Role of the bivalveCorbicula japonica in the nitrogen cycle in a mesohaline lagoon

The effect of the bivalveCorbicula japonica on the nitrogen cycle in Lake Shinji, a mesohaline brackish lagoon in Japan, was examined quantitatively based on field surveys and laboratory experiments carried out in the summer of 1982 and 1983. The biomass of the flesh ofC. japonica comprised 97% of the total biomass of the macrozoobenthos in summer. Total biomass ofC. japonica in the lake was estimated 30 986 t fresh wt. The concentration of suspended solids immediately above the lake bottom, whereC. japonica filters the water, was 1.5 to 4 times higher than that in the surface water. In laboratory experiments, the filtration rate was 5.0 litres g dry flesh wt^-1 h^-1, the excretion rates of ammonia and of feces and pseudofeces ofC. japonica were 200×10^-6 g N g dry flesh wt^-1 h^-1 and 33.4 mg dry wt g dry flesh wt^-1 h^-1, respectively at 27°C, the average summer water temperature in the lake. From our study, we estimated thatC. japonica filters almost the same amounts of particulate organic nitrogen produced in the lake by phytoplankton and that supplied from rivers, and that it excretes 30% of filtered nitrogen as feces or pseudofeces and 18% as ammonia.     

Nitrogen fixation by blue-green algae associated with the siphonous green seaweed Codium decorticatum: effects on ammonium uptake

Nitrogen fixation (acetylene reduction) at rates of up to 1.2 g N₂ g dry wt^-1 h^-1 was measured for the siphonous green seaweed Codium decorticatum. No nitrogenase activity was detected in C. isthmocladum. The nitrogenase activity was light sensitive and was inhibited by the addition of DCMU and triphenyl tetrazolium chloride. Additions of glucose did not stimulate nitrogen fixation. Blue-green algae (Calothrix sp., Anabaena sp., and Phormidium sp.) were implicated as the organisms responsible for the nitrogenase activity. They occurred in a reduced microzone within the C. decorticatum thallus where nitrogen fixation was optimized. Nitrogen fixation did not affect the kinetic constants for ammonium uptake in C. decorticatum (K_s=12.0 M, V_max=13.4 mol NH₃ g dry wt^-1 h^-1) determined using the perturbation method. Nevertheless, C. decorticatum thalli which fixed nitrogen had internal dissolved nitrogen concentrations which were over 1.4 times higher than in non-fixing thalli. This suggests that if C. decorticatum does derive part of its nitrogen requirement from the blue-green algae which it harbors, the transfer does not involve competition between this process and the uptake of ambient ammonium.     

Respiration and excretion rates of Calanus glacialis in arctic waters of the Barents Sea

The respiration and excretion rates of Calanus glacialis (Jaschnov) Copepodite Stages III, IV, V, and adult females from the drift-ice area east of Svalbard (Barents Sea) were measured in shipboard experiments during the period from 27 May to 13 June, 1983. The phytoplankton biomass and abundance varied considerably between localities, but these variations were not generally reflected in the respiration and excretion rates of the copepod. The respiration and excretion rates of C. glacialis at the ambient temperature of-1.8°C (average respiration rates of 0.95, 0.73, 0.57, and 0.60 l O₂ mg^-1 dry wt h^-1 for Copepodite Stage III, IV, V, and adult females, respectively) were similar to those previously reported for other large-sized copepods from cold or temperate areas. Average respiration and excretion rates tended to decrease with incubation time or time after capture. Measurements on ten occasions within a period of 27 h after capture revealed excretion rates of ammonium ranging between 2.9 and 16.8 for C III, 3.7 and 21.1 for C IV, 1.3 and 28.4 for C V, and 1.6 and 18.7 for adult females, all expressed as nmol mg^-1 dry wt h^-1. In all experiments, excretion rates of inorganic phosphate varied between 0.7 and 1.5 (C III), 0.5 and 1.1 (C IV), 0.2 and 0.8 (C V), and 0.3 and 1.0 (adult females) nmol mg^-1 dry wt h^-1. Ratios of O:N, O:P, and N:P indicated that much of the metabolic energy was derived from catabolism of proteins. Comparison of the turnover rate of carbon and nitrogen showed, however, that nitrogen turnover was between 2.6 and 8.9 times higher than that of carbon. This may indicate that the copepods deaminate ingested protein, with the carbon skeleton of the amino acids subsequently being used in the synthesis of lipid compounds, possibly wax esters.     

Studies on short-term variations of heterotrophic activity in the Kiel Fjord

In summer, 1973, a short-term study with frequent sampling was performed in the Kiel Fjord (FRG). Temperature, salinity, number of viable bacteria, maximum uptakevelocity of glucose and acetate, concentrations of chlorophyll a, labile organic substances and ammonia and phosphate were measured. Strong variations in all parameters during short periods of time could be demonstrated. The number of viable bacteria fluctuated between 26x10⁶ and 195x10⁶ bacteria/l. Changes in the maximum uptake-velocity (V _m ) of glucose between 0.23 and 0.85 μg C l^-1 h^-1 and in the V _m of acetate between 0.36 and 2.40 μg C l^-1 h^-1 were recorded. A highly significant correlation between the number of viable bacteria and the maximum uptake-velocity of the two solutes was found. It is assumed that, at least for this area, at certain times the plate counts represent a major part of the active bacteria flora.     

Metabolism of two dominant epibenthic echinoderms measured at bathyal depths in the Santa Catalina Basin

Metabolism of two abundant echinoderm species constituting 99.6% of the epibenthic megafauna in the Santa Catalina Basin, off southern California, USA was measured at 1 300 m during the 1979 “Bathyal Expedition”. Specimens of the ophiuroid Ophiophthalmus normani and the holothurian Scotoplanes globosa, collected by the submersible “Alvin”, were individually placed in respirometers, and measured in situ for O₂ consumption and ammonium excretion rates. For O. normani, weight-specific O₂ consumption rates decreased with increasing weight and were of comparable magnitude to rates of deep-sea and shallow-water ophiuroids; excretion rates were highly variable. Population O₂ consumption and excretion rates for O. normani (estimated from size-frequency distribution, abundance, and rate regression equations) were 1 129.28 μl O₂ m^-2 h^-1 and 27.30 nmol NH ₄ ⁺ m^-2 h^-1. Weight-specific O₂ consumption and ammonium excretion rates of S. globosa decreased with increasing weight and were of comparable magnitude to rases of shallow-water holothurians. Population O₂ constimption and excretion rates of S. globosa were 1.38 μl O₂ m^-2 h^-1 and 4.86 nmol NH ₄ ⁺ m^-2 h^-1. Combined population O₂ consumption rates for O. normani and S. globosa are of comparable magnitude to that of the sediment community and plankton in the benthic boundary layer (sediment and overlying 50 m water column) of the Santa Catalina Basin.     

Obligate mixotrophy inLaboea strobila,a ciliate which retains chloroplasts

The planktonic ciliateLaboea strobila Lohmann sequesters photosynthetically functional chloroplasts derived from ingested algae. The chloroplasts lie free in the cytoplasm and are most abundant just under the pellicle of the ciliate. The maximum rate of photosynthesis (P_max) was 925 pg C ciliate^-1h^-1 (3.7 pg C pg chl.a ^-1h^-1). At saturating irradiance, the amount of carbon fixed h^-1 equaled 12.6% of the body carbon of the ciliate. To grow,L. strobila requires both light and algal food. In the absence of food, survival ofL. strobila is significantly longer in the light than in the dark. Based on ingestion rate and photosynthetic rate, we calculate that photosynthesis can make an important contribution to this ciliate's carbon budget even when algal food is plentiful.     

Nitrogen fixation on a coral reef

Acetylene reduction was used to assess nitrogen fixation on all major substrates at all major areas over a period of 1 to 6 yr (1980–1986) at One Tree Reef (southern Great Barrier Reef). Experiments using ¹⁵N₂ gave a ratio of 3.45:1.0 for C₂H₂ reduced:N₂ fixed. Acetylene reduction was largely light-dependent, saturated at 0.15 ml C₂H₂ per ml seawater, and linear over 6 h. High fixation was associated with two emergent cyanophyte associations, Calothrix crustacea and Scytonema hofmannii, of limited distribution. Subtidally, the major contribution to nitrogen fixation came from well-grazed limestone substrates with an epilithic algal community in the reef flat and patch reefs (3 to 15 nmol C₂H₄ cm^-2 h^-1). Similar substrates from the outer reef slope showed lower rates. Nitrogen fixation on beach rock, intertidal coral rubble, reef crest and lagoon sand was relatively small (0.3 to 1.0 nmol C₂H₄ cm^-2 h^-1). Seasonal changes in light-saturated rates were small, with slight reduction only in winter. Rates are also reported for experimental coral blocks (13 to 39 nmol cm^-2 h^-1) and for branching coral inside and outside territories of gardening damselfish (3 to 28 nmol cm^-2 h^-1). This work supports the hypothesis that the high nitrogen fixation on the reef flat and patch reefs of the lagoon (34 to 68 kg N ha^-1 yr^-1) is because these subtidal areas support highly disturbed communities with the greatest abundance of nitrogen-fixing cyanophyte algae. It is calculated from a budget of all areas that One Tree Reef has an annual nitrogen fixation rate of 8 to 16 kg N ha^-1 yr^-1.     

Release of dissolved organic carbon from the estuarine intertidal macroalga Enteromorpha prolifera

The estuarine macroalga Enteromorpha prolifera was collected from Coos Bay, Oregon, USA during 1981, and its release of photosynthate as dissolved organic carbon (DOC) was studied using ¹⁴C as a tracer. During photosynthesis in 30 S sea water, with a fixation rate averaging 7.37 mg C g^-1 dry wt h^-1, release ranged from 0.13 to 0.57 mg C g^-1 dry wt h^-1 and from 1.65 to 6.23% of total fixed carbon. Release of DOC appears to be linear with time over 3 h. As exposed algae become increasingly desiccated, their photosynthetic rates decline dramatically, but upon reimmersion the highly desiccated algae lose a larger fraction of their fixed carbon than the slightly desiccated algae. This loss comes in a pulse release of DOC over the initial 15 min, followed by declining release rates. The pulse loss due to rainfall is 5 times greater than that due to tidal resubmergence, and may briefly exceed the prior photosynthetic rate. Although lowering the salinity from 30 to 5 does not substantially alter photosynthetic rates, it does increase the DOC release range up to 1.02 mg C g^-1 dry wt h^-1 and 16.10% of fixed carbon. Heterotrophic microbes from the algal habitat readily use the available DOC at about 15% h^-1.     

Unbalanced growth of Acinetobacter CS 13 in glucose-limited chemostats caused by sublethal CdCl2 concentrations

The marine cadmium-sensitive Acinetobacter CS 13 was grown in glucose-limited chemostats to study the chronic toxicity of CdCl₂. Addition of 25, 50 or 100 μg Cd²⁺ l^-1 caused disturbances of balanced growth. At the dilution rate (D) of 0.05 h^-1 the washout rate (A) became constant at 84, 78 or 66 h after addition of 25, 50 or 100 μg Cd²⁺ l^-1 The washout rates were calculated to be respectively 0.037 h^-1, 0.039 h^-1 and 0.077 h^-1. At D=0.1 h^-1, A became constant after 48, 42 and 36 h, and were the same as D for all three Cd concentrations. Acridine orange staining revealed that 99% of the cells fluoresced green (viable cells) and 1% red or orange (dead cells). However, under Cd stress the percentage of red and orange fluorescing cells increased up to 20%. At D=0.1 h^-1, viable and direct cell counts always gave the same result. At D=0.05 h^-1, however, viable cell counts were lower than direct cell counts after 2–3 d of Cd exposure. It is assumed that at D=0.05 h^-1, a part of the cells appearing green was already Cd injured, but the true yellowish green colour was not identified. Decreasing cell densities were accompanied by increasing glucose concentrations. Cells accumulated Cd (0–25 μg Cd²⁺ l^-1) in relation to exposure time and Cd concentration in the medium, but maximum Cd contents were the same, increasing from 5.5 or 5.7 μg g^-1 dry wt in the control to 15 μg g^-1 after 2–3 d (D=0.05 h^-1) or 16 μg g^-1 after 1–2 d (D=0.1 h^-1).     

Ecological growth strategies in the seaweeds Gracilaria foliifera (Rhodophyceae) and Ulva sp. (Chlorophyceae): Soluble nitrogen and reserve carbohydrates

The seaweeds Gracilaria foliifera (Rhodophyceae) and Ulva sp. (Chlorophyceae) were grown in an outdoor continuous-flow system at both ambient incident light (I₀) and 0.13 I₀. During the winter, both species accumulated substantial soluble nitrogen reserves (up to 1020 g-at N·g dry wt^-1 in G. foliifera and 630 g-at N·g dry wt^-1 in Ulva sp.). The rate at which these N reserves were depleted was proportional to the growth rate. Seaweeds grown at 0.13 I₀ had lower growth rates and higher levels of soluble tissue N than plants grown at I₀. During the spring-summer growing season, peaks in tissue N followed nutrient peaks in the ambient seawater. Ulva sp. had higher nutrient uptake and growth rates than G. foliifera and showed greater fluctuations in soluble tissue N. This may characterize opportunistic seaweed species with high biomass turnover rates. At I₀, the levels of starch (up to 340 mg·g dry wt^-1 in G. foliifera and 170 mg·g dry wt^-1 in Ulva sp.) were highest during the spring and summer. During this period, fluctuations in starch content were inversely related to growth rate and soluble tissue N. Seaweeds grown at 0.13 I₀ did not accumulate starch. Neither species was found to overwinter with starch reserves.     

Filter-feeding ethology of benthic invertebrates (ascidians). V. Influence of temperature on pumping,filtration and digestion rates and rhythms in Phallusia mamillata

The influence of temperature has been studied simultaneously on the pumping, filtration, and digestion rates of Phallusia mammillata (Cuvier, 1815). Eighteen experiments were made between 7° and 25°C on 5 individuals. The average velocities of the water current varied between 3.37 and 9.65 cm sec^-1 (maximum 34.90 cm sec^-1). No recognizable rhythm emerged; pumping was continuous except at 7°C, where it soon ceased. Above 20°C, the curves were irregular and reflected the high sensitivity of the ascidian. The pumping rate was highest at 15°C (mean=5,788 ml h^-1 g^-1 dry weight of organs). At 10°C, the mean was 3,560; at 20°C, 2,629 ml h^-1 g^-1 dry weight of organs. At 20°C, the coefficients of variation displayed higher values, indicating a more irregular pumping at this temperature. Although there was no filtration rhythm, the variability of the results was higher at 20°C and above. As for pumping, maximum values were observed at 15°C (mean=4,286 ml h^-1 g^-1 dry weight of organs) decreasing with lower and higher temperatures, such decreases being more marked at the higher temperatures. Means were 352 ml h^-1 g^-1 dry weight of organs at 7°C; 2,935 at 10°C; 1,995 at 20°C; 973 at 25°C. The mean temperature coefficients for the filtration rates were: Q₁₀ for 7° to 15°C=11.86, Q₁₀ for 10° to 20°C=0,66, Q₁₀ for 15° to 25°C=0.22. The filtering efficiency was fairly constant throughout an experiment; the pumping and filtration curves were in fact almost parallel. The filtering efficiency of the branchial sac was high (75 to 85%), with constant values at 10° and 15°C; it became smaller (59%) at 20°C, with a higher coefficient of variation. The digestion rate also displayed maximum values at 15°C (mean=5.47 mg of albumin equivalent 24 h^-1 g^-1 dry weight of organs). It was lower at 10°C (mean=3.60 mg) and reached its minimum at 20°C (mean=1.71 mg). The higher temperature affected the percentage of food utilization, which showed smaller values at 20°C (59%) than at 10°C (89%) and 15°C (87%).     

Relationship of oxygen consumption to swimming speed in Euphausia pacifica

Oxygen consumption rate was measured as a function of swimming velocity for the vertically migrating euphausiid Euphausia pacifica at two temperatures (8° and 12°C) and pressures (1 and 40 atm) typical of its bathymetric distribution. Increased swimming speed (x; mh^-1) required increased oxygen consumption (y; μl O₂ mg dry weight^-1 h^-1), described by the equation y = 0.012x + 0.64 at 8°C, and by y = 0.020x + 0.85 at 12° C. The current concept of low swimming costs of zooplankton, based on determinations of dead drag in copepods, is contradicted by our measurements. Temperature had a more profound effect on metabolism at higher swimming speeds (112 m h^-1; Q₁₀=2.8) than on standard metabolism (O m h^-1; Q₁₀=2.0), indicating that activity is more costly at higher temperatures. Pressure caused a small but significant (P>0.05) rise in the relationship of respiratory rate to swimming speed at both temperatures. The energy cost of vertical migration for E. pacifica was estimated by applying our data on oxygen consumption vs swimming speed to published observations on sonic scattering layer movement and the day-night distribution pattern of this species. Results indicate that the cost of a diel migration of 254 m, through a temperature change of 4 °C (8° to 12° C), would cancel any energetic benefit gained by time spent at the lower temperature typical of daytime depth. The act of vertical migration is energetically expensive; its cost should be thoroughly considered in attempts to describe the energetics of vertically migrating species.     

Carbon metabolism and strobilation in Cassiopea andromedea (Cnidaria: Scyphozoa): Significance of endosymbiotic dinoflagellates

Scyphopolyps and scyphomedusae of Cassiopea andromeda Forskål (Cnidaria, Scyphozoa) containing dinoflagellate endosymbionts (zooxanthellae) were investigated for rates and pathways of carbon fixation. Photosynthesis by the algae, accounting for 80 and 15 mol C h^-1 on a dry weight basis in medusae and polyps, respectively, by far exceeds dark incorporation of inorganic carbon by the intact association. Photosynthetic carbon fixation is operated via C₃ pathway of carbon reduction. DCMU-treatment (1×10^-6 M and 1×10^-5 M) completely inhibits light-dependent carbon assimilation. Major photosynthates presumably involved in a metabolite flow from algal symbionts to animal tissue are glycerol and glucose. A total of 5–10% net algal photosynthate appears to be seleased in vivo to the host. This is probably less than the energy supply ultimately required for the nutrition of the polyps and medusae. The presence of zooxanthellae proved to be indispensable for strobilation in the scyphopolyps. However, photosynthesis by algal symbionts as well as photosynthate release is obviously not essential for the initiation of ephyrae as is shown by DCMU-treatment, culture in continous darkness, and aposymbiotic controls. It is therefore concluded that strobilation is supported, but not triggered by algal photosynthetic activity. The induction of strobilation thus seems to depend on a more complex system of regulation.     

Metabolism and elemental composition of zooplankton from the Barents Sea during early Arctic summer

Rates of oxygen consumption, ammonia excretion and phosphate excretion were measured on a hydromedusae (Aglantha digitale), pteropods (Limacia helicina, Clione limacina), copepods (Calanus finmarchicus, C. glacialis, C. hyperboreus, Metridia longa), an amphipod (Parathemisto libellula), a euphausiid (Thysanoessa inermis) and a chaetognath (Sagitta elegans), all of which were dominant species in the Barents Sea during early summer 1987. Water and ash contents and elemental composition (C and N) were also analysed on the specimens used in these metabolic experiments. Between species variations were 67.8% to 94.7% of wet weight in water content, 6.4% to 56.5% of dry weight in ash content, 16.7% to 61.0% of dry weight in carbon content, and 4.3% to 11.2% of dry weight in nitrogen content. Oxygen consumption rates ranged from 0.33 to 13.8 l O₂ individual^-1 h^-1, ammonia excretion rates, from 0.0072 to 0.885 gN individual^-1 h^-1 and phosphate excretion rates, from 0.0036 to 0.33 g P individual^-1 h^-1. In general, higher rates were associated with larger species, but considerable differences were also seen between species. The ratios between the rates (O : N, N : P, O : P) exhibited a wide species-specific variation, indicating differences in dominant metabolic substrates. Typical protein oriented metabolism was identified only in S. elegans. From the results of metabolic rate measurements and elemental analyses, daily losses of body carbon and nitrogen were estimated to be 0.50 to 4.15% and 0.084 to 1.87%, respectively, showing faster turnover rates of carbon than that of nitrogen. Comparison of daily loss of body carbon of the Barents Sea zooplankton with that of the Antarctic zooplankton indicated reduced rates of the former (63% on average).     

Primary production and release of assimilated carbon by Chlamydomonas provasolii in culture

Radionuclide labeled sodium bicarbonate was used to measure the rates of primary production and release of metabolites by Chlamydomonas provasolii. The total carbon fixed was 23.18 nM (mg dry wt)^-1 h^-1 and approximately 5.56% of the total was released in organic form into the medium. Thin layer chromatography indicated that the materials released were the amino acids, glycine and leucine. These substances are similar to those identified in certain coral and flatworm symbioses with algae.     

Ammonium regeneration and assimilation in eelgrass (Zostera marina) beds

Regeneration and assimilation of ammonium in the water column and in sediments of eelgrass (Zostera marina L.) beds of Izembek Lagoon and Crane Cove, Alaska, USA and Mangoku-Ura, northeastern Japan, were investigated by using a ¹⁵N isotope dilution technique. In the water column of Mangoku-Ura, ammonium was regenerated at a rate of 12 nmol l^-1 h^-1 and assimilated at a rate of 74 nmol l^-1 h^-1. The ammonium regeneration rate in sediments ranged from 2 to 150 nmol g^-1 h^-1, and with one exception, exceeded ammonium assimilation in sediments (0.3 to 77 nmol g^-1 h^-1). The ammonium regeneration in the water column was of little significance for the nitrogen supply to the eelgrass bed ecosystem. Net ammonium production (regeneration minus assimilation) in the sediment of Izembek Laggon met nitrogen demand for eelgrass growth, suggesting that ammonium regeneration in the sediments was very important for the nitrogen cycle in the eelgrass bed ecosystem.     

Glycolate metabolism by Pseudomonas sp., strain S227, isolated from a coastal marine sediment

Glycolate excreted by phytoplankton is a potentially important nutrient for bacteria in coastal and estuarine environments. The metabolism of glycolate by Pseudomonas sp., strain S227, originally isolated from the New York Bight Apex, has been studied. The specific growth rate for this strain on glycolate is 0.156 doublings h^-1. The apparent V_max and K_m for glycolate uptake are 83.6 nmol min^-1 mg cell protein^-1 and 7.4x10^-8 M, respectively. The preferential respiration of the carboxyl carbon (C-1) and the incroporation of the hydroxymethyl carbon (C-2) suggest that the glycerate pathway is used for growth on glycolate. Alternatively, another pathway can be utilized which results in the complete catabolism of glycolate. Glycolate and lactate metabolism are also closely linked either by a common metabolic pathway or a common transport system other than the monocarboxylate transport system. The magnesium ion concentration is also important in glycolate metabolism. The characteristics of glycolate metabolism observed in Pseudomonas sp., strain S227, are advantageous in coastal and estuarine environments where glycolate production is intermittent, and the concentrations are low.     

Dynamique du phytoplancton et matière organique particulaire dans la zone euphotique de l'Atlantique Equatorial

At two fixed stations in the Equatorial Atlantic Ocean (0°–4° W), the physical, chemical and biological properties of the euphotic layer were determined for 14 d (Station A: 5–18 February, 1979) and 13 d (Station B: 20 October–7 November, 1979), respectively. The stability of the water column allowed comparison of 3 different “systems”: (i) a well-illuminated and nitrate-depleted mixed layer; (ii) a chlorophyll maximum layer (chl a _max) in the thermocline which is poorly illuminated (6.3% of surface irradiance); (iii) a well-illuminated but nitrate-rich (>0.9 μg-at l^-1) mixed layer. In each layer the particulate organic carbon (COP), nitrogen (NOP) and phosphorus (POP) contents were measured and compared with the phytoplankton biomass. In the chlorophyll maximum layer, the phytoplankton biomass contributed significantly to the total particulate organic matter (between 55 and 75%). In the nitrate-depleted mixed layer, the results varied according to whether the regression technique [COP=f(chl a)] was used, or the chl a synthesis during the incubation of the samples. With the former technique, the phytoplankton carbon (C _p) content appeared minimal, because the y intercept, computed using all the data of the water column, was probably overestimated for this layer. POP would be more associated with living protoplasm than with carbon and nitrogen in the three layers. In the chlorophyll a maximum layer it constitutes a valuable detritus-free biomass measurement, since 80% of the POP consist of phytoplankton phosphorus. The assimilation numbers (NA=μg C μg chl a ^-1 h^-1) were high in all three layers, but the highest values were recorded in the nitrate-depleted mixed layer (NA=15 μg C μg chl a ^-1 h^-1). In the chlorophyll maximum layer, light would be a limiting factor during incubation: between 10²⁵ and 8.10²⁴ quanta m^-2 d^-1 NA and light are positively correlated independant of nitrate concentration. The growth rates of phytoplankton (μ) were estimated and compared to the maximum expected growth rate. Our main conclusion was that despite very low biomass and nutrient content, the mixed layer was in a highly dynamic state, as evidenced by high rates of phytoplankton growth and short nutrient turnover times (1 d or less for PO-P₄ in the mixed layer versus 3 d in the thermocline). The presence of nitrate in the water column allows the development of a higher phytoplankton biomass but does not increase growth rate.     

Effects of feeding frequency and symbiosis with zooxanthellae on nitrogen metabolism and respiration of the coral Astrangia danae

Colonies of the temperate coral Astrangia danae occur naturally with and without zooxanthellae. Basal nitrogen excretion rates of nonsymbiotic colonies increased with increasing feeding frequency [average excretion rate was 635 ng-at N (mg-at tissue-N)^-1 h^-1]. Reduced excretion rates of symbiotic colonies were attributed to N uptake by the zooxanthellae. Nitrogen uptake rates of the zooxanthellae averaged 8 ng-at N (10⁶ cells)^-1 h^-1 in the dark and 21 ng-at N (10⁶ cells)^-1 h^-1 at 200 Ein m^-2 s^-1. At these rates the zooxanthellae could provide 54% of the daily basal N requirement of the coral if all of the recycled N was translocated. Basal respiration rates were 172 nmol O₂ cm^-2 h^-1 for starved colonies and 447 nmol O₂ cm^-2 h^-1 for colonies fed three times per week. There were no significant differences between respiration rates of symbiotic and nonsymbiotic colonies. N excretion and respiration rates of fed (symbiotic and nonsymbiotic) colonies increased greatly soon after feeding. N absorption efficiencies decreased with increasing feeding frequency. A N mass balance, constructed for hypothetical situations of nonsymbiotic and symbiotic (3×10⁶ zooxanthellae cm^-2) colonies, starved and fed 15 g-at N cm^-2wk^-1, showed that the presence of symbionts could double the N growth rate of feeding colonies, and reduce the turnover-time of starved ones, but could not provide all of the N requirements of starved colonies. Rates of secondary production, estimated from rates of photosynthesis and respiration were similar to those estimated for reef corals.