Experimental studies on elimination of zinc-65, cesium-137 and cerium-144 by euphausiids

The elimination of 3 radionuclides from Euphausia pacifica was measured over a 5 month period. The biological half-lives for ⁶⁵Zn, ¹³⁷Cs, and ¹⁴⁴Ce, calculated after the euphausiids had ingested radioactive Artemia nauplii, were found to be 140 days, 6 days, and 7.5 h, respectively. The percentages of body burdens lost in molts were greatest for the fission products, ¹⁴⁴Ce (21%) and ¹³⁷Cs (7%), and least for ⁶⁵Zn (1%). Elimination of the isotopes in the feces could not be followed because of the difficulty in collecting fecal material for analysis; however, 1 sample collected 2 months after the beginning of the elimination experiment had no measurable radioactivity. Loss of ⁶⁵Zn from molts and time to disintegration of the molts were found to be temperature dependent over a 5° to 15°C range, and the sinking rate of molts was both temperature and salinity dependent. Calculations showed that, in areas in the North Pacific outside the influence of upwelling, percentage ⁶⁵Zn loss from sinking molts (before disintegration of the molts) was likely to be the same throughout the year, since the molts would be exposed to about the same mean temperature in the water column in all seasons. Even though temperature structure in the upper layers changes with season, mean temperatures change very little when calculated over the sinking distance of intact molts. Intact molts would sink to slightly over 400 m in the absence of turbulence, and would lose 87% of their ⁶⁵Zn by the time they reached this depth. Sinking molts thus might contribute substantially to the vertical transport of ⁶⁵Zn in the sea. If loss of ⁶⁵Zn in fecal pellets is assumed to be small under our experimental conditions, and molting loss is only 1% of ⁶⁵Zn body burden, the major mechanism of ⁶⁵Zn loss from euphausiids feeding on non-radioactive food must be isotopic exchange with the water. Approximately 96% of the initial body burden was eliminated over a period of 5 months.Supported by USAEC research contract AT (45-1)1830, PHS grant ES00026, and a Richland Graduate Fellowship to S. W. Fowler.     

Genetic differentiation in a pelagic crustacean (Meganyctiphanes norvegica: Euphausiacea) from the North East Atlantic and the Mediterranean Sea

Meganyctiphanes norvegica (M. Sars)is a pelagic crustacean that plays a key role in marine food webs of North Atlantic Ocean and marginal seas. We studied eight population samples collected in the European Atlantic and Mediterranean Sea. By means of single strand conformation polymorphism analysis (SSCP) and direct sequencing, we investigated a segment of 158 base pairs of the mitochondrial gene coding for the subunit 1 of NADH dehydrogenase. We found 12 sequence variants among the 385 individuals studied. Analysis of molecular variance (AMOVA) showed that 14.75% of the total genetic variability was explained by differences between populations, thus indicating absence of panmixia for these populations. Pairwise comparisons revealed three distinct genetic pools: the first one represented by Cadiz Bay, the second one by the Ligurian Sea, and the third one included all the NE Atlantic samples. We also investigated one population from the Alboran Sea (within the Mediterranean basin, east of the Strait of Gibraltar). This population was found to be genetically intermediate between the NE Atlantic samples and the Ligurian sample, suggesting that the restriction to the gene flow is not associated with the Strait of Gibraltar, but possibly with the Oran–Almeria oceanographic front. The present work indicates that M. norvegica, although endowed with a high dispersal capacity because of its pelagic habit, can develop separate breeding units inside the same oceanic basin (the Atlantic). Furthermore, the Ligurian sample should be considered as a distinct evolutionary entity, separated from the Atlantic population. Received: 2 May 1999 / Accepted: 26 November 1999     

Continuous plankton records: Geographical variations in numerical abundance,biomass and production of euphausiids in the North Atlantic Ocean and the North Sea

Geographical variations in the numbers, biomass and production of euphausiids and the contribution of common species to the total are described from samples taken during 1966 and 1967 in the North Atlantic Ocean and the North Sea by the Continuous Plankton Recorder at 10 m depth. Euphausiids were most abundant in the central and western North Atlantic Ocean and the Norwegian Sea. Thysanoessa longicaudata (Krøyer) was numerically dominant. Biomass was greatest in the Norwegian Sea and the north-eastern North Sea where Meganyctiphanes norvegica (M. Sars) accounted for 81 and 59%, respectively, of the total biomass. Production was highest off Nova Scotia and in Iberian coastal waters; the dominant species were T. raschi (M. Sars) in the former area and Nyctiphanes couchi (Bell) in the latter. The mean P:B ratios were correlated with temperature.     

Distribution of euphausiid assemblages in the Mediterranean Sea

Eleven species of euphausiids from 24 Isaac-Kidd Midwater Trawl (IKMT) night collections taken at stations throughout the Mediterranean Sea were counted. The frequency of occurrence and dominance of individual species and percent similarity faunal analysis of the euphausiid community were used to describe changes in faunal composition between geographical areas and differences in vertical distribution. Although most species were widespread, three distinct patterns of abundance were apparent: Euphausia krohnii, Nematoscelis megalops, Meganyctiphanes norvegica, and Stylocheiron abbreviatum predominated in western basin areas west of the Tyrrhenian Sea; Euphausia hemigibba, Thysanopoda aequalis, and Stylocheiron longicorne predominated in the Tyrrhenian Sea and eastern Mediterranean Sea; Euphausia brevis and Stylocheiron suhmii predominated in the eastern Mediterranean Sea. Percent similarity analysis of data from the IKMT samples and data from Ruud (1936) indicates the Tyrrhenian Sea fauna at the time of the collections was more similar to eastern Mediterranean areas than to most other areas in the western basin, although the degree of similarity was dependent, to some extent, on the depth at which the samples were collected. The composition of euphausiids living above 150 m at night in this area was more similar to eastern basin areas, while the composition of deeper living forms was more similar to those of the rest of the western basin. Comparison of euphausiids collected at three points over a 60 year time-span in the Balearic Sea shows the similarity in composition to be greater within the area over time than between adjacent areas in the western Mediterranean Sea.Contribution No. 2732 from the Woods Hole Oceanographic Institution, Woods Hole, Massachusetts, USA. This research was supported by the Atomic Energy Commission Contract AT (30-1)-3862, ref. NYO-3862-49, National Science Foundation Grant GA 29303 and Office of Naval Research Contract N00014-71-C-0284.     

Interactions of marine plankton with transuranic elements

In a series of laboratory experiments, the biokinetics of ²⁴¹Am, an important transuranium element, was studied in Meganyctiphanes norvegica, a euphausiid common in the northwestern Mediterranean. The euphausiids accumulated Am from water by passive adsorption onto exoskeletons, achieving wet weight concentration factors on the order of 10² after 1 wk exposure; concentration factors varied inversely with the size of the euphausiids and linearly with their surface area:wet weight ratios. Essentially all (96±10%) of the Am taken up from water was associated with the exoskeleton, so that negligible Am was retained by the euphausiids after molting. The retention half-time of Am in molts was 2.9 d. Euphausiids could also concentrate Am from feeding suspensions by ingesting Am-labelled diatom cells, although there was negligible Am assimilation (3±2% after 4 d feeding); after passage through the gut, virtually all (99%) of the ingested Am was defecated within 1 wk. The retention half-time of Am in fecal pellets was 41 and 51 d at 13° and 5°C, respectively. In oceanic waters, where the preponderance of ²⁴¹Am is in the dissolved phase, uptake of Am from water by euphausiids would be the dominant route of bioaccumulation. The results underscore the importance of sinking biogenic debris from zooplankters in mediating the vertical transport of Am in the sea. Given their retention half-times for ²⁴¹Am and their rapid sinking rates, fecal pellets and discarded molts have the potential to deliver most of their Am to the sediments.     

Flux of cadmium through euphausiids

Flux of the heavy metal cadmium through the euphausiid Meganyctiphanes norvegica was examined. Radiotracer experiments showed that cadmium can be accumulated either directly from water or through the food chain. When comparing equilibrium cadmium concentration factors based on stable element measurements with those obtained from radiotracer experiments, it is evident that exchange between cadmium in the water and that in euphausiid tissue is a relatively slow process, indicating that, in the long term, ingestion of cadmium will probably be the more important route for the accumulation of this metal. Approximately 10% of cadmium ingested by euphausiids was incorporated into internal tissues when the food source was radioactive Artemia. After 1 month cadmium, accumulated directly from water, was found to be most concentrated in the viscera with lesser amounts in eyes, exoskeleton and muscle, respectively. Use of a simple model, based on the assumption that cadmium taken in by the organism must equal cadmium released plus that accumulated in tissue, allowed assessment of the relative importance of various metabolic parameters in controlling the cadmium flux through euphausiids. Fecal pellets, due to their relatively high rate of production and high cadmium content, accounted for 84% of the total cadmium flux through M. norvegica. Comparisons of stable cadmium concentrations in natural euphausiid food and the organism's resultant fecal pellets indicate that the cadmium concentration in ingested material was increased nearly 5-fold during its passage through the euphausiid. From comparisons of all routes by which cadmium can be released from M. norvegica to the water column, it is concluded that fecal pellet deposition represents the principal mechanism effecting the downward vertical transport of cadmium by this species.     

Predation on euphausiids by cod,Gadus morhua,in winter in Icelandic subarctic waters

The euphausiids consumed by cod (Gadus morhua Linnaeus, 1758) taken in November 1983 and February–March 1984 on feeding grounds in the subarctic shelf area north-west, north-east, and south-east of Iceland have been examined. A total of 2 714 euphausiids occurred in the 1 029 stomachs with food which were analysed; of these 1 640 (60%) could be identified to species. Thysanoëssa inermis (Krøyer, 1846) and Meganyctiphanes norvegica (M. Sars, 1857) were most numerous, constituting 58 and 40%, respectively, of the euphausiids which could be identified. The other euphausiids were, in declining numbers of abundance, T. raschii (M. Sars, 1864), T. longicaudata (Krøyer, 1846), and Nematoscelis megalops G. O. Sars, 1883. The mean number of euphausiids occurring per stomach showed considerable variation between both sampling areas and time of sampling. In each area, the mean number of M. norvegica occurring per stomach was similar at both sampling times, while for T. inermis it was greater in February–March than in November. The mean number of euphausiids occurring per stomach increased with increasing size of cod, being 1 to 2 and 4 to 6 in 10 to 30 cm and 50 to 70 cm cod, respectively. An examination of the diel variation in the occurrence of the euphausiids in the stomachs indicated two peaks, one in the morning before sunrise and another in the evening around and after sunset.     

Flux of 141Ce through a euphausiid crustacean

The role of the Mediterranean euphausiid Meganyctiphanes norvegica in the cycling of radiocerium (¹⁴¹Ce) was examined. When uptake of ¹⁴¹Ce occurs directly from the water, a dynamic population equilibrium is reached at a concentration factor of about 250. Molting was responsible for up to 99% loss of total body burden at first molt, and about 45% of the remaining activity at second molt, thus denying true longterm equilibrium to individual animals. Fecal pellets did not contain measureable ¹⁴¹Ce activity when the euphausiids accumulated the isotope from water, thus proving that surface adsorption was the key accumulating process from water. When radiocerium was taken in through ingestion of labelled Artemia, about 99% of the body burden was voided as fecal pellets. Excretion by this route was accelerated when euphausiids were fed non-radioactive Artemia during loss phase. Radioactive counts of the pellets confirmed that all ingested ¹⁴¹Ce was lost through defecation. When ¹⁴¹Ce was ingested as labelled phytoplankton, a substantial fraction of the total body burden occurred in the molts, which indicated that the phytoplankton lost ¹⁴¹Ce to the water and the radioactivity was subsequently adsorbed to outer surfaces of the euphausiids. Molts, fecal pellets, and freshly-killed euphausiids lost ¹⁴¹Ce to the water exponentially, the rates being similar to the exponential portions of the loss curves for live, non-molting individuals. It is suggested that M. norvegica, and probably other pelagic zooplankters, can greatly accelerate radiocerium transport to the ocean floor by packaging the isotope as fecal pellets. In coastal areas subject to low-level radioactive waste disposal, ¹⁴¹Ce might be ionic (or at least soluble) to a great extent, in which case euphausiids could take up the isotope rapidly and accelerate its vertical transport via molting.     

Distribution and feeding of the carnivorous copepod Paraeuchaeta norvegica in habitats of shallow prey assemblages and midnight sun

Paraeuchaeta norvegica was found to be widely distributed in the Norwegian Sea. They were least abundant in north-western areas, but otherwise no clear horizontal patterns appeared with respect to latitude, longitude or water mass. Females and males had similar vertical distributions. The highest concentrations of adults occurred at 400-500 m depth; they largely avoided the upper 50-100 m, even at night. Stages CIV and CV lived shallower in the water column than the adults, with the highest concentration between 100 and 300 m. Stages CII-CIII were most abundant at 50-100 m, while CI was distributed slightly deeper (maxima at 100-200 m). Potential prey were most abundant in the upper 100 m; i.e. shallower than P. norvegica. Numbers of fecal pellets produced by freshly collected adult females were relatively low (estimated at 0.7 pellets per individual on average for the entire sea), with maximal numbers for individuals captured in shallow waters. This suggests food limitation during summer, when food is concentrated in upper waters, and short and light nights limit nocturnal access to the shallow food resources. Pellets mainly contained copepod remains.     

Observations sur le développement larvaire de Meganyctiphanes norvegica (Crustacea: Euphausiacea) au laboratoire

Larvae of Meganyctiphanes norvegica (M. Sars), caught with a plankton net, were reared in the laboratory from the calyptopis phase onwards, under different temperature and trophic conditions. Eyestalk ablation was performed on the first furciliae. Data are presented on growth, moulting rate and ontogenesis of the larvae as a function of the varying experimental conditions, and on larval morphology, pigmentation and diet. Recent modifications of larval nomenclature in euphausiids are discussed.

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Observations sur le développement larvaire de Meganyctiphanes norvegica (Crustacea: Euphausiacea) au laboratoire

     

Polonium-210 in euphausiids: A detailed study

A detailed study of ²¹⁰Po, the predominant alpha-emitting nuclide found in most marine organisms, has been undertaken in a particular zooplanktonic species, the euphausiid Meganyctiphanes norvegica. The purpose was to obtain information concerning the origin, the localization and the flux of the nuclide in and through this organism. Measurements of ²¹⁰Po were made in euphausiids of different sizes, in dissected organs and tissues, and in excretion products. The results show higher concentrations in the smaller specimens; this fact cannot be explained on the basis of surface adsorption, but is probably related to the ingestion of food. Dissection results show that the distribution of ²¹⁰Po in euphausiids is not homogeneous, but that the majority is concentrated in the internal organs, the alimentary tract and the hepatopancreas in particular. The natural radiation dose received by these organs is in consequence much higher than that received by the whole animal. Use of a dynamic model allowed the flux of ²¹⁰Po through M. norvegica to be calculated. The calculations confirm that food is the principal source of ²¹⁰Po for this species, and clearly show that fecal pellets constitute the major elimination route. Extrapolation of the data to zooplankton in general leads to the conclusion that zooplankton metabolic activity plays an important role in transporting ²¹⁰Po from the surface layers of the ocean to depth.     

Flux of different forms of 106Ru through a marine zooplankter

The dynamics of accumulation and loss of different physico-chemical forms of ¹⁰⁶Ru were measured in the euphausiid Meganyctiphanes norvegica. The accumulation of ¹⁰⁶Ru was directly related to the concentration of the radioisotope in solution, as evidenced by similar concentration factors for euphausiids in the low and high activity ¹⁰⁶Ru chloride solutions. The chemical form of the radioisotope in solution had a pronounced effect on the uptake, with ¹⁰⁶Ru chloride fractions being accumulated at a faster rate than ¹⁰⁶Ru nitrosyl-nitrato complexes. Euphausiids lost ¹⁰⁶Ru, previously accumulated from the ¹⁰⁶Ru chloride complexes, at a faster rate than ¹⁰⁶Ru which had been accumulated from ¹⁰⁶Ru nitrosyl-nitrato forms. Also, in the case of the ¹⁰⁶Ru chloride complexes, the loss rate was inversely proportional to the time allowed for isotope accumulation. The process of molting greatly accelerated the loss of ¹⁰⁶Ru from euphausiids, with first molts shed during the loss phase accounting for 70 to 80% of the total ¹⁰⁶Ru body burden. When euphausiids accumulated ¹⁰⁶Ru from the food chain, the initial-loss rate was rapid due to large amounts of the radioisotope associated with fecal pellets; however, no relationship was found between loss rate and the number of food rations received. Molts from these individuals did not contain ¹⁰⁶Ru, thus, loss from euphausiids obtaining this radioisotope through the food chain is mainly due to fecal pellet deposition and other excretion or exchange processes.     

Cod end feeding by the euphausiid Meganyctiphanes norvegica

Daytime surface swarms of Meganyctiphanes norvegica in the Bay of Fundy were sampled by hand-held dip nets and by plankton nets. A comparison between the food baskets of the animals from samples obtained by the two collection methods showed that those from plankton nets contained many items whereas those from dip nets rarely contained anything. Samples from towed-nets also had significantly more animals with fuller stomachs than those obtained by dip nets at approximately the same time. It is concluded that the observed differences are due to feeding by the euphausiids in the cod end of plankton nets.     

Effects of temperature and size on molting of euphausiid crustaceans

The effects of temperature and body size on the intermolt periods (molting frequencies) of the North Pacific euphausíid Euphausia pacifica and the Mediterranean forms of Meganyctiphanes norvegica, Euphausia krohnii, Nematoscelis megalops, and Nyctiphanes couchii were studied under controlled conditions in the laboratory. Mean intermolt periods for E. pacifica and M. norvegica were inversely and linearly related to temperature, over temperature ranges which the euphausiids normally encounter in the sea. At higher temperatures there was a tendency for three size groups of M. norvegica to approach a minimum intermolt period independent of temperature. M. norvegica cycled for different time periods between 13° and 18°C molted regularly at mean frequencies which would be expected if the animals had been held constantly at the timeweighted means of the two experimental temperatures. The increase in mean intermolt period per unit weight was faster in small, fast-growing M. norvegica than in large, slow-growing adults. This relationship was corroborated by following the changes in the intermolt period of an actively growing individual N. couchii over an 11 month period. Neither feeding nor the time of year of collection affected the molting frequency as long as temperature and animal weight were held constant. No tendency was found for euphausiids of the same species and/or size, and from the same collection, to molt on the same night. Molting occurred at night 80 to 90% of the time for all species, over the temperature ranges normally experienced by the euphausiids in the sea, and over all animal weights tested. There appeared to be a weakening of the night-time molting rhythm at low temperatures. Although neither temperature nor anímal weight substantially affected the night-time molting rhythm, both affected the mean intermolt period. Therefore, both temperature and body size apparently act together to adjust the length of the intermolt period of each individual in increments of whole days, but they exert little control over time of molting within any 24h period. No information was obtained regarding the factors which specify night-time molting over daytime molting within any 24 h period; however, regulation of certain hormone activities is probably involved.     

Diel vertical migrations and nocturnal feeding of a dense coastal krill scattering layer (Thysanoessa raschi and Meganyctiphanes norvegica) in stratified surface waters

To study the nocturnal feeding of euphausiids during vertical migrations and its impact on the phytoplankton, a phytoplankton-rich water mass (drogue marked) drifting over a dense krill scattering layer (acoustic 104kHz) in the lower St. Lawrence estuary was monitored over 46 h in July 1982. Phytoplankton >20 m was abundant and mostly concentrated at the bottom of the photic layer above the pycnocline. Less than 42% of the particulate carbon was due to phytoplankton. The krill scattering layer was about 2 to 3 km in width, elongated along the 100-m bathymetric contour, and absent when the bottom was shallower than 50 m. Its upper day depth was 50 m. At deeper depths, its vertical distribution frequently changed from unimodal to polymodal shapes and viceversa, often with large concentrations of zooplankton just above the bottom. Typical vertical migrations were observed on both days. At night the scattering layer had a lower scattering strength. Most of it was below the thermocline but net catches showed that large concentrations of euphausiids (up to 57 individuals m^-3) crossed it. Stomach pigment content of Thysanoessa raschi was generally low, but mean stomach fullness was always high. They were more opportunistic than herbivorous. From stomach fullness and the presence of a food bolus in mouth parts, feeding in surface waters appeared to be intensive, but gut content indicated that food was not processed there. It is therefore suggested that individuals underwent vertical interchanges across the thermocline while feeding during the night. Meganyctiphanes norvegica had significant herbivorous activity during the night. The grazing pressure impact of the scattering layer on phytoplankton was negligible.Contribution to the program of GIROQ (Groupe interuniversitaire de recherches océanographiques du Québec)     

Occurrence of large numbers of carcasses of the large,grazing copepod Calanus cristatus from the Japan Sea

Carcasses of Calanus cristatus were discovered in plankton samples collected from the Japan Sea throughout the year from 1970 to 1985. Many carcasses of copepodite Stages IV and V occurred in the layer between 15 and 300 m below a distinct thermocline. The number of copepodite Stage V carcasses also peaked in the layer between 1 500 and 2 000 m. The highest density of copepodite Stages IV and V carcasses was 169 individuals per 1 000 m³ and 1 573 individuals per 1 000 m³, respectively. Carcasses of adults occurred at depths below 500 m and numbers of males and females per 1 000 m³ were 1 to 16 and 1 to 42, respectively. Living males were larger in catch number than living females, but the relationship for carcasses was the opposite. Weight of carcasses was 15 to 25% of living C. cristatus. Carcasses contained about 51% carbon and 8% nitrogen by weight. Carcasses may have been drifting for more than one year in the epipelagic layer under the thermocline because of their slow decomposition rate.     

Metabolic properties of Northern krill, Meganyctiphanes norvegica, from different climatic zones. II. Enzyme characteristics and activities

Activities and characteristics of two metabolic key enzymes, citrate synthase (CS) and pyruvate kinase (PK), were studied in the Northern krill, Meganyctiphanes norvegica, with respect to adaptive properties under different thermal conditions. Krill were sampled during late winter/spring and summer from the constantly warm Ligurian Sea (12-13°C below the thermocline), the colder but also comparatively constant Clyde Sea (7-8°C), and the variable Kattegat (2-16°C). Both enzymes showed distinct tissue- and organ-specific activities, which were highest in the pleopods - the principal locomotive organs. The fourth and fifth abdominal segments, however, were used for routine investigation due to lowest variability. Specific activity of CS and PK did not differ between seasons in krill from the Kattegat or the Clyde Sea. In the Ligurian Sea, in contrast, specific CS activities were significantly lower during summer. Analysis of individual data illustrated a decrease of CS activity with size and an increase of PK activity with size. Taking these allometric effects into account, as emphasized by calculating the ratio between both enzymes, variation of CS and PK activities turned out to be solely dependent on body size, which differed between locations and seasons. Ligurian krill from the summer, however, were unique in that they showed a lower CS/PK ratio than would be predicted by the scaling effect. Thermal characteristics of each enzyme were similar between locations and seasons. During the winter, in Kattegat and Clyde Sea krill, K_m values (Michaelis-Menten constant) of CS towards acetyl-coenzyme A exhibited an almost constant level over the experimental temperature range of 4-16°C. During summer, however, K_m values were lower at 8°C in the Clyde Sea and at 12°C in the Kattegat. In Ligurian krill from the summer, K_m values were consistently lower than those of winter krill over the entire experimental temperature range. In conclusion, Kattegat and Clyde Sea krill show only minor adaptations to their respective thermal environments in terms of CS and PK characteristics. Ligurian krill, in contrast, exhibited decreased specific CS activity during summer, which might be compensated by elevated enzyme-substrate affinity as indicated by lower K_m values. Since temperature was constant during both seasons, this effect cannot be explained as a reaction to thermal conditions. Consequently, oligotrophic conditions in the Ligurian Sea during summer may entail a reduction in the somatic performance of krill, which is reflected by lower CS activity.     

Population dynamics and production of euphausiids

Seasonal changes in abundance, size and aspects of the population structure of Meganyctiphanes norvegica (M. Sars) and Nyctiphanes couchi (Bell) are described from samples taken with the Continuous Plankton Recorder at 10 m depth over a 2 yr period (1966 and 1967) in the North Atlantic Ocean and the North Sea. M. norvegica lived for a maximum of just over 2 yr, and adults of both year-classes spawned during a limited breeding season in the spring or summer. N. couchi spawned over a prolonged breeding season, giving rise to a complex of cohorts with overlapping size ranges. It was concluded that 3 or 4 cohorts were spawned in each year and that the maximum life span was probably greater than 1 yr, although maturity may be attained in less than a year. Estimated annual production at 10 m depth for M. norvegica ranged from 0.80 to 18.74 mg m^-3yr^-1 and for N. couchi from 0.67 to 8.23 mg m^-3yr^-1. P:B ratios ranged from 1.3:1 to 6.3:1 for M. norvegica and 4.0:1 to 5.5:1 for N. couchi.     

Ovarian development and spawning in relation to the moult cycle in Northern krill, Meganyctiphanes norvegica (Crustacea: Euphausiacea), along a climatic gradient

Adaptive processes linked to reproduction were studied comparatively for three populations of Northern krill, Meganyctiphanes norvegica (M. Sars, 1857), sampled during winter and summer cruises in the Clyde Sea (W Scotland), the Kattegat (E Denmark), and the Ligurian Sea (NW Mediterranean). The aim was to investigate the functional relationship between egg production and moulting under contrasted climatic and environmental conditions. A staging system for female sexual development established for live krill was complemented by a histological study of the ovary at various developmental steps. During the reproductive season, all adult female krill were engaged in cyclical egg production. During experiments, female krill released one batch of mature oocytes in one or two spawning events. The ovary of postspawn female krill still contained developing oocytes for another egg batch. In the non-reproductive period, all female krill had a resting ovary. Ovarian structure and pattern of egg production were identical in the three populations, but seasonal timing of egg production was different. The model proposed for the Ligurian population of the annual cycle of ovarian development can be extended to the other two populations, taking into account the seasonal characteristics of each site. Random field samples were staged simultaneously for moult cycle and for sexual development. Moult stages and the seasonal variation of the intermoult period were studied for the Kattegat population using multi-year data and compared to data obtained during summer/winter cruises in the Clyde and the Ligurian Sea. At the three sites, intermoult period was shorter and temperature-dependent during the reproductive period, concurrent with the season of greatest food availability. During most of the year and the period of sexual rest, moulting activity was reduced. The relationship between spawning and the moult cycle was studied comparatively for the three populations. Eggs were released during the premoult phase of a “spawning moult cycle”, in one or two spawnings associated with apolysis and Moult Stage D1, respectively. Yolk accumulation for the next egg batch was completed during an alternating “vitellogenic moult cycle”. A model for the timing of cyclical egg production in relation to moulting, as proposed for the Kattegat, can be extended to the other populations, taking into account intermoult period variation with temperature. Temperature appeared to be the principal environmental factor controlling growth (through moulting) and egg production during the reproductive season, in connection with favourable trophic conditions. Received: 22 December 1997 / Accepted: 29 August 1998     

Diet of 0-group stages of capelin (<Emphasis Type="Italic">Mallotus villosus</Emphasis>), herring (<Emphasis Type="Italic">Clupea harengus</Emphasis>) and cod (<Emphasis Type="Italic">Gadus morhua</Emphasis>) during spring and summer in the Barents Sea

Recruitment of capelin in the Barents Sea fail when juvenile herring and cod are abundant and the potential for feeding competition of wild sympatric capelin and herring larvae and small cod juveniles were investigated. The frequency of gut evacuation after capture of capelin larvae were also studied in mesocosms. Small capelin larvae (<35 mm length) fed on small prey including phytoplankton, invertebrate eggs and nauplii, bivalves, other invertebrate larvae and small copepods. Calanus copepodites were only observed in large capelin larvae (>26 mm length). Calanus copepodites were the major food sources for contemporary herring larvae (25–35 mm length) and Calanus and euphausiids were the major prey for small juvenile herring (37–60 mm length) and cod (18–40 mm length). Capelin larvae reared in mesocosms evacuated the guts shortly after capture. Capelin larvae had a smaller mouth and fed on smaller prey than herring and cod of the same length. This implies that the small capelin larvae, in contrast to sympatric small herring and cod, are not tightly linked to the food chain involving Calanus and euphausiids. Thus, exploitative competition between capelin larvae and planktivorous fish that rely on Calanus and euphausiids in the Barents Sea may be relaxed.