Availability of two forms of dissolved nitrogen to the coral Pocillopora damicornis and its symbiotic zooxanthellae

The relative contribution of dissolved nitrogen (ammonium and dissolved free amino acids DFAAs) to the nitrogen budget of the reef-building coral Pocillopora damicornis was assessed for colonies growing on control and ammonium-enriched reefs at One Tree Island (southern Great Barrier Reef) during the ENCORE (Enrichment of Nutrient on Coral Reef; 1993 to 1996) project. P. damicornis acquired ammonium at rates of between 5.1 and 91.8 nmol N cm⁻² h⁻¹ which were not affected by nutrient treatment except in the case of one morph. In this case, uptake rates decreased from 80.5 to 42.8 nmol cm⁻² h⁻¹ (P < 0.05) on exposure to elevated ammonium over 12 mo. The presence or absence of light during measurement did not influence the uptake of ammonium ions. Nitrogen budgets revealed that the uptake of ammonium from concentrations of 0.11 to 0.13 μM could completely satisfy the demand of growing P. damicornis for new nitrogen. P. damicornis also took up DFAAs at rates ranging from 4.9 to 9.8 nmol N cm⁻² h⁻¹. These rates were higher in the dark than in the light (9.0 vs 5.1 nmol m⁻² h⁻¹, P < 0.001). Uptake rates were highest for the amino acids serine, arginine and alanine, and lowest for tyrosine. DFAA concentrations within the ENCORE microatolls that received ammonium were undetectable, whereas they ranged up to 100 nM within the control microatolls. The contribution of DFAAs to the nitrogen budget of P. damicornis constituted only a small fraction of the nitrogen potentially contributed by ammonium under field conditions. Even at the highest field concentrations measured during this study, DFAAs could contribute only ≃11.3% of the nitrogen demand of P.␣damicornis. This contribution, however, may be an important source of nitrogen when other sources such as ammonium are scarce or during periods when high concentrations of DFAAs become sporadically available (e.g. cell breakage during fish-grazing). Received: 22 April 1998 / Accepted: 3 November 1998     

Amino acid uptake by the toxic dinoflagellate Alexandrium fundyense

A toxic axenic strain of Alexandrium fundyense is shown to be capable of removing dissolved free amino acids (DFAAs) until concentrations are similar (low nM) to those found in natural waters. Uptake is greatest during exponential growth, rather than during C and/or N-stress as is usual in diatoms and other flagellates. A wide range of amino acids can be taken up, their concentration being decreased within a few hours to the levels observed prior to DFAA addition. The maximum rate of DFAA-N uptake, during early exponential phase, was 0.8 pmol-N cell⁻¹ h⁻¹, equivalent to ≃20% of the total N requirement. More typically, the contribution of DFAA-N was only ≃5%. However, these uptake rates are not sustainable. It is apparent that this organism cannot use amino-N to support significant growth, even though it can take up DFAAs. This, and the fact that the composition of the internal amino acid pool differed from that externally, is further evidence that the N-physiology of this genus is abnormal (differences to other dinoflagellates include an abnormally high concentration of glutamine and arginine, an effective absence of amine X, and release of nitrite during the␣concurrent assimilation of nitrate and ammonium in␣darkness). There is no evidence that the use of DFAAs enhance toxin content, except when cells are supplied with very high (unnatural) concentrations of arginine. Received: 8 May 1989 / Accepted: 14 September 1998     

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     

Organic nitrogen uptake and growth by the chrysophyte Aureococcus anophagefferens during a brown tide event

The quantitative importance of light-mediated, dissolved organic nitrogen (DON) utilization in relation to overall nitrogen-assimilation in Aureococcusanophagefferens Hargraves et Sieburth was assessed during a brown tide event in Shinnecock Bay, Long Island, 24 through 26 July 1995. The growth response of A. anophagefferens was maximal in organic-rich Bay water and decreased proportional to the organic:inorganic nutrient ratio of the water. Short-term uptake measurements with six nitrogenous substrates revealed that reduced nitrogen could potentially represent 95% of overall nitrogen uptake of which 70% was due to organic nitrogen alone. Potential uptake of urea by the A. anophagefferens-dominated bloom was substan tially greater than uptake of the other substrates tested during the study, contributing the largest percentage of total nitrogen uptake (58 to 64%; ρ^′ _max(urea) 4.4 μg  atom N l⁻¹ h⁻¹), followed by NH₄ ⁺ (18 to 26%; ρ′_max(NH4+) 2 μg atom N l⁻¹ h⁻¹). The combined rates of uptake of algal extract, lysine and glutamic acid contributed between 11 and 16% of total uptake, whereas NO₃ ⁻contributed 5 to 8%. Based on the kinetic determinations from this study we suggest an ecological framework for the events leading to the dominance and abundance of A. anophagefferens in coastal bays. Received: 29 March 1997 / Accepted: 24 April 1997     

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     

Production and biomass accumulation of periphytic diatoms growing on glass slides during a 1-year cycle in a subtropical estuarine environment (Bay of Paranaguá, southern Brazil)

Production rates, chlorophyll concentrations and general composition of periphytic diatom communities growing on glass slides were studied in relation to environmental parameters during one seasonal cycle in the Bay of Paranaguá, southern Brazil. Slides were routinely submersed at 1, 2 and 3 m depth and recovered weekly for microscopic examinations, analyses of chlorophyll, cell counts and in situ photosynthetic incubations using the Winkler titration method. Water samples were also collected at surface and bottom layers for determinations of temperature, salinity, nutrients and chlorophyll in the water. The periphytic community was mainly formed by epipelic and epipsammic species, dominated by Navicula phyllepta, Cylindrotheca closterium, Navicula spp. and Amphora sp. Weekly chlorophyll a and cell accumulations on slides varied from <1–32 mg m⁻² and up to 31 × 10⁸cells m⁻², respectively. Photosynthetic rates varied from <1 to 35 mg oxygen mg chlorophyll a ⁻¹ h⁻¹, with higher values in summer. Daily production varied from 5 to 3,600 mg oxygen m⁻² day⁻¹ (<0.01–1.4 g carbon m⁻² day⁻¹). Multiple regression analysis revealed that vertical differences in light conditions and grazing pressure jointly affected the influence of temperature on the seasonal patterns of cell densities and chlorophyll concentrations according to depth. Received: 27 April 2000 / Accepted: 16 August 2000     

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     

Abundance and variability of microorganisms and transparent exopolymer particles across the ice–water interface of melting first-year sea ice in the Laptev Sea (Arctic)

The distribution and abundance of transparent exopolymer particles (TEP) was determined in and below pack ice of the Laptev Sea from July to September 1995. Samples were collected from the lowermost 10 cm of ice floes and at 10 cm below the ice–water interface. Abundance of bacteria, protists and TEP was determined, and the sea ice–water boundary layer was characterized using temperature, salinity and molecular viscous shear stress. TEP, with a distinct size distribution signal, were found in highest concentrations inside the sea ice, ranging from not detectable to 16 cm² l⁻¹ (median: 2.9 cm² l⁻¹). In the water, concentrations were one order of magnitude lower, ranged from below detection to 2.7 cm² l⁻¹ (median: 0.2 cm² l⁻¹) and decreased after the middle of August, whereas abundances of autotrophic flagellates (AF), diatoms, heterotrophic flagellates (HF) and ciliates increased. The abundance of TEP decreased with its size in all samples following a power law relationship. The relation of TEP to the microbial community differed between the sea ice and water, being positively correlated with bacteria and diatoms in the ice and negatively correlated with HF in the sea water. The presence of a pycnocline significantly influenced the abundance of organisms, diatom composition and TEP concentrations. Pennate diatoms dominated by Nitzschia frigida were most abundant inside the ice. Though bacteria have the potential to produce exopolymeric substances (EPS), the results of this study indicate that the majority of TEP at the ice–water interface in first-year Arctic summer pack ice are produced by diatoms. Received: 19 August 1999 / Accepted: 4 July 2000     

Trophic ecology of Rimicaris exoculata : a combined lipid abundance/stable isotope approach

The alvinocaridid shrimp Rimicaris exoculata is an abundant component of the biota of Mid-Atlantic Ridge hydrothermal vents. To determine the nutritional strategy of this organism, we analysed the molecular abundance and carbon isotopic composition of its phospholipid fatty acids. High abundances of n-7 fatty acids (>40% total fatty acids) were observed in R. exoculata muscle tissues, in bacterial epibionts scraped from its gill bailers, and from the bacterially infested metal sulphides that the shrimp ingest. The phospholipid fatty acid abundance data indicates that the bacteria in the sulphides are closely related to the bacterial epibiota inhabiting the shrimp gill bailers, carapace and other body parts. Compound specific δ¹³C analyses of the phospholipid fatty acids gave average values of −12‰ for the epibiont bacteria and −21‰ for the sulphide bacteria. This difference may be largely due to the expression of different forms of RuBisCO (Forms I and II) which fractionate against ¹³C to different extents. Carbon limitation within the shrimp epibiont population may be an additional factor. The δ¹³C values (mean = −13‰) of the saturated and monounsaturated fatty acids isolated from the muscle tissues of R. exoculata were very close to those of the epibionts, indicating that the predominant source of dietary carbon for the shrimp is their epibionts, with a lesser contribution from free-living bacteria. The δ¹³C values (−26‰) of shrimp cholesterol were much more negative than those of the fatty acids, and this cholesterol is likely to have derived from the oceanic photic zone. Received: 26 June 1997 / Accepted: 6 November 1998     

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     

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     

Silica uptake kinetics of Halichondria panicea in Kiel Bight

Besides diatoms Demospongiae are the most important consumers of dissolved silica in the sea. They can play an important role for the silica budget especially in the shallow water areas of the Baltic Sea. The dependence of the silica uptake rate on the silica concentration of the seawater was measured for the sponge Halichondria panicea (Pallas, 1766). The sponges were collected in Kiel Bight. The uptake conformed to Michaelis–Menten kinetics with a half-saturation constant of 46.41 μM and a saturated uptake rate of 19.33 μmol h⁻¹ g⁻¹ ( p < 0.01). In the red algae zone of Kiel Bight the sponges depend on silica supply from the surrounding waters and may be silica-limited rather than food-limited in growth. Because of the much faster uptake of silica by diatoms and their lower saturation point, as well as the difference in spatial distribution of the two main silica consumers, a competition for silica between sponges and diatoms seems unlikely. Received: 21 June 1997 / Accepted: 15 July 1997     

Tissue distribution and neurotoxic effects of domoic acid in a prominent vector species, the northern anchovy Engraulis mordax

The planktivorous northern anchovy is a prominent vector of the phycotoxin domoic acid (DA) to organisms at higher trophic levels, including fish-eating seabirds and mammals. Although there are abundant data reporting DA-induced excitotoxic symptoms in higher vertebrates, to date there has been no reported evidence of neurotoxic effects in lower vertebrate vectors such as fish. To explain this apparent lack of toxicity, it has been suggested that DA may not reach the brain in anchovies and/or that fish are not as sensitive neurologically to DA. In the present study, intracoelomic (IC) injection of DA, at doses ranging from 1 to 14 μg DA g⁻¹ total fish weight, resulted in severe neurotoxic symptoms such as spinning, disorientation, inability to school, and mortality, indicating that anchovies are neurologically susceptible and that DA crosses the blood–brain barrier in fish. An ED₅₀ of 3.2 μg DA g⁻¹ total body weight was determined via IC injection of DA in 83 anchovies. Comparable intraperitoneal injection studies with mice, rats, and monkeys report similar DA-induced neurotoxic symptoms at doses near 3.2 μg DA g⁻¹, suggesting a similar neurologic sensitivity and mechanism of toxicity between anchovies and mammals. DA tissue distribution measurements from freshly collected field-exposed anchovies and orally gavaged anchovies indicate that DA uptake from the gastrointestinal tract does occur. Levels as high as 1,175 μg DA g⁻¹ were measured in anchovy viscera, while muscle and brain tissue DA levels were 3 orders of magnitude lower, indicating low but measurable DA uptake. Further evidence is needed to confirm that uptake is sufficient during field events to induce symptoms in anchovies. Our work provides the first reported evidence of neurotoxic symptoms in fish and suggests that anchovies may be affected by DA during toxic diatom blooms. If sufficient uptake occurs, DA-induced neurotoxic symptoms and mortality may make fish easier prey targets, thereby selecting for the highest toxin levels transferred, as well as providing a possible pathway for the transfer of DA to benthic communities. Received: 19 May 2000 / Accepted: 29 November 2000     

Concentrations of inorganic elements in 20 municipal waters in Sweden before and after treatment – links to human health

The water chemistry of 20 municipal water treatment plants in southern Sweden, representing various bedrock situations, and water qualities, were investigated. Four water samples, raw and treated, were collected from each plant and analyzed by predominantly ICP-OES and ICP-MS at four occasions from June to December, 2001. The concentrations of Ca, Mg, K, Na, HCO₃ and a number of micronutrients, varied considerably in treated waters from the studied plants (ranges; Ca: 9.1–53.7 mg L⁻¹, Mg: 1.4–10.9 mg L⁻¹, K: 1.1–4.8 mg L⁻¹, Na; 5.4–75.6 mg L⁻¹, HCO₃: 27–217 mg L⁻¹). The elimination of Fe and Mn from raw water was efficient in all treatments investigated, giving concentrations in treated waters below the detection limits at some plants. Softening filters gave waters with Ca-concentrations comparable to the softest waters in this study. Adjustment of pH by use of chemicals like lye, soda or lime, modified the consumer water composition significantly, besides raising the pH. It was estimated that drinking water contributed to approximately 2.2–13% of the daily Ca uptake, if the gastrointestinal uptake efficiency from food and water was estimated to be around 50%. The corresponding figures for Mg was 1.0–7% and for F 0–59%. None of the studied elements showed any significant time trends in raw or treated waters during the follow-up period. The concentrations of potentially toxic metals such as Al, Pb and U were low and did not indicate risks for adverse health effects (ranges; Al: 0.5–2.3 μg L⁻¹, Pb: 0–0.3 μg L⁻¹, U: 0.2.5 μg L⁻¹).     

Feeding and assimilation kinetics of Artemia franciscana fed Isochrysis galbana (clone T. Iso)

Artemia franciscana was grown on Isochrysis galbana Green (clone T. Iso) at saturated food concentrations (13 to 20 mg C l⁻¹) for 11 d at 26 to 28 °C, and 34 ppt salinity. Three groups of brine shrimp were used in the feeding experiments: metanauplius III and IV (Group 1), post-metanauplius II and III (Group 2) and post-metanauplius VIII (Group 3), corresponding to 4-, 7- and 11-d-old animals, respectively. The ingestion rate, clearance rate and carbon balance were estimated for these stages at different concentrations of ¹⁴C-labeled I. galbana ranging from 0.05 to 30 mg C l⁻¹. The handling time of algae was determined for all three groups. The ingestion rate (I, ng C ind⁻¹ h⁻¹) increased as a function of animal size and food concentration. In all three groups, the ingestion rate increased to a maximum level (I _max) and remained constant at food concentrations ≥10 mg C l⁻¹ (saturated food concentrations). The clearance rate (CR, μl ind⁻¹ h⁻¹) increased with increasing food concentration up to a maximum rate (CR _max), after which it decreased for even higher food concentrations. The functional response of A. franciscana was most consistent with Holling's Type 3 functional response curve (sigmoidal model), which for the two oldest groups (Group 2 and 3) differed significantly from a Type 2 response (p < 0.05). The gut passage time for the three groups of A. franciscana, feeding on saturated food concentration (20 mg C l⁻¹), varied between 24 and 29 min. As the nauplii developed to pre-adult stage the handling time of the algae increased as a function of animal size. The assimilation rate (ng C ind⁻¹ h⁻¹) in the youngest stages (Group 1 and 2) increased with increasing food concentrations, reaching a maximum level close to 10 mg C l⁻¹. At higher food concentrations the assimilation rate decreased, and the proportions of defecated carbon increased, reaching 60 to 68% in the post-metanauplius stages (Group 3). The assimilation efficiency (%) was high at the lowest food concentrations in all three groups (89 to 64%). At higher concentrations, the assimilation efficiency decreased, reaching 56 to 38% at the highest concentrations. Received: 2 February 2000 / Accepted: 25 March 2000     

Phytoplankton production patterns in Massachusetts Bay and the absence of the 1998 winter–spring bloom

The seasonal productivity cycle and factors controlling annual variation in the timing and magnitude of the winter–spring bloom were examined for several locations (range: 42°20.35′–42°26.63′N; 70°44.19′–70°56.52′W) in Boston Harbor and Massachusetts Bay, USA, from 1995 to 1999, and compared with earlier published data (1992–1994). Primary productivity (mg C m⁻² day⁻¹) in Massachusetts Bay from 1995 to 1999 was generally characterized by a well-developed winter–spring bloom of several weeks duration, high but variable production during the summer, and a prominent fall bloom. The bulk of production (mg C m⁻³ day⁻¹) typically occurred in the upper 15 m of the water column. At a nearby Boston Harbor station a gradual pattern of increasing areal production from winter through summer was more typical, with the bulk of production restricted to the upper 5 m. Annual productivity in Massachusetts Bay and Boston Harbor ranged from a low of 160 g C m⁻² year⁻¹ to a high of 787 g C m⁻² year⁻¹ from 1992 to 1999. Mean annual productivity was higher (mean=525 g C m⁻² year⁻¹) and more variable near the harbor entrance than in western Massachusetts Bay. At the harbor station productivity varied more than 3.5-fold (CV=40%) over an 8 year sampling period. Average annual productivity (305–419 g C m⁻² year⁻¹) and variability around the means (CV=25–27%) were lower at both the outer nearfield and central nearfield regions of Massachusetts Bay. Annual productivity in 1998 was unusually low at all three sites (<220 g C m⁻² year⁻¹) due to the absence of a winter–spring phytoplankton bloom. Potential factors influencing the occurrence of a spring bloom were investigated. Incident irradiance during the winter–spring period was not significantly different (P > 0.05) among years (1995–1999). The mean photic depth during the bloom period was significantly deeper (P < 0.05) in 1998, signifying greater light availability with depth. Nutrients were also in abundance during the winter–spring of 1998 with stratified conditions not observed until May. In general, the magnitude of the winter–spring bloom in Massachusetts Bay from 1995 to 1999 was significantly correlated with winter water temperature (r ²=0.78) and zooplankton abundance (r ²=0.74) over the bloom period (typically February–April). The absence of the 1998 bloom was associated with higher than average water temperature and elevated levels of zooplankton abundance just prior to, and during, the peak winter–spring bloom period. Received: 3 July 2000 / Accepted: 6 December 2000     

Strong variability in bacterioplankton abundance and production in central and western Bay of Bengal

With large influx of freshwater that decreases sea-surface salinities, weak wind forcing of <10 m s⁻¹ and almost always warm (>28°C) sea-surface temperature that stratifies and shallows the mixed layer leading to low or no nutrient injections into the surface, primary production in Bay of Bengal is reportedly low. As a consequence, the Bay of Bengal is considered as a region of low biological productivity. Along with many biological parameters, bacterioplankton abundance and production were measured in the Bay of Bengal during post monsoon (September–October 2002) along an open ocean transect, in the central Bay (CB, 88°E) and the other transect in the western Bay (WB). The latter representing the coastal influenced shelf/slope waters. Bacterioplankton abundances (<2 × 10⁹cells l⁻¹) were similar to those reported from the HNLC equatorial Pacific and the highly productive northern Arabian Sea. Yet, the thymidine uptake rates along CB (average of 1.46 pM h⁻¹) and WB (average of 1.40 pM h⁻¹) were less than those from the northwestern Indian Ocean. These abundances and uptake rates were higher than those in the oligotrophic northwestern Sargasso Sea (<7 × 10⁸ cells l⁻¹; av 1.0 pM h⁻¹). Concentrations of chlorophyll a (chl a), primary production rates and total organic carbon (TOC) were also measured for a comparison of heterotrophic and autotrophic production. In the WB, bacterioplankton carbon biomass equaled ∼ 95% of chl a carbon than just 31% in the CB. Average bacterial:primary production (BP:PP) ratios accounted for 29% in the CB and 31% in the WB. This is mainly due to lower primary productivity (PP) in the WB (281 mg C m⁻² d⁻¹) than in the CB (306 mg C m⁻² day⁻¹). This study indicates that bacteria–phytoplankton relationship differs in the open (CB) and coastal waters (WB). Higher abundance and contrastingly low bacterial production (BP) in WB may be because of the riverine bacteria, brought in through discharges, becoming dormant and unable to reproduce in salinities of 28 or more psu. Heterotrophic bacteria appear to utilize in situ DOC rather rapidly and their carbon demand is ∼50% of daily primary production. It is also apparent that allochthonous organic matter, in particular in the western Bay, is important for meeting their carbon demand.     

Effects of ultraviolet radiation and water motion on the reef coral Porites compressa Dana: a flume experiment

The effects of water flow and ultraviolet radiation (UVR, 280–400 nm) on the reef coral Porites compressa Dana were explored in a manipulative flume experiment. The aim of this study was to determine whether this coral responds to changes in the UVR environment by adjusting the tissue concentration of UV-absorbing compounds (mycosporine-like amino acids, MAAs), and to see whether such an acclimation is affected by water flow. Also, calcification rate and chlorophyll-a concentration were measured during the experiment to estimate the potential costs (in the form of slowed growth and/or reduced photosynthetic capacity) to the coral–alga symbiosis of being exposed to UVR and producing MAAs. Branches of P. compressa from a single male colony were exposed to high or low flow (15 cm s⁻¹ and 3 cm s⁻¹, respectively) and ambient or no UVR in an outdoor, continuous-flow seawater system. Chlorophyll-a and MAA concentrations were determined after zero, 3 and 6 weeks of exposure to the experimental conditions. Increase in buoyant weight during the two 3-week periods was used to calculate calcification rate. The presence of UVR had a significant positive effect on total MAA concentration in the P. compressa colonies; however, there were significant interactions present. In colonies exposed to UVR, MAA concentration increased and then decreased to initial levels in high water flow, and increased steadily in low water flow. In colonies receiving no UVR, MAA concentration decreased steadily, declining 23% in 6 weeks. The absence of UVR did not result in higher chlorophyll-a concentrations, but the calcification rate was slightly affected by UVR. This study supports the putative photoprotective role of MAAs in P. compressa, and suggests that the costs of mitigating the effects of ambient UVR are detectable, but they are very small. Received: 29 February 2000 / Accepted: 20 September 2000     

Free amino acids and energy metabolism in eggs and larvae of seabass, Lates calcarifer

Contents of free amino acids (FAA), protein and ammonium ions together with rates of ammonia excretion and oxygen consumption were measured in order to study the role of FAA as an energy substrate in developing eggs and larvae of seabass (Lates calcarifer) maintained in seawater (30 ppt) at 28 °C without feeding. Initially eggs contained 25.3 nmol ind⁻¹ of FAA of which 21.5 nmol was rapidly utilised by the developing eggs and larvae during the period up to 40 h post spawning (PS) when nearly all the yolk had been resorbed. During the same period, a net increase in protein content of 1.7 μg ind⁻¹ was observed, indicating that the major part of the amino acids lost from the free pool had been polymerised into body proteins. Assuming that the balance of the FAA after protein synthesis was used entirely for energy metabolism, FAA appeared to be an important energy substrate during the embryonic stages (2 to 16 h PS); after hatching, the contribution of FAA to energy metabolism was less significant. From 50 h PS until the end of the study period at 100 h PS, amino acids derived from somatic protein were used for energy metabolism. For the overall period from just after spawning up to 100 h PS, the data indicate that ca. 14% of the total aerobic energy metabolism was derived from amino acid catabolism. Received: 26 September 1997 / Accepted: 1 April 1998     

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