Microsensor studies of photosynthesis and respiration in the symbiotic foraminifer Orbulina universa

Oxygen and pH microelectrodes were used to investigate the microenvironment of the planktonic foraminifer Orbulina universa and its dinoflagellate endosymbionts. A diffusive boundary layer surrounds the foraminiferal shell and limits the O₂ and proton transport from the shell to the ambient seawater and vice versa. Due to symbiont photosynthesis, high O₂ concentrations of up to 206% air saturation and a pH of up to 8.8, i.e. 0.5 pH units above ambient seawater, were measured at the shell surface of the foraminifer at saturating irradiances. The respiration of the host–symbiont system in darkness decreased the O₂ concentration at the shell surface to <70% of the oxygen content in the surrounding air-saturated water. The pH at the shell surface dropped to 7.9 in darkness. We measured a mean gross photosynthetic rate of 8.5 ± 4.0 nmol O₂ h⁻¹ foraminifer⁻¹. The net photosynthesis averaged 5.3 ± 2.7 nmol O₂ h⁻¹. In the light, the calculated respiration rates reached 3.9 ± 1.9 nmol O₂h⁻¹, whereas the dark respiration rates were significantly lower (1.7 ± 0.7 nmol O₂ h⁻¹). Experimental light–dark cycles demonstrated a very dynamic response of the symbionts to changing light conditions. Gross photosynthesis versus scalar irradiance curves (P vs E _o curves) showed light saturation irradiances (E _k) of 75 and 137 μmol photons m⁻² s⁻¹ in two O. universa specimens, respectively. No inhibition of photosynthesis was observed at irradiance levels up to 700 μmol photons m⁻² s⁻¹. The light compensation point of the symbiotic association was 50 μmol photons m⁻² s⁻¹. Radial profile measurements of scalar irradiance (E _o) inside the foraminifera showed a slight increase at the shell surface up to 105% of the incident irradiance (E _d). Received: 26 January 1998 / Accepted: 11 April 1998     

Diel cycles of activity,metabolism, and ammonium concentration in tropical holothurians

Movement rate, oxygen consumption, and respiratory tree ammonium concentration were measured in situ in the holothurians Pearsonothuria graeffei and Holothuria edulis in the Agan-an Marine Reserve, Sibulan, Philippines (9°20′30″N, 123°18′31″E). Measurements were made both day and night for both species during April–July 2005. P. graeffei had significantly higher movement rate during the day than at night (1.14 and 0.27 m h⁻¹, respectively; three-way ANOVA, P < 0.05) while H. edulis had higher movement rate at night compared to the day (0.83 and 0.07 m h⁻¹, respectively), spending the daylight hours sheltering under coral. More than 80% of H. edulis had movement rate of zero during the day. Oxygen consumption of P. graeffei was significantly higher during the day than at night (1.61 and 0.83 μmol O₂ g⁻¹ h⁻¹, respectively; two-way ANCOVA, P < 0.05), but the reduction at night was not as pronounced as the reduction in movement. H. edulis had a 75% reduction in oxygen consumption during the day compared to night (0.51 and 1.96 μmol O₂ g⁻¹ h⁻¹, respectively), matching this species’ reduced movement rates during the day. Ammonium concentration in water withdrawn from the respiratory trees of P. graeffei during the day (12.0 μM) was three times higher than in respiratory tree water sampled at night (4.3 μM) and 15 times higher than ambient seawater (0.8 μM; three-way ANOVA, P < 0.05). Ammonium concentration in the respiratory tree water of H. edulis was six times higher at night (14.6 μM) than during the day (2.2 μM) and 16 times higher than that of ambient seawater (0.9 μM). Even though H. edulis and P. graeffei are found within the same coral reef environment, they may affect different substrates and reef organisms due to their different habitats and distinct but opposite diel cycles.     

Metabolism and elemental composition of four oncaeid copepods in the western subarctic Pacific

Respiration rates and elemental composition (carbon and nitrogen) were determined for four dominant oncaeid copepods (Triconia borealis, Triconia canadensis, Oncaea grossa and Oncaea parila) from 0–1,000 m depth in the western subarctic Pacific. Across the four species of which dry weight (DW) varied from 2.0 to 32 μg, respiration rates measured at in situ temperature (3°C) increased with DW, ranging from 0.84 to 7.4 nl O₂ individual⁻¹ h⁻¹. Carbon (C) and nitrogen (N) composition of the four oncaeid species ranged from 49–57% of DW and 7.0–10.3% of DW, respectively, and the resultant C:N ratios were 4.8–8.3. The high C contents and C:N ratios were reflected by large accumulation of lipids in their body. Specific respiration rates (SR, a fraction of body C respired per day) ranged between 0.5 and 1.3% day⁻¹. Respiration rates adjusted to a body size of 1 mg body N (i.e. adjusted metabolic rates, AMR) of the four oncaeid species [0.6–1.1 μl O₂ (mg body N)^−0.8 h⁻¹ at 3°C] were significantly lower than those (1.7–5.1) reported in the literature for oithonid and calanoid copepods at the same temperature. The present results indicate that lower metabolic expenditure due to less active swimming (pseudopelagic life mode) together with rich energy reserve in the body (as lipids) are the characters of oncaeid copepods inhabiting in the epi- and mesopelagic zones of this region.     

Resource acquisition in two intertidal fucoid seaweeds, Fucus serratus and Himanthalia elongata : seasonal variation and effects of reproductive development

Seasonal variations and the effect of reproductive development on resource acquisition by two intertidal fucoid species, the iteroparous Fucus serratus L. and the semelparous Himanthalia elongata (L.) S. F. Gray were examined. The oxygen-exchange characteristics of vegetative apical tissue of both non-fertile and fertile plants and receptacle tissue were compared at monthly intervals throughout reproductive development. Respiratory rates in non-fertile F. serratus varied seasonally between 1.5 and 8.0 μmol g⁻¹ fresh wt h⁻¹; in fertile plants the receptacle had a significantly lower respiratory rate than the vegetative tissue. The respiratory rate of the vegetative button of fertile H. elongata displayed less seasonal variation and was lower than that of the receptacle, which varied from a maximum of 9.5 μmol g⁻¹ fresh wt h⁻¹ at receptacle initiation in October to a minimum of 2.0 μmol g⁻¹ fresh wt h⁻¹ in February. The maximum photosynthetic rate (P _max) of non-fertile plants of both species did not vary in a distinct seasonal manner (∼60 μmol g⁻¹ fresh wt h⁻¹ for F. serratus and ∼12 μmol g⁻¹ fresh wt h⁻¹ for H. elongata). In fertile plants, the P _max of the receptacle tissue was (∼50% lower in F. serratus, and at its peak three times higher in H. elongata, than that of vegetative tissue. The stable carbon-isotope ratio (δ¹³C) did not differ between different tissue types in F. serratus, but values did vary seasonally, being less negative in the summer than in the winter (−13.5‰ compared to −18‰). The receptacle tissue of H. elongata also displayed a distinct seasonal variation in δ¹³C values (−12‰ in summer, −16‰ in winter), whilst the δ¹³C of the vegetative button did not vary seasonally. The rate of uptake of inorganic nitrogen by the vegetative thallus was lower in H. elongata than in F. serratus. The receptacle tissue of F. serratus had lower uptake rates than the vegetative tissue, whilst the uptake rate by H. elongata receptacle tissue was higher than that of the vegetative button. Received: 14 March 1997 / Accepted: 22 April 1997     

Effects of mussel (Perna canaliculus) biodeposit decomposition on benthic respiration and nutrient fluxes

Suspension-feeding bivalves increase the quantity and quality of sedimenting organic matter through the production of faeces and pseudofaeces that are remineralised in coastal sediments and thus increase sediment oxygen demand and nutrient regeneration. Bivalves are intensively cultivated worldwide; however, no bivalve biodeposit decay rates are available to parameterise models describing the environmental effects of bivalve culture. We examined sediment biogeochemical changes as bivalve biodeposits age by incubating coastal sediments to which we added fresh mussel (Perna canaliculus) biodeposits and measured O₂ and nutrient fluxes as well as sediment characteristics over an 11-day period. Biodeposits elevated organic matter, chlorophyll a, phaeophytin a, organic carbon and nitrogen concentrations in the surface sediments. Sediment oxygen consumption (SOC) increased significantly (P=0.016) by ∼1.5 times to 1,010 μmol m⁻² h⁻¹ immediately after biodeposit addition and remained elevated compared to control cores without additions for the incubation period. This increase is in the range of observed in situ oxygen demand enhancements under mussel farms. To calculate a decay rate for biodeposits in sediments we fitted a first-order G model to the observed increase in SOC. The significant model fit (P=0.001, r ²=0.72) generated a decay rate of 0.16 day⁻¹ (P=0.033, SE=0.05) that corresponds to a half-life time of 4.3 day. This decay rate is 1–2 orders of magnitude higher than published decay rates of coastal sediments without organic enrichment but similar to rates of decaying zooplankton faecal pellets. NH₄⁺ release increased rapidly on the day of biodeposit addition (P=0.013) and reached a maximum of 144 μmol m⁻² h⁻¹ after 5 days which was 3.6 times higher compared to control cores. During this period NH₄⁺ release was significantly (P<0.001 to P=0.043) higher in the cores with biodeposit additions than in control cores.     

Effect of temperature fluctuations and food supply on the growth and metabolism of juvenile sea scallops (Placopecten magellanicus)

On the eastern shore of Nova Scotia late summer atmospheric systems cause upwelling of shelf water; the associated temperature variations of 10 °C with a 6 to 8 d period are comparable in magnitude to the seasonal variation. A laboratory study was undertaken to assess the effects of these temperature fluctuations on sea scallop (Placopecten magellanicus) growth and metabolism. In a factorial design, scallops were subjected to constant (10 °C) or a variable (6 to 15 °C) 8 d temperature cycle, and either a low (seston in filtered seawater) or high (seston supplemented with cultured phytoplankton) food diet. During the 48 d experiment scallop mortality was low and growth positive in all treatments. Shell and total tissue growth rate did not differ between temperature treatments, but growth in the high food treatments was 40 to 50% higher than in the low food treatments. However, soft tissue (excluding adductor) growth did show a temperature treatment effect; growth rates were significantly higher in the fluctuating temperature treatment, due in part to greater gonad development. Weight-standardized rates of scallop oxygen consumption (V _sO₂ , μmol O₂ g⁻¹ h⁻¹) were 20 to 25% higher in high food than in low food treatments, consistent with the expected increase in respiration due to the higher growth rates. Scallop metabolism did not acclimate to the fluctuating temperature cycle; V _sO₂ and ammonium excretion (V _sNH⁺ ₄, μmol O₂ g⁻¹ h⁻¹) remained dependent on ambient temperature throughout the experiment. V _sNH⁺ ₄ Q₁₀ (2.77) was higher than V _sO₂ Q₁₀ (2.01) which was reflected in a decrease in the O:N ratio at 15 °C, indicating a shift toward increased protein catabolism and a stressed state. At 10 °C, V _sO₂ and V _sNH⁺ ₄ in the variable temperature treatments were 15 to 18% lower than in the constant temperature treatments, a difference that was not detected in growth measurements. Results demonstrate that the metabolism of Placopecten magellanicus, unlike some bivalve species, is tightly coupled to fluctuations in ambient temperature. Although an absence of compensatory acclimation had a minimal effect on growth in this study, if high temperatures were combined with low food conditions a reduction in scallop production could result. Received: 23 June 1998 / Accepted: 8 February 1999     

Diurnal cycle and kinetics of ammonium assimilation in the green alga <Emphasis Type="Italic">Ulva pertusa</Emphasis>

The kinetics of ammonium assimilation was investigated in Ulva pertusa (Chlorophyceae, Ulvales) from northeastern New Zealand. Ammonium assimilation exhibited Michaelis–Menten kinetics with a maximum rate of assimilation (V _max) of 54 ± 5 μmol g⁻¹ dry weight h⁻¹ and half-saturation constant (K _m) of 23 ± 8 μM. In contrast, values for ammonium uptake were considerably higher with a V _max of 316 ± 59 μmol g⁻¹ dry weight h⁻¹ and K _m of 135 ± 46 μM. At environmentally relevant ammonium concentrations (5 μM), assimilation accounted for most (70%) of the ammonium taken up. Darkness decreased the maximum rate of ammonium assimilation by 83%. We investigated the hypothesis that rates of biosynthetic processes are greater in the early part of the day in Ulva. Consistent with this hypothesis, the maximum rate of ammonium assimilation in U. pertusa peaked in the morning and coincided with low levels of the photosynthetic product sucrose, which peaked in the afternoon. There was a diurnal cycle in the rate of ammonium uptake and assimilation in light and dark, but the amplitude was much greater for assimilation than uptake. Moreover, our data suggest that net ammonium assimilation only occurs during the day in U. pertusa. We suggest that two major roles for diurnal cycles are minimisation of interspecific competition for resources and metabolic costs.     

Oxygen dynamics and transport in the Mediterranean sponge Aplysina aerophoba

The Mediterranean sponge Aplysina aerophoba kept in aquaria or cultivation tanks can stop pumping for several hours or even days. To investigate changes in the chemical microenvironments, we measured oxygen profiles over the surface and into the tissue of pumping and non-pumping A. aerophoba specimens with Clark-type oxygen microelectrodes (tip diameters 18–30 μm). Total oxygen consumption rates of whole sponges were measured in closed chambers. These rates were used to back-calculate the oxygen distribution in a finite-element model. Combining direct measurements with calculations of diffusive flux and modeling revealed that the tissue of non-pumping sponges turns anoxic within 15 min, with the exception of a 1 mm surface layer where oxygen intrudes due to molecular diffusion over the sponge surface. Molecular diffusion is the only transport mechanism for oxygen into non-pumping sponges, which allows total oxygen consumption rates of 6–12 μmol cm⁻³ sponge day⁻¹. Sponges of different sizes had similar diffusional uptake rates, which is explained by their similar surface/volume ratios. In pumping sponges, oxygen consumption rates were between 22 and 37 μmol cm⁻³ sponge day⁻¹, and the entire tissue was oxygenated. Combining different approaches of direct oxygen measurement in living sponges with a dynamic model, we can show that tissue anoxia is a direct function of the pumping behavior. The sponge-microbe system of A. aerophoba thus has the possibility to switch actively between aerobic and anaerobic metabolism by stopping the water flow for more than 15 min. These periods of anoxia will greatly influence physiological variety and activity of the sponge microbes. Detailed knowledge about the varying chemical microenvironments in sponges will help to develop protocols to cultivate sponge-associated microbial lineages and improve our understanding of the sponge-microbe-system.     

Microsensor studies of photosynthesis and respiration in larger symbiotic foraminifera. I The physico-chemical microenvironment of Marginopora vertebralis, Amphistegina lobifera and Amphisorus hemprichii

 The physico-chemical microenvironment of larger benthic foraminifera was studied with microsensors for O₂, CO₂, pH, Ca²⁺ and scalar irradiance. Under saturating light conditions, the photosynthetic activity of the endosymbiotic algae increased the O₂ up to 183% air saturation and a pH of up to 8.6 was measured at the foraminiferal shell surface. The photosynthetic CO₂ fixation decreased the CO₂ at the shell down to 4.7 μM. In the dark, the respiration of host and symbionts decreased the O₂ level to 91% air saturation and the CO₂ concentration reached up to 12 μM. pH was lowered relative to the ambient seawater pH of 8.2. The endosymbionts responded immediately to changing light conditions, resulting in dynamic changes of O₂, CO₂ and pH at the foraminiferal shell surface during experimentally imposed light–dark cycles. The dynamic concentration changes demonstrated for the first time a fast exchange of metabolic gases through the perforate, hyaline shell of Amphistegina lobifera. A diffusive boundary layer (DBL) limited the solute exchange between the foraminifera and the surrounding water. The DBL reached a thickness of 400–700 μm in stagnant water and was reduced to 100–300 μm under flow conditions. Gross photosynthesis rates were significantly higher under flow conditions (4.7 nmol O₂ cm⁻³ s⁻¹) than in stagnant water (1.6 nmol O₂ cm ⁻³ s⁻¹), whereas net photosynthesis rates were unaffected by flow conditions. The Ca²⁺ microprofiles demonstrated a spatial variation in sites of calcium uptake over the foraminiferal shells. Ca²⁺ gradients at the shell surface showed total Ca²⁺ uptake rates of 0.6 to 4.2 nmol cm⁻² h⁻¹ in A. lobifera and 1.7 to 3.6 nmol cm⁻² h⁻¹ in Marginopora vertebralis. The scattering and reflection of the foraminiferal calcite shell increased the scalar irradiance at the surface up to 205% of the incident irradiance. Transmittance measurements across the calcite shell suggest that the symbionts are shielded from higher light levels, receiving approximately 30% of the incident light for photosynthesis. Received: 6 July 1999 / Accepted: 28 April 2000     

Photosynthetic utilisation of carbon and light by two tropical seagrass species as measured in situ

In situ measurements of seagrass photosynthesis in relation to inorganic carbon (Ci) availability, increased pH and an inhibitor of extracellular carbonic anhydrase were made using an underwater pulse amplitude modulated (PAM) fluorometer. By combining the instrument with a specially designed Perspex chamber, we were able to alter the water surrounding a leaf without removing it from the growing plant. Responses to Ci within the chamber showed that subtidal plants of the seagrasses Cymodocea serrulata and Halophila ovalis had photosynthetic rates that were limited by the ambient Ci concentration depending on the irradiance that was available during short-term photosynthesis–irradiance trials. Relative electron transport rates (RETRs) at light saturation (up to 500 μ mol photons m⁻² s⁻¹) increased by 66–100% when the Ci concentration was increased from ca. 2.2 to 6.2 mM. On the other hand, intertidal plants of the same species exhibited a much lesser limitation of photosynthesis by Ci at any irradiance (up to 1500 μ mol photons m⁻² s⁻¹). Both species were able to use HCO⁻ ₃ efficiently, and there was stronger evidence for direct uptake of HCO⁻ ₃ rather than extracellular dehydration of HCO⁻ ₃ to CO₂ prior to Ci uptake. Subtidally, H. ovalis and C. serrulata grew to 10 and 12 m, respectively, where ambient irradiances were approximately 16 and 11% of those at the surface. Maximum RETRs (at light saturation) were lower for these deep-growing plants than for the intertidally growing ones. For both species, the onset of light saturation of photosynthesis (E _k) occurred at approximately 100 μ mol photons m⁻² s⁻¹ for the deep water populations, which was four and two times lower than for the shallow populations of C. serrulata and H. ovalis, respectively. This, and the differences in maximal photosynthetic rates (RETR _max), reflects an acclimation of the deep-growing populations to the lower light environment. The results presented here show that photosynthesis, as measured in situ, was limited by the availability of Ci for the deeper growing plants in Zanzibar, while the intertidally growing plants photosynthesised at close to Ci saturation. The latter result is contrary to previous conclusions regarding Ci limitations for these intertidal plants, and, in general, our findings highlight the need for performing similar experiments in situ rather than under laboratory conditions. Received: 4 April 2000 / Accepted: 31 August 2000     

Spring sea ice photosynthesis,primary productivity and biomass distribution in eastern Antarctica, 2002–2004

While it is known that Antarctic sea ice biomass and productivity are highly variable over small spatial and temporal scales, there have been very few measurements from eastern Antarctic. Here we attempt to quantify the biomass and productivity and relate patterns of variability to sea ice latitude ice thickness and vertical distribution. Sea ice algal biomass in spring in 2002, 2003 and 2004 was low, in the range 0.01–8.41 mg Chl a m⁻², with a mean and standard deviation of 2.08 ± 1.74 mg Chl a m⁻² (n = 199). An increased concentration of algae at the bottom of the ice was most pronounced in thicker ice. There was little evidence to suggest that there was a gradient of biomass distribution with latitude. Maximum in situ production in 2002 was approximately 2.6 mg C m⁻² h⁻¹ with assimilation numbers of 0.73 mg C (mg Chl a)⁻¹ h⁻¹. Assimilation numbers determined by the ¹⁴C incubations in 2002 varied between 0.031 and 0.457 mg C (mg Chl a)⁻¹ h⁻¹. Maximum fluorescence quantum yields of the incubated ice samples in 2002 were 0.470 ± 0.041 with E _k indices between 19 and 44 μmol photons m⁻² s⁻¹. These findings are consistent with the shade-adapted character of ice algal communities. In 2004 maximum in situ production was 5.9 mg C m⁻² h⁻¹ with an assimilation number of 5.4 mg C (mg Chl a)⁻¹ h⁻¹. Sea ice biomass increased with ice thickness but showed no correlation with latitude or the time the ice was collected. Forty-four percent of the biomass was located in bottom communities and these were more commonly found in thicker ice. Surface communities were uncommon.     

Feeding, metabolism and growth in the Antarctic limpet, Nacella concinna (Strebel 1908)

Post-prandial increases in metabolism, the specific dynamic action of feeding (SDA), were evaluated in the Antarctic limpet Nacella concinna. O₂ consumption rose to a peak value 2.3 times higher than pre-feeding standard metabolic rates. This peak rise is low for marine ectotherms, but is typical of polar species. There were three peaks in the SDA, the first lasted only for the 1st day, was caused by handling, and was minor. The second was the major peak. It lasted from post-prandial days 4–9 inclusive, and accounted for around 70% of the SDA response. The third peak lasted from day 11 to day 15 and accounted for 30% of the total SDA. A 15-day SDA is much longer than values for temperate species, but is again typical for polar marine ectotherms. NH₃ excretion declined post-prandially from around 0.4 μmol animal⁻¹ h⁻¹ to values between 0.025 and 0.223 μmol animal⁻¹ h⁻¹ throughout the SDA. The total O₂ consumed in the SDA was 90.2 μmol O₂, which converts to 44.7 J of energy. This was 45–50% of the energy consumed in the meal (93.5 J). Pre-feeding O:N ratios, after 26 days without food, were around 1, indicating protein as the sole metabolic substrate prior to initiating the SDA. After feeding, O:N ratios rose to between 2.5 and 19, indicating significant use of lipid or carbohydrate from the food. Experiments were conducted in ambient seawater with enhanced levels of Sr (SrCl added at 800 mg kg⁻¹), and limpets were fed microalgal films also grown in enhanced Sr media. Sr incorporated in the shells during the experiment allowed the measurement of shell increments deposited during the SDA. Between five and eight microgrowth bands were present in the Sr-enhanced increments, which was similar to the number of days in the second SDA peak. The mean shell increment laid down was 17.6 μm. Estimating tissue deposition from measured growth increments and published ash-free dry mass (AFDM) to length relationships produced a value of 0.81 mg AFDM, which converted to 26.4 J of energy, or 25–30% of the energy ingested in the meal. Estimates of growth increments associated with a single SDA have not previously been possible. Overall energy used in the SDA and tissue deposition accounted for 75–80% of the energy ingested; the remainder was probably accounted for by unmeasured costs such as mucus production. Received: 6 June 2000 / Accepted: 20 September 2000     

Irradiance and temperature regulation of oxygenic photosynthesis and O2 consumption in a hypersaline cyanobacterial mat (Solar Lake, Egypt)

 Short-term effects of temperature and irradiance on oxygenic photosynthesis and O₂ consumption in a hypersaline cyanobacterial mat were investigated with O₂ microsensors in a laboratory. The effect of temperature on O₂ fluxes across the mat–water interface was studied in the dark and at a saturating high surface irradiance (2162 μmol photons m⁻² s⁻¹) in the temperature range from 15 to 45 °C. Areal rates of dark O₂ consumption increased almost linearly with temperature. The apparent activation energy of 18 kJ mol⁻¹ and the corresponding Q ₁₀ value (25 to 35 °C) of 1.3 indicated a relative low temperature dependence of dark O₂ consumption due to mass transfer limitations imposed by the diffusive boundary layer at all temperatures. Areal rates of net photosynthesis increased with temperature up to 40 °C and exhibited a Q ₁₀ value (20 to 30 °C) of 2.8. Both O₂ dynamics and rates of gross photosynthesis at the mat surface increased with temperature up to 40 °C, with the most pronounced increase of gross photosynthesis at the mat surface between 25 and 35 °C (Q ₁₀ of 3.1). In another mat sample, measurements at increasing surface irradiances (0 to 2319 μmol photons m⁻² s⁻¹) were performed at 25, 33 (the in situ temperature) and 40 °C. At all temperatures, areal rates of gross photosynthesis saturated with no significant reduction due to photoinhibition at high irradiances. The initial slope and the onset of saturation (E _k = 148 to 185 μmol photons m⁻² s⁻¹) estimated from P versus E _d curves showed no clear trend with temperature, while maximal photosynthesis increased with temperature. Gross photosynthesis was stimulated by temperature at each irradiance except at the lowest irradiance of 54 μmol photons m⁻² s⁻¹, where oxygenic gross photosynthesis and also the thickness of the photic zone was significantly reduced at 40 °C. The compensation irradiance increased with temperature, from 32 μmol photons m⁻² s⁻¹ at 25 °C to 77 μmol photons m⁻² s⁻¹ at 40 °C, due to increased rates of O₂ consumption relative to gross photosynthesis. Areal rates of O₂ consumption in the illuminated mat were higher than dark O₂ consumption at corresponding temperatures, due to an increasing O₂ consumption in the photic zone with increasing irradiance. Both light and temperature enhanced the internal O₂ cycling within hypersaline cyanobacterial mats. Received: 30 November 1999 / Accepted: 11 April 2000     

Photosynthetic performance of giant clams, Tridacna maxima and T. squamosa, Red Sea

Two species of giant clams, Tridacna maxima and T. squamosa, coexist in the Red Sea, but exhibit distinctly different depth distributions: T. maxima mostly occurs in shallow waters (reef flat and edge), while T. squamosa may occur down to the lower fore-reef slope. Giant clams have been described as mixotrophic, capable of both filter-feeding and photosynthesis due to algal symbionts (zooxanthellae), therefore, observed depth preferences were investigated in relation to possible differences in autotrophy vs. heterotrophy. This study was conducted from April to June 2004, at the reef near the Marine Science Station, Aqaba, Gulf of Aqaba, Red Sea, and in May 2007, at a reef near Dahab, Sinai Peninsula, Egypt. In situ measurements using a submersible pulse amplitude modulated fluorometer (Diving PAM), revealed no significant differences in effective PSII quantum yield (ΔF/Fm′) and relative electron transport rates (ETR) between the two species; but rapid light curves (ETR vs. light, photosynthetically active irradiance, PAR) showed significant differences in maximum photosynthetic rates (ETR_max), with 20% higher values in T. maxima. Chamber incubations displayed higher net and gross oxygen production by T. maxima (88.0 and 120.3 μmol O₂ cm⁻² mantle area day⁻¹) than T. squamosa (56.7 and 84.8 μmol O₂ cm⁻² mantle area day⁻¹); even under shading conditions (simulated depth of 20 m) T. maxima still achieved 93% of the surface gross O₂ production, whereas T. squamosa reached only 44%. A correlation was found between ETR and net photosynthesis measured as oxygen production (T. maxima: R ² = 0.53; T. squamosa: R ² = 0.61). Calculated compensation depth (CD) (gross photosynthesis equals respiration) in T. maxima (16 m) matches the maximum depth of occurrence in this study (17 m). By contrast, the CD of T. squamosa (9 m) was much shallower than the maximum vertical range (42 m). Findings suggest T. maxima is a strict functional photoautotroph limited by light, whereas T. squamosa is a mixotroph whose photoautotrophic range is extended by heterotrophy. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Habitat use,urea production and spawning in the gulf toadfish <Emphasis Type="Italic">Opsanus beta</Emphasis>

Field studies were conducted in Johnson Key Basin, Florida Bay, USA from September 2002 through September 2004 to examine physiological, ecological, and behavioral characteristics of the gulf toadfish, Opsanus beta (Goode and Bean in Proc US Natl Mes 3:333–345, 1880), in relation to nitrogen metabolism, habitat usage, and spawning. Fish collected 5 cm above sediments in experimental shelters (epibenthic) were compared with those collected by throw traps which were found on or burrowing within sediments. The relationship between microhabitat ammonia and urea excretion, as determined by the enzymatic activity of glutamine synthetase (GS), was examined. The hypothesis tested was that O. beta occupying epibenthic nests were less ureotelic with lower GS activities than non-nesting individuals on/in sediments, due to a decreased environmental ammonia burden. Porewater total ammonia (T _Amm) concentrations at a sediment depth of 5 cm, i.e., the approximate depth of burrowing toadfish, ranged from 0 to 106.5 μmol N l⁻¹ while the pH ranged from 7.48 to 9.14. There was a weak but significant correlation between environmental ammonia (NH₃) concentration and hepatic GS activity for epibenthic toadfish (P < 0.001, r ² = 0.10), but not for burrowing toadfish. Mean urea-N and T _Amm concentrations within shelters occupied by toadfish (n = 281) were 9.8 ± 0.83 μmol N l⁻¹ and 13.0 ± 0.7 μmol N l⁻¹, respectively. As predicted, hepatic GS activity was significantly lower in epibenthic toadfish captured in shelters (4.40 ± 0.24 μmol min⁻¹ g⁻¹; n = 281) as compared to individuals on/in sediments (6.61 ± 0.47 μmol min⁻¹ g⁻¹; n = 128). Glutamine synthetase activity generally peaked in March (spawning season) and was lowest in July. Gender differences in hepatic and branchial GS activity were also found during the spawning season, which is attributable to the fact that males brood and guard offspring in their epibenthic nests while females often rest on or burrow into the sediments. Finally, hepatic and branchial GS appeared to have different patterns of enzymatic activity suggesting functional differences in gene expression.     

Phytoplankton community structure and primary production in small intertidal estuarine-bay ecosystem (eastern English Channel,France)

From May 2002 to October 2003, a fortnightly sampling programme was conducted in a restricted macrotidal ecosystem in the English Channel, the Baie des Veys (France). Three sets of data were obtained: (1) physico-chemical parameters, (2) phytoplankton community structure illustrated by species composition, biovolume and diversity, and (3) primary production and photosynthetic parameters via P versus E curves. The aim of this study was to investigate the temporal variations of primary production and photosynthetic parameters in this bay and to highlight the potential links with phytoplankton community structure. The highest level of daily depth-integrated primary production Pz (0.02–1.43 g C m⁻² d⁻¹) and the highest maximum photosynthetic rate P ^B _max (0.39–8.48 mg C mg chl a ⁻¹ h⁻¹) and maximum light utilization coefficient α^B [0.002–0.119 mg C mg chl a ⁻¹ h⁻¹ (μmol photons m⁻² s⁻¹)] were measured from July to September. Species succession was determined based on biomass data obtained from cell density and biovolume measurements. The bay was dominated by 11 diatoms throughout the year. However, a Phaeocystis globosa bloom (up to 25 mg chl a m⁻³, 2.5 × 10⁶ cells l⁻¹) was observed each year during the spring diatom bloom, but timing and intensity varied interannually. Annual variation of primary production was due to nutrient limitation, light climate and water temperature. The seasonal pattern of microalgal succession, with regular changes in composition, biovolume and diversity, influenced the physico-chemical and biological characteristics of the environment (especially nutrient stocks in the bay) and thus primary production. Consequently, investigation of phytoplankton community structure is important for developing the understanding of ecosystem functioning, as it plays a major role in the dynamics of primary production.     

Activity and inhibitor sensitivity of ATPases in the hydrothermal vent tubeworm Riftia pachyptila : a comparative approach

Phosphorylated ATPases may be involved in the effective pH regulation seen in the hydrothermal vent tubeworm Riftia pachyptila. R. pachyptila appears not only to have a large concentration of ATPases, but the main function of these ATPases seems to have shifted from other types of transport, such as Na⁺ and K⁺ movement, to the facilitation of H⁺ elimination. Plume and trophosome ATPase activity for R. pachyptila measured 646.2 ± 29.5 and 481.4 ± 32.0 μmol Pi (inorganic phosphate) g⁻¹ wet wth⁻¹, respectively. Plume tissue ATPase activity (both mass-specific and protein-specific) in R. pachyptila was higher (between 7% and 55%) than the activity measured in any tissue for 7 other shallow- and deep-living species, in this study. This supports the hypothesis that R. pachyptila regulates acid/base balance via high concentrations of H⁺-ATPases, including Na⁺/H⁺ and K⁺/H⁺ exchangers and possibly electrogenic H⁺-ATPases, as evidenced by a higher total ATPase concentration (646 μmol Pi g⁻¹ wet wt h⁻¹), lesser Na⁺/K⁺-ATPase activity (13% of the total, as compared to 20−40% found in other animals), and higher H⁺-ATPase activity (226–264 μmol Pi g⁻¹wet wt h⁻¹). Overall, R. pachyptila appears to demonstrate elevated ATPase activity, with a greater fraction of the enzymes devoted to proton elimination, in order to effectively control its extracellular pH in the face of processes acting to acidify the internal environment. Received: 9 May 2000 / Accepted: 4 October 2000     

Growth and grazing responses of tintinnid ciliates feeding on the toxic dinoflagellate Heterocapsacircularisquama

Growth and feeding rates of two tintinnid species, Favellaazorica and Favellataraikaensis, were determined under various concentrations of the dinoflagellate Heterocapsacircularisquama which has been reported as highly toxic to shellfish. Mean growth rates of F. azorica and F. taraikaensis on a diet of H.circularisquama (ca. 10² cells ml⁻¹) were 2.15 and 1.97 doublings d⁻¹, respectively. These values are similar to those on a diet of Heterocapsatriquetra which is suitable food for various zooplankton. However, growth rates of both tintinnid species decrease with increasing concentrations of >10³ cells ml⁻¹ of H. circularisquama. In particular, H. circularisquama under conditions of >10³ cells ml⁻¹ caused mortality in F.taraikaensis, probably due to toxins. Clearance and ingestion rates of F. azorica on H. circularisquama were 4.1 to 27.5 μl ind⁻¹ h⁻¹ and 1.5 to 28.7 cells ind⁻¹ h⁻¹, respectively, at concentrations of <10⁴ cells ml⁻¹ and those of F. taraikaensis were 0.9 to 22.1 μl ind⁻¹ h⁻¹ and 0.1 to 13.0 cells ind⁻¹ h⁻¹, respectively, at concentrations of <10³ cells ml⁻¹. Both clearance and ingestion rates on H.circularisquama were higher for replicates fed on H.triquetra. Daily grazing impact of the two species of Favella on the initial stage of a bloom of H.circularisquama were estimated to reach 6 to 50% of H. circularisquama at a concentration of 540 cells ml⁻¹, indicating that grazing by tintinnids such as Favella spp. may significantly regulate the initial stages of blooms of H. circularisquama. Received: 3 January 1997 / Accepted: 27 January 1997     

Developmental physiology of Antarctic asteroids with different life-history modes

Rates of respiration and protein synthesis were measured during embryonic and larval development of Antarctic asteroids with different life-history modes (non-feeding and feeding larvae: Acodontaster hodgsoni, Porania antarctica, Odontaster meridionalis). Patterns of respiration for these species all show an increase during embryogenesis, with subsequent maintenance of routine respiration (“starvation resistance”), even in the absence of food for ~4 months (O. meridionalis). Fractional rates of protein synthesis (i.e., rate per unit mass of whole-body protein content) in the Antarctic larvae are essentially identical to those of temperate species. Larvae of O. meridionalis had an average fractional synthesis rate of 0.52% ± 0.05 h⁻¹ at −1.0°C, which is comparable to the temperate asteroid Asterina miniata at 0.53% ± 0.14 h⁻¹ at 15°C. For embryos of the asteroids A. hodgsoni and P. antarctica, fractional rates of protein synthesis (~0.2% h⁻¹) also are comparable to those reported for embryos of temperate echinoderm species. While rates of synthesis are high, rates of protein deposition are relatively low (percent of protein synthesized that is retained for growth). During a ~4 month growth period for larvae of O. meridionalis, the average protein depositional efficiency was 5.2%. This contrasts with higher rates of depositional efficiency reported for similar developmental stages of temperate echinoderm species. The biological significance of maintaining high rates of macromolecular synthesis for species with low rates of cell division and low protein depositional efficiencies is intriguing in the context of understanding the mechanistic bases of extended life spans and dispersal potential in response to changing Antarctic environments.     

In situ net primary productivity and photosynthesis of Antarctic sea ice algal, phytoplankton and benthic algal communities

Primary production at Antarctic coastal sites is contributed from sea ice algae, phytoplankton and benthic algae. Oxygen microelectrodes were used to estimate sea ice and benthic primary production at several sites around Casey, a coastal area in eastern Antarctica. Maximum oxygen export from sea ice was 0.95 mmol O₂ m⁻² h⁻¹ (~11.7 mg C m⁻² h⁻¹) while from the sediment it was 6.08 mmol O₂ m⁻² h⁻¹ (~70.8 mg C m⁻² h⁻¹). When the ice was present O₂ export from the benthos was either low or negative. Sea ice algae assimilation rates were up to 3.77 mg C (mg Chl-a)⁻¹ h⁻¹ while those from the benthos were up to 1.53 mg C (mg Chl-a)⁻¹ h⁻¹. The contribution of the major components of primary productivity was assessed using fluorometric techniques. When the ice was present approximately 55–65% of total daily primary production occurred in the sea ice with the remainder unequally partitioned between the sediment and the water column. When the ice was absent, the benthos contributed nearly 90% of the primary production.