Microheterotrophic utilization of mucus released by the Mediterranean coral Cladocora cespitosa

The amount of mucus released by the Mediterranean coral Cladocora cespitosa (L.) was determined in laboratory experiments and the incorporation of mucus into bacterial biomass was investigated by means of incubation experiments in 1984. Mean mucus release was 8.5 g (mucus dry wt) pclyp^-1 h^-1 and amounted to 44% of the respiratory carbon losses of the coral since mean organic carbon content of freshly collected mucus was 102.2g C mg (mucus dry wt)^-1. Due to the abundance of C. cespitosa in the shallow littoral of the Bight of Piran, the energy content of mucus released is estimated to correspond to about 20% of the phytoplankton primary production in this area. Furthermore, the carbon conversion efficiency of 20% obtained from the bacterial population during decomposition of mucus indicates the high nutritional value of C. cespitosa mucus, although bacterial carbon onto mucus particles contributes less than 0.1% to the total organic carbon pool of the mucus.     

Mucous sheet formation on poritid corals: An evaluation of coral mucus as a nutrient source on reefs

The production, release, and subsequent consumption of coral mucus on reefs has been portrayed as a potential pathway for the transfer of coral and zooxanthellae production to other reef organisms. However, reported mucus production rates and analyses of nutritional value vary widely. Poritid corals provide a test system to measure mucus production because they produce mucous sheets which can be collected quantitatively. Fluid mucus and mucous sheets were collected fromPorites astreoides, P. furcata, P. divaricata during 1986 and 1987 on reefs in the San Blas Islands, Panama, La Parguera, Puerto Rico and the Florida Keys, USA. Mucus samples were collected from Indo-pacific poritids (P. australiensis, P. lutea, P. lobata, andP. murrayensis) on the Great Barrier Reef during 1985. Biochemical analyses of the fluid mucous secretions, and the derivative mucous sheet, indicate that the mucus is primarily a carbohydrateprotein complex.Porites fluid mucus had a mean caloric content of 4.7 cal mg^–1 ash-free dry weight (AFDW), while mucous sheets contained 3.5 cal mg^–1 AFDW. Sixty-eight percent of the mucous sheet was ash, while fluid mucus was 22% ash. The high ash and low organic contents suggest that mucous sheets have a low nutritional value. C:N ratios varied (range 6.9 to 13.7 for fluid mucus, and 4.8 to 5.9 for mucous sheets), but were similar to typical C:N ratios for marine organisms. Bacterial numbers and chlorophyll a concentrations were higher on mucous sheets than in the surrounding water. Although bacteria aggregate on mucous sheets, bacteria accounted for less than 0.1% of the carbon and nitrogen content of the mucous sheet. Lower C:N ratios in aged mucus, i.e. mucous sheets versus fluid mucus, were attributed to a loss of carbon rather than an increase in nitrogen. Mucous sheet production accounts for a small proportion (< 2% gross photosynthesis) of published values for coral production. In the San Blas Islands, Panama,P. astreoides produced mucous sheets at a rate of 1.5 g C m^–2 y^–1 and 0.3 g N m^–2 y^–1.P. astreoides andP. furcata produced mucous sheets with a lunar periodicity and may provide approximately monthly pulses of carbon and nitrogen to the reef food-web. However, the low annual production rates suggest that mucous sheets make a small contribution to overall energy flow on coral reefs.     

Grazing of the mixotrophic dinoflagellate Gymnodinium sanguineum on ciliate populations of Chesapeake Bay

In situ grazing rates for the mixotrophic dinoflagellate Gymnodinium sanguineum Hirasaka feeding on nanociliate populations of Chesapeake Bay were determined in June and October of 1990 using a gut clearance/gut fullness approach. Recently ingested prey were digested beyond the point of recognition at a rate of 23% h^-1. Estimates of in situ ingestion and clearance ranged from 0 to 0.06 prey dinoflagellate^-1 h^-1 and 0 to 5.8 l dinoflagellate^-1 h^-1, respectively, with daily removal of ciliate biomass representing 6 to 67% of the 20-m oligotrich standing stock. Daily consumption of ciliate biomass by G. sanguineum averaged 2.5% of body carbon and 4.0% of body nitrogen with maximal values of 11.6 and 18.5%, respectively. Ingestion of ciliates may help balance nitrogen requirements for G. sanguineum and give this species an advantage over purely photosynthetic dinoflagellates in nitrogen limited environments. By preying on ciliates, these dinoflagellates reverse the normal flow of material from primary producer to consumer and thereby influence trophodynamics of the microbial food web in Chesapeake Bay.     

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.     

Decarboxylation of environmental L-leucine by marine red algae

The uptake and metabolism of ¹⁴C-L-leucine by Cystoclonium purpureum (Huds) Batt. (Rhodophyllidaceae) at concentrations of 0.5–50 μM were studied. Leucine was taken up from seawater at rates of 10–160 nmol·g^-1 fresh weight·h^-1. At low leucine levels ( 5 μM) the amino acid was preferably incorporated into cellular protein; amine formation was less than 5%. However at leucine concentrations of 50 μM, more than 30% of the total leucine taken up was decarboxylated to 3-methylbutylamine. This effect is expressed even more in Dumontia increassata (O. F. Müll) Lamour. (Dumontiaceae). In both species some of the amine occurred as a non-volatile derivative (bound amine) from which free amine could quantitatively be released by acid hydrolysis. Leucine and amine uptake were inhibited by CCCP. However, from cells preloaded with both leucine and amine, only free amine is released in the presence of CCCP. The results indicate that decarboxylase activity will be switched on at increased levels of amino acids in seawater which may be expected frequently in habitats of littoral algae (tidal zone). A possible function of amine formation and release within the scope of chemical defence of benthic algae is discussed.     

Calorific,ash, carbon and nitrogen content in relation to length and dry weight of Parathemisto gaudichaudi (Amphipoda: Hyperiidea) in the North East Atlantic Ocean

The calorific, ash, carbon and nitrogen content, length and dry weight were determined for the hyperiid Parathemisto gaudichaudi (Guerin). Regression equations for all these variables were determined so that they can be estimated by calculation from measurements of length of the hyperiid. Mean values for total nitrogen and carbon were 7.79±0.85% and 36.80±4.18% of the dry weight, respectively. The carbon to calorific equivalent for P. gaudichaudi was 10.37 kcal g^-1 carbon (9.13 kcal g^-1 when corrected for nitrogen). The calorific value for ash-free adult P. gaudichaudi was 5.138 kcal g^-1±1.309 (4.510 kcal g^-1 when corrected for nitrogen). This large variation in the calorific content (coefficient of variation of 25.84%) can be accounted for largely by variation in the ash content (coefficient of variation of 21.84%). The calorific value determined for P. gaudichaudi is similar to that measured for other carnivorous crustaceans and adds support to the hypothesis that animals with high calorific content have a low fecundity and an energy-rich store which can be used as a buffer during unfavourable periods in their life.     

Elemental and biochemical composition of salps (Tunicata: Thaliacea)

Carbon and nitrogen content have been measured in the solitary and aggregate generations of 11 species of salps. Regression equations for each species and generation permit estimation of carbon or nitrogen content as a function of length of live individuals. Different species of the same length may have nearly tenfold differences in carbon content. Fractionation and biochemical analysis of some samples revealed that the organic content of salps is approximately 80% protein. Ash-free dry weights average 27% of dry weights; mean carbon content is 29% of ash-free dry weight. Excess ash-free dry weight not accountable as organic material is thought to be water of hydration.     

Feeding of Pseudocalanus minutus on living and non-living particles

A seasonal study of carbon content of living and of carbon and nitrogen content of non-living particulate material in seawater is presented. Grazing by Pseudocalanus minutus on living and non-living particles has been investigated over 1 year. Seasonal variations in the food uptake were associated with seasonal variations of each chemical component of the particles in the water. The amount of non-living carbon constituted the major part of the food ingested, irrespective of season. The ingested living carbon always accounted for a small fraction of the total copepod body carbon. The proportion of living carbon ingested could be equivalent to or even higher than non-living carbon at times during the late spring, summer and fall. The concentration of both living and non-living material within each particle peak of the spectra in the water seemed to affect the balance between non-living and living particle uptake. Non-living particles cannot be considered only as a supplementary food source for small copepods; they are a basic food for P. minutus at all times.     

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     

Feeding biology and carbon budget of the sediment-burrowing heart urchin Brissopsis lyrifera (Echinoidea: Spatangoida)

This study of the burrowing heart urchin Brissopsis lyrifera includes measurements on feeding and food selection, ingestion rate, absorption efficiency, ventilation and respiration. B. lyrifera regulated feeding depth, ingestion rate and absorption efficiency in relation to food source. When food was added to the top layer of sediment, B. lyrifera burrowed closer to the surface and ingested mainly enriched surface material, whereas it burrowed deeper and ingested deep-seated sediment in the controls. In non-enriched sediment, the feeding rate was 0.04 g sediment DW h^-1 ind.^-1, and, in macro- and microalga-enriched sediment the feeding rate was 0.06 and 0.08 g sediment DW h^-1 ind.^-1, respectively. Absorption efficiency of TOC was about 43% in non-enriched sediment and in microalga-enriched sediment, but was significantly lower (34%) in macroalga-enriched sediment. Absorption efficiency of N varied between 48% and 55%, and was independent of food source. B. lyrifera feed selectively, enriching the gut TOC content about 2-fold and N content about 2.5-fold. The C/N ratio was therefore lower within the gut compared to the surrounding sediment, while the faecal C/N ratio was almost equal to the surrounding sediment. The faeces were, however, slightly richer in TOC and N compared to the surrounding sediment. For 3-5 cm long B. lyrifera, water current rate varied between 4 and 24 ml water h^-1, with a mean of 11 ml h^-1. Mean respiration rate was 205 µl O₂ h^-1 ind.^-1. The water current rate was not sufficient for B. lyrifera to sustain itself by filter feeding only. However, organic-rich particles from the surface are suggested to be an important contribution to the diet. A carbon budget was calculated for B. lyrifera from measured values of consumption, absorption efficiency and respiration, in order to estimate annual production of B. lyrifera. Compared to literature values, growth was overestimated about tenfold in the budget. A large proportion of the absorbed carbon was suggested to leave the animal as dissolved carbon, through mucus production or through anaerobic pathways, either by the heart urchin or by micro-organisms in the gut.     

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).     

Amino acid uptake by the comatulid crinoid Cenometra bella (Echinodermata) following evisceration

Changes in the rates of utilization of dissolved compounds during the period of visceral mass regeneration were examined in the crinoid Cenometra bella (Hartlaub). Rates of respiraton and incorporation of labelled amino acids increase, reaching a maximum 2 d after evisceration and returning to normal after 14 d. Rates of incorporation of radioisotope into the organic components of the arms and cirri decrease, while incorporation rate into the visceral mass increases. Incorporation rates of amino acid-derived radioactivity into skeletal carbonate and the ash-free dry weight:protein ratio of arms, cirri and oral disc decrease, reaching a minimum 2 d following evisceration.     

Relationships between porewater nutrients and seagrasses in a subtropical carbonate environment

The feeding, diet and egg production of the copepod Acartia tonsa were dermined during ten experiments in Los Angeles Harbor, California, between November 1986 and October 1987. Copepods were incubated in situ, in quasi-natural food environments. Water temperatures ranged from 14.6 to 21.5°C. Particulate organic carbon and nitrogen (POC and PON) were high (534 to 3710 g Cl^-1, 51 to 459 Nl^-1) but dominated by small (<8 m diam) particles. Plankton (phytoplankton and microzooplankton) C-biomass composed about 10% of the total POC and was usually dominated by particles >8 m. Plankton biomass was always low. Daily ingestion rates ranged from 3 to 96% of body C; egg production ranged from 4 to 35% of body carbon. Mean ingestion and egg production rates during spring-summer were 1.9 and 1.5 times higher than average for the entire study, respectively. The average gross efficiency of egg production for the study was 80%; the spring-summer mean was 41%. Bivariate and multiple-regression analyses revealed that the ingestion rate was dependent upon both temperature and food availability, but that, below 21°C, egg production depended more upon temperature than upon food concentration. To detect dietary preferences, the composition of diet was compared with that of the food supply. Selective feeding was infrequent, but the diet was often dominated by dinoflagellates and ciliates. It would appear that within metabolic limits governed by temperature, the feeding response of A. tonsa is dependent upon food concentration, while egg production depends more on qualitative attributes of the food supply.     

Sloughed mucus and drag-reduction in a school of Atlantic silversides,Menidia menidia

Atlantic silversides, Menidia menidia, were collected by seine-net from the Newport River estuary, North Carolina, USA and maintained in the laboratory. Direct measurement showed that the amount of mucus sloughed off individual silversides was 567 g h^-1 at a swimming speed of 50 cm s^-1. Individual M. menidia were also allowed to swim in a flowtank filled with 0, 1, and 10 ppm of the synthetic drag-reducer Polyox to determine if any changes in tailbeat frequency or amplitude could be correlated with the level of this sloughed material. Tailbeat measurements between seawater controls and either Polyox concentration did not differ significantly. Calculations based on this evidence indicate that the amount of solubilized mucus sloughed off a school of 10 000 M. menidia would be at least two orders of magnitude less than the highest concentration of Polyox tested. We conclude that solubilized mucus in the water column does not confer drag-reduction to schooling M. menidia.     

Food selection by copepods: discrimination on the basis of food quality

The copepod Acartia tonsa displayed nearly two-fold higher ingestion rates on faster-growing cells of the diatom Thalassiosira weissflogii compared to ingestion rates on slower-growing cells of that species at the same cell concentration. Ingestion rates on slow-growing cells were also enhanced by the addition of cell-free aliquots of algal exudate to the experimental feeding chambers. In addition, the faster-growing algal cells were selectively ingested by the copepod when the two cell types were mixed together in different proportions, indicating that physiological differences between growing cells are a critical factor in the food detection/selection process of zooplankton. Consideration of cell carbon, nitrogen, and protein composition suggests that the copepods are maximizing nitrogenous ingestion (total protein and/or nitrogen). Selectivity for cells with higher protein content results in a higher daily protein ration, even if the selection process results in a decreased rate of ingestion in mixtures of cell types.     

Direct measurement of dissolved organic carbon release by phytoplankton and incorporation by microheterotrophs

It is now possible to divide particulate primary production into algal and heterotrophic components without physical separation. This depends on two innovations, the introduction of isotope in the form of labelled dissolved product(s) of primary production and the employment of a data analysis specifically designed for tracer kinetic incorporation experiments. The ¹⁴C technique described by Steemann Nielsen (1952) is inapplicable in the analyses of certain classes of systems and kinetic tracer incorporation experiments must be employed instead. We show that measurement of PDOC production rate requires such kinetic tracer analyses. Measurements made in the laboratory on water taken from 2 m depth in South West Arm of the Port Hacking estuary showed that: (1) the steady-state rate of PDOC production was 0.10 to 0.13 mg C.m^-3.h^-1; (2) the rate of PDOC incorporation into microheterotroph particulate organic carbon was 0.10 to 0.12 mg C.m^-3.h^-1; (3) the rate at which PDOC was respired to CO₂ was 0.001 to 0.003 mg C.m^-3.h^-1. (4) the PDOC makes up only about 0.1% of the total dissolved organic carbon. The size class of particles associated with PDOC production differed from the size class responsible for uptake of PDOC. More than 50% of the PDOC production was associated with particles having a nominal diameter range of 20 to 63 m, while this fraction was responsible for <10% of the incorporation.     

Flux capacities and acclimation costs in Trichodesmium from the Gulf of Mexico

Phytoplankton function and acclimation are driven by catalytic protein complexes that mediate key physiological transformations, including generation of photosynthetic ATP and reductant, and carbon and nitrogen fixation. Quantitation of capacities for these processes allows estimation of rates for key ecosystem processes, and identification of factors limiting primary productivity. We herein present molar quantitations of PSI, PSII, ATP synthase, RuBisCO and the Fe protein of nitrogenase of Trichodesmium collected from the Gulf of Mexico, in comparison to determinations for a range of cyanobacteria growing in culture. Using these measurements, estimates were generated for Trichodesmium capacities for carbon fixation of 1–3.4 g C g chl a ⁻¹ h⁻¹ and nitrogen fixation of 0.06–0.17 g N g chl a ⁻¹ h⁻¹, with diel variations in capacities. ATP synthase levels show that ATP synthesis capacity is sufficient to support these levels of carbon and nitrogen fixation, and that ATP synthase levels change over the day in accordance with the ATP demands of nitrogenase and RuBisCO activity. Levels of measured complexes indicate that Trichodesmium manifests n-type diel light acclimation through rapid changes in RuBisCO:PSII, supported by significant investment of cellular nitrogen. The plasticity in the levels and stoichiometry of these core complexes show that changes in the abundance of core protein complexes are an important component of acclimation and regulation of metabolic function by Trichodesmium populations. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Energetics of Euphausia pacifica. I. Effects of body carbon and nitrogen and temperature on measured and predicted production

Laboratory production during the life span of Euphausia pacifica was measured directly (as the sum of growth, molting and reproduction) and indirectly (as assimilation minus metabolism and leakage) to test the hypothesis that weight-specific production is a constant for all sizes. Euphausiids were collected in Puget Sound, Washington State, USA, from September 1973 to March 1978. Equations were determined (in terms of carbon and nitrogen at 8° and 12° C) expressing the relationships between body weight and the daily rates of growth, molting, reproduction, ingestion and metabolism. The allometric equation (R=aW ^b ) best related body weight (W) to the rate (R) for growth, molting, ingestion, respiration and excretion for life stages from late larvae through adults. As predicted by the original above hypothesis, the weight-specific coefficient (b) was close to 1.0 for ingestion and excretion; in contrast, b was 0.62 for growth, and 0.77 to 0.85 for molting and respiration. The Q₁₀ s also varied: 3.5 for growth, 2.4 for molting, about 3.0 for ingestion, and 2.0 for respiration and excretion. Assimilation efficiencies, for all weights and at both temperatures, were 81.3% of carbon and 85.9% of nitrogen ingested. The relationships between rate and body weight of early larvae for growth and molting were linear, as was the relationship for reproduction in adults. Weight-specific production was higher by I to 2% at 12° than 8° C for all life stages, and was 2 to 4% for carbon and 2 to 6% for nitrogen in adults, but 13 to 17% for carbon and 14 to 15% for nitrogen in early furcilia larvae. The null hypothesis was rejected for production measured directly, but would have been accepted if only an indirect measurement of nitrogen production had been considered. Clearly, indirect measurement incorporates all errors of measurement and assumption and makes interpretation difficult.     

Effect of the insect growth regulator diflubenzuron (Dimilin®) on the uptake of glucose and N-acetylglucosamine into the cuticle of crab larvae

Incorporation of ³H-labelled glucose and ³H-labelled N-acetylglucosamine (NAGA) — both precursors to chitin —into the cuticle of Rhithropanopeus harrisii (Gould) larvae (Crustacea: Brachyura) has been examined at different stages of the moult cycle in control larvae as well as in larvae treated with the insect growth regulator diflubenzuron (Dimilin^®). As far as the control larvae were concerned, the incorporation of both precursors was high at the postmoult stage when endocuticle was secreted. NAGA appeared to be a more specific precursor of cuticular material than glucose during the premoult stage when exocuticle was produced. Incorporation of both precursors was low immediately before ecdysis and during the intermoult stage when secretion of the cuticle is complete. The results show that incorporation of glucose into chitin was greatly inhibited by the pollutant during the postmoult stage when endocuticle is produced, while incorporation of NAGA was reduced to a lesser extent at this stage. Diflubenzuron treatment markedly affected the incorporation of both NAGA and glucose in the premoult stage during secretion of exocuticle.     

Functional responses of juvenile herring and sprat in relation to different prey types

The relationship between particulate-feeding rates and prey concentrations (functional response) of juvenile herring and sprat (5–9 cm total length) was investigated in controlled feeding experiments monitored by an underwater camera system. A special tank system was developed allowing the regulation and quantification of low prey concentrations (1–160 L^?1). Non-evasive Artemia nauplii was used as prey to estimate the maximum biting rate of both predators. In contrast, Acartia tonsa with a high escape ability was used as a realistic prey type. Herring and sprat showed a type II functional response for both prey types. Nonlinear mixed effects model revealed no significant difference between the functional responses of both predators, except that herring showed significantly higher biting rates than sprat at A. tonsa concentrations below ~40 L^?1. For both predators feeding rates were significantly higher with Artemia nauplii than with A. tonsa. Video analysis indicated that sprat, unlike herring, is an obligate particulate-feeder.