Ecological investigations on the zooplankton community in Balsfjorden,Northern Norway: seasonal changes in the lipid class composition of Meganyctiphanes norvegica (M. Sars), Thysanoessa raschii (M. Sars), and T. inermis (Krøyer)

The lipid class composition of three krill species, Meganyctiphanes norvegica (M. Sars), Thysanoessa raschii (M. Sars), and T. inermis (Krøyer), was determined throughout the year in Balsfjorden, Northern Norway. Meganyctiphanes norvegica had triglycerides as its major neutral lipid, Thysanoessa raschii had mainly triglycerides with small amounts of wax esters, whereas T. inermis had mainly wax esters with lesser amounts of triglycerides. Neutral lipids were generally accumulated by the krill during spring-early summer, maintained at high levels through to mid-winter and then declined to minimal levels during early spring. Differences in rates of deposition and mobilization of neutral lipid between males and females occurred in Meganyctiphanes norvegica and Thysanoessa raschii but not in T. inermis. T. raschii accumulated lipid somewhat later in the year than T. inermis but earlier than Meganyctiphanes norvegica. A major rôle for lipid in the elaboration of gonads is indicated. The accumulation of either wax esters or triglycerides by euphausiids is considered in terms of the feeding habits and ecological niche differentiation of the species.     

Lipolysis post mortem in North Atlantic krill

• 1.1. Three common species of North Atlantic krill, Meganyctiphanes norvegica (M. Sars), Thysanoessa inermis (Krøyer) and T. raschii (M. Sars), have been stored at 0°C post mortem, and the lipolytic activity followed by measuring changes in the lipid composition during storage.
• 2.2. Both phosphoglycerides and triacylglycerols were subjected to extensive hydrolysis with the formation of free fatty acids in all krill species examined, whereas wax esters, constituting a considerable proportion of the lipids in the Thysanoessa species, were not hydrolysed at all.
• 3.3. In M. norvegica the triacylglycerols and phosphoglycerides were hydrolysed at similar rates, whereas in T. inermis and T. raschii the phosphoglycerides were hydrolysed most rapidly.
• 4.4. For all krill species examined, the rate of production of free fatty acids was nearly constant during the initial phase of storage, and subsequently declined on prolonged storage.
• 5.5. At the end of the storage period of 16–24 days, the free fatty acids constituted about 35% of the total lipid in M. norvegica, and about 50% in the Thysanoessa species.
• 6.6. The rate of production of free fatty acids was about the same in all the three species of krill and seemed to be independent of the total lipid content.

     

Lipid composition and trophic relationships of krill species in a high Arctic fjord

Our study deals with the lipid biochemistry of the krill community in the ecosystem of the high Arctic Kongsfjord (Svalbard). During the last decades, Kongsfjord experienced a change in krill species composition due to recent increased advection of Atlantic water masses carrying characteristic boreal as well as subtropical-boreal euphausiids into the ecosystem. The lipid biochemistry and trophic relationships of the species recently inhabiting the Arctic water masses are scarcely known, although a change in a krill population may have a significant impact on the ecosystem. A comparison of nutrition and energy storage strategies, stable isotopes, lipid profiles and fatty acid compositions showed remarkable differences between the krill species. These reflected the diverse feeding behaviours and specific adaptations to the environments of their origin: the boreal Meganyctiphanes norvegica and subtropical Nematoscelis megalops appear more carnivorous and have significantly lower mean lipid contents (29 and 10 %, respectively) and a different energy storage pattern (triacylglycerols and polar lipids, respectively) than the arcto-boreal Thysanoessa inermis, which consists of up to 54 % of lipids mainly stored as wax esters (>40 %). These differences may have significant implications for the rapidly changing marine food web of Kongsfjord—especially for higher trophic levels relying on the nutritional input of animal lipids.     

Understanding the trophic role of the Antarctic ctenophore,Callianira antarctica,using lipid biomarkers

To better understand the trophic role of ctenophores in Antarctica during austral fall and winter, a major species of cydippid ctenophore, Callianira antarctica, was collected during April/May (fall) and August/September (winter) 2002 in the vicinity of Marguerite Bay. Lipid content, lipid classes, fatty acids, fatty alcohols and sterols were analyzed in animals, together with lipid biomarkers in krill and copepod species representing potential ctenophore prey. Lipid content in ctenophores collected in winter was slightly higher than from animals in fall (4.8 and 3.5% of dry weight, respectively). Polar lipids were the dominant lipid class in ctenophores, accounting for over half of the lipid content, with significant amounts of free fatty alcohols (more than 10% of total lipid content) detected. Lipid-class composition, however, differed significantly between seasons, with significant amounts of neutral lipid (wax esters and triacylglycerols) only detected in animals from fall. Although the dominant lipid classes in ctenophores varied between fall and winter, individual lipids (i.e., fatty acids, alcohols and sterols) showed only minor changes between seasons. Specifically, long-chain polyunsaturated fatty acids [20:5(n-3) and 22:6(n-3)] found in high abundance in larval krill were also elevated in ctenophores collected in winter. Very high amounts of monounsaturated fatty alcohols, particularly 20:1(n-9) and 22:1(n-11), known to be important components of wax esters in calanoid copepods, were also observed. Multivariate analysis using the suite of lipids found indicated that copepods are an important diet item for ctenophores in the study area. Results further suggest that C. antarctica feed actively year-round, with larval krill providing a food resource during austral winter.     

Nature,origin and possible roles of lipid deposits in Maurolicus muelleri (Gmelin) and Benthosema glaciale (Reinhart) from Ullsfjorden,northern Norway

Summary The lipid-rich pelagic teleost Maurolicus muelleri has large lipid depots located subcutaneously, intramuscularly and around the digestive tract. The lipid is contained within conventional adipocytes and is composed largely of triacylglycerols rich in 20:1 (n-9) and lipid-rich mesopelagic teleost Benthosema glaciale, except that the lipid is predominantly wax esters whose fatty alcohols and fatty acids are both rich in 20:1 (n-9) and 22:1 (n-11) moieties. An origin for the lipids of both species in the wax esters of calanoid copepods is indicated. The anatomical distribution of the lipids in these teleosts and their intracellular location point to their being fundamentally an energy store. The twin roles of neutral lipids in providing metabolic energy and buoyancy are discussed.     

The lipid composition of two species of serrasalmid fish in relation to dietary polyunsaturated fatty acids

The lipid composition of two species of Serrasalmid fish with different natural feeding habits were compared in relation to the polyunsaturated fatty acids (PUFA) supplied in their diets. Mylossoma aureum , a herbivorous piranha, was maintained on oatmeal flakes in which : 2(n-6) and : 3(n-3) were the only PUFA and accounted for 40–8 and 1.2%, respectively of dietary fatty acids. Serrasalmus nattereri , the carnivorous red piranha, was fed mosquito larvae containing .0-33.4% of their total fatty acids as : 2(n-6)+18 : 3(n-3) and 4.9-8.5% as 20 : 4(n-6)+20 : 5(n-3). The two species had similar lipid class compositions in liver, brain, viscera and carcass, except that lipids from M. aureum were generally richer in triacylglycerols. In both species, visceral and carcass lipid contained high levels of triacylglycerols whose principal PUFA was : 2(n-6). In M. aureum the major PUFA in liver total lipid and triacylglycerols was : 2(n-6) whilst the major PUFA in liver phospholipids were : 4(n-6) and : 5(n-6), with : 6(n-3) being a minor component. The level of : 6(n-3) in ethanolamine glycerophospholipids was significantly greater in brain than liver of M. aureum. Although absent from dietary lipid, : 6(n-3) was the major PUFA in phosphatidylcholine and ethanolamine glycerophospholipids from both the liver and brain of S, nattereri . In both species, the ratio of (n-6)/(n-3)PUFA was consistently lower in tissue lipids than in dietary lipids. The results are consistent with (i) the herbivorous M. aureum converting dietary C18 PUFA to their C20 and C22 homologues, (ii) the carnivorous S, nattereri forming : 6(n-3) from either 18:3(n-3) or 20: 5(n-3) and (iii) both species selectively desaturating and elongating (n-3) rather than (n-6) PUFA.     

Ontogenetic change in the lipid and fatty acid composition of scleractinian coral larvae

Some scleractinian coral larvae have an extraordinary capacity to delay metamorphosis, and this is reflected in the large geographic range of many species. Coral eggs typically contain a high proportion of wax esters, which have been hypothesized to provide a source of energy for long-distance dispersal. To better understand the role of lipids in the dispersal of broadcast spawning coral larvae, ontogenetic changes in the lipid and fatty acid composition of Goniastrea retiformis were measured from the eggs until larvae were 30 days old. Egg biomass was 78.8 ± 0.5% lipids, 86.3 ± 0.2% of which were wax esters, 9.3 ± 0.0% polar lipids, 4.1 ± 0.2% sterols, and 0.3 ± 0.1% triacylglycerols. The biomass of wax esters declined significantly through time, while polar lipids, sterols and triacylglycerols remained relatively constant, suggesting that wax esters are the prime source of energy for development. The most prevalent fatty acid in the eggs was palmitic acid, a marker of the dinoflagellate Symbiodinium, highlighting the importance of symbiosis in coral reproductive ecology. The proportion of polyunsaturated fatty acids declined through time, suggesting that they are essential for larval development. Interestingly, triacylglycerols are only abundant in the propagules that contain Symbiodinium, suggesting important differences in the energetic of dispersal among species with vertical and horizontal transmission of symbionts.     

Functional biodiversity of lipids in Antarctic zooplankton: Calanoides acutus, Calanus propinquus, Thysanoessa macrura and Euphausia crystallorophias

Zooplankton samples were collected in January 1993 off Dronning Maud Land along a transect from open waters to the marginal ice zone close to the Antarctic ice shelf. Thysanoessa macrura was caught in open waters while Calanoides acutus and Calanus propinquus were mainly sampled between ice floes in the marginal ice zone. The “ice-krill”Euphausia crystallorophias was found over the shelf directly associated with ice floes. T. macrura had a lipid content up to 36% of its dry weight with the dominant lipid class, wax ester, accounting for 45–50% of the total lipid. The predominance of 18:1 fatty alcohols is the striking characteristic of the wax esters. Small specimens of E. crystallorophias had lipid levels up to 26% of their dry weight with, unexpectedly, triacylglycerols being the dominant lipid (up to 41% of total lipid). The small levels of wax esters in these animals (3–6% of total lipid) had phytol as a major constituent. Large specimens of E. crystallorophias had up to 34% of their dry weight as lipid, with wax esters (47% of total lipid) dominated by 16:0 and 14:0 fatty alcohols as the major lipid. Calanus propinquus had lipid levels of up to 34% of their dry weight, with triacylglycerols (up to 63% of total lipid) being the dominant lipid. High levels of 22:1 (n-9) fatty acid were present in the triacylglycerols. Calanoides acutus had lipid levels up to 35% of the dry weight with wax esters accounting for up to 83% of total lipid. High levels of (n-3) polyunsaturated fatty acids were recorded with 20:5(n-3), 22:6(n-3) and 18:4(n-3) being the dominant moieties. On the basis of their lipid compositions we deduce that: (1) Calanoides acutus is the strictest herbivore among the four species studied, heavily utilizing the typical spring bloom; (2) T. macrura is essentially omnivorous, probably utilizing the less defined bloom situations found in oceanic waters; (3) E. crystallorophias is an omnivore well adapted to utilize both a bloom situation and to feed on ice algae and micro-zooplankton associated with the ice; (4) Calanus propinquus seems to be the most opportunistic feeder of the four species studied, probably grazing heavily on phytoplankton during a bloom and, during the rest of the year, feeding on whatever material is available, including particulates, flagellates and other ice-associated algae. We conclude that the different biochemical pathways generating large oil reserves of different compositions, enabling species to utilize different ecological niches, are major determinants of biodiversity in polar zooplankton. Accepted: 22 June 1998     

Main lipid classes in some species of deep-sea corals in the Newfoundland and Labrador region (Northwest Atlantic Ocean)

Corals contain large quantities of lipids in their tissues; these lipids may be either structural or for storage. Little information is available about the lipid content of deep-sea corals, as well as ratios of main lipid classes. In this study, lipid percentages of 81 deep-sea specimens were measured and the presence of six major classes, including sterols (STEROLS), free fatty acids (FFA), triacylglycerols (TG), monoalkyldiacyl glycerol (MADAG), wax (WAX), and sterol esters (SE), was assessed. Deep-sea corals had fewer lipids than their shallow water counterparts. Decision-tree analysis revealed a link between coral groups and total lipid percentages, showing that species within the same group were characterized by similar lipid amounts. Depth did not seem to impact the total lipid percentages, suggesting that deep-sea corals adapt to the differential access to food by changing the proportion of lipid classes while maintaining equivalent lipid levels. In deep-sea species, similar to their shallow water counterparts, energy seems to be stored as neutral lipids (wax esters and triacylglycerols), with the notable difference that a high proportion of MADAG is present. These compounds are less rich in energy than TG. Depth trends were found for FFA, TG and SE with an increase in percentages after 800 m suggesting a potential need for storage due to decreased food availability. A subsequent decrease after 1,100 m was observed for FFA and TG but a more detailed investigation is warranted as the number of specimens acquired from these depths was less than 20. It is nonetheless a surprising result as increased storage is expected when food sources are sparse.     

The scheduling of spawning with the moult cycle in Northern krill (Crustacea: Euphausiacea): a strategy for allocating lipids to reproduction

Summary

Euphausiids moult and grow throughout their life, which implies sharing of resources between growth and reproduction for adult krill. In the Northern krill, Meganyctiphanes norvegica (M. Sars), female krill produce eggs cyclically. Spawning moult cycles alternate with vitellogenic moult cycles for lipid yolk accumulation. Histology shows that lipids are associated with the R cells of the digestive gland in both sexes, with the yolk platelets of mature oocytes and with the fat body cell membranes and blood lacunae in reproducing females. Mature female krill can have a total lipid content twice as high as males, mostly due to accumulation in the ovary, the fat body and the haemolymph. In contrast, in males, as well as in non-reproducing females, the highest percentage of lipids is found in the digestive gland and the haemolymph. In Meganyctiphanes norvegica, the most abundant lipid fractions are polar lipids and triglycerides, the latter being relatively low in reproducing female gonad and fat body. Triglycerides are believed to be a pure energy source and polar lipids are essential for membrane development in embryos. The fatty acid content and composition of the triglyceride and polar lipid fractions in females are different from males, related to both reproductive and dietary processes. Higher levels of polyunsaturated fatty acids (PUFA) in the polar lipid fraction were found in reproductive females. During the non-reproductive season, the converse was found, indicating the specific role PUFA and other fatty acids play in growth and egg production. Adaptive processes linked to reproduction were studied comparatively in three populations of the Northern krill—Clyde Sea (W, Scotland), Kattegat (E, Denmark), Ligurian Sea (Mediterranean)—all differing considerably in climatic and trophic conditions. Such adjustments in lipid synthesis and storage are viewed as reproductive strategies developed by the Northern krill in response to different environmental conditions.     

Chemical composition of North Atlantic krill

• 1.1. The main chemical components of Meganyctiphanes norvegica (M. Sars), Thysanoessa inermis (Krøyer) and T. raschii (M. Sars) have been examined.
• 2.2. Protein accounted for 42–47% of the dry weight of M. norvegica and 32–50% of the dry weight of the Thysanoessa species. On a wet weight basis, the protein content was relatively constant and independent of season.
• 3.3. The dominating amino acids in the bulk protein of the krill were glutamic acid/glutamine, aspartic acid/asparagine, glycine, alanine, lysine and leucine.
• 4.4. Lipids were present in amounts of 13–29% of the dry weight in M. norvegica, 15–50% in T. inermis and 12–44% in T. raschii, and the lipid content varied with season.
• 5.5. The main nitrogen extractives in krill, expressed on a dry weight basis, were free amino acids (5–10%), trimethylamine oxide (about 4%), peptides (about 1%) and nucleotides (0.4–1.3%). Trimethylamine and ammonia were present in very low concentrations in living krill.
• 6.6. The amino acids taurine, glycine, proline, arginine, sarcosine and alanine made up 89–93 mol% of the free amino acid pool.
• 7.7. The ash content of krill was in the order of 10–13% of the dry weight, and fluoride represented 1040 and 3200 ppm in the Thysanoessa species and M. norvegioca, respectively.

     

Fatty acid composition of the lipids of Puffinus tenuirostris (Temminck) in relation to its diet

The fatty acid composition of lipids isolated from the depot fat, stomach contents, and proventricular oil of adult and chick Puffinus tenuirostris (Temminck) has been analysed. The diet of both adults and chicks is almost exclusively derived from the euphausiid Nyctiphanes australis Sars, and an attempt was made to determine whether dietary lipid affects the composition of depot fat, and whether individual fatty acids in the stomachs and proventricular oil can be used as markers for the origin of the diet. An apparent selectivity in the deposition of fatty acids in the fat depots can be explained by the conversion of fatty alcohols, derived from the euphausiid wax ester, into fatty acids of equivalent chain length and unsaturation. Hexadecadienoic acid appeared to be the only possible marker fatty acid from the euphausiid, but wide variations in its level limits its usefulness as a reliable index of the diet of Puffinus tenuirostris.     

Lipid composition of positively buoyant eggs of reef building corals

Lipid composition of the eggs of three reef building corals, Acropora millepora, A. tenuis and Montipora digitata, were determined. Sixty to 70% of the egg dry weight was lipid, which consisted of wax esters (69.5–81.8%), triacylglycerols (1.1–8.4%) and polar lipids c/mainly phospholipids (11.9–13.2%). Montipora digitata also contained some polar lipids typical of the thylakoid membrane in chloroplasts, probably due to the presence of symbiotic zooxanthellae in the eggs. The wax esters appeared to be the major contributor to positive buoyancy of the eggs, and specific gravity of wax esters in A. millepora was estimated to be 0.92. Among the fatty acids of the wax esters, 34.9–51.3% was hexadecanoic acid (16:0) while the major fatty acids in polar lipids were octadecenoic acid (18:1), hexadecanoic acid (16:0), eicosapentaenoic acid (20:5) and eicosatetraenoic acid (20:4). The wax ester appears to be the main component of the 4.5 6.0 m diameter lipid droplets which fill most of the central mass of the coral eggs.     

Aliphatic Chains of Esterified Lipids in Isolated Eyespots of Euglena gracilis var. bacillaris

Isolated eyespot granules of Euglena gracilis Klebs var. bacillaris Pringsheim contained approximately 6% lipids (based on protein). Separation of the lipid extracts by thin layer chromatography revealed four major fractions: wax esters, triacylglycerols, free fatty acids, and phospholipids. Methanolysis of each fraction yielded between 27 and 29 different fatty acids ranging from 12:0 to 22:6. Acetates of the fatty alcohols of the wax fraction consisted of 11:0 to 18:0 carbon chains, with 14:0 being the major component; unsaturated alcohols were not detected.     

Distribution of arachidonic and eicosapentaenoic acids in the lipids of mosses.

Lipid classes from four species of mosses, Mnium cuspidatum, and Mnium medium from Minnesota, and Hylocomium splendens and Pleurozium schreberi from Alaska, were analyzed. The total lipids of all species contained 30-40% arachidonic and eicosapentaenoic acids. However, the lipids from the Alaskan mosses contained about 75% neutral lipids (triacylglycerols, steryl esters and wax esters) whereas the lipids of the other species contained only 20% or less of these neutral lipids. Consistently, monogalactosyldiacylglycerols and phosphatidylethanol-amines were enriched in arachidonic acid and the galactolipids in eicosapentaenoic acid. The distribution of these acids in the phospholipids shows some preference for position 2. Together, the highly unsaturated C20 acids represented 80% of acyl groups in steryl esters. In triacylglycerols they were at average levels, while they were much less in sulfolipids and phosphatidylglycerols. Wax esters contained very little of the highly unsaturated acids but appreciable amounts of phytol and phytenic acid were found as wax constituents.     

Lipid and fatty acid composition of the fat body during the female reproductive cycle of Labidura riparia (Insecta Dermaptera)

During the reproductive cycle of the female Labidura riparia, cytological observations show cyclical modifications of lipid droplets in the periovarian adipocyte. Fat body lipids and their constitutive fatty acids are analyzed. The lipids are predominantly triacylglycerols, which increase after adult ecdysis during vitellogenic and non-vitellogenic periods. Small amounts of diacylglycerols and phospholipids are found. Diacylglycerols increase during vitellogenesis and decrease during the non-vitellogenic period. Cytological modifications of lipid droplets are probably related to diacylglycerol fluctuations. Gas-liquid chromatography of fatty acid methyl esters shows oleic acid to be the predominant fatty acid in total lipids and triacylglycerols; unsaturated acids are approximately twice as abundant as saturated acids all along the reproductive cycle. Fatty acid composition of diacylglycerols and phospholipids differs from triacylglycerols and total lipids composition. Palmitic, stearic, oleic and linoleic acids represent the major fatty acids; their relative amounts vary during the different periods of the reproductive cycle. The correlations between fat body lipid changes and ovarian development were discussed and compared with observations made on other insect species. Accepted: 23 April 1997     

Macrophage polarization state affects lipid composition and the channeling of exogenous fatty acids into endogenous lipid pools

Adipose-tissue-resident macrophages (ATMs) maintain metabolic homeostasis but also contribute to obesity-induced adipose tissue inflammation and metabolic dysfunction. Central to these contrasting effects of ATMs on metabolic homeostasis is the interaction of macrophages with fatty acids. Fatty acid levels are increased within adipose tissue in various pathological and physiological conditions, but appear to initiate inflammatory responses only upon interaction with particular macrophage subsets within obese adipose tissue. The molecular basis underlying these divergent outcomes is likely due to phenotypic differences between ATM subsets, although how macrophage polarization state influences the metabolism of exogenous fatty acids is relatively unknown. Herein, using stable isotope-labeled and nonlabeled fatty acids in combination with mass spectrometry lipidomics, we show marked differences in the utilization of exogenous fatty acids within inflammatory macrophages (M1 macrophages) and macrophages involved in tissue homeostasis (M2 macrophages). Specifically, the accumulation of exogenous fatty acids within triacylglycerols and cholesterol esters is significantly higher in M1 macrophages, while there is an increased enrichment of exogenous fatty acids within glycerophospholipids, ether lipids, and sphingolipids in M2 macrophages. Finally, we show that functionally distinct ATM populations in vivo have distinct lipid compositions. Collectively, this study identifies new aspects of the metabolic reprogramming that occur in distinct macrophage polarization states. The channeling of exogenous fatty acids into particular lipid synthetic pathways may contribute to the sensitivity/resistance of macrophage subsets to the inflammatory effects of increased environmental fatty acid levels.     

Lipid dynamics and feeding of dominant Antarctic calanoid copepods in the eastern Weddell Sea in December

Lipid content, fatty acid composition, and feeding activity of the dominant Antarctic copepods, Calanoides acutus, Calanus propinquus, and Metridia gerlachei, were studied at a quasi-permanent station in the eastern Weddell Sea in December 2003. During 3 weeks of the spring phytoplankton development, total lipid levels of females and copepodite stages V (CVs) of C. acutus were almost doubled. Meanwhile, only a slight increase in total lipid content occurred in M. gerlachei, and no clear trend was observed in lipids of C. propinquus females. The pronounced increase of lipids in C. acutus was due to an accumulation of wax esters. The proportion of wax esters in the lipids of M. gerlachei was clearly lower, while triacylglycerols played a more important role. In C. propinquus, triacylglycerols were the only neutral lipid class. There were no pronounced changes in the feeding activity of M. gerlachei, whereas the feeding activity of C. acutus had rapidly increased with the development of the phytoplankton bloom in December, which explains its rapid lipid accumulation. The combination of gut content and fatty acid trophic marker analyses showed that C. acutus was feeding predominantly on diatoms. The typical diatom fatty acid marker, 16:1(n-7), slightly decreased and the tracer for flagellates, 18:4(n-3), increased in females and CVs of C. acutus. This shift indicates the time, when the significance of flagellates started to increase. The three copepod species exhibited different patterns of lipid accumulation in relation to their trophic niches and different duration of their active phases. The investigations filled a crucial data gap in the seasonal lipid dynamics of dominant calanoid copepods in the Weddell Sea in December and support earlier hypotheses on their energetic adaptations and life cycle strategies.     

Density- and sound speed contrasts in sub-Arctic zooplankton

Summary The sound speed was determined for Meganyctiphanes norvegica, for a mixture of Thysanoessa raschii and Thysanoessa inermis and for a mixture of Calanus finmarchicus and Calanus hyperboreus. The sound speed contrasts ranged from 1.014 to 1.044. Seasonal variations in specific density were measured for Thysanoessa inermis, Thysanoessa raschii, Meganyctiphanes norvegica, Calanus finmarchicus and Calanus hyperboreus. The density of 20 mm T. inermis was lowest in November (1.052 g/cm³) and highest in February–March (1.065 g/cm³). For a 20 mm T. raschii the minimal density was determined in December (1.059 g/cm³) and the maximum in February–March (1.074 g/cm³). M. norvegica individuals of 35 mm also had their lowest density in December (1.060 g/cm³), but reached their maximum density in July (1.076 g/cm³).The density of the euphausiids was found to be size dependent. The density increases as the size decreases. C. finmarchicus and C. hyperboreus had densities less than seawater (1.026 g/cm³) during most of the year. Just before spawning the density increased to 1.028 g/cm³ and 1.036 g/cm³ for C. finmarchicus and C. hyperboreus respectively. The seasonal variations of the density were closely related to the lipid content of the animals.     

Lipid dynamics in fish: aspects of absorption, transportation, deposition and mobilization

1. Aspects of lipid metabolism, including absorption and depositional processes, appear quite different in fish as compared to homeothermic vertebrates. 2. Dietary lipids in fish are absorbed as fatty acids and as triacylglycerols aggregated into chylomicra particles. 3. Interorgan transport of lipids, like that of mammals, consists of an exogenous (dietary) loop and an endogenous loop. 4. Fish store lipids among several depot organs, including mesenteric membranes, liver and muscle. 5. Several fast-acting and slow-acting agents modulate depot lipid mobilization. 6. Mobilized lipids may be transported in the serum as free fatty acids bound to specific carrier proteins.