and blue. On the other hand, when suspensions of bottom sediments from Hakata Bay were cultured under the same LEDs and in the dark, vegetative cells of S. costatum appeared under all LEDs except for orange and near-infrared, vegetative cells of T. minima appeared under all LEDs but not in the dark, and vegetative cells of Chaetoceros sp. appeared under violet and blue LEDs. However, vegetative cell densities of the three diatoms increased much more under violet light than under other LEDs within a short period (6 days). Our study indicates that underwater penetration by short-wavelength light, such as violet and blue, may be an important factor in the initiation and development of meroplanktonic diatom blooms.     

Diurnal and circadian rhythms in the turbidity of growing Skeletonema costatum cultures

Skeletonema costatum (Grev.) Cleve grown in batch culture at low light intensity under a 14 h light: 10 h dark photocycle showed exponential cell proliferation (1.1 doublings d^-1) without significant phasing of the cell division by the light: dark cycle. The growth in carbon concentration was, however, restricted to the light period. The turbidity of the culture closely followed the carbon oattern, and was not affected by the increase in the cell number during the dark period. It was found that a trustule suspension had only approximately 1% of the turbidity of the corresponding intact algae. Culture turbidity was therefore regarded as a biomass parameter similar to the carbon concentration, without direct correlation to the timing of the cell division. The short-time variations in the turbidity of growing algal cultures were further studied in a cage culture turbidostat. The growth rate (based on turbidity) increased rapidly during the first half of the light period, decreased slightly towards the evening and was zero throughout the dark period. When transformed to continuous light, the growth of the culture continued to show damped oscillations for up to 1 wk, but with a period of 26.7 h instead of 24 h. The same circadian rhythm was observed in chlorophyll content, and is thus possibly a reflection of a freely oscillating internal biological clock. The cage culture turbidostat was found to be a suitable device for studies of the photocycle related regulation of biosynthesis in S. costatum.     

Influence of size on primary production in the reef coral Pocillopora damicornis and the macroalga Acanthophora spicifera

Size influences the photosynthesis-irradiance (P-I) relationship in colonies of the branched reef-coral Pocillopora damicornis and in intact plants of the branched redmacroalga Acanthophora spicifera. The light saturation constant is proportional to size. Maximum net rate of oxygen production (net photosynthesis) per colony and nocturnal dark oxygen-uptake rate per colony (respiration) increase with increasing size, but the latter increases at a much lower rate. Therefore, the photosynthesis to respiration ratio increases with increasing canopy size. Large increases in chlorophyll per unit reef area also accompany increase in size. The initial slope (alpha) of the chlorophyll-specific P-I curve and assimilation number are inversely related to size. Integrated daytime oxygen production increases with size more rapidly than nighttime oxygen consumption. Consequently, net primary production of an entire colony or plant (or rate per unit area of reef) increases with increasing size of the canopy. Production efficiency also increases with size. The coral is rigid, symmetrical and highly organized. Chlorophyll distribution is more stratified in comparison to the macroalga. The coral shows higher photosynthetic efficiency, as would be expected according to the stratified production model of Odum et al. (1958). This research was conducted on specimens from Kaneohe Bay, Oahu, Hawaii, USA in 1981.     

Temporary cyst formation of<Emphasis Type="Italic"> Heterocapsa triquetra</Emphasis> (Dinophyceae) in natural populations

Heterocapsa triquetra (Ehrenberg) Stein is a phototrophic marine dinoflagellate with wide coastal distribution. It is known to be capable of mixotrophy and diel vertical migration. The species was particularly abundant in the Gulf of Finland (the Baltic Sea) during the summers of 1996 and 1998, leading to discolouration of water on the south-west coast of Finland. Large-scale (50 m³) coastal mesocosm experiments in the north-west Gulf of Finland (the Baltic Sea) in the summers of 1996 and 1998 with daily mineral nutrient additions provoked a biomass increase of phytoplankton dominated by H. triquetra. From the first days of the experiment temporary cysts of H. triquetra were found in the bottom sediment water of the mesocosms. Maximum temporary cyst production rates reached values up to 20×10⁶ cysts m^–2 day^–1, accounting for <1% of the depth-integrated motile population size. The environmental features favouring temporary cyst production remain uncertain; zooplankton grazing and nutrient stress are potential factors. Temporary cysts of H. triquetra were observed in a unialgal culture (f/2 medium) isolated in summer 1999 from Eel Pond (Woods Hole, Mass., USA).Communicated by M. Kühl, Helsingør     

A comparative analysis of the photobiology of zooxanthellae and zoochlorellae symbiotic with the temperate clonal anemone <Emphasis Type="Italic">Anthopleura elegantissima</Emphasis> (Brandt). III. Seasonal effects of natural light and temperature on photosynthesis and respiration

The sea anemone Anthopleura elegantissima hosts two phylogenetically different symbiotic microalgae, a dinoflagellate Symbiodinium (zooxanthellae, ZX) and a chlorophyte (zoochlorellae, ZC). The photosynthetic productivity (P), respiration (R), and contribution of algal carbon translocated to the host (CZAR) in response to a year’s seasonal ambient changes of natural light and temperature are documented for both ZX- and ZC-bearing anemones. Light and temperature both affect photosynthesis, respiration, and CZAR, as well as various algal parameters; while there are evident seasonal differences, for the most part the relative effects on P, R, and CZAR by the two environmental variables cannot be determined. Net photosynthesis (P_n) of both ZX and ZC was significantly higher during spring and summer. During these seasons, the P_n of ZX was always greater than that of ZC. Regardless of algal symbiont, anemone respiration (R) was significantly higher during the spring and summer. The annual net carbon fixation rate of anemones with ZX and ZC was 325 and 276 mg C anemone⁻¹ year⁻¹, respectively, which translates to annual net community productivity rates of 92 and 60 g C m⁻¹ year⁻¹ for anemones with ZX or ZC, respectively. CZAR did not show a clear relationship with season; however the CZAR for ZX was always significantly greater than for ZC. Lower ZX growth rates, coupled with higher photosynthetic rates and higher CZAR estimates, compared to ZC, suggest that if A. elegantissima is simply carbon limited, ZX-bearing anemones should be the dominant symbiont in the field. However ZC-bearing anemones persist in low light and reduced temperature microhabitats, therefore more than the translocation of carbon from ZC must be involved. Given that global climate change will increase water temperatures, the potential for latitudinal range shifts of both ZC and ZX (S. californium and muscatinei) might be used as biological indicators of thermal shifts in the littoral zone of the Pacific Northwest.     

Chlorophyll fluorescence as a proxy for microphytobenthic biomass: alternatives to the current methodology

Pulse amplitude modulated (PAM) fluorescence has been used as a proxy of microphytobenthic biomass after a dark adaptation period of 15 min to stabilise the minimum fluorescence yield (F _o ¹⁵). This methodology was investigated for in situ migratory and ex situ engineered non-migratory biofilms, comparing dark adaptation to low (5% ambient) and far-red light treatments over different emersion periods. Far-red and low light reduced potential errors resulting from light history effects, by reversal of non-photochemical quenching after 5 min of treatment, compared to over 10 min required by conventional dark adaptation. An in situ decline of minimum fluorescence yield over 15 min was observed during the dark adaptation for migratory biofilms, but was not observed in the non-migratory biofilms indicating that the major cause of decline was downward vertical migration of cells into the sediment. This pattern occurred in far-red light after 10 min, but not for the low light treatment, indicating that low light maintained the biomass at the surface of the sediment. It is therefore concluded that low light treatment is a better option than conventional dark adaptation for the measurement of minimum fluorescence as a proxy of microphytobenthic biomass.     

Herring (Clupea harengus) filter-feeding in the dark

The behaviour of juvenile herring (Clupea harengus L.) feeding on Artemia sp. nauplii in both light and dark was recorded using an infra-red-sensitive television-recording technique. In the light, two modes of feeding were observed, particle biting and filtering, but in the dark only filtering was observed. Marked differences in swimming behaviour were seen between light and dark. In the light, the fish continued to school while feeding in both modes; in the dark, the school dispersed, the fish swam slower in tight circular paths and fed only by filtering. In the dark, filtering fish swam faster (0.11 m s^-1) than non-filtering fish (0.07 m s^-1). In the light, no difference in speed was measured between filtering and non-filtering fish (0.34 m s^-1). Owing to the lower filtering speeds in the dark, the removal rate of nauplii from the water was much lower than in light, except in the highest prey concentrations. This suggests that if night-time filter-feeding takes place in the sea, it will be of importance only when exploiting dense patches of food.     

Resting spore formation and biochemical composition of the marine planktonic diatom Chaetoceros pseudocurvisetus in culture: ecological significance of decreased nucleotide content and activation of the xanthophyll cycle by resting spore formation

The biochemical composition of vegetative cells and resting spores of Chaetoceros pseudocurvisetus Mangin was compared in cultures under various nutrient and light conditions. The cellular content of major nucleotides such as AMP, ADP, ATP and UTP decreased in the order: vegetative cells, nutrient-starved (vegetative) cells and resting spores, indicating that the general metabolism of resting spores is relatively inactive. ADP-glucose was only abundant in nutrient-starved vegetative cells, suggesting metabolic imbalance in these cells. The chl a–specific fluorescence yield of vegetative cells grown under all culture conditions was low, but very high in resting spores. The ratios of the cellular contents of diadinoxanthin to chl a and of diatoxanthin to chl a were higher in resting spores and nutrient-starved vegetative cells than in nutrient-replete vegetative cells. The diadinoxanthin–diatoxanthin xanthophyll cycle was active in resting spores; the xanthophyll cycle was synchronized with a 14 h light:10 h dark photoperiod. Also, the ratios of cellular content of diadinoxanthin and diatoxanthin to cellular content of chl a in resting spores were relatively high in high irradiance, and decreased gradually in conditions of darkness over long culture periods. Under conditions of strong light and high temperature, most resting spores survived more than 40 d while nutrient-starved vegetative cells died within 33 d. These results suggest that resting spore formation is a strategy for enhancing protection and lowering metabolic rate for survival. These physiological changes accompanying spore formation enable resting spores not only to overwinter but also to “oversummer” in the coastal euphotic layer. Received: 23 March 1999 / Accepted: 11 August 1999