Diel Feeding Rhythm Research Articles

Diel Feeding Rhythm and Grazing Selectivity of Small-Sized Copepods in a Subtropical Embayment, the Northern South China Sea

Small marine copepods are key components of the pelagic food webs in Chinese coastal waters, but very few studies have addressed their trophodynamics, with even fewer studies addressing their diel feeding rhythms. In this study, the diel feeding rhythm and grazing selectivity of the copepod assemblage in Daya Bay during September 30 to October 2, 2014, were studied based on gut pigment analysis. Small copepods (body length < 1.5 mm) including Paracalanus parvus, Temora turbinata, Acrocalanus gibber, Temora stylifera, Euterpe acutifrons, and Acrocalanus gracilis, accounted for 73.9–100% of the total copepod abundance. The copepod assemblage generally exhibited a diurnal feeding pattern, characterized by a higher gut pigment content and ingestion rate during the daytime, consistent with variation in the ambient Chl α concentration. Fifty-five percent of the phytoplankton standing stock per day was consumed by the copepod assemblage, wherein diatoms, prymnesiophytes, and cyanobacteria were the main prey items with average contributions of 19.4–32.9% to the gut pigment contents. The copepod assemblage showed a strong feeding preference for prymnesiophytes, a weak feeding preference for diatoms, and avoidance of cyanobacteria. These results suggest a strong top-down control on phytoplankton community, especially on small groups from small copepods in the Daya Bay ecosystem.

Predator Chemical Cue Effects on the Diel Feeding Behaviour of Marine Protists.

We have assessed the effect of copepod chemical cues on the diel feeding rhythms of heterotrophic and mixotrophic marine protists. All phagotrophic protists studied exhibited relatively high diurnal feeding rates. The magnitude of the diel feeding rhythm, expressed as the quotient of day and night ingestion rates, was inversely related to the time that phagotrophic protists were maintained in the laboratory in an environment without predators. In the case of the recently isolated ciliate Strombidium arenicola, the rhythm was lost after a few months. When challenged with chemical alarm signals (copepodamides) from the copepod Calanus finmarchicus at realistic concentrations (0.6-6 pM), S. arenicola partially re-established diurnal feeding. Conversely, the amplitude of the diel feeding rhythm for the ciliate Mesodinium rubrum was not affected by copepodamides, although the 24-h integrated food intake increased by approximately 23%. For the dinoflagellates Gyrodinium dominans and Karlodinium armiger, copepodamides significantly reduced the amplitude of their diel feeding rhythms; significant positive effects on total daily ingestion were only observed in G. dominans. Finally, the dinoflagellate Oxyrrhis marina, isolated >20 years ago, showed inconsistent responses to copepodamides, except for an average 6% increase in its total ingestion over 24 h. Our results demonstrate that the predation risk by copepods affects the diel feeding rhythm of marine protists and suggests a species-specific response to predation threats.

Non-lethal effects of the predator Meganyctiphanes norvegica and influence of seasonal photoperiod and food availability on the diel feeding behaviour of the copepod Centropages typicus.

Predators can induce changes in the diel activity patterns of marine copepods. Besides vertical migration, diel feeding rhythms have been suggested as an antipredator phenotypic response. We conducted experiments to assess the non-lethal direct effects of the predator Meganyctiphanes norvegica (northern krill) on the diel feeding patterns of the calanoid copepod Centropages typicus. We also analysed the influence of seasonal photoperiod and prey availability on the intensity of copepod feeding rhythms. We did not detect any large effect of krill presence on the diel feeding behaviour of copepods, either in day-night differences or total daily ingestions. Seasonal photoperiod and prey availability, however, significantly affected the magnitude of copepod feeding cycles, with larger diel differences in shorter days and at lower prey concentrations. Therefore, the role of non-lethal direct effects of predators on the diel feeding activity of marine copepods remain debatable and might not be as relevant as in freshwater zooplankton.

Trophic interactions and diel feeding rhythms of microzooplankton in a productive Swedish Fjord

Abstract Microzooplankton play a pivotal role in the energy transfer between lower and upper trophic levels in marine planktonic food webs. While laboratory data suggest that microzooplankton exhibit higher feeding rates during the daytime, evidence from the field is scarce and contradictory. In this study, we first characterized the nano- and microplanktonic communities of the Gullmar Fjord (Sweden) and its environmental conditions during July and August 2017. Then, we explored the grazing impact of microzooplankton on the phytoplankton community of this ecosystem and assessed their diel grazing activity using the dilution technique. Finally, we evaluated the impact of mesozooplankton at natural concentrations during the experiments. Microzooplankton removed 26% of the phytoplankton standing stock and 96% of the primary production daily, while mesozooplankton did not exert a significant impact on microplankton activity. We did not detect significant diel microzooplankton grazing rhythms during the first experimental period; however, during the second part of the study, after an upwelling event, grazing rates were significantly higher during the night. Therefore, the microzooplankton grazing rhythm in natural systems may vary according to the species composition and abundances of both microzooplankton and prey communities.

Towards an Understanding of Diel Feeding Rhythms in Marine Protists: Consequences of Light Manipulation

Temporal programs synchronised with the daily cycle are of adaptive importance for organisms exposed to periodic fluctuations. This study deepens into several aspects of the exogenous and endogenous nature of microbial grazers. We investigated the diel rhythms of cell division and feeding activity of four marine protists under different light regimes. In particular, we tested if the feeding cycle of protistan grazers could be mediated by a light-aided enhancement of prey digestion, and also explored the consequences of cell division on diel feeding rhythms. Cell division occurred at night for the heterotrophic dinoflagellates Gyrodinium dominans and Oxyrrhis marina. In contrast, the mixotrophic dinoflagellate Karlodinium armiger and the ciliate Strombidium sp. mostly divided during the day. Additionally, a significant diurnal feeding rhythm was observed in all species. When exposed to continuous darkness, nearly all species maintained the cell division rhythm, but lost the feeding cycle within several hours/days (with the exception of O. marina that kept the rhythm for 9.5days). Additional feeding experiments under continuous light also showed the same pattern. We conclude that the feeding rhythms of protistan grazers are generally regulated not by cell division nor by the enhancement of digestion by light. Our study, moreover, indicates that the cell division cycle is under endogenous control, whereas an external trigger is required to maintain the feeding rhythm, at least for most of the species studied here.

Planktonic food web structure and trophic transfer efficiency along a productivity gradient in the tropical and subtropical Atlantic Ocean

Oligotrophic and productive areas of the ocean differ in plankton community composition and biomass transfer efficiency. Here, we describe the plankton community along a latitudinal transect in the tropical and subtropical Atlantic Ocean. Prochlorococcus dominated the autotrophic community at the surface and mixed layer of oligotrophic stations, replaced by phototrophic picoeukaryotes and Synechococcus in productive waters. Depth-integrated biomass of microzooplankton was higher than mesozooplankton at oligotrophic stations, showing similar biomasses in productive waters. Dinoflagellates dominated in oligotrophic waters but ciliates dominated upwelling regions. In oligotrophic areas, microzooplankton consumed ca. 80% of the production, but ca. 66% in upwelling zones. Differences in microzooplankton and phytoplankton communities explain microzooplankton diel feeding rhythms: higher grazing rates during daylight in oligotrophic areas and diffuse grazing patterns in productive waters. Oligotrophic areas were more efficient at recycling and using nutrients through phytoplankton, while the energy transfer efficiency from nutrients to mesozooplankton appeared more efficient in productive waters. Our results support the classic paradigm of a shorter food web, and more efficient energy transfer towards upper food web levels in productive regions, but a microbially dominated, and very efficient, food web in oligotrophic regions. Remarkably, both models of food web exist under very high microzooplankton herbivory.

Diel feeding rhythms, daily ration, and seasonal changes thereof in marbled flounder Pseudopleuronectes yokohamae

This work was partly supported by the Stock Assessment Program of Fisheries Agency and Fisheries Research and Education Agency of Japan and the program “Project on Clarifying the Impact of Nuclear Substances” of the Fisheries Agency of Japan.

The Community Structure and Grazing Rates of Meiobenthos in Chunsu Bay, West Coast of Korea

Min, W.-G.; Kwon, K.-G.; Rho, H.S., and Kim, D., 2018. The Community Structure and Grazing Rates of Meiobenthos in Chunsu Bay, West Coast of Korea. In: Shim, J.-S.; Chun, I., and Lim, H.S. (eds.), Proceedings from the International Coastal Symposium (ICS) 2018 (Busan, Republic of Korea). Journal of Coastal Research, Special Issue No. 85, pp. 401–405. Coconut Creek (Florida), ISSN 0749-0208.The community structure and grazing rates of meiobenthos was studied in Chunsu bay on west coast of Korea. To examine feeding habit of major meiofaunal groups, grazing rate of nematodes and harpacticoid copepods was measured using the isotope-tracer (3H thymidine, 14C amino acid) method at 3 stations in subtidal area and 1 station in intertidal area of Chunsu bay. Time-serial measurement of grazing rate was made to elucidate the effects of day/night and tidal cycle on diel feeding rhythm of the animals. The number of culturable bacteria fluctuated from 1.73±0.25×105 to 18.23±1.17×105 CFU/cm3 during the study period, and was highest in October. The Concentration of Chl.a in the sediment at the study site was 18.0~168.2 mg/m2 in April and 36.6~111.0 mg/m2 in October. The most abundant meiobenthos was nematodes and followed by sarcomastigophorans, harpacticoid copepods. Density of meiobenthos were 5~1632 inds./10cm2 during the study period. The grazing rates of nematode varied 0.5~57.2%/h on bacteria and 0.4~53.3%/h on microalgae at the station in tidal flat. The grazing rates of harpacticoid copepods were 0.2~14.2%/h on bacteria and 0.3~12.2%/h on benthic microalgae at the station in tidal flat. In the subtidal area, the grazing rates of nematode were 33.4~82.5%/h on bacteria and 10.3~71.5%/h on microalgae and harpacticoid copepods were 0.2~14.2%/h on bacteria and 0.3~12.2%/h on benthic microalgae.

Diel feeding rhythms in marine microzooplankton: effects of prey concentration, prey condition, and grazer nutritional history

In this study, we aim at disentangling the causes and consequences of diel feeding rhythms in marine microzooplankton. We focused on the diel feeding activity of two heterotrophic dinoflagellate species, Gyrodinium dominans (one laboratory strain) and Oxyrrhis marina (laboratory cultivated and wild strains). We observed higher ingestion during the day in both dinoflagellate species. Feeding rhythms appeared to be independent of circadian changes in prey biochemical composition. Grazers fed with prey under stationary phase, with equivalent stoichiometric composition between day and night, showed 5 (G. dominans) and 10 (O. marina) times higher ingestion rates during the day. Previous grazer feeding history (starved vs well-fed) did not affect the feeding rhythm. However, prey concentration altered the rhythm; food limiting conditions reduced the amplitude of the rhythms. Our results establish a resource dependence of diel periodicity in microzooplankton grazing, which can have unanticipated consequences for standard field dilution grazing experiments.

Regulation of hemolymph trehalose level by an insulin-like peptide through diel feeding rhythm of the beet armyworm, Spodoptera exigua

Like vertebrate insulins, some insect insulin-like peptides (ILPs) play crucial roles in controlling immature growth, adult lifespan, and hemolymph sugar level. An ILP gene (SeILP1) was predicted from a transcriptome database of Spodoptera exigua. SeILP1 encodes 95 amino acid sequence and shares sequence homologies (33–83%) with other insect ILPs, in which six conserved cysteine residues are found in the predicted B–A chains. SeILP1 was expressed in all developmental stages of S. exigua. However, SeILP1 expression was tissue-specific because the transcript was detected in fat body and epidermis, but not in hemocytes and gut. Its expression increased with feeding activity. Hemolymph trehalose levels of the fifth instar larvae maintained a relatively constant level at 2.31±0.62mM. However, starvation induced a significant increase of the hemolymph trehalose level by more than twofold in 48h, at which few SeILP1 was transcribed. RNA interference of SeILP1 using its specific double-stranded RNA induced a significant increase of hemolymph trehalose level. Interestingly, a bovine insulin decreased hemolymph trehalose level in a dose-dependent manner. These results indicate that SeILP1 plays a role in suppressing hemolymph trehalose level in S. exigua.

Versatile mysids exploit multiple basal resources: implication of the bentho-pelagic habit in estuarine food webs

Mysid feeding ecology was studied in a tropical mangrove estuary and adjacent coastal mudflat, using stomach content and stable isotope analyses. We tested the hypothesis that estuarine mysids are able to exploit the various basal food resources, while their bentho-pelagic habit related to nocturnal feeding explicates their important role as trophic intermediaries in estuarine food webs. Diet composition differed among species, between habitats and between day and night. Acanthomysis thailandica, Notacanthomysis hodgarti, Mesopodopsis orientalis, and Rhopalophthalmus hastatus were generally omnivorous, while R. orientalis tended to carnivory. Although both examined species, A. thailandica and N. hodgati, showed no diel feeding rhythm, heavy feeding on surface copepods occurred at night. The largely mudflat species (A. thailandica, N. hodgarti, M. orientalis, and R. orientalis) showed significantly enriched mean δ13C values compared to the upper estuarine species, R. hastatus. The δ 13C values suggest that coastal species utilized carbon sources from phytoplankton and benthic diatoms, while estuarine species utilized more mangrove carbon. Since mysid shrimps can derive nutrition from all basal sources, they form an important link in the flow of trophic energy from source to the higher consumers of tropical estuarine ecosystems.

Food consumption of five deep-sea fishes in the Balearic Basin (western Mediterranean Sea): are there daily feeding rhythms in fishes living below 1000 m?

Predation and food consumption of five deep-sea fish species living below 1000 m depth in the western Mediterranean Sea were analysed to identify the feeding patterns and food requirements of a deep-sea fish assemblage. A feeding rhythm was observed for Risso's smooth-head Alepocephalus rostratus, Mediterranean grenadier Coryphaenoides mediterraeus and Mediterranean codling Lepidion lepidion. Differences in the patterns of the prey consumed suggest that feeding rhythms at such depths are linked with prey availability. The diets of those predators with feeding rhythms are based principally on active-swimmer prey, including pelagic prey known to perform vertical migrations. The diets of Günther's grenadier Coryphaenoides guentheri and smallmouth spiny eel Polyacanthonotus rissoanus, which did not show any rhythm in their feeding patterns, are based mainly on benthic prey. Food consumption estimates were low (<1% of body wet mass day(-1) ). Pelagic feeding species showing diel feeding rhythms consumed more food than benthic feeding species with no feeding rhythms.

The food supply of the Pacific salmon of the genus Oncorhynchus in the northwestern Pacific Ocean. 2. Comparative characterization and general state

The interannual variations and general state of the food supply of Pacific salmon (Oncorhynchus spp.) in the 2000s in the northwestern Pacific Ocean (including the Bering Sea and the Sea of Okhotsk) were analyzed based on indirect characteristics that indicate the variability of their forage base, feeding habits, growth, and biomass. A new index for the quantitative evaluation of food supply was suggested. The food supply of the Pacific salmon during the 2000s was found to be sufficient to maintain the normal functioning of populations. With high abundance of Pacific salmon, the food supply tended to decrease. However, this caused no negative consequences for the survival of major salmon stocks during the marine period of life and, as a rule, no marked decrease in the food consumption and growth rates of fish. A relative increase in food competition was compensated by adaptive changes in the diet and diel feeding rhythm of salmon. With the shortage of preferred food organisms (amphipods, euphausiids, and pteropods), Pacific salmon changed to consuming minor prey (copepods and chaetognaths), and numerous mesopelagic species of macroplankton and micronekton in the evening hours.

Vertically resolved prey selectivity and competition of Baltic herring Clupea harengus and sprat Sprattus sprattus

Prey selectivity of Baltic herring Clupea harengus and sprat Sprattus sprattus was studied in the Bornholm Basin, Baltic Sea, in June 2001. A total of 165 sprat stomachs (10 to 15 cm total length, TL) and 214 herring stomachs (12 to 30 cm TL) were analysed. The diel vertical distribution of zooplankton prey was analysed by multi-net samples; clupeid distributions were estimated by hydroacoustic measurements. These measurements enabled us to describe the diel feeding rhythm and to estimate vertically resolved selectivity indices for the 2 most important zoo-planktivores in the Baltic Sea. Diet composition of herring and sprat were similar (mainly copepods and cladocerans), resulting in strong competition. Possibly to reduce this competition, both species were partly specializing on certain prey species (sprat: Podon spp.; herring: Evadne nordmanni and Temora longicornis) and copepodite stages (sprat: adult [C6] males of Pseudocalanus acuspes; herring: C6 females of P. acuspes). Sprat and, to some extent, herring exhibited a marked shift in prey preference between day and night. Sprat mainly selected T. longicornis during the day and Podon spp. during the night, while herring mainly selected T. longicornis during the day and E. nordmanni during parts of the night. A comparison of the field stomach contents with the estimated gastric evacuation predicted by parameters based on laboratory experiments indicated that sprat fed during the night, while herring did not or only to a minor extent. Comparison of our zooplankton sampling scheme with commonly used sampling designs revealed that investigations which consider both time and stage are needed to fully understand the feeding selectivity dynamics of herring and sprat in the Baltic Sea. However, the objective of a selectivity study should determine the most appropriate zooplankton sampling scheme.

Diel vertical migration and feeding of three species of chaetognaths (Flaccisagitta enflata, Aidanosagitta delicata and Aidanosagitta neglecta) in two shallow, subtropical bays in Hong Kong

The diel behavior and feeding ecology of three coexisting species of chaetognaths in two shallow subtropical bays in the northeastern part of Hong Kong were studied in summer 2005 and winter 2005–2006. Average densities of adult Flaccisagitta enflata, Aidanosagitta delicata and Aidanosagitta neglecta ranged from 21 to 48 ind. m in the semi-enclosed Tolo Harbour, but densities of the two Aidanosagitta species were low in Mirs Bay, which is deeper and opens into the South China Sea. Adult F. enflata, the largest of the three species, exhibited the most pronounced diel vertical migration (DVM) and diel feeding rhythm. The smaller juvenile F. enflata and adult A. delicata did not exhibit DVM. Although copepods were the most abundant component of the mesozooplankton, they were not preferred prey of chaetognaths. Predation impacts on copepods were generally ,0.3% of the standing stock. Larvaceans, on the other hand, were highly preferred, with predation impact as high as .30%. As prey abundances in Tolo Harbour could be .9 10 ind. m, there was no evidence of resource partitioning behavior by the coexisting chaetognaths.

Effect of Water Temperature on Feeding Rhythm in Common Carp (Cyprinus carpio haematopterus Temminck et Schlegel)

Feeding rhythm of common carp was investigated from 4°cC to 34°C. The results indicated that there was a diel feeding rhythm for both adult (630–850 g) and youth (61–91 g) at all tested temperatures. There were two main activity peaks at 8:00–11:00 a.m. and 19:00–23:00 p.m., during which feeding quantities were 10.68%–32.53% and 16.25%–33.41% of the daily intake, respectively. When water temperature dropped to below 10°C, the feeding peak concentrated at 8:00–11:00 a.m. and 19:00 p.m. to 4:00 a.m. At 6°C, though both adult and youth would still feed, the feeding quantities were only 0.01% and 0.35% of body mass. Daily feeding rate of adult and youth reached 1.21% and 2.63% at 14°C, respectively. Both adult and youth carps reached the maximum daily feeding rate at 28°C, being 2.84% and 12.06% of body mass, respectively. The daily feeding rate of adult and youth reduced suddenly after at 34°C, and the daily feeding rate was only 0.74% and 9.45% of body mass, respectively. There was significant difference in daily feeding rate at different water temperatures (P<0.05).

The seasonal distribution, diel vertical distribution and feeding behavior of Paraeuchaeta concinna in the shallow subtropical coastal waters of eastern Hong Kong

BackgroundPredatory copepods of the family Euchaetidae are widely distributed in polar, temperate, subtropical and tropical oceans. Paraeuchaeta concinna is the most abundant Euchaetidae in the subtropical coastal seas of Hong Kong and southern China. However, compared to Euchaetidae species in temperate and polar regions, relatively little information is available on the ecology of P. concinna and other Euchaetidae species in the subtropical oceans. This paper provides information on the seasonal abundance of P. concinna in the coastal seas of eastern Hong Kong. The diel vertical distribution of P. concinna, feeding behavior, and predation impact on mesozooplankton in eastern Hong Kong were also investigated.ResultsP. concinna is most abundant in winter and spring. Their abundance decreases shoreward, and densities are generally higher in the open waters of eastern Hong Kong than in the inner parts of Mirs Bay and Tolo Harbour. P. concinna exhibits both diel vertical migration and diel feeding rhythms in Mirs Bay. P. concinna females show strong preference for the copepods of the genera Acrocalanus, Paracalanus, and Parvocalanus, and remove ~4% of their standing stocks daily.ConclusionsThe low abundance of P. concinna during most of the year suggests it is not indigenous to coastal seas of eastern Hong Kong. P. concinna performs diel vertical migration, most likely as a strategy to avoid visual predation. Gut content analysis showed that Acrocalanus, Paracalanus, and Parvocalanus are highly preferred prey of P. concinna. A daily predation impact of ~4% of the standing stocks of Acrocalanus, Paracalanus, and Parvocalanus suggests that P. concinna may play an important role in regulating the populations of these small copepods in Mirs Bay, especially during winter and spring.

Copepod feeding in a coastal area of active tidal mixing: diel and monthly variations of grazing impacts on phytoplankton biomass

AbstractThis study examined monthly feeding rates and grazing impact on phytoplankton biomass, as well as diel feeding rhythms of four key copepod species in a tidally well mixed estuary (Asan Bay, Korean Peninsula). Monthly ingestion rates estimated based on gut pigment analysis were closely associated with their peak densities, but not with phytoplankton biomass, implying high ingestion may be related to reproductive output for population growth. The three smaller copepods, Acartia hongi, Acartia pacifica and Paracalanus parvus, showed feeding preference for smaller phytoplankton (&lt;20 μm) with higher clearance rates, whereas the larger Calanus sinicus preferred larger phytoplankton. Acartia pacifica and P. parvus showed distinct increased nocturnal feeding rates as measured with gut fluorescence, whereas A. hongi showed no significant day–night differences. Copepod diel feeding patterns were not associated with food quantity, and endogenous physiological rhythm might be hypothesized as responsible for the observed diel feeding patterns. Grazing impact on phytoplankton biomass by the four copepods in the estuary was on average 8% (range 0.2–29.8%) of the phytoplankton standing stock, similar to values reported in other coastal waters. Very high copepod abundances but low daily carbon ration (&lt;20% for all copepods) provided by feeding on phytoplankton indicate that copepods also grazed on other non‐phytoplankton foods in Asan Bay.

Transpositional feeding rhythm of loach Misgurnus anguillicaudatus from larvae to juveniles and its ontogenesis under artificial rearing conditions

The diel feeding rhythm and ontogenesis during early life stage of loach Misgurnus anguillicaudatus were investigated under experimental conditions (light: L 06:00–18:00, D 18:00–06:00 h). Morphological and behavioral developments of loach from newly hatched to 40 days after hatching were observed. Larvae were able to prey on daphnia 3–4 days after hatching at 23 ± 0.5°C. As the larvae grew, they showed an increasing feeding capacity and a distinct feeding rhythm. Feeding intensity and incidence for day-4 larvae were highest at 10:00 and 16:00 h. The highest levels of feeding intensity for day-12 larvae occurred at 08:00, 12:00, and 18:00 h as did feeding incidence. By day 20, when the larvae metamorphosed, the highest levels of feeding intensity occurred at 06:00, 18:00, and 24:00 h and were concurrent with the highest feeding incidence. After metamorphosis, feeding capacity had again increased considerably and, in contrast to the earlier stages before day 20, feeding intensity for day-30 juveniles peaked at 05:00 and 20:00 h, about 1–2 h after the maximum feeding incidence. The feeding rhythm of loach juveniles at day 40 was almost the same as the day-30 juveniles. The estimated maximum daily feeding rates were 43.1%, 33.4%, 19.0%, 12.8%, and 5.8% of body weight on days 4, 12, 20, 30, and 40, respectively. Thus, loach was found to have different feeding rhythms in the pre- and post-metamorphosis stages, with the highest feeding activity in daytime during the larval planktonic stage before metamorphosis, and intensely nocturnal feeding behavior during the juvenile benthic stage after metamorphosis.

Reproductive activity and physiological status of the calanoid copepods Calanus helgolandicus and Calanoides carinatus under food-limiting conditions

The reproductive activity and the physiological state of the calanoid copepods Calanus helgolandicus and Calanoides carinatus were investigated off the coast of NW Spain during autumn to evaluate the effect of short food resources on both populations. Phytoplankton biomass was low, and neither phytoplankton size distribution nor composition was suitable to support high reproductive rates. Accordingly, egg production rates (EPR) were much lower than maximum rates for both species, pointing to food limitation. The reproductive index (RI), which represents the proportion of females with mature gonads, was < 50% at each of the three zones into which the sampling area was divided (coast, shelf and ocean). Potential recruitment rates were very low except at some nearshore stations, where the highest concentrations of chlorophyll- a (Chl- a), diatoms, dinoflagellates and large cells were found. EPR of C. helgolandicus and C. carinatus were correlated with phytoplankton biomass and unaffected by temperature. Phytoplankton carbon ingestion explained ca. 50% of the variability in EPR for both species. At most of the stations, herbivory was insufficient to cover the carbon requirements for reproduction and respiration, so females probably fed on heterotrophic prey to meet their demands. However, given the low fecundity observed, this omnivorous diet did not seem to be optimum for reproduction, and a severe food limitation is thus suggested. Furthermore, the high C/N values measured point to a notable lipid storage, but given the low EPR found, lipid reserves were probably invested into female maintenance rather than into gonad maturation. C. helgolandicus and C. carinatus populations did not mirror phytoplankton biomass distribution, but they correlated well when considering only copepodites V (CV). The CV could be preparing for the overwintering, storing lipid reserves to ensure a successful diapause, and they could also be advected by the poleward current detected during the study. Females showed a diel feeding rhythm, with highest ingestion rates during night. From our results, it follows that C. helgolandicus and C. carinatus females did not perform diel vertical migrations. We suggest that this behaviour is likely due to the food-limiting conditions, which make it more advantageous to remain at the surface during daytime.