Filtering and feeding rates of cyclopoid copepods feeding on phytoplankton

The algal biomass ingested by omnivorous cyclopoid copepods (Cyclops kolensis and C. vicinus) was measured by two methods in the hypertrophic Heiligensee in Berlin (West Germany). The clearance and ingestion rates inferred from measurements of natural populations of ¹⁴C labelled phytoplankton were compared with those obtained from chlorophyll a determinations using the presence/absence method (observed chlorophyll a content of natural lake phytoplankton with and without addition of cyclopoids). Both methods gave similar results. Nevertheless, the radio tracer method is preferred, mainly because the short feeding duration excludes high variations in both the food composition and food concentration that limit the presence/absence method.     

Complementary impact of copepods and cladocerans on phytoplankton

The differences in the impact of two major groups of herbivorous zooplankton (Cladocera and Copepoda) on summer phytoplankton in a mesotrophic lake were studied. Field experiments were performed in which phytoplankton were exposed to different densities of two major types of herbivorous zooplankton, cladocerans and copepods. Contrary to expectation, neither of the two zooplankton groups significantly reduced phytoplankton biomass. However, there were strong and contrasting impacts on phytoplankton size structure and on individual taxa. Cladocerans suppressed small phytoplankton, while copepods suppressed large phytoplankton. The unaffected size classes compensated for the loss of those affected by enhanced growth. After contamination of the copepod mesocosms with the cladoceran Daphnia , the combined impact of both zooplankton groups caused a decline in total phytoplankton biomass.     

Implication of the feeding limb morphology for herbivorous feeding in some freshwater cyclopoid copepods

1. The mouthparts of five species of adult planktonic cyclopoid copepods (Cyclops vicinus, C. abyssorum, Acanthocyclops robustus, Mesocyclops leuckarti and Thermocyclopscrassus), in particular the distance between setae and setules of the maxilliped (which can indicate the ability to retain small particles), were compared using electron and light microscopy. 2. The mesh‐sizes of the food‐collection grid formed by these setae and setules ranged between 4.6 and 13.2 μm; the area covered by the grid ranged between 6000 and 32 000 μm². 3. Mesh‐size was not simply correlated with body size. Cyclops abyssorum and M. leuckarti have the coarsest meshes and T. crassus the finest, while C. vicinus and A. robustus were intermediate. 4. The results suggest that cyclopoid copepods are able to retain particles in the size range of nanoplankton and that differences in mesh‐sizes between species may explain differences in the ability to subsist and reproduce on a diet of small algae.     

The role of turbulent diffusion for copepods with feeding currents

The encounter rate between predator and prey is frequently modeledin terms of the ‘swept volume’ associated with therelative velocity of the two organisms and an appropriate cross-sectionalarea. For the copepods which use feeding currents, an alternativeconceptual model of the process is that the food particles arediffusing towards the predators. Their feeding currents trapthe prey (even though they are well beyond the range of eithervisual or chemical detection) and entrain them towards theirwaiting arms. The predators thus benefit from the turbulentmotion and diffusion, even though much of it is due to motionswith scales significantly larger than their body's size. Thefeeding currents serve to dramatically increase the flux offood.     

Life history consequences for Daphnia pulex feeding on nutrient-limited phytoplankton

1. The growth and feeding of Daphnia pulex De Geer on different algal species was examined. The green algae Chlamydomonas reinhardtii Dangeard and Scenedesmus acutus Meyen, the diatom Synedra tenuissima Kützing, the cryptophyte Cryptomonas pyrenoidifera Geitler and the cyanobacterium Microcystis aeruginosa Kützing were cultured in non-limiting and in N- or P-limiting medium and used as food for D. pulex.
2. Growth limitations were reflected in the elemental and biochemical composition and the morphological characteristics of the algal resources.
3. The clearance rates of D. pulex feeding on nutrient-limited algae were reduced. This was not observed when nutrient-limited mutant Chlamydomonas cells without cell walls were used as food, indicating that the cell wall may play an important part.
4. In comparison with animals grown on nutrient-sufficient cells, nutrient-limited algae resulted in smaller body length, reduced brood sizes, reduced size at maturity, increased age at first reproduction and, consequently, in reduced Daphnia population growth rates.
5. Daphnia population growth rates ( r ) were negatively correlated with the C : P ratio and the carbohydrate content of the food. Moreover, significant correlations between r and clearance rates were found.
6. The observed differences in the grazing and the life history parameters of Daphnia feeding on non-limited and nutrient-limited algae may be the result of both reduced nutritional value and reduced digestibility of nutrient-limited algae.     

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.     

The feeding strategies of two large marine copepods

We compared the feeding behaviour of the two copepods Paraeuchaetanorvegica and Chiridius armatus, allowing them to prey on othercalanoids in small-scale laboratory experiments. Several differenceswere found. When fed either live, free-swimming or dead, non-movingprey, P.norvegica seemed unable to locate the dead prey itemswhile C.armatus foraged heavily on them. When starved, P.norvegicaincreased its feeding rate while C.armatus reduced its feedingafter an initial increase. None of the predators changed theirfeeding rates when exposed to light. Nighttime versus daytimefeeding was tested only with P.norvegcia, which seemed to possessan endogenous feeding rhythm with increased rates at night.Chiridius armatus infected with epizooic ciliates appeared tohave increased feeding rates.     

Variability due to feeding activity of individual copepods

This paper addresses (the inter- and intraindividual) variabilityin release rates of fecal pellets of early copepodids (Cll)and adult females of the calanoid copepod Eucalanus pileatus.The intervals between releases of fecal pellets of tetheredindividuals were used as a measure of feeding activity. Earlycopepodids represented animals with a known, and adult femalesthose with an unknown, and probably highly variable history.Food concentration had a significant effect on the frequencyof pellet release by adult females, but not by CII. There wereno significant differences in release intervals among femalesat the lowest food level, but significant differences at middleand highest food concentrations. These differences could bepartly attributed to different history (seasonally). The coefficientof variation (CV) as a measure of variability, rarely exceeded30% for females and was always <30% for CII. For most individuals,the CV was <20%. The range in the frequency of pellet releasewas almost always less than factor of two. Our data indicatethat in pellet release frequency, variability in feeding behaviorwithin and between individual E.pilealus is rather low comparedto data on feeding rates, and is attributed largely to the appliedprocedures.     

Pigment gut contents of copepods and deep phytoplankton maximum in the Western Mediterranean

The feeding activity of Centropages typicus in the Western Mediterraneanwas evaluated during the summer stratification period, whena deep phytoplankton and chlorophyll maximum (DCM) develops.The lack of significant day-night differences in gut contentssuggests continuous phytoplankton ingestion, despite nictemeralmigration of this species. Copepods captured in the DCM hadgreater phytoplankton gut contents, both in nocturnal and diurnalsamples. The observed pattern of phytoplankton ingestion andvertical migration implies copepod-mediated upward transportof nutrients, which might contribute to the maintenance of primaryproduction in the photic layers.     

A model for seasonal phytoplankton blooms

We analyse a generic bottom-up nutrient phytoplankton model to help understand the dynamics of seasonally recurring algae blooms. The deterministic model displays a wide spectrum of dynamical behaviours, from simple cyclical blooms which trigger annually, to irregular chaotic blooms in which both the time between outbreaks and their magnitudes are erratic. Unusually, despite the persistent seasonal forcing, it is extremely difficult to generate blooms that are both annually recurring and also chaotic or irregular (i.e. in amplitude) even though this characterizes many real time-series. Instead the model has a tendency to 'skip' with outbreaks often being suppressed from 1 year to the next. This behaviour is studied in detail and we develop analytical expressions to describe the model's flow in phase space, yielding insights into the mechanism of the bloom recurrence. We also discuss how modifications to the equations through the inclusion of appropriate functional forms can generate more realistic dynamics.     

Selective feeding of four zooplankton species on natural lake phytoplankton

Grazing experiments evaluated by microscopical counting were conducted with different size classes of Daphnia hyalina, D. galeata, Eudiaptomus gracilis, and Cyclops sp., all from Lake Constance, using natural lake phytoplankton as food. Species-specific grazing selectivity coefficients were calculated for the dominant phytoplankton species from weekly experiments. Specific selectivities were found to be largely invariant through the growing season. All zooplankters grazed more efficiently on phytoflagellates such as Rhodomonas and Cryptomonas than on coccales such as Chlorella and Scenedesmus, regardless of their relative abundance in the phytoplankton assemblage. Filtering rates did not decrease in the presence of filamentous algae. Certain filamentous species were grazed efficiently, but only by D. hyalina: Anabaena planktonica, Oscillatoria amphigranulata, and Stephanodiscus binderanus. Large diatom colonies like Asterionella formosa and Fragilaria crotonensis were grazed well only by Cyclops sp. Some algal species were consistently selected against: Mougeotia thylespora and Dinobryon sp. The species-specific selectivity coefficients can be used as weighting factors to assess the effective food concentration relative to Rhodomonas minuta, a reference species for optimal food.     

Danger of zooplankton feeding: the fluid signal generated by ambush-feeding copepods

Zooplankton feed in any of three ways: they generate a feeding current while hovering, cruise through the water or are ambush feeders. Each mode generates different hydrodynamic disturbances and hence exposes the grazers differently to mechanosensory predators. Ambush feeders sink slowly and therefore perform occasional upward repositioning jumps. We quantified the fluid disturbance generated by repositioning jumps in a millimetre-sized copepod (Re ∼ 40). The kick of the swimming legs generates a viscous vortex ring in the wake; another ring of similar intensity but opposite rotation is formed around the decelerating copepod. A simple analytical model, that of an impulsive point force, properly describes the observed flow field as a function of the momentum of the copepod, including the translation of the vortex and its spatial extension and temporal decay. We show that the time-averaged fluid signal and the consequent predation risk is much less for an ambush-feeding than a cruising or hovering copepod for small individuals, while the reverse is true for individuals larger than about 1 mm. This makes inefficient ambush feeding feasible in small copepods, and is consistent with the observation that ambush-feeding copepods in the ocean are all small, while larger species invariably use hovering or cruising feeding strategies.     

Herbivory in the tilapia Oreochromis niloticus: a comparison of feeding rates on phytoplankton and periphyton

A quantitative comparison of the grazing behaviour of young Oreochromis niloticus feeding on the planktonic cyanobacterium Microcystis aeruginosa and a periphytic community dominated by the cyanobacterium Oscillatoria sp., determined that biomass ingestion rates of fish filter-feeding on planktonic cyanobacteria were significantly lower than those surface-grazing on periphyton. Comparisons of published laboratory data on filter-feeding with field data on algal ingestion rates suggest that filter-feeding may be a relatively unimportant method of ingesting algae.     

Combining a regional climate model with a phytoplankton community model to predict future changes in phytoplankton in lakes

1. Linking a regional climate model (RCM) configured for contemporary atmospheric greenhouse gas concentrations, with a phytoplankton community model (PROTECH) produced realistic simulations of 20 years of recent phytoplankton data from Bassenthwaite Lake, in the North‐West of England. 2. Meteorological drivers were derived from the RCM to represent a future climate scenario involving a 1% per annum compound increase in atmospheric CO₂ concentrations until 2100. Using these drivers, PROTECH was run for another 20 year period representing the last two decades of the 21st century. 3. Comparison of these present and future simulations revealed likely impacts on the current seasonal phytoplankton development. Under future climate conditions, the simulated spring bloom showed an increase in cyanobacteria dominance caused by greater success of Planktothrix. Also, the summer cyanobacteria bloom declined earlier because of nutrient limitation caused by the increased spring growth. Overall productivity in the lake did not change. 4. Analysis showed that these predicted changes were driven by changes in water temperature, which were in turn triggered by the higher air temperatures predicted by the RCM.     

Mass density contrast in relation to the feeding currents in calanoid copepods

Theoretical analyses show that positively buoyant copepods areable to generate feeding currents by adopting upside-down bodypositions and pushing water upward. Thus, the excess buoyancyacting on the copepods will be balanced and cone-shaped feedingcurrents generated to transport water to the capture areas.The intensities of the feeding currents, which can be measuredin the present modeling study by calculating the volumetricflux going through the capture areas, are proportional to themass density contrasts between the copepods and the ambientseawater. The mass density contrasts may vary spatially andtemporally depending on copepod body contents and on the propertiesof the seawater immediately surrounding them. We focus on thecase where the mass density contrast between a wax ester-richcopepod and its ambient seawater can vary strongly with depthbecause wax esters are more compressible and 6–10 timesmore thermally expansible than seawater. These theoretical analysesshow that the intensities of the feeding currents generatedby wax ester-rich copepods vary strongly with depth. Our conclusionsfrom these theoretical analyses need to be tested by directobservations. This paper is one of six on the subject of the role of zooplanktonpredator–prey interactions in structuring plankton communities.     

Fecundity studies of neritic calanoid copepods Acartia clausi Giesbrecht and A. Steueri Smirnov: A simple empirical model of daily egg production

Fecundity studies of neritic calanoid copepods, Acartia clausi Giesbrecht and A. steueri Smirnov, were made under controlled laboratory conditions. Effects of temperature and food abundance upon daily egg production were intensively investigated. Under food-satiated conditions, the daily reproductive rate was a function of temperature, and, at constant temperature, it was a function of chlorophyll a concentration. A simple model equation describing fecundity was derived from the combined effects of temperature and chlorophyll a on daily egg production. The observed daily reproductive rates of natural populations of Acartia were simulated closely by the simple model.     

Temperature-dependent development in marine copepods: a comparative analysis of models

A comparative analysis was carried out on the several equationsmost commonly used to describe the dependence of the developmentof organisms on temperature. Goodness of fit, number of parameters,ease of fitting data and biological significance were compared.     

Abundance,population structure and vertical distribution of dominant calanoid copepods on the eastern Weddell Sea shelf during a spring phytoplankton bloom

The main emphasis of this study was to analyse the short-term development of abundance, population structure and vertical distribution of the dominant calanoid copepods during a phytoplankton bloom in the coastal area of the eastern Weddell Sea in December 2003. Microcalanus pygmaeus was by far the most abundant calanoid species. Metridia gerlachei, Ctenocalanus citer, Calanoides acutus, Calanus propinquus and the ice-associated Stephos longipes were also present in considerable proportions. The observed changes in the population characteristics and parameters of these species are described in detail and discussed in the context of the spring phytoplankton bloom. A conspicuous event occurring during the final stage of the study was the development of a strong storm. While the results suggest that this storm did not have any considerable influence on the populations of all other investigated copepod species, it very likely caused pronounced changes in the S. longipes population present in the water column. Before the storm, S. longipes was found primarily in the upper 100 m of the water column, and its population was dominated by adults (mean proportion = 41%) and the copepodite stage I (mean proportion = 30%). After the storm, the abundance increased considerably, and the copepodite stage I contributed by far the largest proportion (53%) of the total population indicating that the early copepodite stages probably had been released from the sea ice into the under ice water layer due to ice break-up and ice melt processes caused by the storm.     

The periodically forced Droop model for phytoplankton growth in a chemostat

 It is proved that the periodically forced Droop model for phytoplankton growth in a chemostat has precisely two dynamic regimes depending on a threshold condition involving the dilution rate. If the dilution rate is such that the sub-threshold condition holds, the phytoplankton population is washed out of the chemostat. If the super-threshold condition holds, then there is a unique periodic solution, having the same period as the forcing, characterized by the presence of the phytoplankton population, to which all solutions approach asymptotically. Furthermore, this result holds for a general class of models with monotone growth rate and monotone uptake rate, the latter possibly depending on the cell quota. Received 10 October 1995; received in revised form 26 March 1996