Feeding strategies of planktonic cyclopoids in lacustrine ecosystems

Present knowledge of feeding tactics and behaviour, food composition and consumption, and feeding strategies of planktonic cyclopoid copepods is synthesized. Planktonic freshwater cyclopoids consume both plant and animal food. Predatory feeding is highly selective: prey species differ in their size, defense structures, the distance at which they are recognized by the cyclopoid, defensive behaviour when attacked, and their occurrence in the same space as the predator. Within a prey species, cyclopoids select for smaller individuals. The impact of cyclopoid predation on the other zooplankton may be an important source of mortality. However, algal material is consumed to a large extent by the adult and later developmental stages and is a necessary food source for the youngest stages, which have to compete with other planktonic herbivores. Some implications of these food requirements for life strategies of planktonic cyclopoid species in the seasonally changing environments are discussed.     

Modelling Pseudocalanus elongatus stage-structured population dynamics embedded in a water column ecosystem model for the northern North Sea

This paper outlines an approach to couple a structured zooplankton population model with state variables for eggs, nauplii, two copepodites stages and adults adapted to Pseudocalanus elongatus into the complex marine ecosystem model ECOHAM2 with 13 state variables resolving the carbon and nitrogen cycle. Different temperature and food scenarios derived from laboratory culture studies were examined to improve the process parameterisation for copepod stage dependent development processes. To study annual cycles under realistic weather and hydrographic conditions, the coupled ecosystem–zooplankton model is applied to a water column in the northern North Sea. The main ecosystem state variables were validated against observed monthly mean values. Then vertical profiles of selected state variables were compared to the physical forcing to study differences between zooplankton as one biomass state variable or partitioned into five population state variables. Simulated generation times are more affected by temperature than food conditions except during the spring phytoplankton bloom. Up to six generations within the annual cycle can be discerned in the simulation.     

Fluxes of biogenic carbon in the Southern Ocean: roles of large microphagous zooplankton

The Southern Ocean is an extreme environment, where waters are permanently cold, a seasonal ice cover extends over large areas, and the solar energy available for photosynthesis is severely restricted, either by vertical mixing to considerable depths or, especially south of the Antarctic Circle, by prolonged seasonal periods of low or no irradiance. Such conditions would normally lead to low productivity and a water column dominated by recycling processes involving microbial components of pelagic communities but this does not seem to be the case in the Southern Ocean, where there is efficient export to large apex predators and deep waters. This paper investigates the role of large microphagous zooplankton (salps, krill, and some large copepods) in the partitioning of biogenic carbon among the pools of short- and long-lived organic carbon and sequestered biogenic carbon. Large microphagous zooplankton are able to ingest microbial-sized particles and thus repackage small, non-sinking particles into both metazoan biomass and large, rapidly sinking faeces. Given the wide spatio-temporal extent of microbial trophic pathways in the Southern Ocean, large zooplankton that are omnivorous or able to ingest small food particles have a competitive advantage over herbivorous zooplankton. Krill efficiently transfer carbon to a wide array of apex predators and their faecal pellets are exported to depth during occasional brief sedimentation episodes in spring time. Salps may be a significant link towards some fish (directly) and other apex predators (indirectly) and, at some locations (especially in offshore waters) and time, they may account for most of the downward flux of biogenic carbon. Large copepods are a trophic link towards fish and at least one whale species, and their grazing activity generally impedes the export of organic particles to depth. As a result, biogenic carbon is channelled mainly towards apex predators and episodically into the deep ocean. Without these original interactions, Antarctic waters might well be dominated by microbial components and recycling processes instead of active export from the generally small primary producers towards large apex predators.     

Community structure of mesopelagic copepods (>500 μm) in the Ligurian Sea (Western Mediterranean)

The distribution of mesopelagic copepods between 250 and 1000 m depth was studied along a transect off the coast of Villefranche-sur-Mer in June 1991 by means of a BIONESS multiple-net sampler. Among the most abundant species at all stations were Pleuromamma gracilis, Paraeuchaeta acuta and overwintering Calanus helgolandicus CV copepodites, each species inhabiting different depth layers with maximum abundances in the 250–350 m, 450–550 m and the 700–850 m layer, respectively. Cluster analysis of the vertical distribution of all species caught revealed three distinct strata (‘TOP', ‘INTERMEDIATE', ‘BOTTOM') consisting of characteristic species that occurred in the same stratum at all stations. Differences in total abundances of the species assemblage occupying each layer are discussed with regard to predation pressure by mesopelagic macroplanktonic crustaceans and fish. We show that each stratum is characterized by distinct trophic interactions and life strategies, which play an important role in the structuring of mesopelagic zooplankton communities.     

Study of the seasonal cycle of the biogeochemical processes in the Ligurian Sea using a 1D interdisciplinary model

A one-dimensional coupled physical–biogeochemical model has been built to study the pelagic food web of the Ligurian Sea (NW Mediterranean Sea). The physical model is the turbulent closure model (version 1D) developed at the GeoHydrodynamics and Environmental Laboratory (GHER) of the University of Liège. The ecosystem model contains 19 state variables describing the carbon and nitrogen cycles of the pelagic food web. Phytoplankton and zooplankton are both divided in three size-based compartments and the model includes an explicit representation of the microbial loop including bacteria, dissolved organic matter, nano-, and microzooplankton. The internal carbon/nitrogen ratio is assumed variable for phytoplankton and detritus, and constant for zooplankton and bacteria. Silicate is considered as a potential limiting nutrient of phytoplankton's growth. The aggregation model described by Kriest and Evans in (Proc. Ind. Acad. Sci., Earth Planet. Sci. 109 (4) (2000) 453) is used to evaluate the sinking rate of particulate detritus. The model is forced at the air–sea interface by meteorological data coming from the “Côte d'Azur” Meteorological Buoy. The dynamics of atmospheric fluxes in the Mediterranean Sea (DYFAMED) time-series data obtained during the year 2000 are used to calibrate and validate the biological model. The comparison of model results within in situ DYFAMED data shows that although some processes are not represented by the model, such as horizontal and vertical advections, model results are overall in agreement with observations and differences observed can be explained with environmental conditions.     

Dynamics of suprabenthos-zooplankton communities around the Balearic Islands (western Mediterranean): Influence of environmental variables and effects on the biological cycle of Aristeus antennatus

Dynamics of suprabenthos and zooplankton were analyzed in two areas located in the NW (off Sóller harbour) and S (off Cabrera Archipelago) of Mallorca (Balearic Islands, western Mediterranean) at depths ranging between 135–780 m. Four stations situated respectively at 150 m (shelf-slope break), and at bathyal depths of 350, 650 and 750 m were sampled at bi-monthly intervals during six cruises performed between August 2003 and June 2004. Suprabenthos showed maximum biomass in both areas from late spring to summer (April to August), while minimum biomass was found in autumn (September–November). Though variable, temporal dynamics of zooplankton showed peaks of biomass in late winter and summer (February and June), while minimals occurred in autumn (August–September) and, at bathyal depths, in April. Suprabenthos (abundance; MDS analyses) showed a sample aggregation as a function of depth (3 groups corresponding to the shelf-slope break, upper slope — over 350 m; and the middle, deeper part of the slope — over 650–750 m), without any separation of hauls by season. By contrast, zooplankton samples were separated by season and not by depth. There was evidence of three seasonal groups corresponding to summer (June 2004–August 2003), autumn–winter (September and November 2003, February 2004), and spring (April 2004), being especially well established off Sóller. In general, suprabenthos was significantly correlated with the sediment variables (e.g. total organic matter content (% OM), potential REDOX), whereas zooplankton was almost exclusively dependent on Chl a at the surface, which suggests two different food sources for suprabenthos and zooplankton. The increase of suprabenthos abundance in April–June was paralleled by a sharp increase (ca. 2.8 times) in the %OM on sediment during the same period, coupled ca. 1–2 months of delay with the peak of surface Chl a recorded in February–March (from satellite imagery data). Suprabenthos biomass was also correlated with salinity close to the bottom, suggesting a link between suprabenthos abundance and changes in the oceanographic condition of water masses close to the bottom. It is suggested that a higher suprabenthos biomass recorded off Sóller in comparison to that off Cabrera in June could, in turn, be related to a seasonal inflow of Levantine Intermediate Water (LIW) in April–June in this area at mid bathyal depths (350–650 m). This trend would be based on: 1) it was evident only at mid-slope depths between 350–750 m, coinciding with the LIW distribution, and 2) it was not recorded among zooplankton (collected throughout the water column). The possible effect of the fluctuations of suprabenthos and zooplankton on higher trophic levels has been explored studying the diet and food consumption rates of the red shrimp Aristeus antennatus, as indicator species by its dominance in bathyal communities. A. antennatus increased its food consumption from February to April–June 2004 off Sóller, which in the case of large (CL > 40 mm) specimens was found in both areas. In addition, there was a shift of diet from winter to spring–early summer. In this last period, A. antennatus preyed upon euphausiids and mesopelagic decapods and fish, while benthos (e.g. polychaetes and bivalves) decreased in the diet. This indicates an increase in the food consumption and probably in the caloric content of the diet in pre-spawning females in April–June 2004, which is synchronized with the period when gonad development begins in A. antennatus females (May–June). Anyway, macrozooplankton, and not suprabenthos, is crucial as a high energetic food source in the coupling between food intake and reproduction in the red shrimp.     

Distribution, reproduction and development of Calanus species in the Northeast water in relation to environmental conditions

The vertical and horizontal distribution of mesozooplankton biomass and its composition, together with the reproduction and development of the three dominant herbivorous copepods Calanus finmarchicus, C. glacialis and C. hyperboreus, were studied in the Northeast Water between the end of May and end of July, 1993. Biomass varied between 0.15 and 10.6 g m⁻² and was lower on the shallow banks. Highest biomass was found on the shelf slope and in the entrances to Belgica and Westwind Troughs. There was no seasonal trend during the study period. Among the zooplankton taxa, copepods were dominant, with 84% biomass of all other taxa, followed by chaetognaths with 14%. The large Calanus species made up 91% of copepod biomass. The most abundant species, C. glacialis, inhabited areas of low current speeds on Belgica and Ob Bank, C. hyperboreus dominated shelf slope and trough stations, while C. finmarchicus was most abundant in the Return Atlantic Current along the shelf slope and also eastern Belgica Trough. C. glacialis was the only Calanus species spawning during this period, but young copepodites of the other species were also abundant. Egg production of C. glacialis was at a maximum by our arrival and continued to at least mid August. According to the results from starvation experiments, its egg production was fuelled by food uptake, but was decoupled from phytoplankton chlorophyll until July, indicating ice-algae and microzooplankton as an alternative food source. Only when the polynya approached its maximum extent was a close relationship to phytoplankton established. Due to both spatial and temporal heterogeneity of the occurrence of young copepodite stages it was impossible to follow the growth of cohorts of developmental stages. Herbivorous carbon requirements estimated from egg production rates were mostly less than one third of the phytoplankton stock. From egg production and the distribution of young stages, the outer Westwing Trough seems to be the centre of biological activities. This may be related to the formation of young ice in winter in this area, which was found to carry a large mass of under-ice algae. High biomass but low production in Belgica Trough indicate this as an advective and expatriated community of C. finmarchicus and C. hyperboreus, where grazing is negligible, at least during early summer.     

On the relation of structure, perception and activity in marine planktonic copepods

The goal of this paper is to illustrate how in juvenile and adult subtropical marine planktonic copepods various structures or morphological features function in concert to detect prey and predators. Without motion by either food (e.g. flagellate, ciliate) or feeder (e.g. feeding current) or both (e.g. Acartia spp. and ciliate) few feeding activities will occur. Through motion a food particle is either perceived mechanically or chemically to be followed by appendage activities. A combination of mechano- and chemosensors on their cephalic appendages (and probably on other extremities) serve juvenile and adult copepods to perceive signals. Perception is followed by alternation of motion and sensing by these appendages, or by no motion at all (e.g. behavior by Eucalanus pileatus when perceiving a weak hydrodynamic signal). Non-moving and extended sensors (setae) are best suited for mechanical/hydrodynamic perceptions in those copepods which lack a feeding current and hardly move. Numerous mechanosensors arranged in three dimensions on the first antennae (A1) are required to perceive the precise location of moving prey at a distance (e.g. Oithona feeding on ciliates but also sinking particles). Those copepods which create a weak or intermittent feeding current can supplement nutrition with carnivory, which requires perception by the A1 (e.g. Centropages velificatus adults). These two groups require, in addition to perception of prey motion/location, rapid motion by their appendages (A1, second maxillae M2, etc.) to capture the prey. Nauplii, which satiate at far lower food levels than adults, have one of several means of food acquisition: encounter through forward motion, perception through feeding current, or perception of a moving food particle. The nearly continuous motion of most calanoid nauplii makes them vulnerable to predation because all three pairs of appendages are usually moving. Opposite are nauplii of cyclopoid and a few calanoid species which move only occasionally. Copepodid stages and adults use non-moving and often extended setae on the tips of their A1 to perceive predators at a distance. This structure and their pronounced escape motion may reduce their vulnerability to predation as compared to nauplii.     

Plankton distribution and the impact of copepod grazing on primary production in Fram Strait, Greenland Sea

Two hydrobiological transects across the East Greenland Shelf and the open waters of Fram Strait in summer were chosen to illustrate the distribution and production of phyto- and zooplankton in relation to water masses and ice cover. The parameters used were temperature and salinity, inorganic nutrients, chlorophyll a, primary production, phytoplankton species composition, abundance of the dominant herbivorous copepods Calanus finmarchicus, C. glacialis, C. hyperboreus, Metridia longa and egg production of C. finmarchicus and C. glacialis. Grazing impact of copepodites and adults of these four species was modelled for each station by using egg production rates as an index of growth. Seasonal development of plankton communities was closely associated with the extent of the ice cover, hydrographic conditions and the water masses typical of the different hydrographic domains. Four regions were identified from their biological activities and physical environment: The Northeast Water polynya on the East Greenland Shelf, with a springbloom of diatoms and active reproduction of herbivorous copepods. The pack ice region, dominated by small flagellates and negligible grazing activities. The marginal ice zone, with high variability and strong gradients of autotroph production related to eddies and ice tongues, an active microbial loop and low egg production. The open water, with high station-to-station variability of most of the parameters, probably related to hydrographic mesoscale activities. Here, Phaeocystis pouchetii was a prominent species in the phytoplankton communities. Its presence may at least partly be responsible for the generally low egg production in the open waters. Grazing impact on primary production was always small, due to low zooplankton biomass in the polynya and due to low ingestion in the remaining regions.     

Meroplankton abundance in the Northeast Water Polynya: Insights from oceanographic parameters and benthic abundance patterns

We investigated meroplankton (planktonic larvae of benthic organisms) abundance and distribution in the Northeast Water (NEW) Polynya, located on the northeast coast of Greenland, from July 15 to August 15, 1992. Meroplankton was present at all sites visited (0.03–84.83 individuals per m³); at one station meroplankton comprised 8.28% of total zooplankton. Total meroplankton abundance was correlated with total zooplankton abundance and total benthic infaunal abundance but was not correlated with either microscopic carbon concentration or primary productivity. Examination of distribution data for barnacle nauplii and adults indicated that both adults and larvae were concentrated at the same locations. Patterns of distribution were also examined for stelleroid plutei, polychaete larvae and trochophores. There were distinct geographic patterns in total and class-specific meroplankton distributions, with maximal abundances occurring over the Belgica Bank and in the eastern regions of the Westwind Trough and minimal abundances in the Belgica Trough. The apparent control of meroplankton distribution by the hydrography of the region, coupled with the correlation between meroplankton, zooplankton and adult infaunal abundance, reinforces the hypothesis that hydrography plays a major role in controlling the distribution of biota in the NEW polynya (Ambrose and Renaud, 1995; Ashjian et al., 1995, 1997-this volume; Smith et al., 1995; Piepenburg et al., 1997-this volume).     

Ichthyoplankton in the neritic and coastal zone of Antarctica and Subantarctic islands: A review

Since the article published by Loeb et al. [Loeb, V.J., Kellermann, A., Koubbi, P., North, A.W., White, M., 1993. Antarctic larval fish assemblages: a review. Bull. Mar. Sci. 53(2), 416–449.] about Antarctic ichthyoplankton, many surveys were carried out in different sectors of the Southern Ocean focusing on different aspects of the ecology of fish larvae. Some of these researches were conducted in the Subantarctic Kerguelen Islands and others on the continental shelf off Terre Adélie and Georges V land. Oceanographic and geographic features influence fish larvae ecology such as island mass effects, gyres, canyons. Antarctic fishes show also temporal segregation of spawning which induces temporal succession of early stage larvae. This avoids competition and probably the predation on early stages for species having few recruits. In that case, we have to understand how these larvae can deal with the match–mismatch with their preys and how they find sufficient food to survive. But our knowledge on Antarctic fish larvae is still insufficient as we do not know larvae for quite a lot of species and because of the difficulty to sample during winter.     

Benthic community and food web structure on the continental shelf of the Bay of Biscay (North Eastern Atlantic) revealed by stable isotopes analysis

The North Bay of Biscay continental shelf is a major French demersal fishery, but little was known on the trophic food web of its benthic communities. In order to determine the benthic trophic web, the objectives of this study are to describe the macro- and megafaunal benthic community structure (species richness, abundance and biomass) and to establish the trophic pathways (food sources and trophic levels) by applying carbon and nitrogen stable isotopic analysis to the main benthic and demersal species (invertebrates and fish). Two distinct benthic communities have been identified: a muddy sand community within the central part of the bay, and an outer Bay of Biscay Ditrupa sand community of higher species richness, abundance and biomass than the muddy sand community. Deposit-feeders, suspension feeders and predators, distributed in three main trophic levels, dominate both communities. Large differences in stable carbon ratio values within the primary consumers provide evidence of two different food sources: i) a pelagic food source made up of recent sedimenting particulate organic matter on which zooplankton and suprabenthos feed and ii) a benthic detrital food source supplying deposit feeders and partly benthic suspension feeders. Differences in isotopic signatures were also observed within the upper trophic levels that allowed estimation of the contribution of each food source component to the diet of the upper consumers. Finally, the use of stable isotopic composition together with the species' feeding strategy allow identification of the main differences between the trophic functioning of the two benthic communities and highlight the importance of the role of detrital pathways in the carbon cycling within the continental shelf benthic trophic web.     

Host specificity in copepod parasites of deep-sea fishes

Parasitic copepods belonging to two orders, Siphonostomatoida and Poecilostomatoida, are frequently reported from fish hosts in the deep sea. Three families of copepods are most commonly encountered, Sphyriidae, Lernaeopodidae and Chondracanthidae, but members of another four families, Hatschekiidae, Pennellidae, Philichthyidae and Hyponeoidae, are occasionally recorded. These parasites utilise various deep demersal fishes as hosts, especially species of the most abundant families, Macrouridae, Moridae, Synaphobranchidae and Alepocephalidae. Host specificity levels are variable, as for shallow-water fishes. In contrast, few parasites are regularly reported from fishes inhabiting the pelagic water column away from the bottom and away from the near-surface zone. Only the pennellids Sarcotretes scopeli and Cardiodectes medusaeus appear common on pelagic fishes, in the Atlantic and Pacific respectively. Host specificity levels in these two pennellid species are relatively low. It is speculated that the difficulty of encountering a host in the vast pelagic biome has restricted the diversity of parasitic copepods that have successfully colonized pelagic fishes.     

Estuary Management and Recreational Fishing Benefits

Recognition of the benefits to society supported by estuary ecosystem functions and services, and threats to these benefits posed by human activities, has led to various public programs to restore and protect estuaries at the federal, state, and local levels. As available budgets shrink, program administrators and public elected officials struggle to allocate limited restoration and protection funds to the highest priority areas. Economic benefit and cost information can provide useful inputs into this decision-making process by quantifying estuary restoration and protection benefits and costs in commensurate terms. In this paper, a combined actual and intended travel behavior model is described that can be applied to estimate the recreational fishing benefits of estuary restoration and protection. The model was estimated for recreational fishing in the Lower Atchafalaya River Basin estuary along the Gulf of Mexico, Louisiana, USA coast. Changes in freshwater flows into this estuary may affect redfish and speckled trout game fish populations. The model indicates that changes in catch rates of these two species would have a relatively minor affect on annual fishing trips per angler. However, because total effects may be large when effects per angler are aggregated across total anglers, resource management agencies should consider these changes in recreation benefits when evaluating projects that influence the ecology of coastal estuaries, fish populations, and catch rates. Moreover, in other coastal areas or situations, the responsiveness of angling trips to changes in catch rates may vary because of differences in user populations, environmental conditions, fish populations, and fishing experiences.     

Analysis of the parasitic copepod species richness among Mediterranean fish

The Mediterranean ichthyofauna is composed of 652 species belonging to 405 genera and 117 families. Among these, 182 were studied for their parasitic copepods. The analysis of all the works conducted on these crustacea yielded 226 species distributed in 88 genera and 20 families. For each fish species we have established a file providing the species name of the fish, its family, its geographical distribution within the Mediterranean and some of its bio-ecological characteristics. Within each file, all the parasitic copepod species reported on each host species were listed. This allowed to know the species richness (SR) of these hosts. We thus produced 182 files within which 226 copepod species are distributed. A program was created under the Hypercard software, in order to analyse our data. Two parameters were studied. The first one is the mean species richness (MSR), which corresponds to the mean of the different SR found on the different host species. The second is the parasite–host ratio (P/H), which is the ratio of the number of copepod species by the number of host species. These parameters are calculated by our program for all the 182 species of Mediterranean fishes retained in our investigation, on the first hand, and, on the second hand, for one particular group of fish species. We used the following variables to investigate their correlations with copepod species richness: taxonomy—fish families, genera and species; biometry—maximal size of the adult fish; eco-ethology—mode of life (benthic, pelagic or nectonic), displacements (sedentary, migratory with environmental change, or migratory without environmental change), behaviour (solitary or gregarious). Other variables (colour, food, reproduction, abundance, distribution area) were also analysed but did not reveal any clear correlation. Providing that our study does not rely on quantitative (prevalence, intensity) but qualitative basis our aim was only to reveal some tendencies. These tendencies are as follows: (1) In many cases, parasite and host phylogeny seem to play an important role. There are fish families with copepods and families with few species of these parasites. The phyletic constraints could be due to the morphological characteristics of the habitat (e.g. structure of the gills) or biological/ecological characteristics that we were unable to identify. (2) It appears that the presence in a same environment of related fish species (e.g. several species of the same genus, or numerous genera of the same family) is correlated with high parasite richness. A likely explanation is that such situations favours alternated processes of lateral transfers and speciation. (3) Some eco-ethological criteria seem to favour the establishment of a large parasite species richness. It should be noted for instance that Mediterranean fishes the most often infected with copepods are generally nectonic or pelagic, migratory, and gregarious species.     

Adaptability and vulnerability of zooplankton species in the adjacent regions of the Black and Marmara Seas

During 2005–2008 species composition, abundance patterns, natural mortality rates and salinity effects on zooplankton were studied in the Bosphorus regions of the Black and Marmara Seas. The tendencies to diminish for abundance and biomass of the Black Sea originated zooplankton species and to increase for the proportion of their carcasses in the direction from the Black Sea toward the Marmara Sea were found. The mortality in the Black Sea species increased with depth in the Marmara Sea. The contribution of organic matter of carcasses of the Black Sea originated organisms to bacterial processes in the deep strata of the Marmara Sea was estimated. Different salinity and temperature regimes restrict mutual penetration of the species in these seas.     

A numerical investigation of phytoplankton and Pseudocalanus elongatus dynamics in the spring bloom time in the Gda sk Gulf

A nutrient–phytoplankton–zooplankton–detritus (1D-NPZD) ‘phytoplankton {Phyt} and Pseudocalanus elongatus {Zoop} dynamics in the spring bloom time in the Gda sk Gulf. The 1D-NPZD model consists of three coupled, partial second-order differential equations of the diffusion type for phytoplankton {Phyt}, zooplankton {Zoop}, nutrients {Nutr} and one ordinary first-order differential equation for benthic detritus pool {Detr}, together with initial and boundary conditions. In this model, the {Zoop} is presented by only one species of copepod (P. elongatus) and {Zoop} is composed of six cohorts of copepods with weights (W_i) and numbers (Z_i); where . The calculations were made for 90 days (March, April, May) for two stations at Gda sk Gulf with a vertical space step of 0.5m and a time step of 900 s. The flow field and water temperature used as the inputs in the biological model 1D-NPZD were reproduced by the prognostic numerical simulation technique using hydrographic climatological data. The results of the numerical investigations described here were compared with the mean observed values of surface chlorophyll-a and depth integrated P. elongatus biomass for 10 years, 1980–1990. The slight differences between the calculated and mean observed values of surface chlorophyll-a and zooplankton biomass are ca. 10–60 mg C m⁻³ and ca. 5–23 mg C m⁻², respectively, depending on the location of the hydrographic station. The 1D-NPZD model with a high-resolution zooplankton module for P. elongatus can be used to describe the temporal patterns for phytoplankton biomass and P. elongatus in the centre of the Gda sk Gulf.     

The influence of food quality on the nutritional acclimation of the copepod Acartia clausi

The influence of food quality on the nutritional metabolism of Acartia clausi was studied experimentally using four different diets: (1) diatom cells of Thalassiosira weissflogii, (2) detritus prepared from the same culture, (3) 50 : 50 mix on a protein basis of the two previous diets, and 4) dinoflagellate cells of Prorocentrum micans. For each trophic, ingestion, gut transit time, trypsin activity and K_m (half saturation constant) were measured at limiting and saturating concentration. Assimilation rates were also estimated for both pure diatoms and mixed live–detrital cell diets. Ingestion followed a Holling type 2 response for diets 1 and 4, a linear one for detritus and an intermediate response for diet 3. Gut transit time displayed different adaptive changes with food regime depending on protein concentration. Trypsin activity was lower for detrital food and trypsin K_m significantly decreased with increasing concentration of live diatoms. Assimilation rates were higher for live food than for mixed live–detrital food. Results illustrated that Acartia-type copepods optimize nitrogen or protein uptake. They suggest that besides chemoreception-mediated selectivity, internal controls by digestion and assimilation also regulate ingestion (feed-back).