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     

Larval distribution and reproductive ecology of Thysanoessa macrura (Crustacea: Euphausiacea) in the Scotia Sea

In the Scotia Sea, the larvae of Thysanoessa macrura G.O. Sars, 1885 were scarce in spring (September 27 to October 6, 1971) and mainly present at the Calyptopis I stage, while in summer (December 17, 1974 to January 3, 1975) they were much more abundant and much further developed (up to Furcilia VI). The spawning season commences as early as September and, as indicated by the presence of numerous nauplii and metanauplii in summer, is rather long. T. macrura begins to spawn much earlier in West-Wind-Drift waters than in waters of the Weddell Sea. It starts to spawn simultaneously with Euphausia frigida and E. triacantha, but significantly earlier than E. superba. These timings of the breeding process are related to differential biogeographical characteristics of these species. Plankton collections made at depths of 500 or 1000 m have, for T. macrura, revealed a developmental ascent analogous to that of E. superba. Ontogenetical deep-water migrations are, consequently, characteristic not only for E. superba, but also for other Antarctic euphausiids (in particular E. frigida, but possibly not for inshore E. crystallorophias) and, as correctly postulated by several other investigators, for all oceanic euphausiid species.     

Antarctic shallow-water mega-epibenthos: shaped by circumpolar dispersion or local conditions?

The mega-epibenthos of two different geographic areas, the Antarctic Peninsula and the high Antarctic (eastern Weddell Sea), were investigated using underwater video. The distribution of the marine fauna at shallow depths between 55 and 160 m in these two areas was investigated to determine whether there are any zoogeographic differences at the community level. A total of 237 taxa represented by 85,538 individuals was identified. Multivariate analyses revealed significant faunal differences between northern Marguerite Bay (western Antarctic Peninsula) and the stations from the Weddell Sea, Atka Bay and Four-Seasons Bank. Echinoderms, especially ophiuroids, dominated Marguerite Bay, bryozoans and ascidians were abundant at Atka Bay, and hydroids and gorgonians were well represented at Four-Seasons Bank. These clear differences can mainly be explained by the influence of local environmental conditions that are probably the primary feature responsible in shaping the Antarctic shallow-water epifauna and not an intensive exchange with larger depths or a limited dispersion due to scarce and isolated shallow areas. In addition, modes of reproduction and characteristics of the early life history (e.g. brooding, viviparity or budding) of key taxa may also shape patterns of species distribution in shallow benthic Antarctic communities.Communicated by O. Kinne, Oldendorf/Luhe     

Proximate composition of Antarctic mesopelagic fishes

Eleven mesopelagic fish species from the Weddell/Scotia Sea region of the Antarctic captured during the austral spring 1983, austral fall 1986, and austral winter 1988, were analyzed for proximate composition. Water, ash level, protein, lipid and carbohydrate were examined in relation to depth of occurrence and season. No depth-related trends were evident, primarily due to a low species diversity and minimal differences in those species' vertical distributions. The Antarctic speciesElectrona antarctica showed a significant increase in lipid level (% wet wt and % ash-free dry wt) between spring, fall and winter. The increase may signify an accumulation over the productive season, possibly as a reserve for the winter months. Lipid levels (% wet wt and % ash-free dry wt) were significantly lower in the Weddell Sea specimens examined in this study than in previously examined identical and congeneric species taken during the same season from a more productive near-shore Antarctic region. Comparisons with congeners and confamilials from tropical-subtropical and temperate systems revealed variable trends. The Antarctic speciesE. antarctica andCyclothone microdon had lower water and protein (% wet wt) levels than similar species from tropical-subtropical or temperate regions. Lipid levels of the two species are similar to temperate individuals, while energy levels are slightly higher. In contrast, species of the genusBathylagus show no trends in composition as a function of latitude. Differences in productivity, water-column temperature-structure, and seasonality are important considerations when examining trends among mesopelagic species.     

Symbiontic bacteria in the gut of the blood-sucking Antarctic fish parasite Gnathia calva (Crustacea: Isopoda)

Structure and ultrastructure of the digestive tract of the ectoparasitic stages of the Antarctic isopod Gnathia calva, collected in the Weddell Sea and around the Antarctic Peninsula in the 1984–1985 season, are briefly described. There are only two digestive glands, with an ultrastructure similar to that of other isopods. The gut is divided into six regions: oesophagus, stomach, dilatable reservior for sucked-in blood, sphincter, rectal vesicle with symbiontic bacteria, and rectum. The highly dilatable anterior hindgut (reservoir) takes part in the resorption of nutrients and stores lipids and glycogen. The rectal vesicle has an epithelium with all the features of an organ with high metabolism and the capacity for the transportation of small molecules. The surface is increased by irregular microvilli. A very thin intima and a basal labyrinth are present. The presence of symbiontic bacteria is discussed in correlation with the haematophagous nutrition of G. calva.     

An attempt at a functional analysis of the distributional range of Euphausia superba

The literature on the distribution of different developmental stages of Euphausia superba is discussed. It is suggested that the reason why the mass concentrations of E. superba larvae are confined to the circulations of the Weddell Sea and the Eastern Drift is the high level of the upper boundary of dense bottom water which prevents the sinking of eggs to great depths. Outside these regions the hatched larvae seem unable to cover the great distance separating them from the upper water layers.     

Genetic diversity of Nymphon (Arthropoda: Pycnogonida: Nymphonidae) along the Antarctic Peninsula with a focus on Nymphon australe Hodgson 1902

Sea spiders are conspicuous, and often abundant, members of the Antarctic benthic community. Nymphonidae (Pycnogonida) in Southern Ocean waters comprise over 240 species which are often difficult to assign due to their intraspecific ‘highly variable’ morphology. In particular, Nymphon australe, the numerically dominant species in Antarctic waters is known to have a high level of phenotypic variation in external morphology and is also reported to have a circumpolar distribution. Circumpolarity seems contradictory to the pycnogonid’s brooding lifestyle and presumably limited dispersal. Here we examine the genetic diversity of several Nymphon species collected in the Antarctic Peninsular region. Concomitantly, we assess the genetic structure of N. australe to gain insight into Nymphon dispersal capacity. Cytochrome c oxidase subunit I (COI) and 16S ribosomal gene data suggest a recent common history and/or recent gene-flow of N. australe populations across nearly 800 km of the Antarctic Peninsula. Furthermore, these data support that the Antarctic Peninsula region may hold two previously unrecognized species of Nymphon. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Abundance and feeding ecology of Antarctic phaeodarian radiolarians

Phaeodarian radiolarians were sampled from the upper 200 m along a transect through the ice-edge zone in the Weddell Sea in the austral autumn (March 1986) and at several stations in the western Antarctic Peninsula region in the austral winter (June 1987). Abundances of phaeodarians reached 3 132 m^-3 and were similar to or higher than maximum abundances of polycystine radiolarians, foraminiferans, and acantharians, and similar to or less than those of the heliozoan Sticholonche sp. Phaeodarians varied in abundance and species composition both seasonally and/or geographically. In contrast to the more numerous ciliates and flagellates that were most abundant in the upper 100 m, phaeodarians were most abundant from 100 to 200 m and showed no distinct pattern related to the ice edge. Electron microscopical examination of food vacuoles showed that phaeodarians are omnivorous generalists, feeding on a variety of food ranging in size from bacteria to large protozoans in both regions and seasons. Algal cells consumed in addition to diatoms and dinoflagellates included Chlorella-like cells and members of the recently described chrysophyte order Parmales. Scales of prasinophytes were common. Phaeodarians are consumed by the non-selective particlefeeding salp Salpa thompsoni. Thus, phaeodarians link microbial food webs to macrozooplankton and increase the complexity of the Antarctic food web.     

Growth and reproduction in the Antarctic brooding bivalve Adacnarca nitens (Philobryidae) from the Ross Sea

We present information on the reproductive biology, population structure, and growth of the brooding Antarctic bivalve Adacnarca nitens Pelseneer 1903, from the Ross Sea, Antarctica. Individuals ranging from 0.85 to 6.00 mm were found attached to a hydrozoan colony. This species shows low fecundity and large egg size, common to other brooding species. The minimum size at which oogenesis was detected was 2.3 mm and the minimum size at which brooding was evident was 3.9 mm. Embryos of a full range of developmental stages were brooded simultaneously in females. The population showed a log–normal distribution and results suggest non-periodic reproduction with continuous embryonic development. The reproductive traits of A. nitens are discussed in the context of circum-Antarctic species distribution and limitations to dispersal in brooding benthic invertebrates.     

Physiological development of brooded larvae from two pocilloporid corals in Taiwan

In southern Taiwan, brooded larvae of Pocillopora damicornis and Seriatopora caliendrum are released year-round in synchrony with new moons, and each larval release occurs over multiple days. Using P. damicornis and S. caliendrum as a model system, we describe within-brood variation in larval phenotypes and test for release-day effects that influence larval performance in the pelagic phase. Research was conducted in 2010 using larvae from corals collected in June and July from Nanwan Bay (21°56.179??N, 120°44.85??E). In June, larval phenotypes of both species were characterized immediately following release, and their competency to settle assessed. In July, larvae of P. damicornis were collected on 3?days over the peak release period and incubated for 7?C11?days at 28.0?°C and 320???mol?quanta?m^?2?s^?1; their phenotypes and settlement competency were measured every 2?days. P. damicornis larvae released close to peak release were 1.6 times larger in size, contained twice the number of Symbiodinium larva^?1, and were 44?% more likely to settle in the first 24?h than larvae released early in the brood. In addition, peak-release larvae respired at a lower rate than larvae released late in the brood. Similarly, S. caliendrum larvae released close to peak release were 1.4 times larger in size and were 33?% more likely to settle in the first 5?h than larvae released early in the brood. In July, P. damicornis larvae differed between early (2?days prior to peak), peak, and late (2?days after peak) release. Protein content of early-release larvae was lower than peak- and late-release larvae, and this difference persisted throughout the development. Further, release day affected the way larval respiration varied throughout development. By showing that brooded coral larvae differ between release days and display maternal effects influencing performance in the swimming phase, our results suggest that pocilloporid corals utilize bet-hedging to increase reproductive success.     

Depth utilisation by breeding Adélie penguins,Pygoscelis adeliae,at Esperanza Bay,Antarctica

Miniature depth gauges were attached in December 1987 and January 1988 to Adélie penguins,Pygoscelis adeliae, breeding at Esperanza on the Antarctic Peninsula. Results from 34 birds showed that foraging penguins with eggs and with brooded and crèching chicks spent mean periods away from the nest of 96, 36 and 21 h, respectively, during which time means of 29.0 h (30%), 11.2 h (31%) and 2.7 h (13%), respectively, were spent under water at depths > 5 m. Time under water was positively correlated with time absent from nest. Maximum depth reached was 170 m but overall birds spent most time at shallower depths. Birds foraging for brooded chicks dived deeper than birds foraging for crèching chicks. Stomach-pumping indicated that the principal prey caught at this time was krill,Euphausia superba. Mean mass changes of adults during single foraging trips indicated that krill were caught at a mean rate of 7.2 g min^–1 spent under water.     

Reproduction in Anthelia glauca (Octocorallia: Xeniidae). I. Gametogenesis and larval brooding

Anthelia glauca Lamarck, 1816 is a gonochoric, external-brooding soft coral found in KwaZulu-Natal. It is reproductively active in the summer months. The development of gametes produced in late summer is arrested in winter. Several stages of gametes are found at the base of the polyps, and female polyps produce several cycles of larvae over an extended breeding period of 4 to 5 months. Larvae are brooded in a unique pharyngeal brooding pouch not yet described in other coral species. The brood pouch consists of an expansion of the pharynx with constrictions proximal and distal to the embryos and larvae. Our data suggest that egg transfer and fertilization occur at full moon and the mature larvae are released after new moon. Zooxanthellae are absent in A. glauca oocytes, but zooxanthella infestation commences at the immature larval stage. Received: 15 July 1997 / Accepted: 12 March 1998     

Mega-epibenthic communities in Arctic and Antarctic shelf areas

Mega-epibenthic shelf assemblages were investigated off Northeast Greenland and in the Weddell, Bellingshausen and Amundsen Seas in the Antarctic using underwater video. In the Arctic a total of 94 taxa represented by more than 100 000 individuals were identified. Echinoderms, particularly brittle stars, were the most important elements of the mega-epibenthic fauna on the shelf off Northeast Greenland. Multivariate analyses of the species distribution revealed a conspicuous depth zonation in which an assemblage on the shallow banks can be clearly distinguished from that in the troughs. Between these a transitional zone with a heterogeneous benthic fauna was found. Physical disturbances are supposed to be responsible for the pronounced dominance patterns observed on the shallow banks. The fauna in the troughs, which consists of more than 50% suspension feeders, is diverse but low in numbers of individuals. In the Antarctic more than 115 000 individuals belonging to 169 taxa were analyzed. Obvious faunal differences exist between the stations in the Weddell Sea and the Bellingshausen/Amundsen Seas. Assemblages of suspension feeders dominated by sponges and bryozoans are prevalent on the shelf of the eastern Weddell Sea, but almost absent in the Bellingshausen and Amundsen Seas. These assemblages seem to be restricted to areas where bottom currents provide favourable feeding conditions. However, motile deposit feeders are more abundant in both regions where there is a soft bottom substrate with presumably slow bottom currents and reduced horizontal transport of organic particles. Received: 22 January 1999 / Accepted: 7 June 1999     

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.     

Effect of the receding ice-edge on the condition of mid-water fishes in the northwestern Weddell Sea: results from biochemical assays with notes on diet

Electrona antarctica and Bathylagus antarcticus are dominant fishes in the Southern Ocean pelagic ecosystem with disparate life histories, making them excellent subjects for comparative studies. Twenty-one indices of nutritional condition were compared in both species by observing changes in each index as it varied spatially and temporally in association with the marginal ice-zone bloom in the northwestern Weddell Sea. E. antarctica and B. antarcticus exhibited very different patterns of response to the bloom. Nutritional condition increased in 11 of 21 measures in E. antarctica, suggesting that it was in poorer condition at ice-covered stations and in greatly improved condition in post-bloom, open-water stations. The data, combined with a few observations of larvae, indicate that E. antarctica increases feeding before the spring bloom, rapidly building stores of lipid, which are probably used for spawning. In contrast, only three measures of condition were variable in B. antarcticus, and lipid actually decreased from ice-covered to open water. RNA:DNA values increased in open water, coinciding with an increase in food volume present in their guts. These observations, coupled with observations of larvae in ice-covered stations, suggest that B. antarcticus possesses sufficient energetic stores to spawn prior to the bloom, so that larvae are able to fully utilize available resources associated with the spring bloom. Increased somatic growth, as indicated by RNA:DNA, may not begin until later, when the bloom was beginning to decline. Earlier work, which suggested that the deeper living, non-migrating species, like B. antarcticus, would not be affected by the bloom until after shallower, migratory species, like E. antarctica, is supported. Of the assays tested, RNA:DNA appeared to be the most sensitive. The combined assessment of many measures including biochemical, compositional, dietary, and age estimates from single specimens is possible if the samples are treated with sufficient care. The reliance upon a single measure to help interpret the ecology of a species, especially in nekton-sized species, is not as effective as techniques used in combination. Received: 12 January 2000 / Accepted: 16 August 2000     

Asexual production of planulae in the coral Pocillopora damicornis

The reproduction of scleractinian corals through planular larvae has traditionally been viewed as a strictly sexual process. Here, the results of an electrophoretic study of a ubiquitous Indo-Pacific coral, Pocillopora damicornis, show an exact inheritance of parental genotypes by brooded planulae, demonstrating the existence of an asexual mode of production of planular larvae. Comparisons of the genetical structure of a number of populations with structures predicted for sexual reproduction suggest that, although there is probably also a sexual form of reproduction, asexually produced planulae can be of major importance in the maintenance of populations of this species.     

Lipid components as a measure of nutritional condition in fish larvae (Pleuragramma antarcticum) in East Antarctica

Pleuragramma antarcticum is a key component of the neritic assemblages in the Antarctic coastal waters. Larvae of this species were sampled from 2008 to 2011 in the Dumont d’Urville Sea (East Antarctica). The lipid class composition [triacylglycerols (TAG), cholesterol (Chol) and polar lipids (PL)] of larvae was measured to assess the larval condition. The total amount of lipids was linearly related to the quantity of structural polar lipids, suggesting that growth is favored over lipid storage. The TAG:Chol ratio showed interannual variability in the condition of fish larvae, probably related to prey availability. Nevertheless, the essential fatty acids composition of polar lipids illustrates that larvae with low levels of TAG:Chol could be either growing or under starvation. Only the combination of a low TAG:Chol ratio and low polar lipids content, which can also be mobilized during starvation periods, allowed identification of larvae in poor condition. This lipid condition index should be of great assistance to evaluate the probability of survival of P. antarcticum larvae in long-term monitoring. It has widespread applicability and should also be useful in the diagnosis of nutritional condition in other species.     

Genetic evidence of self-fertilization in the sea anemone Epiactis prolifera

Biochemical genetic variation provided evidence for the mode of reproduction of brooded young in the sea anemone Epiactis prolifera Verrill, 1869. Individuals of E. prolifera are female when small but hermaphroditic when large (i.e., gynodioecious); juveniles are brooded externally on the column. Brooding individuals collected from 6 intertidal sites (5 in central California and 1 in Washington State, USA) in the spring and summer of 1980 were assayed for gene-enzyme variation by starch-gel electrophoresis. Three of 12 enzyme loci were polymorphic; phosphoglucose isomerase appeared to be encoded by two, closely linked loci. Genotypic frequencies deviated markedly from expected random mating proportions. Only three heterozygotes were found; two were heterozygous at all three polymorphic loci, and the other was polymorphic at the two PGI loci. All 158 juveniles from 25 brooding individuals were assayed (2–19 juveniles per parent). Juveniles on homozygous adults were always identical to their parent. However, brooded young of heterozygous individuals were not identical to their parent. but showed 1:2:1 phenotypic segregation ratios consistent with reproduction by self-fertilization. This genetic evidence together with findings of marked heterozygote deficiencies and genetic identity of homozygous adults and their brooded young supports the conclusion that E. prolifera usually reproduces by self-fertilization, and cross-fertilization is rate.     

Reproductive patterns in the Antarctic brachiopod Liothyrella uva

The reproductive cycle of the Antarctic articulate brachiopod Liothyrella uva (Broderip, 1833) is described from monthly samples collected between September 1985 and July 1987 from a population at Signy Island, Antarctica. Spermatogenesis and oogenesis are described for the first time in this species. Surface dried-tissue masses for a standard individual (41 mm shell length) were calculated for the digestive diverticula, gut, lophophore and gonad from monthly dissections of 15 brachiopods. Seasonal patterns, with summer peaks, were observed in the digestive diverticula and gut. The lophophore and gonad masses did not exhibit seasonal trends. Females showed a sharp decrease in proportion of large oocytes between October and November in 1986, suggesting spawning during this period. There was no similar decrease during the same period in 1985. Mean percent spermatozoa measurements revealed a large increase in November 1985 and a rapid decline in December 1985. This suggested a large spawning event for males in 1985 which was not repeated the following year. These data indicate large inter-annual differences in reproductive activity as well as differences between males and females. They also suggest the possibility of sperm storage by females. Brood characteristics were also highly variable. The smallest brooding female was 31.5 mm in length. Some females brooded more than one developmental stage simultaneously, and variation in brood size (numbers of embryos or larvae held in the lophophore) and brood composition between individuals was high. All samples collected throughout the 2 yr period contained some females with broods. The data suggest that the reproductive strategy of L. uva is highly plastic, and that there may be three reproductive periodicities on seasonal, annual and inter-annual time scales. Received: 16 May 1997 / Accepted: 2 March 1998