Lipid storage and composition in tropical surgeonfishes (Teleostei: Acanthuridae)

The lipid composition of tropical marine reef fishes is poorly known, despite their use as food by local human populations and recent interest in health-related benefits of fish lipids. We examined the composition of lipids from epaxial muscle, liver, and two storage sites [mesenteries surrounding the gut (intraperitoneal fat, IPFs) and retroperitoneal fat bodies (FBs) posterior to the peritoneal cavity] in three species of surgeonfishes from Ishigaki Island, Japan: Naso lituratus (Bloch and Schneider, 1801), Acanthurus lineatus (Linnaeus, 1758), and A. bariene (Lesson, 1830). Triacylglycerols dominated all samples of neutral lipid and constituted ≥ 99% of FBs and IPFs. Polar lipids generally contained large fractions of phosphatidylethanolamine and phosphatidylcholine. Quantified fatty acids ranged in length from C₁₄ to C₂₄. C₁₆ fatty acids prevailed (>35% of neutral fatty acids, >23% of polar fatty acids), although C₁₈ (>16 and >14%, respectively) and C₂₀ acids (>8 and >19%, respectively) were also common. Saturated fatty acids, dominated by palmitic acid (16:0), comprised 38.7 to 50.7% of acids from neutral lipids and 30.8 to 41.1% from polar lipids. The most common monounsaturated acids were 18:1n9 and 20:1n9. Polyunsaturated acids were prevalent in polar lipids (especially 20:4n6, 20:5n3, 22:2n3, 22:5n3, 22:5n6 and 22:6n3). Common polyunsaturated acids of neutral lipids were 18:2n6, 18:4n3, several n-3 and n-6 C₂₀ acids, 22:2n3 and 22:5n3. IPF and FB were almost identical across species, and lipids of fat bodies (IPFs, FBs) were more similar to those of muscle than those of liver for all three species. The FBs appear to constitute an accessory storage site, which overcomes constraints on lipid storage imposed by a small, inflexible abdominal cavity that contains both viscera and consistently voluminous gut contents. Fatty acid signatures indicate that largely overlooked epiphytic or epilithic diatoms contribute significantly to lipid acquisition. The combination of large quantities of both saturated and n-3 and other polyunsaturated fatty acids in surgeonfishes, in contrast to low saturates and high polyunsaturated acids in lipids of commercially important cool-water fishes, suggests that a study of dietary effects of fish lipids on human inhabitants of the tropics may be instructive insofar as human health and nutrition are concerned. Received: 16 March 1998 / Accepted: 6 August 1998     

Lipids in atlantic halibut (Hippoglossus hippoglossus) eggs from planktonic samples in Northern Norway

Fertilized Atlantic halibut (Hippoglossus hippoglossus L.) eggs in different developmental stages (Days 0 to 18) were sampled from plankton in North Norway in February 1986 and analysed for lipid classes and fatty acid content. In unfertilized ovulated eggs taken from ripe fish caught in 1983/1984, polar and neutral lipids comprised ca. 71 and 30% of the total lipids, respectively, decreasing and increasing to 67 and 33%, respectively, in Stage III (11 to 18 d old) fertilized eggs. Of the polar lipids, phosphatidylcholine decreased markedly from 62% in unfertilized ovulated eggs to ca. 40% in Stage I (0 to 3 d old) fertilized eggs, while phosphatidylethanolamine increased from ca. 7 to 33%. Triacylglycerols, the major neutral lipids, increased from ca. 13% in unfertilized ovulated eggs to 16% in Stage III fertilized eggs. The total lipid in Stage I fertilized eggs had relatively low levels of polyunsaturated fatty acids (PUFA), with (n-3) PUFA accounting for only ca. 25% of the total fatty acids. The (n-3) PUFA increased to ca. 40% of the total fatty acids in Stage III, while the (n-3): (n-6) ratio increased from 4.1 to 7.0.     

Variation in lipid classes during the molting cycle of the prawn Penaeus japonicus

The variation in the concentration and fatty acid composition of lipid classes during the molting cycle of the prawn Penaeus japonicus was investigated. The lipid concentration of the whole body reached a maximum at mid-premolt (Stage D₂) and then decreased to low level at late premolt (Stage D_3–4). The accumulation of lipids during the premolt period seemed to be attributable to the increase of both polar and neutral lipids. The increase of neutral lipids at Stage D₂ was derived from not only triglycerides but also free sterols and free fatty acids. Regarding the fatty acid composition of every lipid class, a marked variation occurred mainly at the intermolt (Stage C). In this stage, the polar lipids were rich in monoenoic acids such as 18:1 and poor in polyenoic acids such as 20:53 and 22:63. The triglycerides were rich in polyenoic acids at Stage C, but poor in monoenoic acids such as 16:1 and 18:1. The steryl esters contained large amounts of saturated acids such as 16:0 and 18:0 throughout the molting cycle, however the level of polyenoic acids increased at Stage C.     

Differential utilization of biochemical components during larval development of the spider crab Maja brachydactyla (Decapoda: Majidae)

Changes in lipid class, fatty acid, fat-soluble vitamins, amino acid and minerals were studied during larval development of Maja brachydactyla Balss, 1922 in order to provide information of its ontogeny and evaluate possible dietary deficiencies and constraints. Four different batches were analyzed from hatching to metamorphosis using enriched Artemia as food. Cultured larvae were in a good nutritional condition as confirmed by the continuous lipid accumulation throughout ontogeny. A regulation of polyunsaturated fatty acids (PUFA) occurred during development in order to maintain adequate basal levels of ω-3 fatty acids (likely through a retro conversion of C₂₀ and C₂₂ into C₁₈ PUFA). Variations in the tocopherol levels indicated its possible role in PUFA protection against oxidation. Essential amino acid balance during development was not correlated with the Artemia feeding regime, but rather reflected inherent variations of the own species ontogeny. Larval requirements in essential minerals were fully supplied by the enriched Artemia.     

Larval growth and settlement of the European oyster (Ostrea edulis) as a function of food quality measured as fatty acid composition

Larval growth rate and settlement of the European flat oyster Ostrea edulis were experimentally studied as a function of the composition of dietary fatty acids. Diets differing in fatty acid composition were composed by mixtures of the microalgae Isochrysis galbana, Pavlova lutheri and Chaetoceros calcitrans. Fatty acid content in the tissue of the feeding larvae, analyzed by gas chromatography and mass spectrometry, reflected the composition in the diet. Larval growth rate was significantly correlated to the three omega-3 polyunsaturated fatty acids (PUFA) C18:3, C18:4 and C22:6, with minor differences for neutral and polar lipids. No relation between growth rate and the omega-3 PUFA C20:5 was detected, a PUFA often implied as essential for bivalves. It is suggested that naturally occurring variability in fatty acid composition may constrain larval growth. In settlement experiments in both still water and flume flow little substrate selectivity was found for some contrasting substrates. It is concluded that differences in dietary fatty acids may explain as much of settlement success as the variability of substrates. Received: 12 October 1998 / Accepted: 6 April 1999     

Studies on the lipid metabolism of some oceanic crustaceans

Mid-water, oceanic crustaceans were either fed food labelled with (¹⁴C) palmitic acid under controlled conditions, or injected directly with (¹⁴C) palmitic acid. Lipid classes and their constituent fatty acids and fatty alcohols were subsequently separated and assayed for radioactivity. Significant levels of radioactivity were present in the 16:0¹ alcohols and 18:1 acids of wax esters, and in the 16:0/16:1, 18:0/18:1, 20:5 and 22:6 acids of both triglycerides and phospholipids. It was concluded that these crustaceans were capable of biosynthesis of wax esters and higher polyunsaturated fatty acids.     

Sources and major biogeochemical pathways of organic matter in the mangrove system of Rufiji estuary

Fatty acids were extracted from the surface sediments (10 cm) of three sampling sites of Rufiji estuary to infer their sources and biogeochemical pathways. The fatty acids ranging from C₈ to C₂₄ were distinguished from this study, and were broadly classified into saturated (SFAs), monounsaturated (MUFAs) and polyunsaturated fatty acids (PUFAs). SFAs were found to be the major fraction at station 1 and 3 where as at station 2, MUFAs dominated. A total of 19 fatty acids with a total concentration of 64.91 μg/g dry weight were characterised at station 1. C_16:0 was the most abundant fatty acid contributing 21.94% of total fatty acids (TFAs). C_22:2 was the second most abundant, which accounted for 9.46% of TFAs. Fatty acids ranging from C₁₂ to C₂₄ were identified at station 2. C₂₀ fatty acid was the most abundant fatty acid contributing 21.94%, followed by C_16:0. At station 3, fatty acids ranging from C₈ to C₂₄ were obtained. The PUFA C_20:5n-3 was the most abundant fatty acid contributing 21.65%, followed by C_24:0 (15.00% of TFAs). The ratio of lower molecular weight (LMW) to higher molecular weight (HMW) biomarkers was used as an indicators to distinguish higher plants organic matter from algae-derived fatty acids.     

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.     

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     

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.     

Lipid and fatty acids in naturally occurring particulate matter during spring and summer in a high arctic fjord (Kongsfjorden, Svalbard)

The total lipids and fatty acid composition of natural particulate matter and nutritional quality for zooplankton grazers was studied on a seasonal basis in the Arctic fjord Kongsfjorden (Svalbard) during the spring, summer of 2007 and during the early summer of 2006. Both years were abnormally warm, and the study attempted to evaluate the potential impact of the intrusion of North Atlantic waters. Samples were collected in surface layers and at deep chlorophyll maximum (DCM when present). Both years, chlorophyll concentrations were low (<2 μg L^?1) even during bloom periods. Species determination indicated Phaeocystis spp. as main constituent of the May bloom while ciliates and flagellates dominated the rest of the survey period. Total lipids showed similar changes at both depths with maximum values in mid-summer of 2007, while it showed reverse patterns between surface and DCM in 2006. Total fatty acid composition was dominated by saturates and monoenoic acids at both depths with significant percentages of pentaenoic acids and 22:6n-3 (DHA) recorded at all times. The 2007 fatty acid dynamics identified four main successions in term of particulate assemblage related mainly to the succession of living cells versus detrital material and to a lesser extent to phytoplankton community changes (diatoms versus non-diatoms). Redundancy analysis confirmed that live phytoplankton is one of the main drivers in the fatty acid changes. Temperature and density of the surface water are also influential in relation to water mass dynamics. Concentrations of fatty acids available to consumers showed n-3 PUFA ranging from 2 to 15 μg L^?1 and n-6 PUFA ranging from 0.3 to 2 μg L^?1. Concentration of EPA (20:5) and DHA are potentially limiting, suggesting a negative impact of Phaeocystis pouchetti-type phytoplankton linked to advection of Atlantic waters in relation to global warming of Arctic waters.     

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.     

Metabolism of palmitic,linoleic, and linolenic acids in adult oysters,Crassostrea virginica

This study investigated incorporation and metabolism of saturated [(1-¹⁴C) 16:0] and unsaturated [(1-¹⁴C) 18:26 and (1-¹⁴C) 18:33] fatty acids in adult eastern oysters,Crassostrea virginica Gmelin (spawned from parents obtained in 1986 from Mobjack Bay, Virginia, USA), and the influence of temperature on these processes. InC. virginica, incorporation of injected palmitic (16:0) and linolenic (18:33) acids was increased when oysters which had been grown in warm water (22 to 23°C) were transfered to cold water (5 to 7°C) for 8 to 18 d. Incorporation of linoleic acid (18:26) was unchanged under these conditions. The changes in concentration may have been linked to depression of metabolism in these oysters, in particular that of 16:0, which was reduced by 90%. Oxidation of incorporated fatty acids was much higher in warm than in cold water. Cold-temperature conditioning ofC. virginica altered the distribution of fatty acids among the neutral and polar lipid fractions. Long-term exposure to cold water increased the proportion of fatty acids in the polar fraction, which may be related to maintenance of membrane fluidity. Short-term exposure to cold water had the opposite effect, which may be due to increased energy requirements as the oyster adapts to new conditions. Reutilization of¹⁴C-acyl groups demonstrated de novo synthesis of 16:0 and 18:0 fatty acids. Only limited elongation and no desaturation of the administered fatty acids was observed.     

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.     

Variations saisonnières de la composition en acides gras chez Penaeus japonicus (Crustacea: Decapoda)

The lipid and fatty acid composition of male and female Penaeus japonicus Bate was investigated over a period of 11 months to determine the effects of environmental and nutritional factors such as temperature and starvation. The lipid content of the prawns increases from May to November, whereas water content decreases. We observed a time-lag between the highest lipid rate of males (in November) and that of females (in October); this is probably related to ovarian metabolism. From September to March, the high lipid content of the females coincided with a high percentage of monounsaturated fatty acids (especially 16:1 and 18:1). Winter starvation induces a decrease in palmitic acid content. As temperature decreases, the monounsaturated (16:1, 18:1) and polyunsaturated (20:4, 20:5, 22:6) fatty acids increase at the expense of the saturated fatty acids (16:0, 18:0) in the adult prawn. However, in post-larvae and juveniles, this increased rate of polyunsaturated fatty acids would be related to their particular lipid metabolism.     

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.     

Variation in the lipid content of wild-caught females of the marine shrimpPenaeus kerathurus during sexual maturation

Changes in total lipids, lipid classes and their fatty acid contents were studied in the ovaries and midgut glands ofPenaeus kerathurus Forskäl females during sexual maturation. The shrimp were captured in the Gulf of Cádiz (southwest Spain) in 1990. The lipid content and fatty acids, in relative terms, increased during ovarian development. The greatest changes occurred between Maturation Stages III and IV. Ovarian lipids were dominated by polar classes, whereas in the midgut gland the major classes were triacylglycerols and sterol esters. The amounts of major fatty acids in ovaries (16:0, 16:1n-7, 18:1n-9, 18:1n-7, 20:5n-3 and 22:6n-3) increased with increasing maturity, but declined slightly between Stages III and IV. The total polar lipid content of the midgut was 5.7% (by dry weight) and its fatty acid composition remained constant during the whole study period. Total lipid content of the midgut gland showed an upward trend during sexual maturation, except between Stages II and III, when a slight decrease was observed. Predominant fatty acids in the midgut gland (16:1n-7, 20:5n-3 and 22:6n-3) displayed a noteworthy decline between Stages II and III, corresponding with the marked increase in total lipid fatty acid content in the ovaries during the same period.     

The Mediterranean intertidal habitat as a natural laboratory to study climate change drivers of geographic patterns in marine biodiversity

Nylon bags containing yellow leaves of Rhizophora apiculata and Avicennia marina, were immersed for 80 days from August to October, 1996. the decomposing leaves were collected every 10 days and analysed for dry weight loss and six biochemical parameters: tannins, total amino acids, total sugars, total nitrogen, total lipids and fatty acid profile. the leaf weight initially decreased very rapidly by about 50% of the start in two species of mangroves within 10 days. Similar changes were observed with tannins, total amino acids and sugars. However, the concentration of nitrogen increased significantly with decomposition. There was no significant change in total lipid and fatty acid profile. the highest concentration of fatty acid in the decomposing leaves was palmitic acid (16:0). Unsaturated fatty acids such as, 18:1 w7c and 18:1 w9c were found to be present in decomposing leaves of both species.     

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.     

Lipids and fatty acids of the benthic marine harpacticoid copepod<Emphasis Type="Italic"> Heteropsyllus nunni</Emphasis> Coull during diapause: a contrast to pelagic copepods

Many free-living copepods produce and store lipids prior to entering diapause (long-term dormancy). Heteropsyllus nunni Coull is the only marine harpacticoid copepod known to undergo any form of diapause. This study presents the first information on the types of lipids and fatty acids produced for long-term diapause in this benthic species. Sexually immature adults of H. nunni undergo diapause within a pliable self-made cyst. Prior to entering diapause (which lasts 3–4 months), they produce and store large amounts of orange lipid. The lipids apparently are utilized during diapause. Although some residual lipids remain, chiefly around the gonads, after the copepods emerge from their cysts, the lipid stores are visibly reduced. Typically, the copepods mate and produce eggs within 48 h after diapause is terminated. Light level and confocal laser scanning microscopy revealed that the lipid stores are distributed throughout the body in numerous oil droplets and not as a single oil sac, as seen in many marine calanoid copepods prior to overwintering (winter diapause). Transmission electron microscopy showed lipid spheres within the gut epithelium and large droplets of lipids stored extracellularly. Confocal laser scanning microscopy of copepods in pre-diapause, during diapause (encysted), post-diapause (recently excysted), and in reproductive condition, revealed that lipid stores are reduced following diapause, but are not totally absent. Analysis of lipid classes showed that H. nunni store predominantly wax esters/sterol esters (83% of total lipids) during diapause. The predominant lipid is most likely wax esters, as sterol esters typically are found only in small amounts in copepods. Fatty acid (FA) profiles of the copepods in diapause showed 16:0 to be most abundant followed by 16:1n-7 and 18:0; other FA occurred at concentrations <10% of total FA. Three polyunsaturated fatty acids (PUFA), 20:5n-3, 18:2n-6 and 20:4n-6, were found at concentrations <2% of total FA. These PUFA are "essential fatty acids" in H. nunni, obtained through dietary sources. The lipid classes and fatty acids present in H. nunni during diapause are compared to those of other copepods, some in a state of diapause and others not. It appears that lipid class and FA profiles are indicative of genetic makeup, type of diet or amount of food consumed prior to dormancy. Some classic paradigms of lipids and their association with copepod diapause are re-evaluated.Communicated by P.W. Sammarco, Chauvin