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.     

Lipogenesis in the arctic euphausiid Thysanoessa inermis

Biosynthesis of lipids by Thysanoessa inermis collected from Balsfjorden, northern Norway, in May 1980, was examined in vitro. The highest concentration of lipid within the krill was in the hepatopancreas, and this organ was the most active in esterifying free fatty acids into wax esters. The hepatopancreas (i.e., thoracic contents) incorporated (¹⁴C) glucose, (¹⁴C) alanine and ³H₂O into wax esters, with the fatty alcohol moieties being labelled more than the fatty acids. (¹⁴C) fatty acid was incorporated preferentially into the fatty acid moieties of wax esters, this incorporation being markedly stimulated by free fatty alcohol. It is concluded that the fatty alcohols of wax esters are preferentially biosynthesized de novo from dietary protein and carbohydrates, whereas the fatty acids derive preferentially from dietary lipid. On the basis of ³H incorporated from ³H₂O, the hepatopancreas in a 50 mg II-group (2 yr old) individual of T. inermis is capable of biosynthesizing de novo, approximately 0.1 mg of lipid (as fatty acids) per day at 5°C.     

The composition and biosynthesis of lipids in Thysanoessa raschi from the Clyde Estuary,Scotland

The lipid composition and biosynthesising activity of Thysanoessa raschi collected from the Clyde Estuary, Scotland, in May 1981 were examined. Triacylglycerols were the major lipid class present, although 16.7% of the total lipid were wax esters in which phytol was the dominant fatty alcohol. The thoracic contents (hepatopancreas) of the krill were capable of biosynthesising lipids in vitro from various labelled substrates. Radioactivity from [1-¹⁴C] palmitic acid was incorporated into lipids in the order phospholipids>triacylglycerols>wax esters; the bulk of the radioactivity was present in all cases in the fatty acyl moieties of the lipids. [U-¹⁴C] glucose labelled lipids in the order phospholipids>triacylglycerols>free fatty acids> was esters; in the first two lipids the radioactivity was mainly in the glycerol moieties, whereas in was esters it was solely in the fatty acyl moieties. The extent of labelling of these lipids from [U-¹⁴C] alanine was less than that from [U-¹⁴C] glucose, but the pattern of labelling was generally similar. More than 90% of the radioactivity incorporated into total lipid from ³H₂O was present in free fatty acids from which it was calculated that the hepatopancreas of T. raschi can synthesise 2.5 g of fatty acid per hour at 15°C. This value is approximately three times lower than that previously determined for T. inermis from Balsfjorden, northern Norway. The results are discussed in terms of the sources of the dietary lipids of krill and the role of endogenous biosynthesis in contributing to its lipid reserves.     

Ecology of the euphausiidsThysanoessa raschi,T. inermis andMeganyctiphanes norvegica in Ísafjord-deep,northwest-Iceland

The seasonal abundance, distribution, maturity, growth and population dynamics of the euphausiidsThysanoessa raschi (M. Sars, 1864),T. inermis (Krøyer, 1846) andMeganyctiphanes norvegica (M. Sars, 1857) were studied in Ísafjord-deep, a fjord in northwest Iceland, from February 1987 to February 1988. Sampling was made at nine stations along the length of the fjord at approximately monthly intervals, along with hydrographic measurements and water sampling for nutrient analysis and measurements of chlorophylla concentrations. Spring warming of the water began in late May and maximum temperatures (8° to 10°C) were observed in late July–September. The phytoplankton spring-bloom started in early April, and the highest chlorophylla levels were measured in early May (7.0 mg m^–3). A small increase was observed in the chlorophylla content in August. The greatest abundance of juveniles and males and females of all three species was observed during January and February 1988, during which period the euphausiids were concentrated in the middle and inner parts of the fjord. Euphausiid eggs were first recorded in the plankton in mid-May, and the greatest abundance ofThysanoessa spp. larvae occurred at the end of May. Larvae ofM. norvegica were not observed in Ísafjord-deep, indicating that recruitment of this species was occurring from outside the fjord.T. raschi andT. inermis had a life span of just over 2 yr; the life span ofM. norvegica was more difficult to determine. Almost all femaleT. raschi were mature at the age of 1 yr, while mostT. inermis females appeared not to mature until 2 yr of age. Most males of both species took part in breeding at 1 yr of age. The maximum carapace length ofT. raschi andT. inermis was 8 to 9 and 9 to 10 mm, respectively. The largestM. norvegica had a carapace length of 9 to 10 mm. The spawning of the euphausiids in Ísafjord-deep appeared to be closely related to the phytoplankton spring bloom; water temperature appeared to have no influence on spawning.     

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.     

Wax-ester metabolism in the orange roughyHoplostethus atlanticus (Beryciformes: Trachichthyidae)

Freshly caught orange roughy (Hoplostethus atlanticus) obtained at the edge of the Chatham Rise, east of the South Island of New Zealand, in June 1988, were used to investigate tissue-specific lipid synthesis and to obtain blood for plasma lipoprotein analysis. Tissue slices prepared from an intraneurocranial fat deposit, pieces of skull (neurocranium), swimbladder and liver were incubated with radioactive acetate, palmitate and oleate, and the labelled lipids extracted and analysed. All four tissues could incorporate acetate label into fatty acids which were themselves incorporated into triacylglycerols and phospholipids although not into wax esters, the most abundant lipid class in these tissues. Exogenous palmitate and oleate were also incorporated into more complex lipids, with the label from oleate (but not palmitate) being found in wax esters as well as triacylglycerols and phospholipids. The distribution of the label in the wax esters showed that some reduction of the fatty acid to fatty alcohol had occurred. The only exception to this pattern was the swimbladder, which incorporated small amounts of palmitate-label into both the fatty acid and fatty alcohol portions of the wax esters. Lipoproteins were isolated from serum by centrifugation. All lipoprotein classes contained phospholipid, cholesterol, triacylglycerol, cholesteryl esters and wax esters. A very low-density lipoprotein class also contained large amounts of unesterified fatty acids, which are possibly artefacts resulting from storage of the samples. Significant quantities of-3 fatty acids were also found in the serum lipids.     

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.     

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.     

Carbon assimilation and lipid production in phytoplankton in northern Norwegian fjords

Carbon assimilation and lipid production were studied in phytoplankton in Balsfjorden and Ullsfjorden, northern Norway, during the exponential growth phase of the spring bloom in 1983 (6–7 April). In Balsfjorden, phytoplankton biomass was constant with depth and equivalent to 1.5 g chlorophyll a 1^-1. Phytoplankton biomass in Ullsfjorden varied with depth, with a maximum of ca. 7 g chlorophyll a 1^-1 occurring at 5 to 10 m. Particulate carbon-14 assimilation was about 18 mg C per m^-2 h^-1 in Balsfjorden and about 39 mg C per m^-2 h^-1 in Ullsfjorden over the depth range 4 to 8 m. In Balsfjorden, the percentage of total fixed carbon recovered as total lipid was 14.7 and 20.4% at 4 and 8 m depth, respectively. In Ullsfjorden, the corresponding values were 8.8 and 28.1% at 4 and 8 m, respectively. The percentages of total fixed carbon present as fatty acids were 1.1 and 1.6% at 4 and 8 m, respectively, in Balsfjorden, and 0.8 and 6.4% at 4 and 8 m in Ullsfjorden. The majority of the radioactivity in lipid at both locations and at both depths was present as polar lipid, with small percentages present in triacylglycerols and very small percentages present in free fatty acids. On average, about 18% of the total carbon-14 incorporated into phytoplankton over a 6 to 7 h mid-day period was recovered as total lipid and its percentage tended to increase with depth. The relatively low percentage of incorporated carbon-14 present as fatty acids in total lipid implies that most of the radioactivity is present in glyceryl and/or glucosyl moieties and that measurement of total radioactivity in total lipid does not necessarily give an accurate estimation of lipogenesis in phytoplankton. Fatty acid analyses of total phytoplankton in Balsfjorden and Ullsfjorden in 1983 and of a surface slick at the end of a bloom of Phaeocystis pouchetii in Balsfjorden in May 1980 showed an abundance (more than 40% of the total) of (n-3) polyunsaturates in all cases. C-18 polyunsaturates, especially 18:4 and 18:5, were very abundant (about 30% of the total) in the P. pouchetii surface slick in Balsfjorden in 1980. Both P. pouchetii biomass and C-18 polyunsaturates were more abundant in Ullsfjorden than in Balsfjorden (1983). Lipids from the P. pouchetii surface slick were deficient in C-16 polyunsaturates and relatively deficient in C-20 polyunsaturates, but both these classes of fatty acids were abundant in Balsfjorden and Ullsfjorden in 1983. The phytoplankton in both locations in 1983 was dominated by P. pouchetii and diatoms; Chaetoceros socialis was especially abundant in Balsfjorden. The results are discussed in terms of the fatty acids present in herbivorous zooplankton in northern Norwegian fjords.     

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.     

Distribution of lipids between the zooxanthellae and animal compartment in the symbiotic sea anemoneAnemonia viridis: Wax esters,triglycerides and fatty acids

The temperate sea anemoneAnemonia viridis (Forskäl) contained about 11% lipid on a dry weight basis when maintained at light levels of about 10µE m^–2 s^–1 and a temperature of 10°C. Aposymbiotic forms of the anemone had similar lipid levels. These values are very low compared with tropical symbiotic Anthozoa in which lipid levels constitute up to 50% of dry weight. In symbioticA. viridis, <6% of total lipid consisted of the storage lipids, wax esters and triglycerides. Most of the triglyceride was stored in the animal tissues rather than the zooxanthellae. Zooxanthellae contained only small amounts of wax esters. An analysis was made of the wax ester, triglyceride and fatty acid composition of symbiotic anemones, isolated zooxanthellae and aposymbiotic anemones. Wax ester composition was similar in symbiotic and aposymbiotic forms. However, triglyceride composition differed. In particular trimyristin (C42) was found only within the symbiotic association. Fatty acids showed a high degree of unsaturation, and acids with both even and odd numbers of carbon atoms were found. The most abundant fatty acid was 16:0 in all samples, except for the total lipids from zooxanthellae in which the major fatty acid wastrans-18:1.     

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.     

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     

Lipogenesis in the intact coral Pocillopora capitata and its isolated zooxanthellae: Evidence for a light-driven carbon cycle between symbiont and host

Surface tissue of the reef coral Pocillopora capitata contained approximately 34% lipid on a dry weight basis. Of this, 75% was storage lipid (wax ester and triglyceride) and 25% structural (phospholipid, galactolipid, etc.). Based on chlorophyll a: lipid ratios of intact coral and isolated zooxanthellae, it was determined that over 90% of the storage lipid resided in the host tissue. One half of the structural lipids was found in the host and the other in the symbiotic algae. Gentle fractionation of coral tissue indicated that zooxanthellae possessed less than 14% of the total coral protein. Coral tips and isolated zooxanthellae were incubated with sodium acetate-1-¹⁴C in light and dark to obtain lipogenic rates and proportions of fatty acids and lipid classes synthesized. The rate of lipid synthesis from acetate-1-¹⁴C by intact coral was stimulated three-fold in the light (1200 lux), which indicated that the majority of coral lipogenesis occurred in the zooxanthellae. Intact coral triglycerides contained ca. 68% of the ¹⁴C-activity and wax esters ca. 21%. Zooxanthellae isolated by the Water Pik technique synthesized negligible amounts of wax ester, which implied that wax ester synthesis was a property of the animal tissue. Isolated zooxanthellae and intact coral synthesized identical triglyceride fatty acids from acetate-1-¹⁴C. This study provides evidence for a carbon cycle between host and symbiont whereby the zooxanthellae take up host-derived carbon (probably in the form of acetate), synthesize fatty acids using their photosynthetically derived energy, and return the lipid to the host where it appears as wax ester and triglyceride.     

Biosynthesis of wax esters in oceanic crustaceans

Slices from the hepatopancreas of various oceanic curstaceans incorporated radioactivity into wax esters from ¹⁴C glucose and ¹⁴C aspartic acid to a lesser extent and from ¹⁴C palmitic acid to a much greater extent. Radioactivity was incorporated from ¹⁴C palmitic acid into both fatty acid and fatty alcohol moieties of wax esters, the percentage of total radioactivity present in alcohol moieties being greater in deep-living than in shallow-living species. Cell-free preparations from the hepatopancreas but not from muscle, supplemented with ATP and reduced pyridine nucleotides, incorporated radioactivity from 1-¹⁴C palmitoyl Coenzyme A into both fatty alcohol and fatty acid moieties of wax esters. Incorporation into fatty alcohol was NADPH- rather than NADH-specific. Preparations from deep-living species had a greater percentage of total radioactivity in the fatty alcohol moieties of wax esters than preparations from shallow-living species. We conclude that the level of wax esters in a given species is correlated with the rate at which the species biosynthesises these lipids de novo; deep-living species have higher rates of wax ester biosynthesis and higher levels of wax esters than shallow-living species. The results support the thesis that wax esters in oceanic crustaceans are derived largely from the animals' internal biosynthetic activities, presumably in response to particular biochemical and/or physiological requirements, rather than from their diets.     

Feeding ecology and energetics of the Antarctic chaetognaths Eukrohnia hamata, E. bathypelagica and E. bathyantarctica

Ecological and physiological studies focused on dietary preferences, lipid biochemistry and energetics within the three Antarctic chaetognaths Eukrohnia hamata, E. bathypelagica and E. bathyantarctica from meso- and bathypelagic depths. Eukrohnia hamata and E. bathypelagica respired 0.15 μL O₂ mg dry mass (DM)⁻¹ h⁻¹, which translates to an average metabolic loss of only <1.1% of body carbon per day. Lipid storage was not substantial in E. bathypelagica (mean 11.5 ± 6.5% DM) and E. bathyantarctica (mean 15.4 ± 4.1% DM) during summer and winter, suggesting year-round feeding of these predators mainly on copepods. In E. bathypelagica, total fatty acids were dominated by the fatty acids 16:0, 20:5(n-3) and 22:6(n-3) and in E. bathyantarctica also by 18:1(n-9), a fatty acid usually found in storage lipids. Only the latter species was characterized by significant amounts of wax esters, consisting largely of the common fatty alcohols 16:0, 20:1(n-9) and the unusual fatty alcohol isomer 22:1(n-9).     

Studies on the mechanism of biosynthesis of wax esters in Euchaeta norvegica

Preparations of Euchaeta norvegica catalyse the incorporation of (1-¹⁴C) hexadecanol, (9,10-³H₂) oleic acid, (U-¹⁴C) glucose, (U-¹⁴C) alanine and (U-¹⁴C) aspartic acid into wax esters. Both the fatty alcohol and fatty acid moieties of the wax esters are labelled, indicating de novo biosynthesis. A limited interconversion of fatty acid and fatty alcohol has been demonstrated. Incorporation of glucose into wax esters is diminished by prior starving of E. norvegica; it is also reduced in the presence of ecdysterone. Formation of wax esters from glucose is inhibited by increased oxygen tensions, rotenone and dinitrophenol, and is relatively insensitive to decreased oxygen tensions and cyanide. The data are discussed with particular reference to the reasons why wax esters are biosynthesised by calanoids.     

Assimilation of the wax esters of marine zooplankton by herring (Clupea harengus) and rainbow trout (Salmo gairdnerii)

Herring (Clupea harengus L.) and rainbow trout (Salmo gairdnerii) fed frozen-thawed zooplankton for 7 days assimilated more than 90% of the dry matter in the zooplankton consumed. Less than 5% of the total lipid ingested, largely wax esters, was excreted in the faeces. Faecal lipids from both the herring and the trout, especially the latter, were rich in wax esters and free fatty alcohols. Faecal fatty alcohols were enriched in 22:1 and deficient in 16:0 and 16:1 alcohols compared to zooplankton fatty alcohols. The growth of rainbow trout fed frozen-thawed zooplankton over 3 months was about one-half of that of trout fed commercial pellets. The fish fed on zooplankton appeared normal and were more pigmented than the fish fed on commercial pellets.     

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.     

Population dynamics and production of euphausiids II. Thysanoessa inermis and T. raschi in the North Sea and American Coastal Waters

Results from the Continuous Plankton Recorder (CPR) survey for 1966 and 1967 are used to describe seasonal changes in abundance, size and aspects of the population structure of Thysanoessa inermis (Krøyer) and T. raschi (M. Sars) at a depth of 10 m in the North Sea and in American coastal waters from the Grand Banks to the Gulf of Maine. Production and dry weight were estimated from these data. Two year-groups were usually present in the breeding population, the proportion surviving into a second year being higher in American waters than in the North Sea. Annual production for each species was within the range 0.69 to 4.66 mg m^-3 and the ratio between production and biomass (P:B) was between 1.3 and 4.2; values outside these ranges were obtained only for American coastal waters in 1967, when the frequency of sampling was low.