Impact of copepod grazing on the red-tide flagellate Chattonella antiqua

Although planktonic copepods are major suspension feeders in the sea, the impact of their grazing pressure upon red-tide flagellates has not been fully investigated. In the present study, the grazing of adult females of several copepod species is examined using three food types: viz. natural suspended particles, natural suspended particles mixed with cultured Chattonella antiqua, and cultured C. antiqua. The functional response on C. antiqua was investigated for five species of copepods (Acartia erythraea, Calanus sinicus, Centropages yamadai, Paracalanus parvus and Pseudodiaptomus marinus). Ingestion rates increased linearly with increasing cell concentrations until a maximum level was reached, beyond which the rates were constant. This cell concentration was higher for larger copepods. The weight-specific maximum ingestion rates were higher in the small species. In general, copepods tended to feed selectively on larger particles when feeding on natural particles. This tendency was strongest in a simulated red-tide environment. Thus, it can be surmised that copepods may selectively graze on C. antiqua during the outbreak of a red tide. Grazing pressure by the natural copepod community in Harima Nada, the Inland Sea of Japan, was calculated by integration of the laboratory determined feeding rates and field measurements of zooplankton biomass. The daily removal rate was 3.4 to 30.8% (mean: 12.3%) of C. antiqua biomass at 20 cells ml^-1 and decreased to 0.6–4.3% (mean: 1.8%) at 500 cells ml^-1. Therefore, the grazing pressure by the copepod community is important at the initial stage of the red tide.     

Short-term effects of sucralose on Calanus finmarchicus and Calanus glacialis in Disko Bay,Greenland

The potential effects of sucralose on the Arctic copepods Calanus finmarchicus and Calanus glacialis were studied in Disko Bay, Greenland. Sucralose is a non-calorie sweetener and chlorine derivate of sucrose containing three chlorine atoms. Scandinavian screening studies of sucralose in 2007, revealed sucralose in all effluent samples. To investigate whether sucralose is harmful to the Arctic aquatic ecosystems, possible short-term effects were investigated on egg production, hatching rate, food intake and mortality of two species of Arctic copepods. The copepods were exposed to six different concentrations (0–50,000 ng · L^?1) of sucralose, which spans the range of concentrations found in the screening studies. Exposure led to no mortality among the copepods. Food intake by C. glacialis increased with increasing concentrations of sucralose. In C. finmarchicus, food intake did not differ with increasing concentrations. No effect of sucralose was observed on egg production of C. finmarchicus. Despite increased food intake with increasing concentrations of sucralose, C. glacialis did not increase its egg production. The results show that both species responded weakly to sucralose, but with C. glacialis being possibly slightly more sensitive to sucralose than C. finmarchicus.     

Diets of calanoid copepods on the West Florida continental shelf: Relationships between food concentration,food composition and feeding activity

Bottle incubations were conducted in March, July/August and October 1992. to measure the daily rations (R) and objectively characterize the diets of the calanoid copepodsEucalanus elongatus, Undinula vulgaris, Centropages velificatus andTemora stylifera from the west Florida continental shelf. Daily rations,R, were clustered around two, order-of-magnitude different means, 1.3 and 11.2% of body C d^–1, representative of quiescent and active feeding modes, respectively. The food concentration at which the transition from quiescent to active mode occurred was influenced by food particle size. In the quiescent mode, diets were dominated by nanoplankton, whereas no food type dominated the diet in the active mode. Selective feeding, defined as a statistically significant difference between the frequency distributions of foods in the diet and environment, occurred in both quiescent and active copepods. However, what appeared to be selective feeding in quiescent copepods could be explained by processes that passively modified the distribution of the diet relative to that of the food supply. Conversely, selective feeding in active copepods apparently resulted from foraging for particles >5 m in diameter in food environments dominated by nanoplankton (<5 m).     

Feeding of marine planktonic copepods on mixed phytoplankton

The feeding of juveniles and adults of the copepods Eucalanus pileatus, Temora stylifera and T. turbinata fed a mixture of the phytoplankton Skeletonema costatum, Leptocylindrus danicus and Rhizosolenia alata f. indica was studied at 20°C. E. pileatus nauplii, copepodids and adult females ingest similar percentages of the 3 algae in terms of carbon. Temora juveniles younger than CII ingest mainly S. costatum; at more advanced developmental stages, the ingestion rate on S. costatum remains constant whereas feeding on L. danicus and R. alata f. indica increases with increasing body weight. Feeding on high concentrations of large particles reduces the grazing pressure on small particles, thus favoring zooplankton which require small-sized food.     

Etude de la respiration chez des copépodes de niveaux bathymétriques variés dans la région sud marocaine et canarienne

The results of 281 respiration analyses of pelagic copepods sampled in the South Moroccan and Canarian area are described. The copepods used in experimentation were collected from various bathymetric levels: hyponeustonic species (Anomalocera patersoni, Pontellopsis villosa, P. regalis, Labidocera wollastoni, Pontella lo biancoi), epipelagic species (Centropages typicus, Temora stylifera, Calanus helgolandicus, Acartia clausi), meso and bathypelagic species (Pleuromma xiphias, P. abdominalis, Euchaeta acuta, Undeuchaeta plumosa and some other rare species). Analysis of the metabolismtemperature curve makes it possible to describe different modes of adaptation in relation to geographical distribution, altitudinal repartition, and dietary characteristics. When all species are studied together, a strong correlation exists between the log of respiration and the log of weight. The regression coefficient varies with temperature; it is strongest at a temperature near that of the natural thermic conditions.     

Copepod grazing during a declining spring phytoplankton bloom in the Northern North Sea

Grazing rates of larger (Calanus finmarchicus) and smaller (Acartia clausii Pseudocalanus elongatus etc.) copepods on naturally occurring phytoplankton populations were measured during a declining spring phytoplankton bloom. During the initial period, dominated by Chaetoceros spp. diatoms, constant ingestion rates were observed in Calanus finmarchicus at suspended particulate concentrations above 300 g carbon l^-1. Average daily intake during this time amounted to 35 to 40% of body carbon and reached a maximum of 50%. The feeding response of the smaller copepods was not so well defined, although a maximum daily intake of 56% body carbon was recorded. In both groups, feeding thresholds were at particulate concentrations around 50 g C l^-1. The feeding response of C. finmarchicus was correlated with both a change in their own population and in the food cell type. Linear regressions describing the concentration-dependent feeding response were: ingestion rate (IR)=1.16 total particulate volume (TPV)-36.15 during the initial part of the period compared with IR=0.41 TPV-12.18 for the latter period. C. finmarchicus filtered out slightly larger (x 1.2 diameter) particles than the small copepods and, in both groups, some filtering adjustment was made to accomodate to modal changes in the phytoplankton population from 20–30 m to 10 m diameter cells. Particle production during feeding was frequently evident in the smallest size ranges of particles and the ratio of particle production to ingestion rate was greater at low feeding rates.     

Zooplankton feeding ecology: contents of fecal pellets of the copepods Temora turbinata and T. stylifera from continental shelf and slope waters near the mouth of the Mississippi River

In-situ feeding habits of the copepods Temora turbinata and T. stylifera were investigated by scanning electron microscope examination of fecal pellets, the contents of which reflected copepod gut contents upon capture. Pellet contents were compared with assemblages of available phytoplankton in the water column at the times of zooplankton sampling. Samples were collected in continental shelf and slope waters of the Gulf of Mexico near the mouth of the Mississippi River. Both species ingested a wide size range and taxonomic array of phytoplankters, and to a lesser extent, other crustaceans. Fecal pellets contained primarily the remains of the phytoplankters that were most abundant in the water at times of collection. There was considerable overlap in the food items ingested by adult females of both copepod species, or two stages of T. turbinata copepodites. Thus, T. turbinata and T. stylifera are omnivores, but primarily opportunistic herbivores.     

In situ feeding rhythms of herbivorous copepods,and the effect of starvation

In situ diel variations in gut pigment contents of neritic (Acartia omorii andPseudocalanus minutus) and oceanic copepods (Calanus plumchrus andC. cristatus) were analyzed.A. omorii andP. minutus were sampled in Onagawa Bay on the east coast of Japan in May and August 1987, andC. plumchrus andC. cristatus were sampled in the Bering Sea in June 1986. Gut pigments were generally high at night, and bimodal feeding rhythms were observed in all species. The first peak of gut pigments occurred between sunset and midnight and was followed by a midnight decrease in gut pigment levels, resulting in eventual evacuation of the gut. The second peak was observed a few hours after sunrise. Incubation experiments indicated that ingestion rates of starved copepods were higher than those of acclimated copepods. This phenomenon was most notable at high food concentrations. Gut pigments of starved copepods rapidly increased after exposure to high concentrations of culturedThalassiosira decipiens. These findings suggest that in situ feeding behavior of herbivorous copepods includes periods of cessation or reduction in feeding during the night, and consequently, feeding activity is periodically enhanced with starvation. Starvation enhanced feeding behavior is most obvious in the large oceanic species,C. plumchrus andC. cristatus and is not distinct in small coastal species such asA. omorii.     

Etude expérimentale de la ponte chez trois espèces de copépodes pélagiques (Centropages typicus,Acartia clausi et Temora stylifera)

Egg-laying of 3 common copepod species from the Gulf of Marseilles (Centropages typicus Kröyer, Acatia clausi Giesbrecht, Temora stylifera Dana) has been studied under various trophic conditions, at different periods of the year. The role of phytoplankton abundance in the induction and importance of egg-laying is indicated. The specific quality of the algal suspension used for feeding affected also fertility. A seasonal variation in the importance of egg-laying is established. Some observations are made on egg deposition and development.     

Under-ice feeding and diel migration by the planktonic copepodsCalanus glacialis andPseudocalanus minutus in relation to the ice algal production cycle in southeastern Hudson Bay,Canada

The marine planktonic copepodsCalanus glacialis Jaschnov andPseudocalanus minutus (Kroyer) typically dominate the copepod biomass in spring under the ice in southeastern Hudson Bay, Canada. Females of both species exhibited significant diel feeding cycles, as measured by gut pigment content, throughout a bloom of ice algae at the ice-water interface in 1986. Periods of grazing correlated well with a nighttime vertical migration by females to within 0.2 m of the ice-water interface, suggesting that feeding took place at or just below the thermohaline boundary between seawater and the interfacial layer containing the ice algae. Seasonal melting of the ice bottom in mid-May resulted in freshening of the surface layer and release of the ice algae into the water column. FemaleC. glacialis andP. minutus responded by ceasing migration to the interface. Gut pigment content, and by reasonable assumption, feeding activity in the water column, increased substantially immediately after this event. In mid-May, the water column phytoplankton consisted of flagellates, sedimenting ice algal cells, and diatoms (Navicula pelagica andChaetoceros sp.) previously found at the interface and then growing in the water column. We conclude that algae growing at the ice-water interface, and sedimenting or actively growing algae derived from this interfacial layer, are a regular and principal source of nutrition for these pelagic copepods during and immediately after the ice algal bloom.     

Food selection capabilities of the estuarine copepod Acartia clausi

Existing viewpoints and theories of selective grazing by copepods are briefly reviewed in order to formulate explicit hypotheses to be tested experimentally. Based on these hypotheses, a series of grazing experiments was run to determine (1) the extent of the selective ingestion capabilities of Acartia clausi and (2) how these capabilities were affected by previous feeding histories. Groups of copepods were separately preconditioned on a small diatom (Thalassiosira pseudonana), a large diatom (T. fluviatilis), or a plastic sphere. The ingestive behavior was then examined on various combinations of spheres and food particles. Spheres offered alone were not ingested. In mixtures of diatoms and spheres, the copepods avoided ingesting spheres intermediate in size between the sizes of the diatoms. The copepods either ingested particles on either side of the spheres, or ignored all particles less than the size of the largest spheres. The pattern observed depended upon the size of the preconditioning food. However, if the spheres were larger than the largest food particles, the copepods still selectively ingested the food particles. The above results demonstrate that A. clausi has a complex grazing behavior consisting of (1) more efficient grazing on larger particles within its particle-size ingestion range; (2) the ability to alter effective setal spacing to optimize feeding behavior (i.e., the ability to increase efficiency of capture of food particles, and to avoid non-food particles); and (3) the ability for post-capture rejection of non-food particles when they interfere with the ingestion of food particles on which the copepod has been preconditioned. The behavioral patterns observed depend heavily on the food preconditioning and the presence or absence of non-food particles. These results clearly indicate that a simple mechanistic explanation of selective grazing is insufficient.     

Egg production of Calanus finmarchicus : effect of temperature, food and season

The use of the egg production rate of herbivorous copepods as an important parameter for understanding population dynamics and as an index of secondary production requires knowledge of the regulatory mechanisms involved and of the response to changes in food concentrations and temperature. Furthermore, the effects of season and generation on egg production have to be studied. In this context data are presented for Calanus finmarchicus from the northern North Atlantic. Prefed and prestarved females were exposed to different concentrations of the diatom Thalassiosira antarctica over 1 to 2 wk at 0 or 5 °C, and egg deposition was controlled daily. Egg production increased with higher food concentrations, but much less when prestarved. The effect of temperatures between −1.5 and 8 °C on egg production was studied in females maintained at optimum feeding conditions. Egg production rate increased exponentially over the whole temperature range by a factor of 5.2, from 14.2 to 73.4 eggs female⁻¹ d⁻¹, and carbon-specific egg production by 4, from 2.1 to 8.5% body C d⁻¹. The response to starvation was also temperature dependent. In both the temperature and feeding experiments egg production rate was regulated mainly by changes of the spawning interval, while changes of clutch size were independent of experimental conditions. Different responses to optimum feeding conditions were observed in females collected in monthly intervals on three occasions between March and May. The March females deposited more clutches than the April and May females. In May, >50% of the females did not spawn at all. Maximum egg production rates were never >25% of the rate expected at 5 °C, indicating endogenous control of egg production in addition to food and temperature effects. Received: 4 August 1996 / Accepted: 11 September 1996     

Influence of cultivation and food concentration on body length of calanoid copepods

The cephalothorax length of the marine pelagic Copepoda Acartia clausi Giesbrecht, Temora longicornis (Müller), Centropages hamatus (Lilljeborg) and Pseudocalanus sp. was monitored at 15°C during prolonged cultivation through up to 55 filial generations and at different concentrations of food. The length of T. longicornis decreased considerably during the first 15 generations and remained rather constant thereafter. In the other species, body length increased slightly or remained almost constant. Genetic changes are probably involved. Food concentration influenced body size of all species, particularly C. hamatus, in which 80% of the natural size range may be explained by differences in food concentration. The idea that temperature is a dominant factor in determining the length of copepods should be reconsidered.     

Zooplankton grazing on the coccolithophore Emiliania huxleyi and its role in inorganic carbon flux

Grazing and faecal pellet production by the copepods Calanus helgolandicus and Pseudocalanus elongatus, feeding on the coccolithophore Emiliania huxleyi, were measured under defined laboratory conditions, together with the chemical characteristics and sinking rates of the faecal pellets produced. Ingestion rates of both copepods were equivalent at comparable cell concentrations, the relationship between ingestion rate (I, cells copepod^-1 h^-1) and food concentration (C, cells ml^-1), being I=0.558C for both species. P. elongatus produced a larger number of smaller faecal pellets than C. helgolandicus, but egested a larger volume of material per individual. Only between 27 and 50% of the ingested coccolith calcite was egested in the faecal pellets, and it is possible that acid digestion in the copepod gut is responsible for these considerable losses. Average sinking rates of faecal pellets containing E. huxleyi coccoliths, produced by both species, were >100 m d^-1. The implications of the quantitative laboratory estimates for the vertical flux of inorganic carbon are considered using recently studied shelf-break and oceanic E. huxleyi blooms in the N. E. Atlantic as examples.     

Fate of domoic acid ingested by the copepod <Emphasis Type="Italic">Acartia clausi</Emphasis>

Two important issues in the studies of harmful algae include ecological role of the toxic compounds and their fate through the food web. The aims of this study were to determine whether the production of domoic acid is a strategy evolved to avoid predation and the role of copepods in the fate of this toxic compound through the food web. The copepod Acartia clausi was fed with single and mixed cultures of the toxic diatom Pseudo-nitzschia multiseries and the non-toxic diatom Pseudo-nitzschia delicatissima. Ingestion rate as a function of diatom abundance was the same for the toxic and non-toxic Pseudo-nitzschia species, indicating no selective feeding behaviour against P. multiseries. The toxins ingested by the copepods did not affect mortality, feeding behaviour, egg production and egg hatching of the copepods. Copepods assimilated the 4.8% of the total domoic acid ingested. Although the amount of toxins daily detoxificated by the copepods was 63.6%, the copepods accumulated domoic acid in their tissues. We conclude that domoic acid is not toxic for copepods and, probably for this reason, this toxin does not act as feeding deterrent for copepods. However, even though the production of domoic acid has apparently not evolved to deter predation, copepods may play an important role on the fate of this toxic compound through the marine food web.     

Comparison of gut-evacuation rates of feeding and non-feeding Calanus marshallae

Gut-evacuation rates were compared for feeding and non-feeding Calanus marshallae collected near Yaquina Bay, Oregon (44°37N; 124°04W) from June 1986 through May 1987. Evacuation rates were measured at four concentrations of Thalassiosira weissflogii from the decline of gut-pigment fluorescence following transfer of copepods to filtered seawater and from the loss of cells labelled with 68-germanium, a radioactive analog of silicon. There was no significant difference between gut-evacuation rates of feeding and non-feeding copepods over both short-term (20 min) and long-term (90 min) evacuation times. Furthermore there was no difference between rates obtained using either pigment or ⁶⁸Ge as tracers of gut food-passage. These results are discussed in the light of possible dilution of tracer in the guts of feeding copepods due to mixing with unlabelled food. Gut-evacuation rates measured at food concentrations ranging from 500 to 4000 cells ml^-1 were not significantly different, regardless of the technique employed.     

Biologie d'un copépode des mares temporaires du littoral méditerranéen français: Eurytemora velox

The feeding activity of Eurytemora velox, a brackish copepod from temporary lakes of the south of France, was studied in 1978–1979 using various foods (natural particles, monospecific algal cultures, and artificial food) under different conditions of temperature and salinity. Experiments with Amphidinium sp. or Tetraselmis maculata as food showed that the ingestion rate increased with food concentration according to an asymptotic or a linear relationship. Although of slightly smaller size, T. maculata was ingested at a higher rate than Amphidinium sp. Large maximum daily rations (up to 150% of body carbon with Amphidinium sp. and up to 250% with T. maculata) were attained. These values, which greatly exceed those generally obtained with marine copepods, could result from adaptation of the feeding processes of this copepod to its very rich trophic environment. A significant correlation was demonstrated between ingestion rate and fecal pellet production using T. maculata as food. Therefore, daily fecal production was used as an index of feeding activity in experiments carried out with natural food, T. maculata cultures and artifical food (Tetramin). Increased temperature generally resulted in an activation of grazing and filtration rates and of fecal production at low temperatures (10° to 15°C), but a strong decrease was observed over 22°C. Differences of 10 S over or under the natural salinity level led to a decrease in fecal production, suggesting unachieved acclimatization to salinity variation due to a too short acclimation period before the experiments. Fecal pellet production was higher during the day than during the night. It depended also on the quality of food used: high values were obtained with T. maculata, Phaeodactylum tricornutum, Rhodomonas sp. and Chlamydomonas sp., low values with Chlorella sp. and Amphidinium sp., and medium values with natural food material. The assimilation rate (A) was calculated by Conover's methods. A significant negative correlation was obtained between A and the ash content of the food. High assimilation rates were attained with chlorophycean algae, while natural particulate food produced variable assimilation rates, depending on the amount of inorganic material present.

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Biologie d'un copépode des mares temporaires du littoral méditerranéen français: Eurytemora velox

     

Feeding ecology of the lanternfish Benthosema pterotum from the Indian Ocean

The feeding ecology of lanternfish Benthosema pterotum (Alcock) from the north Arabian Sea, Mozambique and the Bay of Bengal was studied. Samples were collected on cruises carried out by R.V. Dr. Fridtjof Nansen during the period 1978 to 1983. A wide variety of zooplankton organisms were identified in the diet of B. pterotum with crustanceans dominating the diet. Copepods constituted ca. 40 to 90% of the diet. Dry weight analyses of the stomach contents from the Gulf of Oman in February 1983 showed copepods to be 35 to 55% in weight (average in samples). Ontogenetic differences were observed in the diet. Prey size increased as the fish length increased, but the largest fish did not exclude the smaller prey organisms from their diet. Regional variation in diet was also shown in B. pterotum. The degree of filling and the state of digestion of stomach contents revealed that this species feeds intensively at night in the epipelagic layer. All copepods indentified were epipelagic species, providing additional evidence of diurnal pattern in the feeding chronology of B. pterotum. Identification of copepods from the Gulf of Oman in February 1983, revealed that herbivorous species dominated in biomass. Quantitative analyses show that B. pterotum probably have a daily food intake of ca. 4.5% of the body weight.     

Phytoplankton pigments in the gut concents of planktonic copepods from coastal waters off southern California

Phytoplankton xanthophylls in the gut contents of the copepods Calanus pacificus, Corycaeus anglicus, and Paracalanus parvus, collected from 5 stations off San Onofre, California, in June 1982, were measured by reverse phase, high-performance liquid chromatography (HPLC). The dinoflagellate pigment, peridinin, was usually the most abundant xanthophyll in the guts of all three species of copepods. Evidently, feeding was principally on dinoflagellates (which dominated the phytoplankton biomass). The level of feeding activity, rather than the class of phytoplankton ingested, seemed to differentiate the behaviors of the copepods. Xanthophyll content per unit copepod wet weight was higher in Corycaeus anglicus and Paracalanus parvus than in Calanus pacificus. Chlorophyll a fluorescence of the copepod gut contents was measured in conjunction with the analysis of gut xanthophylls. The xanthophyll content of the gut varied directly with the concentration of chlorophyll a in the gut. Xanthophyll content was not related to the concentration of pheopigments in the gut. Apparently, the xanthophylls that were detected were due to the presence of recently ingested phytoplankton biomass.     

Effects of food-level acclimation on digestive enzyme activities and feeding behavior of Calanus pacificus

A recent hypothesis in the zooplankton literature states that zooplankton acclimate to ambient food concentrations such that higher digestive enzyme activities and, consequently, higher maximum ingestion rates are achieved at higher food levels. To test this hypothesis, adult female Calanus pacificus, collected from the main basin of Puget Sound, Washington, USA, in August 1979 and May 1982, were conditioned for 2 wk at different concentrations of the diatom Thalassiosira weissflogii (=fluviatilis). Ingestion rates and the activity of the digestive enzymes laminarinase, maltase, and cellobiase were measured periodically during acclimation and in a block-designed feeding experiment at the end of acclimation. Consistent with the hypothesis, maximum ingestion rate and digestive enzyme activity were positively correlated. However, in contrast to the hypothesized mechanism, this result arose because both maximum ingestion rate and digestive enzyme activity were negatively correlated with food concentration during acclimation. The enhanced ingestion of copepods following long-term (12 to 14 d) acclimation to low food is similar to that previously described for short-term (e.g. 1 d) starvation. It might be energetically optimal for copepods experiencing a patchy food environment to maintain higher levels of digestive enzymes at low food concentrations in order to exploit high concentrations of food when encountered.