Lipid composition of the copepod Calanus hyperboreas from the Arctic Ocean. Changes with depth and season

A build-up of reserve lipid, predominantly wax esters, occurred during the summer in the copepod Calanus hyperboreas, collected off an Arctic ice-island. This lipid storage was correlated with a phytoplankton bloom and was followed by a progressive decrease of lipid from 2.1 mg per individual in September to 0.4 mg in June. There was a rapid decrease in lipid utilization between October and December, followed by much slower decreases until June. Lowered respiration rates or the availability of different types or quantities of food in the winter and spring are possible explantions for the slower rate of lipid use. Laboratory starvation experiments for up to 90 days correlated with results from the field. Gas-liquid chromatographic studies of the lipids showed that both the alcohols and fatty acids of the wax esters were highly variable with season and depth, whereas the phospholipid fatty acids were not affected by changes in these parameters. Only summer samples bad wax esters with a phytoplankton-like fatty acid composition, and upper water winter copepods had wax esters with little polyunsaturation. The deep-water winter copepods had a very different wax-ester composition from the upper water samples, with a predominance of hexadecanol (all other copepod samples had 20:1 and 22:1¹ as the principal alcohols) and a high content of polyunsaturated acids. Deep-water C. hyperboreas may differ in food habits and life history from those in the upper water community.     

On the lipid biochemistry of polar copepods: compositional differences in the Antarctic calanoids Euchaeta antarctica and Euchirella rostromagna

During austral summer of 1985 different developmental stages (CIII, CIV, CV, females, males) of the Antarctic copepod Euchaeta antarctica and females of Euchirella rostromagna were collected in the southeastern Weddell Sea to determine their lipid contents and compositions. For E. antarctica the analyses revealed a strong ontogenetic accumulation of lipids towards the older copepodids with highest lipid contents in late CV stages and adults. The females of E. rostromagna had moderate lipid levels. The most striking difference between these two species concerns their lipid class compositions. E. antarctica deposited predominantly wax esters, whereas in E. rostromagna the major lipid class consisted of triacylglycerols, an unusual storage lipid in polar marine copepods. Principal fatty acids in E. antarctica were the monounsaturates 18:1(n-9) and 16:1(n-7), especially in the lipid-rich stages, while the polyunsaturated fatty acids 20:5(n-3) and 22:6(n-3), usually membrane lipids, dominated in the lipid-poor stages. The wax ester moieties in E. antarctica consisted almost entirely of 14:0 and 16:0 fatty alcohols. Major components in E. rostromagna were the fatty acids 18:1(n-9), 16:0, 20:5(n-3) and 22:6(n-3). The potential of fatty acids and alcohols as typical trophic markers is rendered largely insignificant in the two species due to catabolic processes.     

Ecological investigations on the zooplankton community in balsfjorden,northern Norway: Lipids and fatty acids in Meganyctiphanes norvegica,Thysanoessa raschi and T. inermis during mid-winter

Total lipid of Meganyctiphanes norvegica (M. Sars) contained 53% triacylglycerols and traces of wax esters, that of Thysanoessa raschi (M. Sars) contained 44% triacylglycerols and 10% wax esters and that of T. inermis (Krøyer) contained 28% triacylglycerols and 40% wax esters. The triacylglycerols of M. norvegica were relatively rich in 20:1 and 22:1 fatty acids and its traces of wax esters resembled those of calanoid copepods. The triacylglycerols of both Thysanoessa species were deficient in 20:1 and 22:1 fatty acids but were richer in 16:1(n-7) and 18:1 (n-7) acids than those of M. norvegica. The wax esters of T. raschi contained phytol as almost the only fatty alcohol and were rich in 16:0 and 18:1 (n-9) fatty acids. The wax esters of T. inermis contained mainly 16:0 and 14:0 fatty alcohols with lesser amounts of phytol and their dominant fatty acid was 18:1, especially the (n-9) isomer. The triacylglycerols of T. inermis had 18:4 (n-3) as the major polyunsaturated fatty acid. From these and other aspects of fatty acid and fatty alcohol analyses it is concluded that a major foodstuff of M. norvegica in Balsfjorden is wax ester-rich calanoid copepods. T. raschi and especially T. inermis are concluded to have much more preference for phytoplanktonic food. Results are discussed in terms of current knowledge of the lipid chemistry of krill in the northern and southern hemispheres.     

Importance of wax esters and other lipids in the marine food chain: Phytoplankton and copepods

Wax esters, which function as reserve fuels, account for 25 to 40% of the lipid of the pelagic copepod Calanus helgolandicus (Copepoda, Calanoida). In laboratory experiments with these crustaceans, diatoms (Lauderia borealis, Chaetoceros curvisetus, and Skeletonema costatum) and dinoflagellates (Gymnodinium splendens), which contained no wax esters, were used as food. Changes in the food concentration affected both the amount of lipid and the composition of the wax esters. Since the fatty acids of the triglycerides and wax esters of C. helgolandicus resembled the dietary fatty acid composition, it appeared that copepods incorporated their dietary fatty acids largely unchanged into their wax esters. The polyunsaturated alcohols of the wax esters did not correspond in carbon numbers or degrees of unsaturation to the dietary fatty acids. We postulate two different metabolic pools to explain the origin of these long chain alcohols. The phospholipid fatty acids were not affected by changes in the amount or type of food, probably because of their structural function.     

Species-specific differences in lipid composition and omnivory indices in Arctic copepods collected in deep water during autumn (North Water Polynya)

To discern species-specific patterns in omnivory indices in copepods from the North Water Polynya, lipid classes and fatty acids were determined in Calanus hyperboreus Krøyer copepodite stage 5 (CV), C. glacialis Jaschnov (CV) and Metridia longa Lubbock (females) sampled below the mixed layer during autumn 1999. Generally, M. longa contained higher proportions of triacylglycerols, polar lipids and 18:1(n-9) than the other species. M. longa also had lower relative amounts of wax esters (WE), polyunsaturated fatty acids (PUFA), 20:1(n-9), 22:1(n-11), and lower absolute (g lipid copepod^-1) and relative (% lipid) lipid levels. Unsaturation coefficients (UC; ratio of polyunsaturated to total WE) were usually lowest in M. longa. These differences probably relate to a lesser dependence on phytoplankton in M. longa, and hence more ingestion of PUFA-poor prey. Moreover, levels of the carnivory index 18:1(n-9)/18:1(n-7) were highest in M. longa. Our data support the widespread contention that M. longa is more omnivorous than C. hyperboreus and C. glacialis. Proportions of bacterial fatty acids [odd-numbered and/or branched (OBFA); 18:1(n-7)] and PUFA/SFA (SFA: saturated fatty acids) ratios were largely unrelated to feeding strategy. Relationships between relative and absolute amounts of 16:1(n-7) and 18:1(n-7) in copepods suggest that the latter fatty acid is formed in vivo by chain elongation of the former. However, elevated levels of 18:1(n-7) and OBFA in M. longa at stations dominated by the microbial loop imply that these indices can be used to track ingestion of PUFA-poor protozoans when diatoms are scarce.Communicated by J.P. Grassle, New Brunswick     

The fate of dietary lipids in the Arctic ctenophore Mertensia ovum (Fabricius 1780)

Lipids of the Arctic ctenophore Mertensia ovum, collected from Kongsfjorden (Svalbard) in 2001, were analysed to investigate seasonal variability and fate of dietary lipids. Total lipids, lipid classes and fatty acid and alcohol compositions were determined in animals, which were selected according to age-group and season. Changes in lipids of age-group 0 animals were followed during growth from spring to autumn. Total lipids increased from May to September. Lipids as percentage of dry mass were lowest in August indicating their use for reproduction. Higher values occurred in September, which may be due to lipid storage for overwintering. Wax esters were the major lipid class accounting for about 50% of total lipids in age-group 0 animals from July and August. Phospholipids were the second largest lipid fraction with up to 46% in this age-group. The principal fatty acids of M. ovum from all age-groups were 22:6(n-3), 20:5(n-3) and 16:0. Wax ester fatty alcohols were dominated by 22:1(n-11) and 20:1(n-9) followed by moderate proportions of 16:0. The unique feature of M. ovum lipids was the high amount of free fatty alcohols originating probably from the dietary wax esters. In May, free alcohols exhibited the highest mean proportion with 14.6% in age-group 0 animals. We present the first data describing a detailed free fatty alcohol composition in zooplankton. This composition was very different from the alcohol composition of M. ovum wax esters because of the predominance of the long-chain monounsaturated 22:1(n-11) alcohol accounting for almost 100% of total free alcohols in some samples. The detailed lipid composition clearly reflected feeding of M. ovum on the herbivorous calanoid species, Calanus glacialis and C. finmarchicus, the abundant members of the zooplankton community in Kongsfjorden. Other copepod species or prey items seem to be less important for M. ovum.     

The Arctic pteropod<Emphasis Type="Italic"> Clione limacina</Emphasis>: seasonal lipid dynamics and life-strategy

During various seasons from May 2001 until July/August 2003, the lipid dynamics of the pteropod Clione limacina from Kongsfjorden, Svalbard, were investigated with respect to ontogenetic development and life-cycle. Polytrochous larvae, which were dominant in spring (April, May), composed the lipid-richest specimens of the population, with total lipid of about 50% of dry mass (%DM). Major lipid classes were triacylglycerols (TAG) and 1-O-alkyldiacylglycerol ethers (DAGE), accounting on average for 53.1 and 21.9% of total lipid, respectively. Until summer, larvae grew to adults by utilising their storage lipids. In July/August, lipids were depleted to about 10%DM due to maturation and reproduction. Almost all animals in autumn (September) were mature and able to replenish their lipid deposits by accumulating DAGE (26.7%) and TAG (39.6%). This is probably the prerequisite for successful overwintering.Principal component analysis (PCA), based on the fatty acid compositions, revealed ontogenetic differences between polytrochous larvae, and small and full-grown adults. Higher proportions of 18:4(n-3) and 14:0 were found in polytrochous larvae and smaller adults during spring. Both fatty acids were highly significantly correlated with the proportions of TAG, which were used for growth and development because they are presumably easier to metabolise. PCA also divided C. limacina specimens into DAGE-rich and DAGE-poor. We suggest that DAGE are a long-term energy store and hypothesise that they are necessary during periods of food scarcity, but may also serve as an energy source for reproduction. The fatty acids 17:1(n-8), 15:0, 16:1(n-7) and 18:1(n-7) were significantly correlated with the proportion of DAGE but not with TAG. These fatty acids, which do not originate from their only prey, Limacina helicina, are synthesised de novo. Their abundance reflects an efficient lipid production by C. limacina. Based on the results of lipid biosynthesis and accumulation in combination with the population structure, we suggest that C. limacina has at least a 2-year life-cycle in Svalbard waters.Communicated by M. Kühl, Helsingør     

Lipid classes and their content of n-3 highly unsaturated fatty acids (HUFA) in Artemia franciscana after hatching, HUFA-enrichment and subsequent starvation

The distribution of n-3 highly unsaturated fatty acids (HUFA) over the major neutral and polar lipid classes was determined for two predominant types of live food used in the larviculture of marine fish and shrimp, i.e. freshly hatched and HUFA-enriched Artemia, and compared with data reported in the literature for wild copepods, representing the natural diet of these larvae. Lipid class composition and their content of n-3 HUFA, particularly eicosapentaenoic acid (EPA, 20:5n-3) and docosahexaenoic acid (DHA, 22:6n-3), were assessed in freshly hatched, HUFA-enriched and subsequently starved Artemia franciscana. The n-3 HUFA enrichment was based on feeding Artemia a lipid emulsion in which either fatty acid ethyl esters (EE, diluted with olive oil) or triacylglycerol (TAG) provided a level of 30% n-3 HUFA. Enrichment of Artemia with either type of the lipid emulsions resulted in an increase of total lipid content from 20.0 to 28.2–28.7% of dry matter mainly due to the accumulation of neutral lipid, primarily TAG (from 82 to 158 mg g⁻¹ dry wt in freshly hatched and 24-h enriched Artemia). Enriched brine shrimp utilized up to 27–30% of their TAG content during 72 h of starvation at 12 °C. The absolute tissue concentrations of polar lipids remained constant at 71 to 79 mg g⁻¹ dry wt throughout the enrichment and subsequent starvation. The level of n-3 HUFA increased drastically during enrichment from 6.3% of total fatty acids (8.2 mg g⁻¹ dry wt) in freshly hatched nauplii to between 20.4 and 21.8% (40.4 to 43.2 mg g⁻¹ dry wt) in 24-h enriched Artemia and was not significantly affected by the source of n-3 HUFA. During starvation, 18:0, 20:4n-6 and 20:5n-3 were retained, whereas 18:4n-3, 22:5n-3 and 22:6n-3 were specifically catabolized. The major polar lipids, phosphatidylethanolamine (PE) and phosphatidylcholine (PC), of freshly hatched Artemia showed very low levels of DHA (<0.1% of total fatty acids) and carried about 45% of the total EPA present. Enrichment with either of the emulsions resulted in an increase of the neutral lipid fraction which concentrated >64% of the EPA and >91% of the total DHA present. This is in sharp contrast with the high levels of n-3 HUFA, in particular DHA, in the polar lipid fraction reported for wild copepods. The contrasting distribution of DHA in the neutral and polar lipid fractions of enriched brine shrimp compared to the natural diet may influence the efficacy of this essential fatty acid for marine fish larvae in aquaculture systems. Received: 10 June 1997 / Accepted: 8 August 1997     

Trophodynamics and seasonal cycle of the copepod Pseudocalanus acuspes in the Central Baltic Sea (Bornholm Basin): evidence from lipid composition

Seasonal lipid dynamics of the copepod Pseudocalanus acuspes were studied in the Bornholm Basin (Central Baltic Sea) on a monthly basis from March 2002 until March 2003 and were interpreted in light of life cycle strategies and diet selection. The individual total lipid content of females ranged from 0.9 to 1.8 μg, with relative wax ester content reaching a significant maximum in May (44% of total lipids) and minimum (17% of total lipids) in April and November. Significant changes in size, lipid content, lipid classes and fatty acid composition of structural as well as storage lipids suggested five characteristic seasonal phases that were induced by different feeding histories and environmental conditions. Storage lipids were characterized by 18:1(n−9) as major component, which ranged between 44% of total fatty acids in June and 23% in February. The strong coherence between 18:1(n−9) in the seston lipids and the occurrence of ciliates emphasized the importance of ciliates in the diet of P. acuspes. As indicated by changes in the amounts of fatty acid markers, other food sources varied over the year, suggesting an opportunistic feeding behavior. The spring period was characterized by an increase in typical diatom and dinoflagellate markers, whereas other sources, potentially cyanobacteria, became more important during summer. The life cycle strategy is discussed with respect to extant adaptations to high latitudinal habitats.     

Ontogenetic and seasonal changes in lipid and fatty acid/alcohol compositions of the dominant Antarctic copepods Calanus propinquus,Calanoides acutus and Rhincalanus gigas

Lipid compositions of the dominant Antarctic copepods Calanoides acutus, Rhincalanus gigas and Calanus propinquus from the Weddell Sea have been investigated in great detail. Copepods were collected during summer in 1985 and late spring/early winter in 1986. The analyses revealed specific adaptations in the lipid biochemistry of these species which result in very different lipid components. The various copepodite stages of C. acutus synthesize wax esters with long-chain monounsaturated moieties and especially the alcohols consisted mainly of 20:1(n-9) and 22:1(n-11). R. gigas also generates wax esters, but with moieties of shorter chain length. The fatty alcohols consisted mainly of 14:0 and 16:0 components, while the major fatty acids were 20:5, 18:4 and 22:6, of which 18:4 probably originated from dietary input. In contrast, C. propinquus accumulates triacylglycerols, a very unusual depot lipid in polar calanoid copepods. Major fatty acids in C. propinquus were the long-chain monounsaturates 22:1(n-9) and 22:1(n-11), which may comprise up to 50% of total fatty acids. In C. acutus and C. propinquus there was a clear increase of long-chain fatty acids with increasing developmental stage. In contrast, the fatty acid and alcohol composition of the R. gigas copepodite stages were characterized by the dominance of the polyunsaturated fatty acids as well as high amounts of the monounsaturates 18:1(n-9) and 16:1(n-7). There was a considerable decrease of the dietary fatty acid 18:4(n-3) towards the older stages during summer; in late winter/early spring 18:4 was only detected in very low amounts. This tendency was also found in the other two species, but was less pronounced. In all three species dry weight and lipid content increased exponentially from younger to older stages. The highest portion of wax esters, or of triacylglycerols in C. propinquus, was found in the adults. Dry weight and lipid content were generally higher during summer. In late winter/early spring the variability was more pronounced and lipid-rich specimens showed a selective retention of long-chain monounsaturated fatty acids, whereas in lipid-poor specimens these fatty acids were very much depleted.     

Changes in lipids during the development of Calanus finmarchicus s.l. from Copepodid I to adult

The total lipid and wax ester content as well as the fatty acid and alcohol composition of all copepodid stages and adults of Calanus finmarchicus s.l. were investigated at different locations in the North Sea in 1983 and 1984. Total lipid and the wax ester proportion increased exponentially until Copepodid V. The females were sometimes lower in lipids than the Stage V. The wax ester proportion reached about 90% of total lipids in males and Copepodid V and up to 40% in Copepodid I. The major fatty acids were 16:0, 20:5, and 22:6 and the major fatty alcohols were 16:0, 20:1 and 22:1. At one station the 18:4 acid became one of the dominant acids, because of a Phaeocystis sp. bloom, indicating that the fatty acids of the diet are incorporated mostly unchanged into the lipids of the copepods. The other main fatty acids 20:1 and 22:1 are probably synthesized de novo, serving as precursors for the principal alcohols 20:1 and 22:1. Their levels decreased in the younger stages due to increases in 16:0 alcohol. The fatty alcohol-forming enzyme seems to be specific for saturated and monounsaturated acids, which may be synthesized de novo or derived from diet.     

Fatty-acid composition of ovulated eggs from wild and cultured turbot (Scophthalmus maximus) in relation to yolk and oil globule lipids

The fatty-acid composition of lipids from ovulated eggs of wild and cultured turbot was investigated in order to estimate the nutritional requirements during embryonic and early larval development. Lipid comprised 13.8±0.5% (n=5) and 13.2±0.7% (n=7) of the egg dry weight in wild and cultured turbot, respectively. Polyunsaturated fatty acids (PUFA) of the (n-3) series accounted for 39% of total fatty acids in total lipid of both wild and cultured fish. The predominant (n-3) FUFA was docosahexaenoic acid (22:6 n-3), which also was the most abundant fatty acid in turbot eggs and comprised 24 and 23% of the total egg fatty acids in wild and cultured fish, respectively. Phospholipids, triacylglycerols and cholesterol-wax esters of turbot eggs all exhibited a specific fatty-acid profile distinctly different from that of total lipid. The general pattern of the fatty-acid distribution in lipids of eggs from wild and cultured turbot was similar, but the relative amount of 18:2(n-6) was considerably higher and 20:1(n-9) slightly higher in cultured fish. These differences were extended to all lipid classes and probably reflect the dietary intake of certain vegetable and marine fish oils. Calculations based on light microscopical studies showed that 55 to 60% of the total lipids in cultured turbot eggs are confined to the oil globule. The size of the oil globule remained constant during embryogenesis, and a reduction in size occurred first after hatching and mainly after yolk depletion. This implies that the total amount of lipids utilised during the embryonic development is considerably less than the total lipids present in ovulated turbot eggs. Comparison of the fatty-acid composition of total lipids from eggs and vitellogenin of wild turbot reveals that egg lipids contained a lower level of saturated and a higher level of monounsaturated fatty acids. Eggs also contained wax esters, which were not detected in vitellogenin, suggesting that vitellogenin is not the only source of lipids for turbot eggs.     

Strain-related physiological and behavioral effects of <Emphasis Type="Italic">Skeletonema marinoi</Emphasis> on three common planktonic copepods

Three strains of the chain-forming diatom Skeletonema marinoi, differing in their production of polyunsaturated aldehydes (PUA) and nutritional food components, were used in experiments on feeding, egg production, hatching success, pellet production, and behavior of three common planktonic copepods: Acartia tonsa, Pseudocalanus elongatus, and Temora longicornis. The three different diatom strains (9B, 1G, and 7J) induced widely different effects on Acartia tonsa physiology, and the 9B strain induced different effects for the three copepods. In contrast, different strains induced no or small alterations in the distribution, swimming behavior, and turning frequency of the copepods. 22:6(n-3) fatty acid (DHA) and sterol content of the diet typically showed a positive effect on either egg production (A. tonsa) or hatching success (P. elongatus), while other measured compounds (PUA, other long-chain polyunsaturated fatty acids) of the algae had no obvious effects. Our results demonstrate that differences between strains of a given diatom species can generate effects on copepod physiology, which are as large as those induced by different algae species or groups. This emphasizes the need to identify the specific characteristics of local diatoms together with the interacting effects of different mineral, biochemical, and toxic compounds and their potential implications on different copepod species.     

Biosynthesis of lipds in zooplankton from Saanich Inlet,British Columbia,Canada

Lipid analyses were carried out on various species of zooplankton captured in Saanich Inlet, British Columbia, Canada, during September, 1972. The amphipod Cyphocaris challengeri had the highest level of lipid, consisting mainly of wax esters. The copepod Calanus pacificus had moderate amounts of lipid, with triglyceride as the major neutral lipid. The euphausiid Thysanoessa raschii contained mainly triglyceride in its moderate levels of neutral lipid, while wax esters and, to a lesser extent, triglycerides were present in the very small levels of neutral lipid in the chaetognath Sagitta elegans. The major fatty alcohols in wax esters of both Cyphocaris challengeri and Calanus pacificus were 20:1 and 22:1, although notable differences were found in the major fatty acids. Biosynthetic studies showed that phospholipids were labelled faster than neutral lipids in all species with both (U-¹⁴C) glucose and (1-¹⁴C) palmitic acid as precursors. Only species containing significant amounts of wax esters in their neutral lipids incorporated substantial amounts of radio-activity from (1-¹⁴C) palmitic acid into wax esters in (i) living animals, (ii) preparations containing fragments of tissue, (iii) cell-free systems. All species incorporated added fatty alcohols into wax esters in preparations containing tissue fragments and in cell-free preparations. Both the fatty acid and fatty alcohols of the wax esters of both Calanus pacificus and S. elegans were labelled from (1-¹⁴C) palmitic acid, consistent with de novo biosynthesis of the esters. (1-¹⁴C) hexadecanol was incorporated into wax esters almost entirely in the fatty alcohol moiety. It is concluded that all species examined formed was esters when presented with preformed fatty alcohols, but only those species that had wax esters as a major component of the neutral lipids were capable of de novo biosynthesis of the lipids.     

Trophic upgrading of food quality by protozoans enhancing copepod growth: role of essential lipids

 Protozoa are known for their intermediary trophic role in transferring organic matter from small size planktonic particles to mesozooplankton. This study concentrates on the possible addition of biochemical value during this transfer, by new production of compounds that are essential in copepod food. In laboratory experiments, copepods could not be raised on a diet of the chlorophycean Dunaliella sp., though they readily consumed this alga. Dunaliella sp. contained all essential amino acids, but was deficient in highly unsaturated fatty acids and in sterols. In contrast to copepods, the heterotrophic dinoflagellate Oxyrrhis marina grew well on Dunaliella sp., producing significant amounts of the long-chain fatty acids docosahexaenoic acid and eicosapentaenoic acid, in addition to cholesterol and brassicasterol. Using this O. marina grown on Dunaliella sp. to feed Temora longicornis and Pseudocalanus elongatus, both copepod species rapidly developed from young nauplius larvae to maturity on the dinoflagellate diet. Hence, in this experimental food-chain the inadequate chlorophycean food was biochemically upgraded by the protozoan to high-quality copepod food. The results indicate that highly unsaturated fatty acids and/or sterols are essential compounds, which can be produced by protozoans. Due to their intermediate size, the mechanism of trophic upgrading by protozoans may bridge the gap of essential nutrients between the microbial loop and higher trophic levels. Received: 11 January 1999 / Accepted: 3 June 1999     

Seasonal lipid dynamics of the copepods <Emphasis Type="Italic">Pseudocalanus minutus</Emphasis> (Calanoida) and <Emphasis Type="Italic">Oithona similis</Emphasis> (Cyclopoida) in the Arctic Kongsfjorden (Svalbard)

Seasonal lipid dynamics of various developmental stages were investigated in Pseudocalanus minutus and Oithona similis. For P. minutus, the dominance of 16:1(n−7), 16:4(n−3) and 20:5(n−3) fatty acids indicated a diatom-based nutrition in spring, whereas 22:6(n−3), 16:0, 18:2(n−6) and 18:1(n−9) pointed to a flagellate-based diet during the rest of the year as well as omnivorous/carnivorous low-level feeding during winter. The shorter-chain fatty alcohols 14:0 and 16:0 prevailed, also reflecting biosynthetic processes typical of omnivores or carnivores. Altogether, the lipid signatures characterized P. minutus as an opportunistic feeder. In contrast, O. similis had consistently high amounts of the 18:1(n−9) fatty acid in all stages and during all seasons pointing to a generally omnivorous/carnivorous/detritivorous diet. Furthermore, the fatty alcohol 20:1(n−9) reached high percentages especially in adult females and males, and feeding on Calanus faecal pellets is suggested. Fatty alcohols, as wax ester moieties, revealed significant seasonal variations in O. similis and a seasonal trend towards wax ester accumulation in autumn in P. minutus. P. minutus utilized its lipid deposits for development in the copepodite stages III and IV and for gonad maturation in CV and females during the dark season. However, CVs and females depended on the spring phytoplankton bloom for final maturation processes and reproduction. O. similis fueled gonad maturation and egg production for reproduction in June by wax esters, whereas reproduction in August/September co-occurred with the accumulation of new depot lipids. Both species revealed significantly higher wax ester levels in deeper (>50 m) as compared to surface (0–50 m) dwelling individuals related to a descent prior to overwintering.     

Lipids and fatty acids as indicators of egg condition,larval feeding and maternal effects in Cape hakes (<Emphasis Type="Italic">Merluccius paradoxus</Emphasis> and <Emphasis Type="Italic">M. capensis</Emphasis>)

Cape hakes, Merluccius paradoxus and M. capensis, are important gadoid fish that are commercially harvested in the Benguela Current system off Namibia and South Africa. The aim of this study was to elucidate the nutritional condition and feeding preferences of their larvae. Hake eggs and larvae were sampled in austral spring of two consecutive years, 2007 and 2008, off the west coast of South Africa. They were identified to species using genetics, and total lipid content and fatty acid (FA) composition were analysed for each individual egg and larva to compare the condition of different early life stages of both hake species. Higher abundances of M. paradoxus eggs and larvae were consistently found compared to M. capensis. In both species, eggs contained wax esters (WE) and had significantly higher lipid content per dry mass than larvae. Lipid content as well as FA composition changed with the developmental stage of larvae. Quantities of essential fatty acid (EFA) increased with feeding of larvae due to dietary lipid incorporation. In 2007, yolk-sac larvae contained significantly lower total lipids than in 2008. It is argued that this was due to reduced lipid transfer by the spawning females to the eggs. These findings indicate that maternal effects are important in determining condition of hake larvae and that this may have an effect on their survival and subsequent recruitment.     

Lipids of some Caribbean and Red Sea corals: total lipid,wax esters,triglycerides and fatty acids

The Caribbean reef-building corals Porites porites (Pallas) and Montastrea annularis (Ellis and Solander) and the Red Sea corals Pocillopora verrucosa (Ellis and Solander), Stylophora pistillata (Esper) and Goniastrea retiformis (Lamark) were analysed for total lipid, wax ester and triglyceride content, and fatty acid composition. M. annularis contained about 32% of dry weight as total lipid, whereas much lower values of between 11 and 17% were recorded for the other four species. It is concluded that there is greater variation in coral total lipid than hitherto thought. The total lipid contained a substantial proportion of wax ester (22 to 49%) and triglyceride (18 to 37%). The storage lipids (wax esters and triglycerides) accounted for between 6 and 20% of the dry weight and between 46 and 73% of the total lipid. Variation in lipid content between species could not be attributed to geographical location, but the low values for total lipid in Red Sea corals may in part be due to environmental factors as these samples were collected in winter. All corals analysed contained high levels of saturated fatty acids, the most abundant fatty acids being 16:0, 18:0 and 18:1n-9. Marked differences were observed in polyunsaturated fatty acid (PUFA) content between species, with comparatively low levels of 10 and 11% of fatty acids being recorded in M. annularis and G. retiformis, respectively. The values for the other species ranged between 21 and 37%. Fatty acid composition may vary according to the proportions of fatty acids obtained from diet, algal photosynthesis and synthesis by the animal tissues.     

Fatty acid signature analysis documents the diet of five myctophid fish from the Southern Ocean

Fatty acid (FA) and fatty alcohol (FAlc) compositions of both total lipid and neutral lipid fractions were studied for five myctophid species sampled in Kerguelen waters. Both qualitative and quantitative FA signature analyses were then performed to investigate their diet over longer time scales than the conventional stomach content analysis. Regarding their lipid class, FA and FAlc compositions, the five species could be discriminated into two groups: wax-ester-rich species (Electrona antarctica, Krefftichthys anderssoni) characterised by large amounts of monounsaturated FAs (>73% of total FAs) and triacylglycerol-rich species (Electrona carlsbergi, Gymnoscopelus nicholsi, Protomyctophum bolini) with major amounts of saturated and monounsaturated FAs (>29 and >46% of total FAs, respectively). Qualitative and quantitative FA analyses showed that K. anderssoni mainly preyed upon copepods, E. antarctica upon copepods and more euphausiids and P. bolini and E. carlsbergi mainly upon euphausiids with some copepods, while G. nicholsi had a more diverse diet. This study shows the usefulness of quantitative statistical analysis to determine the diet of Antarctic and sub-Antarctic predators and stresses the need of increasing the lipid and FA analyses of more zooplanktonic and micro-nektonic marine species.     

Characterizing dietary variability and trophic positions of coastal calanoid copepods: insight from stable isotopes and fatty acids

The spring zooplankton community in the Strait of Georgia (British Columbia, Canada) is characterized by the presence of several calanoid copepod species which collectively make up ~90% of the mezozooplankton biomass. Here, we investigate interspecific, interannual, and geographic variability in the diets and trophic positions of these copepods using a combination of fatty acids and stable isotopes. To characterize geographic variability in diet, we compare our findings from the Strait of Georgia with similar data from Ocean Station P in the subarctic northeast Pacific. Both fatty acid and stable isotope signatures indicate the existence of three trophic levels, even within the limited size range of these copepods: Neocalanus plumchrus and Calanus marshallae are primarily omnivorous, while Euchaeta elongata is carnivorous and Eucalanus bungii is herbivorous. Fatty acid markers of trophic position (e.g., DHA/EPA, 18:1n-9/18:1n-7) correlate significantly with δ¹⁵N, while markers indicating the proportion of diatoms to flagellates in the diet (e.g., 16PUFA/18PUFA and DHA/EPA) correlate significantly with δ¹³C, after the effect of lipid concentration on δ¹³C is accounted for. Despite the general correlation between stable isotopes and fatty acids, the former are not sensitive enough to capture the range of interannual variability observed in the latter, and can only capture substantial shifts in the diet over geographic scales. However, regardless of variability in food quality, the relative trophic positions of these copepods do not change significantly either spatially or temporally.