Lipid metabolism of the Antarctic krill Euphausia superba and its ecological implications

 Various developmental stages (early larvae to adults) of Euphausia superba have been collected in different seasons in the Weddell Sea, the Lazarev Sea and off the Antarctic Peninsula to investigate the role of lipids and fatty acids in the life cycle of the Antarctic krill. The total-lipid data for E. superba exhibited seasonal variations, with low lipid levels in late winter/early spring and the highest levels in autumn. Seasonal changes were most pronounced in the immature and adult specimens, increasing from about 10% lipid of dry mass to more than 40%. The fatty-acid compositions of the younger stages were dominated by 20:5(n-3), 22:6(n-3) and 16:0. These are typical phospholipid fatty acids, which are major biomembrane constituents. The phospholipid composition was similar in the older stages. With increasing storage of triacylglycerols in the lipid-rich immature and adult stages, the fatty acids 14:0, 16:0 and 18:1(n-9) prevailed, comprising about 70% of total triacylglycerol fatty acids. The trophic-marker fatty acids 16:1(n-7) and 18:4(n-3), indicating phytoplankton ingestion, were less abundant. They reflected, however, the dependence of the larvae on phytoplankton as well as the seasonal changes in algal composition. The generally close linear relationships between fatty acids and lipid suggest that the fatty-acid compositions of the collected specimens were largely independent of the respective developmental stage, season and region. The linear fit indicates that triacylglycerol accumulation started at a level of about 5% of total lipid. Considering the various overwintering scenarios under discussion, the life cycle and reproductive strategies of krill are discussed in the context of the lipid metabolism and fatty-acid composition of E. superba. Lipid production is effective enough to accumulate large energy reserves for the dark season, but E. superba does not exhibit the sophisticated biosynthetic pathways known from other Antarctic euphausiids and copepods. Although important, lipid utilisation appears to be just one of several strategies of E. superba to thrive under the extreme Antarctic conditions, and this pronounced versatility may explain the success of this species in the Southern Ocean. Received: 16 June 2000 / Accepted: 18 December 2000     

Variable within-brood maternal provisioning in newly extruded embryos of Homarus gammarus

Offspring quality of decapod crustaceans has been widely studied, with special emphasis on the sources of variability determining embryonic and larval quality. Nevertheless, maternal provisioning has commonly been overlooked as a potential source of offspring within-brood variability. In the present study, the existence of variable maternal provisioning was assessed through the analysis of the fatty acid (FA) profile of newly extruded embryos from different regions of the brooding chamber of Homarus gammarus. Significant differences in the FA profile of embryos sampled from different pleopods and sides of the brooding chamber were recorded. Significant deviations of the overall mean of each surveyed female were also observed for essential FA, particularly 20:4n-6, 20:5n-3 and 22:6n-3. Lipid energy available to fuel embryogenesis also varied among embryos sampled from different regions of the brooding chamber. Results suggest variable female investment at oocyte production, which may be amplified during the incubation period of developing embryos by differential lipid catabolism. For the first time, maternal provisioning is evidenced as an additional source for within-brood variability in the FA profile of embryos.     

Diet composition and quality for <Emphasis Type="Italic">Calanus finmarchicus</Emphasis> egg production and hatching success off south-west Iceland

Egg production and hatching success of the copepod Calanus finmarchicus was measured during spring and summer in the waters south-west of Iceland. Egg-production rates varied greatly, both temporally and spatially, with highest average rates found at a station with low chlorophyll-a concentrations (0.4 mg m^-3). Excluding this high production rate from statistical analysis, the remaining egg-production rates were found to be positively correlated with phytoplankton biomass, as well as with parameters representing healthy phytoplankton condition, food quality and diatom-type fatty acids. Hatching success of eggs was negatively correlated with some saturated and monounsaturated fatty acids related to phytoplankton senescence.     

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.     

Spatio-temporal variability in fatty acid trophic biomarkers in stomach contents and muscle of Iberian sardine (<Emphasis Type="Italic">Sardina pilchardus</Emphasis>) and its relationship with spawning

Temporal variation in the fatty acid (FA) composition of stomach contents of Iberian sardines was compared to the relative contribution to dietary carbon made by different prey types for fish from two areas off Portugal. The effect of the FA content of the diet on sardine muscle FA composition was also studied, aiming at (1) analysing if FA biomarkers can be used as a complementary technique for the study of sardine diet and (2) to relate spatial and temporal variations of prey FA content with sardine condition and reproduction. Significant spatial differences in the FA composition of sardine diet occurred with concentrations of n-3 polyunsaturated FA, namely eicosapentaenoic acid [EPA, 20:5n-3] and linolenic acid 18:3n-3, being significantly higher in the diet of sardines from the west coast, whilst the diet of sardines from the south coast was richer in monounsaturated fatty acids (MUFA), namely the carnivory biomarker oleic acid 18:1n-9. These results are in agreement with the higher contribution made by diatoms and dinoflagellates to the diet of sardines off the west coast. Spatial variation in sardine dietary FA was also detected in their muscle composition, specifically for EPA, and the eicosapentaenoic/docosahexaenoic acid and (n-3)/(n-6) ratios, which were higher in sardines from the west coast. No difference in FA composition was detected between sexes, and the seasonal variability in sardine total FA concentration was primarily related to the seasonality of spawning. Sardines accumulate high concentrations of FAs during the resting stage of reproduction when the feeding intensity is similar or lower to that observed during the spawning season. Additionally, sardines show a high selective retention of MUFA and polyunsaturated FA (PUFA) throughout the year except at the beginning of the spawning season, when these FAs are largely invested in the formation of the gonads. Therefore, temporal and regional differences of prey environments are strong enough to be reflected in fish body composition, namely on the accumulation of essential FAs, which can have a strong impact on reproduction success for this species.     

Ontogenic variations in fatty acid and alcohol composition of the pelagic amphipod <Emphasis Type="Italic">Themisto libellula</Emphasis> in Kongsfjorden (Svalbard)

The fatty acid and alcohol composition of the pelagic amphipod, Themisto libellula, was monitored during the 5 first months of its life cycle (4–20 mm length) in an Arctic fjord, Kongsfjorden, Svalbard. Fatty acids of the three major lipid classes, polar lipids (PL), triacylglycerol (TAG), and wax esters (WE), were analyzed to highlight ontogenic changes in their diet and metabolism. The PL composition of T. libellula did not show any strong variations along their growth except during the first month where an important increase of 20:5(n-3) (EPA) and 22:6(n-3) (DHA) was observed. The TAG composition revealed a clear gradient corresponding to a diet shift from omnivorous juveniles toward carnivorous sub-adults and adults. Indeed, fatty acid trophic markers of diatoms were dominant in the juveniles, whereas 20:1(n-9) and 22:1(n-11), the Calanus sp. trophic markers, overwhelmed in the older stages. The WE composition highlighted the same general trend, however, differences were found with the TAG and are discussed as a result of differences in turnover rates and assimilation pathways between the two lipid classes.     

Seasonal patterns in fatty acids of <Emphasis Type="Italic">Calanus hyperboreus</Emphasis> (Copepoda,Calanoida) from Cumberland Sound,Baffin Island,Nunavut

The marine copepod Calanus hyperboreus accumulates large quantities of lipids and essential fatty acids during summer months in Northern oceans. However, few data exist regarding their winter fatty acid profiles, which could be informative regarding the use of lipids by C. hyperboreus to successfully survive and reproduce during times of ice-cover and limited food. The present study compared fatty acids of C. hyperboreus between summer (August 2007 and 2008) and winter (early April 2008 and 2009) in Cumberland Sound, Canada. Summer samples from both years had significantly higher ∑polyunsaturated fatty acids and unsaturation indices (based on μg fatty acid mg dry tissue⁻¹) than winter samples and separated on a principal component analysis due to higher 18:2n-6, 18:4n-3, and 20:5n-3, consistent with phytoplankton consumption. Winter C. hyperboreus had significantly higher ∑monounsaturated fatty acids (MUFA) versus summer samples and separated on the principal component analysis due to higher proportions of 16:1n-7, 20:1n-9, and 22:1n-9, suggesting they were not actively feeding. Based on the seasonal fatty acid comparison, C. hyperboreus was catabolizing specific fatty acids (e.g. 20:5n-3), conserving others (e.g. 22:6n-3), and maintaining or increasing biosynthesis of certain MUFA (e.g. 18:1n-9) during winter. These findings provide insight into the seasonal strategy of acquisition (summer) and utilization (winter) of specific fatty acids by a key Arctic organism and could become important for monitoring changes in fatty acids associated with decreased ice-cover duration due to climate warming.     

Energy substrates for eggs and prefeeding larvae of the dolphinCoryphaena hippurus

Changes in the chemical composition of developing dolphin (Coryphaena hippurus) eggs and prefeeding yolksac larvae were determined in order to estimate probable dietary requirements of first-feeding larvae. Daily dry matter, protein nitrogen (PN), non-protein nitrogen (NPN), lipid, gross energy content, fatty acid and amino acid profiles from Day 1 to Day 2 eggs and Day 1 to Day 3 larvae were compared. Lipid was the primary endogenous energy source accounting for the daily caloric deficit through both the egg and larval stages, except over the day of hatching. The catabolism of lipid by embryos (0.078 cal d^–1) was greater than that by yolksac larvae (0.036 cal d^–1). The higher demand for energy by embryos was related to a greater rate of protein synthesis during the egg stage. The ratio of PN:NPN increased during egg development without change in total nitrogen content, but was constant throughout the yolksac larvae period. The lipid content per embryo did not decrease over the hatching period (Day 2 to 3, postspawning). However, there was a loss in amino acid content not totally accounted for by sloughing of the chorion at hatching. This loss, as protein, accounted for 0.053 cal of gross energy, which represented 70% of the total estimated energy needs of the fish over this period. Loss of non-essential amino acids (25%) was higher than that of essential amino acids (13%). Proline and tyrosine accounted for 32% of the total loss of amino acids at this time. The only preferential use of fatty acids over any period was a small but significant drop in the content of C22:6n-3 prior to the onset of feeding (Day 5, postspawning). It is speculated that the pattern of energy-substrate use of first-feeding dolphin larvae will reflect the pattern of endogenous energy use during the egg and prefeeding yolksac larval stages. Diets or feeding regimens with lipid as the primary energy source, and containing a fatty acid profile similar to that of eggs or yolksac larvae, should be useful in culturing this species, at least during the early feeding stages.     

Comparative lipid dynamics of euphausiids from the Antarctic and Northeast Pacific Oceans

To better understand the feeding and reproductive ecology of euphausiids (krill) in different ocean environments, lipid classes and individual lipid components of four different species of euphausiids from Northeast Pacific (temperate species) and Southern Ocean (Antarctic species) were analyzed in animals from multiple life stages and seasons. The dominant krill species in the Northeast Pacific Euphausia pacifica and Thysanoessa spinifera, were compared to the two major Antarctic species, Euphausia superba and E. crystallorophias. Analysis comprised total lipid and lipid classes together with individual fatty acid and sterol composition in adults, juveniles, and larvae. Antarctic krill had much higher lipid content than their temperate relatives (10–50 and 5–20% of dry mass for Antarctic and temperate species, respectively) with significant seasonal variations observed. Phospholipids were the dominant lipid class in both temperate krill species, while neutral storage lipids (wax esters and triacylglycerols for E. crystallorophias and E. superba, respectively) were the major lipid class in Antarctic krill and accounted for up to 40% of the total lipid content. Important fatty acids, specifically 16:0, 18:1ω9, 20:5ω3, and 22:6ω3, were detected in all four krill species, with minor differences between species and seasons. Detailed lipid profiles suggest that krill alter their lipid composition with life stage and season. In particular, larval Antarctic krill appear to utilize alternate food resources (i.e., sea-ice associated organisms) during austral winter in contrast to juveniles and adults (i.e., seston and copepods). Lipid dynamics in krill among krill in both systems appear closely linked to their life cycle and environmental conditions including food availability, and can provide a more complete comparative ecology of euphausiids in these environmentally distinct systems.     

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.     

Fatty acid composition of organs and tissues of the tiger prawn Penaeus esculentus during the moulting cycle and during starvation

The fatty acids (FA) in neutral lipid (NL) and polar lipid (PL) of digestive gland, muscle and integument of Penaeus esculentus Haswell were analysed and compared during the moulting cycle and during starvation. The prawns were collected from Moreton Bay, Queensland, Australia, by trawling during 1985–1987, and were fed with a standard semi-purified diet. Compared with a natural diet, the artificial diet had much higher levels of 18:1n-9 and 18:2n-6, but only trace amounts of 20:4n-6, but there was no evidence of dietary imbalance. The fatty acid composition (percentage of total lipid) of the digestive gland changed markedly during the moulting cycle and during starvation, but the small changes observed in both muscle and integument suggested that these tissues maintained their composition under both conditions. When the fatty acids were calculated as absolute amounts, muscle composition, as well as that of the digestive gland, changed significantly. In the digestive gland, saturated FA (SFA), monounsaturated FA (MUFA), diunsaturated FA (DUFA) and polyunsaturated FA (PUFA) all increased up to the middle of the moulting cycle and then declined; with starvation all groups decreased. In muscle, SFA, MUFA and DUFA all increased during the moulting cycle; starvation caused SFA, MUFA and PUFA to decrease, whereas DUFA did not vary. Starvation caused both 18:2n-6 (linoleic) and 18:3n-3 (linolenic) in the digestive gland to reach or almost reach zero. The other essential PUFA, 20:4n-6 (arachidonic), 20:5n-3 (eicosapentaenoic) and 22:6n-3 (docosahexaenoic), decreased during the moulting cycle, but during starvation 20:4n-6 did not decrease as much. In muscle, the levels of 18:2n-6 and 18:3n-3 increased, while 20:4n-6, 20:5n-3 and 22:6n-3 remained approximately constant during the moulting cycle. Starvation reduced 20:5n-3 and 22:6n-3 to about 60%. The data suggest that levels of 18:3n-3, 20:4n-6, 20:5n-3 and 22:6n-3 are regulated, and that 20:4n-6 can be synthesised from 18:2n-6. There is no clear evidence that 20:5n-3 and 22:6n-3 are essential in P. esculentus, but tissue catabolism of cell membranes during starvation may have provided sufficient amounts for maintenance.     

Phytoplankton food quality determines time windows for successful zooplankton reproductive pulses

Recruitment success at the early life stages is a critical process for zooplankton demography. Copepods often dominate the zooplankton in marine coastal zones and are prey of the majority of fish larvae. Hypotheses interpreting variations of copepod recruitment are based on the concepts of "naupliar predation," "nutritional deficiency," and "toxic effect" of diatom diets. Contradictory laboratory and field studies have reached opposite conclusions on the effects of diatoms on copepod reproductive success, blurring our view of marine food-web energy flow from diatoms to higher consumers by means of copepods. Here we report estimates of copepod feeding selectivity and reproduction in response to seasonally changing phytoplankton characteristics measured in a highly productive coastal upwelling area off the coast of central Chile. The variable phytoplankton diversity and changing food quality had a strong and highly significant impact on the feeding selectivity, reproduction, and larval survival of three indigenous copepod species. Seasonal changes in copepod feeding behavior were related to the alternating protozoan-diatom diets, mostly based on dinoflagellates and ciliates during winter and autumn (low highly unsaturated fatty acids [HUFA]/polyunsaturated fatty acids [PUFA] availability), but switched to a diet of centric and chain-forming diatoms (high HUFA/PUFA availability) during the spring/summer upwelling period. Ingestion of diatom cells induced a positive effect on egg production. However, a negative relationship was found between egg hatching success, naupliar survival, and diatom ingestion. Depending on the phytoplankton species, diets had different effects on copepod reproduction and recruitment. In consequence, it seems that the classical marine food web model does not apply to some coastal upwelling systems.     

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.     

Lipids and fatty acid composition of particulate matter in the North Atlantic: importance of spatial heterogeneity,season and community structure

Lipid class profiles and total fatty acid composition of particulate matter were studied in the northeast Atlantic during the spring bloom and fall. Eddies of known physical and chemical properties were sampled at different depths. HPLC pigment data were used to characterize the phytoplankton communities. In spring, a dominance of prymnesiophytes was recorded at all depths, while in fall prochlorophytes dominated near the surface and prymnesiophytes only at deep chlorophyll maximum. Lipid classes included triglycerides, sterols, glycolipids and phospholipids. A differential relationship between phytoplankton abundance and lipid accumulation was observed: spring lipid concentrations were positively related to phytoplankton biomass, while fall particulate lipid did not show any relationship. The main feature was a northward increase in lipid concentrations unrelated to the mesoscale hydrological structures. Polar lipids dominated over neutral acyl-glycerols with phospholipids dominating over glycolipids in spring, while glycolipids dominated in fall. This resulted from different nutrient availability with a dominance of flagellates associated with mesotrophy in spring and of picophytoplankton associated with oligotrophy in fall. In terms of fatty acids, factorial correspondence analyses illustrate the influence of seasonally changing assemblages: (1) in spring, the main source of variability was the bloom with an opposition between bloom sites characterized by n-3 and n-6 PUFA, and more detrital deep samples characterized by saturated, monoenoic and branched acids; (2) fall fatty acid profiles were similar at all depths and very close to those observed for spring deep samples. Comparison of pigment and fatty acids using redundancy analysis suggested that pelagophytes were linked to saturated and branched acids. It also showed that prymnesiophytes and prochlorophytes were significantly associated with n-6 and n-3 PUFA. The spring period illustrated the complexity of these relationships with dinoflagellates and prymnesiophytes linked with n-3 PUFA, diatoms linked with palmitoleic and myristic acids, and pelagophytes linked with n-6 PUFA and higher-chain-length monoenes.     

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.     

δ13C and fatty acid composition of mesopelagic fishes in the South China Sea and their influence factors

Study of the ecology of mesopelagic fishes is central for assessing the active biological pump in the ocean, especially in the mesopelagic layers. The use of δ¹³C and fatty acid analysis can help to analysis the ecology of mesopelagic fishes. Here, we analysed the fatty acid composition of mesopelagic fishes from the continental northern slope of the South China Sea (NSSCS) and compared with nearshore SCS fishes and mesopelagic fishes collected from the Southern Ocean. The mesopelagic fishes had unusually high lipids, which resulted in Δδ¹³C values exceeding 1‰, more than the enrichment factor in the food web. The mesopelagic fishes had higher C18:1n-9/C18:1n-7 and C20:1n-9/ C18:1n-7 ratios compared with other fishes in the SCS, which confirmed that plankton were their main dietary source. The mesopelagic fishes from SCS and Southern Ocean had different ratios of C20:5n-3/C22:6n-3 (EPA/DHA), suggesting geographical locations and diet sources had obvious influence on their fatty acid composition. The SCS mesopelagic fishes had higher C20:4n-6/C22:6n-3 (ARA/DHA) and C20:4n-6/C20:5n-3 (ARA/EPA) ratios than mesopelagic fishes in the Southern Ocean, indicating the influence of physical factors on fatty acid composition. Thus, future studies of the fatty acids in mesopelagic fishes should consider both dietary sources and physical environments.     

Inter-individual and within-brood variability in the fatty acid profiles of Norway lobster, <Emphasis Type="Italic">Nephrops norvegicus</Emphasis> (L.) embryos

The present study investigated the existence of inter-individual and within-brood variability in the fatty acid (FA) profile of developing embryos of Nephrops norvegicus. In all surveyed females (n = 5), the quantitatively most important FAs were as follows: 22:6n-3 (20.8 ± 3.9% average of total FA ± standard error), 18:1n-9 (19.5 ± 2.0%), 16:0 (15.2 ± 3.4%), 20:5n-3 (10.2 ± 1.4%), 16:1n-7 (8.9 ± 1.6%), and 18:1n-7 (5.7 ± 1.3%). Differences in FA profiles of embryos in the same clutch were assessed using brooding chamber side (left and right) and pleopod (1st and 2nd, 3rd and 4th, and 5th) as predictive factors. There were no significant differences in the FA composition of embryos sampled from both sides of the brooding chamber in 4 of the 5 surveyed females. However, all females exhibited significant differences in the FA profiles of embryos sampled from different pleopods. Both saturated FA (SFA) and highly unsaturated FA (HUFA) present in developing embryos exhibited marked differences along the breeding chamber. Overall, FA reserves appeared to vary significantly within broods, which can ultimately be reflected on early larval survival. A potential cause for the within-brood variation recorded in the FA profile of developing embryos include (1) differential female investment during ovarian maturation, mainly due to variation in food quality/availability; (2) differential lipid catabolism during the incubation period of developing embryos, as a consequence of embryos position within the female’s brooding chamber; or (3) differential female investment during ovarian maturation amplified by differential lipid catabolism during the incubation period.     

A study of the variations in lipid levels,lipid class composition and fatty acid composition in the first stages ofArtemia sp.

The lipid class composition and the fatty acid composition of total lipids of the cysts, newly hatched nauplii and 24-h-old metanauplii of a Spanish parthenogenetic diploid strain ofArtemia sp. were studied. Substantial differences in the total lipid level occurred among these stages, with a marked increase from the cyst to the nauplii being followed by a decrease in the metanaupliar stages. This variation affected the absolute levels (mg/g dry wt) of the total lipid classes and individual fatty acids, although the percent composition of the fatty acids in total lipid was essentially unchanged. An exception occurred during hatching in that the percentages of 16:0 and 16:1n-7 in total lipid decreased whereas that of 20: 5n-3 increased. The lipid classes showed higher variation than the fatty acids both in absolute and in relative terms, and in particular, the ratio of phosphatidylcholine:phosphatidylethanolamine decreased progressively from cysts to nauplii and metanauplii. The implications of these findings for the use ofArtemia sp. as a larval feed in aquaculture are considered.     

The ecological significance of lipid/fatty acid synthesis in developing eggs and newly hatched larvae of Pacific cod (Gadus macrocephalus)

The lipid/fatty acid composition of marine fish eggs and larvae is linked with buoyancy regulation, but our understanding of such processes is largely restricted to species with pelagic eggs. In this study, we examined developmental changes in the lipid/fatty acids of eggs and embryos of Pacific cod (Gadus macrocephalus), a species that spawns demersal eggs along coastal shelf edges, but as larvae must make a rapid transition to the upper reaches of the water column. Adult Pacific cod were collected in the Gulf of Alaska during the spawning season and eggs of two females were artificially fertilized with sperm from three males for each female. The eggs were subsequently reared in the laboratory to determine (1) how lipids/fatty acids were catabolized during egg and larval development, and (2) whether lipid/fatty acid catabolism had measurable effects on egg/embryo density. Eggs incubated at 4°C began hatching after 3-weeks and continued to hatch over a 10-day period, during which there was a distinct shift in lipid classes (phospholipids (PL), triacyglycerols (TAG), and sterols (ST)) and essential fatty acids (EFAs: 22:6n-3 (DHA), 20:5n-3 (EPA), and 20:4n-6 (AA)). In the egg stage, total lipid content steadily decreased during the first 60% of development, but just prior to hatch we observed an unexpected 2–3-fold lipid increase (~6–9 μg individual⁻¹) and a significant drop in egg density. The increase in lipids was largely driven by PL, with evidence of long-chained fatty acid synthesis. Late-hatching larvae had progressively decreasing lipid and fatty acid reserves, suggesting a shift from lipogenesis to lipid catabolism with continued larval development. Egg density measures suggest that lipid/fatty acid composition is linked to buoyancy regulation as larvae shift from a demersal to a pelagic existence following hatch. The biochemical pathway by which Pacific cod are apparently able to synthesize EFAs is unknown, therefore representing a remarkable finding meriting further investigation.     

Changes in lipid class and fatty acid contents in the ovary and midgut gland of the female fiddler crab Uca tangeri (Decapoda,Ocypodiadae) during maturation

Changes in biochemical composition, lipid class and fatty acid contents were studied in the ovaries and midgut glands of the fiddler crabs Uca tangeri Eydoux during maturation. Wild females were caught during spring and early summer of 1992 in the Bay of Cádiz (southwest Spain), near the mouth of the San Pedro river. Protein and total lipid contents in the ovaries increased significantly from Stages III to IV, at the expense of total carbohydrate, which showed a large decrease during the same period. In the midgut gland, the protein content did not present any significant variation, whereas total lipids and total carbohydrates presented opposite up and down trends during maturation. In the ovary, total polar lipids increased significantly during the final phase of maturation (Stages III to IV), mainly due to the significant contribution of the phosphatidylcholine and phosphatidylethanolamine fractions. In contrast, total neutral lipids showed an upward trend throughout the whole maturation period, mainly due to significant increases of the triacylglycerol fraction. In the midgut gland, total polar lipids (mainly phosphatidylcholine) and total neutral lipids (mainly triacylglycerol) presented significant decreases from Stages II to III, the phase which preceded major increases in both polar and neutral lipids in the ovaries. Cholesterol content did not vary during maturation in either organ, in the ovary or midgut gland. Major fatty acids in the ovaries [16:0, 16:1 (n-7), 18:1 (n-9), 18:1 (n-7), 18:2 (n-6), 18:3 (n-3), 20:4 (n-6), 20:5 (n-3) and 22:6 (n-3)] did, however, accumulate significantly at later stages of maturation. It is noteworthy that arachidonic acid [20:4 (n-6)] content remained constant during all stages of maturation but decreased significantly in total polar lipids in the later phases of maturation. In contrast, eicosapentaenoic acid [20:5 (n-3)] increased significantly in all lipid fractions in the later stages, and docosahexaenoic acid [22:6 (n-3)] remained constant in the polar lipids and increased during later stages in the triacylglycerol fraction. Major fatty acids in the midgut gland lipids showed significant decreases from Stages II to III, just before the final period of maturation.