The physiological response of Northern krill (Meganyctiphanes norvegica) to temperature gradients in the Kattegat

The Alkor-Deep (140 m), which forms part of a depression system in the northern Kattegat channel east of the island of Læsø (Denmark), is the location of a self sustaining population of Northern krill, Meganyctiphanes norvegica (Euphausiacea). This population is exposed to one of the most pronounced thermal gradients within the distributional range of this pelagic crustacean. During summer, the temperature of the water column ranges between 4 and 6 in the deep to 16 °C near the surface which results in the krill being exposed to temperature differences of 8–10 °C during diel vertical migration. Oxygen consumption rates were used to investigate the physiological adaptation of the animal to such gradients in temperature. The rates were found to increase exponentially from 31 mol O₂ h^-1 g_dw ^-1 at 4 °C to 72 mol O₂ h^-1 g_dw ^-1 at 16 °C, giving a Q ₁₀-value of 2.0, and indicating that physiological adaptation to varying thermal conditions does not take place. Behavioural adaptations are discussed which may help the krill to cope with large temperature gradients in their environment.     

Diel vertical migrations ofMeganyctiphanes norvegica in the Kattegat: Comparison of net catches and measurements with Acoustic Doppler Current Profilers

Diel vertical migration of a stable and well-defined population of Nordic krill,Meganyctiphanes norvegica (Crustacea, Euphausiacea) was investigated during eight days in August 1989, in the Läsö-Deep, East of the Danish island Läsö. Net catches with a multi-net (MOCNESS) and measurements with a moored and a shipboard Acoustic Doppler Current Profiler (ADCP) were compared. Backscattered energy as a measure for biomass gave good correlations to the dry weight ofM. norvegica and smaller zooplankton from net catches. Diel migratory patterns matched well, as determined, parallel with both methods. Migratory vertical velocity was determined with ADCP at 2–3 cm sec^?1. The potential for the use of ADCPs for biological investigation is discussed. Vertical migration was dependent on environmental parameters. The krill did not cross a temperature barrier of 14°C, although rich food sources were situated beyond it. Differences in salinity did not play a role. Currents were involved in plankton distribution. Light was an important Zeitgeber (synchronizer) and determined the density of the krill aggregations. Feeding behaviour did not interfere with the light-induced migratory pattern of Nordic krill at the Läsö-Deep.     

Studies on metabolic properties in the Northern Krill, Meganyctiphanes norvegica (Crustacea, euphausiacea): influence of nutrition and season on pyruvate kinase

The specific activity and the kinetic properties of partly purified pyruvate kinase (PK) (EC 2.7.1.40) from the Northern Krill, Meganyctiphanes norvegica, were investigated in relation to varying food resources. In order to evaluate the effect of starvation on the total energy metabolism, the respiration rates of fed and unfed krill were determined. The FPLC-elution profile of PK displayed two distinct peaks - PK I and II. The first isoform represented 80% of the total PK activity in the organism, and 20% was contributed by the second isoform. PK I was inhibited by ATP but was not influenced by fructose-1,6-bisphosphate (FBP). In contrast, PK II showed ATP inhibition and up to 2.5-fold increased activity by addition of 17 micromol.l(-1) FBP. The Michaelis-Menten constants of both isoforms were 2-10-fold higher for ADP than for phosphoenolpyruvate (PEP). Alanine showed no regulatory effect on PK I and II. In specimens starved for 7 days oxygen consumption decreased by 20%. Neither the feeding experiments nor the animals captured in the field during low and high productive seasons indicate that PK properties of M. norvegica are modified in relation to food supply. Accordingly, alternative mechanisms are involved in the depression of the metabolic rate in terms of oxygen consumption.     

Below-halocline oxygen consumption in the Kattegat

Sediment and seston oxygen consumption rates below the sharp halocline in the south-eastern part of the shallow Kattegat were measured and compared to calculated rates of carbon addition through the halocline. The mean rate of decrease in deep-water oxygen concentrations between March and September 1988 was 1.0 ml O₂ M^–3 h^–1. Measurements of benthic oxygen uptake using laboratory-incubated sediment cores from depths 30 m gave a mean value of 7.8 ml O₂ m^–2 h^–1. Below-halocline water (from 20 m, 30 m and 1 m above bottom) incubated in bottles showed oxygen consumption rates varying from 0.5 ml O₂ m ^–3 h^–1 in March to 2.8 ml O₂ M^–3 h^-1 in late August. The sum of benthic and deep-water oxygen consumption was equivalent to a mean oxygen decrease rate of 1.7 ml O₂ m^–3 h^–1 below the halocline. Of the total oxygen consumption below the halocline 65% was due to oxygen up-take in the water and 35% was due to benthic oxygen consumption. The sum of oxygen consumption measured in sediment cores and in bottles corresponds to a carbon utilisation of 80.1 g C m^–2 (respiratory quotient (RQ), assumed 1.0 and 1.4 for water and sediment, respectively), while the decrease in deep-water oxygen concentration was equivalent to 43.0 g C m^–2 (RQ assumed = 1.0). Using published values for the external N loading (including deep-water supply), ¹⁵NO₃-uptake, ¹⁴CO₂-uptake in combination with % ¹⁵NO₃-uptake of total ¹⁵N-uptake (nitrate, ammonia and urea) and a Redfield C/N ratio of 6.6, rates of carbon addition (new or export production) through the halocline were calculated to 31.9, 46.7 and 36.3 g C m^–2, respectively, with a mean value of 38.3 g C m^–2 for the 8 month period March–September. This is somewhat less than the value (50.5 g C m^–2) calculated from a published empirical relationship between total and export production. The fact that the calculated carbon addition through the halocline was appreciably less than the carbon equivalent of the measured below-halocline respiration may be an effect of sediment focusing (horizontal transport of sedimenting material to deeper areas), since the bottom area below the halocline is much smaller than the total area of the Kattegat. A lower observed decrease in the oxygen concentration below the halocline compared to the sum of measured sediment and deep-water oxygen consumption on the other hand indicates oxygen supply to below-halocline waters through advection and/or vertical entrainment.     

On the physical oceanography of the Kattegat and Clyde Sea area, 1996–98, as background to ecophysiological studies on the planktonic crustacean, Meganyctiphanes norvegica (Euphausiacea)

 The euphausiid, Meganyctiphanes norvegica (M. Sars), has an extremely wide distribution around the North Atlantic, from Canada to the Arctic to the western Mediterranean. It appears to be adaptable to life under quite different oceanographic conditions, yet no morphological features have been identified that distinguish any of the populations occurring in different areas between which exchange must be supposed to be restricted. The genetic, physiological, behavioural and ecological investigations that are being carried out on M. norvegica in the Kattegat, the Clyde Sea area and the Ligurian Sea (the EU MAST III PEP Project) in order to study its adaptability must be closely related to the physical environment within which the respective populations occur. This paper presents a summary of the hydrographic conditions found in the Kattegat and the Clyde Sea area in summer and in winter, during each research cruise, and places them in the context of current understanding of the hydrography in each area. An account of the hydrography of the Ligurian Sea is in preparation (Dallot et al., in preparation). The data will be used in the further analysis of genetic, physiological and behavioural adaptability of M. norvegica, and will be available for use as appropriate to supplement other hydrographic studies in the regions concerned. Received: 2 February 1999 / Accepted: 6 April 1999     

Studies on the biochemistry and fine structure of silica-shell formation in diatoms

Summary The distribution of radioactivity in ethanol-water-soluble compounds after short periods of photosynthetic incorporation of ¹⁴CO₂ is consistent with the operation of the photosynthetic carbon reduction (PCR) cycle in the fresh water diatom Navicula pelliculosa. Incorporation of ¹⁴CO₂ for extended time periods established the presence of the intermediates of the PCR and tricarboxylic acid (TCA) cycles, amino acids, and organic acids; free sugars were not observed. The main labelled soluble carbohydrate was a glucan. Hydrolysis of the radioactive insoluble material indicated the presence of carbohydrates containing several distinct sugars, and proteins with the usual amino-acid composition. During silicon starvation of exponentially growing cultures, rates of incorporation of both ³²P _i and ¹⁴CO₂ decreased. Incorporation into the lipid increased, with a corresponding decrease into protein and carbohydrate. Reintroduction of Si to staryed cells led to an increased rate of incorporation of both isotopes, and transient changes in the radioactivity in most metabolic intermediates investigated. After 30 min the radioactivity in all PCR cycle intermediates, except phosphoglyceric acid, had increased by about 300%. The radioactivity of citrate and -keto-glutarate increased, whereas that of other TCA-cycle intermediates decreased. An initial decrease in the levels of glutamate, aspartate and glutamine was apparently reversed by cleavage of glutamate-aspartate peptides, as radioactivity of other amino acids increased. Incorporation into the soluble glucan and into protein increased markedly although the rate of incorporation into insoluble carbohydrates remained constant.     

Sediment toxicity in the Kattegat and Skagerrak

Sediments were sampled from 62 sites in the Kattegat and Skagerrak, which are located between the Baltic and the North Sea in the Western Atlantic, during autumn 1989 and spring 1990. From each site 5 to 6 samples were taken wit ha box-corer. After mixing to composite samples on board, transport and storage (at 4 °C for 2 to 4 weeks), the samples were tested for toxicity to Daphnia magna and Nitocra spinipes. Immobility in Daphnia after exposure to 16 percent sediment (wet wt) in reconstituted standardized water (ISO, 1982) ranged from 0 to 88 percent after 24 h and from 3 to 95 percent after 48 h. For Nitocra the toxicity, determined as the 96-h LC50 (% wet wt) at 7 salinity, ranged from > > 32 percent (nontoxic) to 1.8 percent (most toxic). All exposures were made in duplicates and the effects obtained in the duplicates with the same sediment were correlated to each other. However, sediment toxicity to Daphnia and Nitocra was not. The test with Nitocra, which was made at several concentrations of sediment, was considered to give the most reliable picture of sediment toxicity in the Kattegat and Skagerrak. This ambient toxicity assessment identified three areas with toxic sediment, (1) the Göta älv estuary (outside the city of Göteborg) and its surroundings, (2) the Bay of Laholm in southern Kattegat, which is an area with periodic oxygen depletion and where repeated mussel kills have occurred during the last decade, and (3) an area in the open Skagerrak northwest of Skagen (the tip of the Jutland peninsula). Sediments, which had been stored at 4 °C, were tested again after 6 to 13 mos with the Nitocra test. Stored sediment toxicity was poorly correlated with fresh sediment toxicity. The average detoxification during storage was 5 times, but the range was 3 orders of magnitude, from 17 times more toxic to 73 times less toxic. The reasons for the observed areal and storage differences in sediment toxicity are so far not understood.     

Variation of lipid classes among organs of the Northern krill Meganyctiphanes norvegica, with respect to reproduction.

Lipid content and class in the Northern krill Meganyctiphanes norvegica (digestive gland, stomach, gonad, fat body, abdomen) was investigated and correlated with sex and reproductive stage. Ready to spawn females, have high lipid content in ovaries, while in males and spent females, the major site of lipid deposits was the digestive gland, followed by the fat body. These differences among spawning and spent females are indicative of strong interactions between the ovary and digestive gland and the ovary and fat body during vitellogenesis. Triacylglycerols (TAG) were the major neutral lipid class with high levels in the digestive gland. The major phospholipid was phosphatidylcholine (PC) particularly in the muscular tissue of the abdomen. Phosphatidyl-ethanolamine (PE) and -serine-inositol (PS-PI), were present at intermediate levels. Reproductive males were depleted in TAG and diacylglycerols (DAG) in the digestive gland, gonad and fat body, and had 4 times lower cholesterol in the gonad than ready to spawn females. Furthermore, ready to spawn females had in the ovary higher amounts of TAG, DAG and phospholipids (PC, PE, PS-PI) than spent females. Linear relationships between lipid content and main lipid class (TAG, PC, PE, PS-PI) in different fractions of males and ready to spawn females showed that: (1). TAG was stored for both sexes in all cephalothorax fractions with highest values in the digestive gland and ovary fluid; (2). PC was accumulated for both sexes in the fat body and the gonad with a higher slope for females, with the highest values in the ovary fluid and in the abdomen of males and that (3). PS-PI was stored only in the ovary and abdomen of mature females. These results are discussed in terms of the strategy developed by Meganyctiphanes norvegica to allocate lipids to the next generation for optimised embryogenesis.     

Studies on the biochemistry and fine structure of silica shell formation in diatoms

Summary Cell division in Navicula pelliculosa (Bréb.) Hilse, strain 668 was synchronized with an alternating regime of 5 h light and 7 h dark. Cell volume and dry weight increased only during the light period. DNA synthesis, which began during the third h of light, was followed sequentially by mitosis, cytokinesis, silicic acid uptake, cell wall formation, and cell separation. Silicification and a small amount of net synthesis of DNA, RNA and protein occurred during the dark at the expense of carbohydrate reserves accumulated during the light period. Cells kept in continuous light, after synchronization with the light-dark regime, remained synchronized through a second division cycle; the sequence of morphological events was the same as that in the light-dark division cycle, but the biosynthesis of macromolecular components changed from a stepwise to a linear pattern. The silicon-starvation synchrony was improved by depriving light-dark synchronized cells of silicic acid at the beginning of their division cycle, then resupplying silicic acid to cells blocked at wall formation.Abbreviation L light - D dark Portions based on a thesis submitted by W.M.D. to the University of California, San Diego in partial fulfillment of the requirements for the PH.D degree     

Studies on the biochemistry and fine structure of silica shell formation in diatoms

Summary The fine structure and pigmentation of an apochlorotic diatom isolated from decaying Macrocystis pyrifera is described. The morphology of acid cleaned shells suggests that the isolate is Nitzschia alba Lewin and Lewin. Light microscope observations indicated a centrally located nucleus and numerous highly refractile bodies which stained differentially with Nile blue and Sudan black B. The stained globules could be correlated in thin-sectioned profiles with either electron dense or lucent areas depending on the fixation technique. In the electron microscope the nucleus, Golgi complex, and mitochondria were similar in appearance to those described for other diatoms. Proplastid-like organelles, delimited by a double membrane, and containing small vesicles were also observed. Neither carotenoids nor chlorophylls could be detected by spectroscopic or spectrofluorometric analysis in vivo or in organic solvent extracts. Deposition of new walls was initiated by formation of silicon deposition-vesicles in the central region of dividing cells. The acentric raphes were deposited last. The genesis and interrelationship of the old plasmalemma, silicalemma, and newly formed plasmalemma are discussed.     

Studies on the biochemistry and fine structure of silica shell formation in diatoms

Summary The morphogenesis of the different types of labiate processes is compared among three species of centric diatoms [Stephanopyxis turns (Greville) Ralfs,Odontella sinensis (Greville) Grunow, andOdontella aurita (Lyngbye) Agardh]. In all species, a cytoplasmic structure,i.e., the labiate process apparatus, situated close to the developing labiate process, appears before the labiate process begins to form and disappears upon its maturation. The possibility that the labiate process apparatus is implicated in the phylogeny of the labiate process is discussed.     

Studies on the biochemistry and fine structure of silica shell formation in diatoms

In diatoms, the siliceous cell walls are enveloped by an organic component which includes 4-hydroxyproline and 3,4-dihydroxy-L-proline. The formation of these two amino acids were studied in Nitzschia angularis in Si-starvation synchrony. Both appear to arise from peptidyl proline. Its conversion to peptidyl hydroxyproline was shown in cell-free extracts and in kinetic studies using [¹⁴C]proline. Two lines of evidence indicate that dihydroxyproline does not arise from the further hydroxylation of peptidyl hydroxyproline: First, there was a lag of several minutes between the incorporation of [¹⁴C]proline into protein and the appearance therein of [¹⁴C]hydroxyproline but no such lag for the appearance of dihydroxyproline. Second, ,-dipyridyl blocked the formation of hydroxyproline, but not of dihydroxypyroline, from peptidyl proline. Cell walls made in the presence of dipyridyl differed little in overall chemical composition from walls made in its absence and were morphologically identical. [¹⁴C]dehydroproline was rapidly metabolized in the cells, with [¹⁴C]dihydroxyproline a prominent product. Studies of the conversion of [¹⁴C]proline to [¹⁴C]hydroxyproline at various stages of wall formation showed an increased synthesis of [¹⁴C]dihydroxyproline at the end of cell separation.     

Studies on the biochemistry and fine structure of silica shell formation in diatoms

Summary The sequential wall formation in the centric diatom,Ditylum brightwellii (West) Grunow, is described. The silica deposition vesicle is formed by the coalescence of small vesicles. Silicification of the new valve starts from the central labiate process area prior to the completion of cytokinesis, and the developing valve grows in a centrifugal direction. The initiation of the structures on the valve follows the sequence: labiate process, marginal ridge, and rota. A novel labiate process apparatus, which is situated in the cytoplasm close to the developing labiate process, appears prior to the initiation of the labiate process and disappears upon its maturation. Segments of the girdle bands are formed in individual silica deposition vesicles after the valve matures and is exocytosed. Three morphological forms of deposited silica have been determined: thin base layers, microfibrils, and hexagonal columns. The involvement of cytoplasmic structures in the patterning of the siliceous wall is discussed.     

The life-cycles of four amphipod species in the Kattegat

Populations of four species of Gammaridae (Crustacea, Amphipoda) were sampled over 15 months in the south-western Kattegat at Sangstrup Klint (Denmark). Chaetogammarus marinus, C. stoerensis and Gammarus oceanicus had annual life-cycles, while G. salinus most likely had two generations per year. Mature C. marinus females were found from March to July, mature C. stoerensis females from March to June, mature G. oceanicus females from December to early June and mature G. salinus females from late November to late July. The theoretical maximum number of clutches were estimated at five for C. marinus , five plus for C. stoerensis , and six for G. oceanicus and both winter and summer reproducing G. salinus . Spring was the period of main recruitment in all populations, with additional recruitment during summer in G. salinus .
The geographical patterns of the reproductive strategies are discussed in relation to environmental factors and interspecific competition.