Role of infochemical mediated zooplankton grazing in a phytoplankton competition model

Infochemicals released by marine phytoplankton play important roles in food web interactions by influencing the feeding behavior and selectivity of zooplanktonic predators. Recent modeling efforts have focused on the role of such chemicals as toxic grazing deterrents in phytoplankton competition. However, infochemicals may also be utilized as grazing cues, leading predators to profitable foraging patches. Here we investigate the role of infochemical mediated zooplankton grazing in a standard 3-species phytoplankton competition model, with the aim of further elucidating the ecological role of phytoplankton derived infochemicals. We then extend this to consider a more realistic 4-species model. The models produce a range of solutions depending on the strength of competition and microzooplankton grazing selectivity. Our key result is that infochemical chemoattractants, which increase the susceptibility of the producer to grazing, can provide a refuge for both competing phytoplankton species by attracting carnivorous copepods to consume microzooplankton grazers in a multi-trophic interaction. Our results indicate that infochemicals potentially have important consequences for the dynamics of marine food webs.     

Effects of food diversity on diatom selection by harpacticoid copepods

The diversity of species assemblages which occupy a basal position in the trophic pyramid (typically unicellular algae in aquatic environments) is known to influence the interaction with organisms of higher trophic levels. A laboratory feeding experiment was conducted with cultures of three benthic diatom species (Navicula phyllepta, Grammatophora marina and Cylindrotheca closterium) as primary producers and three harpacticoid copepod species (Harpacticus obscurus, Paramphiascella fulvofasciata and Tigriopus brevicornis) as grazers to evaluate the effects of food diversity (and concomitant food availability) on grazing selectivity. This kind of selectivity experiments is singular for benthic harpacticoid copepods as so far, information on food selection of harpacticoids is scarce.Uptake of a unispecific food source by a single copepod species decreased as food diversity (and concomitant overall food concentration) increased. All three consumers reacted similarly to changing food diversity, but exhibited strong species-specific responses to food identity i.e. which diatom was added was crucial. Irrespective of level of food diversity, H. obscurus took up high amounts of G. marina, whereas both P. fulvofasciata and T. brevicornis preferred C. closterium when given the choice between different diatoms. As for zooplanktonic taxa, this experiment showed that in lower benthic marine food webs both prey organisms (primary producers) and grazers play a very specific role. Diversity of food and its identity are of critical importance at the base of the trophic pyramid, influencing trophic transfer from primary producers over grazers to higher trophic levels.     

Effect of food preservation on the grazing behavior and on the gut flora of the harpacticoid copepod Paramphiascella fulvofasciata

Harpacticoids owe their ‘reputation’ as primary consumers in aquatic food webs to their substantial grazing on diatoms, thus assuring an efficient energy flow to higher trophic levels. Due to the complex feeding behavior of harpacticoids, the nature and dynamics of diatom-harpacticoid trophic interactions remain poorly understood. In addition, there is a growing interest from aquaculture industry in mass-culturing harpacticoids with algal foods but the labor costs of maintaining algal stock cultures are high. This study focuses on the palatability of preserved diatoms for copepods and considers the possible role of bacterially mediated effects on diatom food uptake.The grazing of Paramphiascella fulvofasciata on a preserved freeze-dried diatom diet was tested and compared to the grazing on fresh cells. P. fulvofasciata assimilated the preserved diet, but assimilation of fresh cells was higher. When both cell types were mixed, no selective feeding was observed. Community fingerprinting of the bacteria associated with diatoms and fecal pellets suggests that the copepod gut flora was modified depending on the food source. Furthermore, the results suggest that the egestion of gut bacteria enriches the microenvironment and this can have an additional influence on the feeding behavior of the copepod.Experimental research using preserved foods must take into account that copepod grazing assimilations of fresh foods are likely to be significantly higher. Yet, the stated high assimilation of the mixed diet, encourages further exploration of the application of preserved ‘balanced’ foods for harpacticoid mass-culturing.     

PRELIMINARY RESULTS ON SECONDARY METABOLITES FROM ANTARCTIC BROWN ALGAE AND THEIR ECOLOGICAL RELEVANCE

Ianora, A. Stazione Zoologica A. Dohrn, Villa Comunale 80121 Naples, Italy Traditionally, diatoms have been regarded as providing the bulk of the food that sustains the marine food chain to top consumers and important fisheries. However, this view has recently been challenged on the basis of laboratory studies showing that these small, unicellular algae possess anti-mitotic properties similar to the cytotoxic compounds isolated from numerous marine and terrestrial higher plants. In fact, when copepods, the principal predators of diatoms, are fed diatom diets, they produce abnormal eggs that either fail to develop to hatching or hatch into malformed nauplii that die soon afterwards. The aldehydes responsible for this anti-cell growth activity have recently been isolated and these compounds have been shown to arrest not only the development of copepod and sea urchin embryos, but also the proliferation of human carcinoma cells. In terrestrial environments, there are many reports of secondary metabolites produced by plants that interfere with the reproductive capacity of grazing animals, and which act as a form of population control. But this type of biological model is new for the marine environment where most of the attention on plant-animal interactions has focused on feeding deterrents and poisoning compounds. Such “birth-control” compounds may discourage herbivory by sabotaging future generations of grazers, thereby allowing diatom blooms to persist when grazing pressure would normally have caused them to crash.     

Inhibition of copepod feeding by exudates and transparent exopolymer particles (TEP) derived from a Phaeocystis globosa dominated phytoplankton community

We investigated if (1) dissolved compounds excreted by Phaeocystis globosa and (2) transparent exopolymer particles (TEP) formed from carbohydrates excreted into the water affect the feeding of nauplii and females of the calanoid copepod Temora longicornis during a P. globosa bloom. Copepod grazing on the diatom Thalassiosira weissflogii in the presence of these possible grazing deterrents was measured during three successive weeks of a mesocosm study, simulating the development of a P. globosa bloom. Our results demonstrate no indication for the presence of feeding deterrents in the dissolved phase, but a strong inhibitory effect of transparent exopolymer particles (TEP) on the consumption of algae by both nauplii and adult copepods. The inhibitory effect of TEP was connected to the accumulation of DOM during the progress of the bloom. We suggest that a reduction in the grazing pressure of zooplankton may increase the survival of the liberated single cells during disruption of colonies and allow seeding populations to persist. Furthermore, P. globosa reduces the trophic efficiency of the food web not only by withdrawal of its colonies from grazing but also by a relaxation of the grazing pressure on co-occurring phytoplankton and by alteration of the food web structure via TEP production.     

Food selectivity and grazing impact on toxic Dinophysis spp. by copepods feeding on natural plankton assemblages

Food selectivity and grazing impact by Acartia bifilosa, Temora longicornis and Centropages typicus on Dinophysis spp. plankton assemblages were experimentally investigated in the Baltic Sea. Toxin analyses were carried out on phyto- and zooplankton-dominated size fractions from field-collected samples to assess if toxins produced by Dinophysis spp. would end up in the zooplankton. All copepod species fed actively on toxic Dinophysis spp. (Dinophysis acuta and Dinophysis norvegica). Despite the non-selective feeding behaviour by T. longicornis and C. typicus, selectivity coefficients on D. acuta progressively decreased as food availability increased. Similar response was not observed for A. bifilosa, which displayed an even less selective behaviour. A. bifilosa had no significant negative effect on the net growth of D. norvegica at the lowest food concentration offered, whereas T. longicornis and C. typicus had significant negative effects on the net growth of D. acuta at low concentrations, similar to those observed in situ. Both species could potentially contribute as a substantial loss factor for Dinophysis spp. provided they are abundant at the onset of the blooms. The estimated grazing impact by the copepod populations was only considerable when C. typicus abundance was high and D. acuta population in sharp decline. Our results suggest that when high abundance of grazers and poor growth condition of prey populations prevail, grazing impact by copepods can contribute considerably to prevent Dinophysis spp. populations to grow or to cause the populations to decline. Okadaic acid was detected in the zooplankton size fraction at one occasion, but the concentration was far lower than the one expected from the ingested toxins. Thus, even if copepods may act as vectors of DSP-toxins to higher trophic levels, the amount of these toxins transported in the food web by copepods seems limited.     

Algal toxins alter copepod feeding behavior

Using digital holographic cinematography, we quantify and compare the feeding behavior of free-swimming copepods, Acartia tonsa, on nutritional prey (Storeatula major) to that occurring during exposure to toxic and non-toxic strains of Karenia brevis and Karlodinium veneficum. These two harmful algal species produce polyketide toxins with different modes of action and potency. We distinguish between two different beating modes of the copepod's feeding appendages-a "sampling beating" that has short durations (<100 ms) and involves little fluid entrainment and a longer duration "grazing beating" that persists up to 1200 ms and generates feeding currents. The durations of both beating modes have log-normal distributions. Without prey, A. tonsa only samples the environment at low frequency. Upon introduction of non-toxic food, it increases its sampling time moderately and the grazing period substantially. On mono algal diets for either of the toxic dinoflagellates, sampling time fraction is high but the grazing is very limited. A. tonsa demonstrates aversion to both toxic algal species. In mixtures of S. major and the neurotoxin producing K. brevis, sampling and grazing diminish rapidly, presumably due to neurological effects of consuming brevetoxins while trying to feed on S. major. In contrast, on mixtures of cytotoxin producing K. veneficum, both behavioral modes persist, indicating that intake of karlotoxins does not immediately inhibit the copepod's grazing behavior. These findings add critical insight into how these algal toxins may influence the copepod's feeding behavior, and suggest how some harmful algal species may alter top-down control exerted by grazers like copepods.     

Feeding response of a benthic copepod to ciliate prey type,prey concentration and habitat complexity

1. Protozoans are important consumers within microbial food webs and, in turn, they represent potential prey for small metazoans. However, feeding interactions within these food webs are rarely characterised and this is especially true for freshwater sediments. 2. We aimed to quantify the feeding links between a freshwater meiofaunal copepod and ciliates in two laboratory experiments. The first experiment addressed the response of Eucyclops serrulatus towards ciliate density and type (two ciliate species of the same genus differing in terms of body size). A second experiment assessed the effect of habitat structure on feeding rates by introducing different structural complexity into the feeding arena. In contrast to the first experiment, which was run only for one time period, this experiment also tested three different total feeding times (4, 7 and 9 h). 3. Eucyclops serrulatus exhibited high ingestion rates, with 3–69 ciliates copepod^?1 h^?1 consumed depending on food concentration, food type and habitat complexity. Copepods exhibited a preference for the smaller ciliate when total ciliate concentration was low, but selected both ciliates equally when food concentrations were medium or high. However, at very high food concentration, Eucyclops preferred the larger ciliate (which was 1/3 of its own body size), suggesting that the longer handling times of the larger prey are rewarding when the large prey is present in high numbers. In terms of total numbers consumed, copepods fed on more small ciliates, but in terms of carbon units both ciliates were selected equally when total prey concentration was low or medium. However, copepods derived more carbon from the larger prey at high and very high prey concentrations (up to 0.7 μgC out of a maximum of 1.1 μgC copepod^?1 h^?1). Habitat complexity influenced the feeding of copepods when it was observed over time. 4. The copepod–ciliate link is well known from the pelagic zone of both marine and freshwater habitats. We have shown its potential importance within the benthos, where it can be influenced by food identity, food quantity and possibly by habitat complexity.     

Copepod grazing during an iron-induced diatom bloom in the Antarctic Circumpolar Current (EisenEx): I. Feeding patterns and grazing impact on prey populations

Feeding activity, selective grazing and the potential grazing impact of two dominant grazers of the Polar Frontal Zone, Calanus simillimus and Rhincalanus gigas, and of copepods < 2 mm were investigated with incubation experiments in the course of an iron fertilized diatom bloom in November 2000. All grazers were already actively feeding in the low chlorophyll waters prior to the onset of the bloom. C. simillimus maintained constant clearance rates and fed predominantly on diatoms. R. gigas and the small copepods strongly increased clearance and ingestion of diatoms in response to their enhanced availability. All grazers preyed on microzooplankton, most steadily on ciliates, confirming the view that pure herbivory appears to be the exception rather than the rule in copepod feeding. The grazers exhibited differences in feeding behavior based on selectivity indices. C. simillimus and R. gigas showed prey switching from dinoflagellates to diatoms in response to the phytoplankton bloom. All grazers most efficiently grazed on large diatoms leading to differences in daily losses for large and small species, e.g. Corethron sp. or Thalassionema nitzschioides. Species-specific diatom mortality rates due to grazing suggest that the high feeding activity of C. simillimus prior to and during the bloom played a role in shaping diatom population dynamics.     

Chemical Response of the Toxic Dinoflagellate Karenia mikimotoi Against Grazing by Three Species of Zooplankton

We investigated the toxicity of Karenia mikimotoi toward three model grazers, the cladoceran Moina mongolica, the copepod Pseudodiaptomus annandalei, and the crustacean Artemia salina, and explored its chemical response upon zooplankton grazing. An induction experiment, where K. mikimotoi was exposed to grazers or waterborne cues from the mixed cultures revealed that K. mikimotoi might be toxic or nutritionally inadequate toward the three grazers. In general, direct exposure to the three grazers induced the production of hemolytic toxins and the synthesis of eicosapentaenoic acid (EPA). Both EPA and the hemolytic toxins from K. mikimotoi decreased the survival rate of the three grazers. In addition, the survival rates of M. mongolica, P. annandalei, and A. salina in the presence of induced K. mikimotoi that had previously been exposed to a certain grazer were lower than their counterparts caused by fresh K. mikimotoi, suggesting that exposure to some grazers might increase the toxicity of K. mikimotoi. The chemical response and associated increased resistance to further grazing suggested that K. mikimotoi could produce deterrents to protect against grazing by zooplankton and that the substances responsible might be hemolytic toxins and EPA.     

Copepod feeding in the ocean: scaling patterns,composition of their diet and the bias of estimates due to microzooplankton grazing during incubations

Here, we report insights from the compilation and analysis of data on marine calanoid copepod feeding rates in the ocean. Our study shows that food availability and body weight are major factors shaping copepod feeding rates in the field, with a relatively minor role of temperature. Although the maximal feeding rates of copepods that are observed in the field agree with the well-known 3/4 of body size scaling rule for animals, copepod feeding in the oceans is typically limited and departs from this rule. Ciliates and dinoflagellates appear to be highly relevant in the composition of copepod diets, and this represents an indirect increase in the flux of primary production that is likely to reach the upper trophic levels; this contribution is higher in the less productive systems and may help to explain accounts of proportionally higher standing stocks of copepods supported per unit of primary producer biomass in oligotrophic environments. Contrary to common belief, diatoms emerge from our dataset as small contributors to the diet of copepods, except in some very productive ecosystems. We have also evaluated the bias in the estimation of copepod grazing rates due to within-bottle trophic cascade effects caused by the removal of microheterotrophs by copepods. This release of microzooplankton grazing pressure accounts for a relevant, but moderate, increase in copepod grazing estimates (ca. 20–30%); this bias has an effect on both the carbon flux budgets through copepods and on our view of their diet composition. However, caution is recommended against the indiscriminate use of corrections because they may turn out to be overestimates of the bias. We advise that both uncorrected and corrected grazing rates should be provided in future studies, as they probably correspond to the lower and upper boundaries of the true grazing rates.     

Fitness consequences for copepods feeding on a red tide dinoflagellate: deciphering the effects of nutritional value, toxicity, and feeding behavior

Phytoplankton exhibit a diversity of morphologies, nutritional values, and potential chemical defenses that could affect the feeding and fitness of zooplankton consumers. However, how phytoplankton traits shape plant–herbivore interactions in the marine plankton is not as well understood as for terrestrial or marine macrophytes and their grazers. The occurrence of blooms of marine dinoflagellates such as Karenia brevis suggests that, for uncertain reasons, grazers are unable to capitalize on, or control, this phytoplankton growth—making these systems appealing for testing mechanisms of grazing deterrence. Using the sympatric copepod Acartia tonsa, we conducted a mixed diet feeding experiment to test whether K. brevis is beneficial, toxic, nutritionally inadequate, or behaviorally rejected as food relative to the palatable and nutritionally adequate phytoplankter Rhodomonas lens. On diets rich in K. brevis, copepods experienced decreased survivorship and decreased egg production per female, but the percentage of eggs that hatched was unaffected. Although copepods showed a 6–17% preference for R. lens over K. brevis on some mixed diets, overall high ingestion rates eliminated the possibility that reduced copepod fitness was caused by copepods avoiding K. brevis, leaving nutritional inadequacy and toxicity as remaining hypotheses. Because egg production was dependent on the amount of R. lens consumed regardless of the amount of K. brevis eaten, there was no evidence that fitness costs were caused by K. brevis toxicity. Copepods limited to K. brevis ate 480% as much as those fed only R. lens, suggesting that copepods attempted to compensate for low food quality with increased quantity ingested. Our results indicate that K. brevis is a poor food for A. tonsa, probably due to nutritional inadequacy rather than toxicity, which could affect bloom dynamics in the Gulf of Mexico where these species co-occur.     

Mesozooplankton Grazing on Picocyanobacteria in the Baltic Sea as Inferred from Molecular Diet Analysis

Our current knowledge on the microbial component of zooplankton diet is limited, and it is generally assumed that bacteria-sized prey is not directly consumed by most mesozooplankton grazers in the marine food webs. We questioned this assumption and conducted field and laboratory studies to examine picocyanobacteria contribution to the diets of Baltic Sea zooplankton, including copepods. First, qPCR targeting ITS-1 rDNA sequence of the picocyanobacteria Synechococcus spp. was used to examine picocyanobacterial DNA occurrence in the guts of Baltic zooplankton (copepods, cladocerans and rotifers). All field-collected zooplankton were found to consume picocyanobacteria in substantial quantities. In terms of Synechococcus quantity, the individual gut content was highest in cladocerans, whereas biomass-specific gut content was highest in rotifers and copepod nauplii. Moreover, the gut content in copepods was positively related to the picocyanobacteria abundance and negatively to the total phytoplankton abundance in the water column at the time of sampling. This indicates that increased availability of picocyanobacteria resulted in the increased intake of this prey and that copepods may rely more on picoplankton when food in the preferred size range declines. Second, a feeding experiments with a laboratory reared copepod Acartia tonsa fed a mixture of the picocyanobacterium Synechococcus bacillaris and microalga Rhodomonas salina confirmed that copepods ingested Synechococcus, even when the alternative food was plentiful. Finally, palatability of the picocyanobacteria for A. tonsa was demonstrated using uptake of ¹³C by the copepods as a proxy for carbon uptake in feeding experiment with ¹³C-labeled S. bacillaris. These findings suggest that, if abundant, picoplankton may become an important component of mesozooplankton diet, which needs to be accounted for in food web models and productivity assessments.     

An ecological and economic review of meiofauna as food for fish*

After briefly reviewing the evidence that meiofauna are an integral part of the complex food web of invertebrate infaunal and epifaunal organisms, it is shown that, at least in intertidal and shallow subtidal habitats, meiofauna are an important food for the small (30–60 mm) juveniles of some commercially important fishes (flatfish and salmonids) or for fish which are themselves food for commercial species. The evidence that fish feed preferentially on a few species of harpacticoid copepod rather than on the numerically dominant meiofaunal taxa is presented and discussed. It is argued that, in addition to benthic copepods being energetically efficient as food for fishes, it is differences in the activity and availability between major taxa of meiofauna as they relate to the feeding strategies of the fish which go a long way to explaining this apparent general preference of fish for harpacticoid copepods rather than nematodes. From literature estimates of consumption rates, it is concluded that the supply of meiofaunal food to juvenile fish is unlikely to be limiting although this may not be the case in some shallow subtidal muddy sand habitats. Whilst it is recognized that fish predation may influence the magnitude of the seasonal peaks in numbers, predation is not the principal cause of seasonal fluctuations in harpacticoid copepod species abundances. The effects of predation on other aspects of community structure is unknown. Finally, the possible role of meiofauna in mariculture systems is briefly discussed.     

Copepods induce paralytic shellfish toxin production in marine dinoflagellates

Among the thousands of unicellular phytoplankton species described in the sea, some frequently occurring and bloom-forming marine dinoflagellates are known to produce the potent neurotoxins causing paralytic shellfish poisoning. The natural function of these toxins is not clear, although they have been hypothesized to act as a chemical defence towards grazers. Here, we show that waterborne cues from the copepod Acartia tonsa induce paralytic shellfish toxin (PST) production in the harmful algal bloom-forming dinoflagellate Alexandrium minutum. Induced A. minutum contained up to 2.5 times more toxins than controls and was more resistant to further copepod grazing. Ingestion of non-toxic alternative prey was not affected by the presence of induced A. minutum. The ability of A. minutum to sense and respond to the presence of grazers by increased PST production and increased resistance to grazing may facilitate the formation of harmful algal blooms in the sea.     

Population regulation and role of mesozooplankton in shaping marine pelagic food webs

Copepods constitute the majority of the mesozooplankton in the oceans.By eating and being eaten copepods have implications for the flow of matterand energy in the pelagic environment. I first consider populationregulation mechanisms in copepods by briefly reviewing estimates of growthand mortality rates and evidence of predation and resource limitation. Theeffects of variations in fecundity and mortality rates for the demography ofcopepod populations are then examined by a simple model, which demonstratesthat population growth rates are much more sensitive to variations inmortality than to variations in fecundity. This is consistent with theobserved tremendous variation in copepod fecundity rates, relatively low andconstant mortality rates and with morphological and behavioralcharacteristics of pelagic copepods (e.g., predator perception and escapecapability, vertical migration), which can all be considered adaptations topredator avoidance. The prey populations of copepods, mainly protozoa(ciliates) and phytoplankton, may be influenced by copepod predation tovarying degrees. The highly variable morphology and the population dynamics(e.g., bloom formation) of the most important phytoplankton prey populations(diatoms, dinoflagellates) suggest that predation plays a secondary role incontrolling their dynamics; availability of light and nutrients as well ascoagulation and sedimentation appear generally to be more important. Thelimited morphological variation of planktonic ciliates, the well developedpredator perception and escape capability of some species, and the oftenresource-unlimited in situ growth rates of ciliates, on the other hand,suggest that copepod predation is important for the dynamics of theirpopulations. I finally examine the implications of mesozooplankton activityfor plankton food webs, particularly their role in retarding vertical fluxesand, thus, the loss of material from the euphotic zone. This revised version was published online in July 2006 with corrections to the Cover Date.     

Food web complexity affects stoichiometric and trophic interactions

The stoichiometry of trophic interactions has mainly been studied in simple consumer–prey systems, whereas natural systems often harbour complex food webs with abundant indirect effects. We manipulated the complexity of trophic interactions by using simple laboratory food webs and complex field food webs in enclosures in Lake Erken. In the simple food web, one producer assemblage (periphyton) and its consumers (benthic snails) were amended by perch, which was externally fed by fish food. In the complex food web, two producer assemblages (periphyton and phytoplankton), their consumers (benthic invertebrates and zooplankton) and perch feeding on zooplankton were included. In the simple food web perch affected the stoichiometry of periphyton and increased periphyton biomass and the concentration of dissolved inorganic nitrogen. Grazers reduced periphyton biomass but increased its nutrient content. In the complex food web, in contrast to the simple food web, perch affected periphyton biomass negatively but increased phytoplankton abundance. Perch had no influence on benthic invertebrate density, zooplankton biomass or periphyton stoichiometry. Benthic grazers reduced periphyton biomass and nutrient content. The difference between the simple and the complex food web was presumably due to the increase of pelagic cyanobacteria ( Gloeotrichia sp.) with fish presence in the complex food web, thus fish had indirect negative effects on periphyton biomass through nutrient competition and shading by cyanobacteria. We conclude that the higher food web complexity through the presence of pelagic primary producers (in this case Gloeotrichia sp.) influences the direction and strength of trophic and stoichiometric interactions.     

Copepod Population-Specific Response to a Toxic Diatom Diet

Diatoms are key phytoplankton organisms and one of the main primary producers in aquatic ecosystems. However, many diatom species produce a series of secondary metabolites, collectively termed oxylipins, that disrupt development in the offspring of grazers, such as copepods, that feed on these unicellular algae. We hypothesized that different populations of copepods may deal differently with the same oxylipin-producing diatom diet. Here we provide comparative studies of expression level analyses of selected genes of interest for three Calanus helgolandicus populations (North Sea, Atlantic Ocean and Mediterranean Sea) exposed to the same strain of the oxylipin-producing diatom Skeletonema marinoi using as control algae the flagellate Rhodomonas baltica. Expression levels of detoxification enzymes and stress proteins (e.g. glutathione S-transferase, glutathione synthase, superoxide dismutase, catalase, aldehyde dehydrogenases and heat shock proteins) and proteins involved in apoptosis regulation and cell cycle progression were analyzed in copepods after both 24 and 48 hours of feeding on the diatom or on a control diet. Strong differences occurred among copepod populations, with the Mediterranean population of C. helgolandicus being more susceptible to the toxic diet compared to the others. This study opens new perspectives for understanding copepod population-specific responses to diatom toxins and may help in underpinning the cellular mechanisms underlying copepod toxicity during diatom blooms.     

The importance of protozooplankton as prey for copepods in the coastal areas of the central Irish Sea

This study evaluates food supply for copepods, highlighting the trophic relationship between copepods and protozooplankton. To test the hypotheses that protozooplankton prey are capable of sustaining the copepod standing stock in the western Irish Sea, the taxonomic and size composition of these two groups and the size-specific predation of copepods on protozooplankton were investigated. Protozooplankton and copepod samples were collected off the southwest coast of the Isle of Man using 1.7 l Niskin water bottles and two nets (64 and 280 μm meshes), respectively. Copepod predation on protozooplankton was calculated using weight-specific clearance rates from the literature, considering the availability of prey that was accessible to a given size of copepod. Low protozooplankton biomass was dominated by small cells (<60 μm), and high copepod biomass was dominated by small species, which were more efficiently collected by a 64-μm mesh net. However, large copepods were only collected by a 280-μm mesh net, suggesting that the combination of the two nets provided a better estimate of copepod biomass. Predation by the copepod assemblage in the Irish Sea removed 1–47% and 0.5–22% of ciliates and dinoflagellates standing stock, respectively, resulting in 1–40% of the copepod feeding requirement per day. Contrary to our hypothesis, copepods could not meet their feeding requirements by grazing only on the microzooplankton prey (15–200 μm), and other food sources (i.e. nanoplankton) must be important additional dietary components to copepods in the Irish Sea. Handling editor: S. M. Thomaz