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Microalgae are the main source of the omega‐3 fatty acids eicosapentaenoic acid (EPA) and docosahexaenoic acid (DHA), essential for the healthy development of most marine and terrestrial fauna including humans. Inverse correlations of algal EPA and DHA proportions (% of total fatty acids) with temperature have led to suggestions of a warming‐induced decline in the global production of these biomolecules and an enhanced importance of high latitude organisms for their provision. The cold Arctic Ocean is a potential hotspot of EPA and DHA production, but consequences of global warming are unknown. Here, we combine a full‐seasonal EPA and DHA dataset from the Central Arctic Ocean (CAO), with results from 13 previous field studies and 32 cultured algal strains to examine five potential climate change effects; ice algae loss, community shifts, increase in light, nutrients, and temperature. The algal EPA and DHA proportions were lower in the ice‐covered CAO than in warmer peripheral shelf seas, which indicates that the paradigm of an inverse correlation of EPA and DHA proportions with temperature may not hold in the Arctic. We found no systematic differences in the summed EPA and DHA proportions of sea ice versus pelagic algae, and in diatoms versus non‐diatoms. Overall, the algal EPA and DHA proportions varied up to four‐fold seasonally and 10‐fold regionally, pointing to strong light and nutrient limitations in the CAO. Where these limitations ease in a warming Arctic, EPA and DHA proportions are likely to increase alongside increasing primary production, with nutritional benefits for a non‐ice‐associated food web. [ABSTRACT FROM AUTHOR]

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Global warming causes dramatic environmental change to Arctic ecosystems. While pelagic primary production is initiated earlier and its intensity can be increased due to earlier ice melt and extended open‐water periods, sea‐ice primary production is progressively confined on a spatio‐temporal scale, leading to unknown consequences for the ice‐associated (sympagic) food web. Understanding ecological responses to changes in the availability and composition of pelagic and sympagic food sources is crucial to determine potential changes of food‐web structure and functioning in Arctic marine communities under increasingly ice‐free conditions. Focus was placed on the importance of suspended particulate organic matter vs. sympagic organic matter for 12 zooplankton species with different feeding modes covering five taxonomic groups (copepods, krill, amphipods, chaetognaths, and appendicularians) at two ice‐covered, but environmentally different, stations in the north‐western Barents Sea in August 2019. Contributions of diatom‐ and flagellate‐associated fatty acids (FAs) to total lipid content and carbon stable isotopic compositions of these FAs were used to discriminate food sources and trace flows of organic matter in marine food webs. Combination of proportional contributions of FA markers with FA isotopic composition indicated that consumers mostly relied, directly (herbivorous species), or indirectly (omnivorous and carnivorous species), on pelagic diatoms and flagellates, independently of environmental conditions at the sampling locations, trophic position, and feeding mode. Differences were nevertheless observed between species. Contrary to other studies demonstrating a high importance of sympagic organic matter for food‐web processes, our results highlight the complexity and variability of trophic structures and dependencies in different Arctic food webs. [ABSTRACT FROM AUTHOR]

In polar regions, the microphytobenthos has important ecological functions in shallow-water habitats, such as on top of coastal sediments. This community is dominated by benthic diatoms, which contribute significantly to primary production and biogeochemical cycling while also being an important component of polar food webs. Polar diatoms are able to cope with markedly changing light conditions and prolonged periods of darkness during the polar night in Antarctica. However, the underlying mechanisms are poorly understood. In this study, five strains of Antarctic benthic diatoms were isolated in the field, and the resulting unialgal cultures were identified as four distinct species, of which one is described as a new species, Planothidium wetzelii sp. nov. All four species were thoroughly examined using physiological, cell biological, and biochemical methods over a fully controlled dark period of 3 months. The results showed that the utilization of storage lipids is one of the key mechanisms in Antarctic benthic diatoms to survive the polar night, although different fatty acids were involved in the investigated taxa. In all tested species, the storage lipid content declined significantly, along with an ultrastructurally observable degradation of the chloroplasts. Surprisingly, photosynthetic performance did not change significantly despite chloroplasts decreasing in thylakoid membranes and an increased number of plastoglobules. Thus, a combination of biochemical and cell biological mechanisms allows Antarctic benthic diatoms to survive the polar night. [ABSTRACT FROM AUTHOR]

During the productive polar day, zooplankton and sea‐ice amphipods fulfill a critical role in energy transfer from primary producers to higher trophic‐level species in Arctic marine ecosystems. Recent polar night studies on zooplankton and sea‐ice amphipods suggest higher levels of biological activity than previously assumed. However, it is unknown if these invertebrates maintain polar night activity on stored lipids, opportunistic feeding, or a combination of both. To assess how zooplankton (copepods, amphipods, and krill) and sea‐ice amphipods support themselves on seasonally varying resources, we studied their lipid classes, fatty acid compositions, and compound‐specific stable isotopes of trophic biomarker fatty acids during polar day (June/July) and polar night (January). Lipid storage and fatty acid results confirm previously described dietary sources in all species during polar day. We found evidence of polar night feeding in all species, including shifts from herbivory to omnivory. Sympagic‐, pelagic‐, and Calanus spp.‐derived carbon sources supported zooplankton and sea‐ice amphipods in both seasons. We provide a first indication of polar night feeding of sea‐ice amphipods in the pelagic realm. [ABSTRACT FROM AUTHOR]

Sea ice continues to decline across many regions of the Arctic, with remaining ice becoming increasingly younger and more dynamic. These changes alter the habitats of microbial life that live within the sea ice, which support healthy functioning of the marine ecosystem and provision of resources for human-consumption, in addition to influencing biogeochemical cycles (e.g. air–sea CO2 exchange). With the susceptibility of sea ice ecosystems to climate change, there is a pressing need to fill knowledge gaps surrounding sea ice habitats and their microbial communities. Of fundamental importance to this goal is the development of new methodologies that permit effective study of them. Based on outcomes from the DiatomARCTIC project, this paper integrates existing knowledge with case studies to provide insight on how to best document sea ice microbial communities, which contributes to the sustainable use and protection of Arctic marine and coastal ecosystems in a time of environmental change. [ABSTRACT FROM AUTHOR]

Over the past decades, two key grazers in the Southern Ocean (SO), krill and salps, have experienced drastic changes in their distribution and abundance, leading to increasing overlap of their habitats. Both species occupy different ecological niches and long-term shifts in their distributions are expected to have cascading effects on the SO ecosystem. However, studies directly comparing krill and salps are lacking. Here, we provide a direct comparison of the diet and fecal pellet composition of krill and salps using 18S metabarcoding and fatty acid markers. Neither species' diet reflected the composition of the plankton community, suggesting that in contrast to the accepted paradigm, not only krill but also salps are selective feeders. Moreover, we found that krill and salps had broadly similar diets, potentially enhancing the competition between both species. This could be augmented by salps' ability to rapidly reproduce in favorable conditions, posing further risks to krill populations. In this manuscript, Pauli et al. directly compare the diets of two key Southern Ocean grazers, krill and salps. The 18S profiles of diet and pellets from both species do not match the planktonic profile, suggesting that both are selective grazers with a similar diet. [ABSTRACT FROM AUTHOR]

The hydrography of the Arctic Seas is being altered by ongoing climate change, with knock‐on effects to nutrient dynamics and primary production. As the major pathway of exchange between the Arctic and the Atlantic, the Fram Strait hosts two distinct water masses in the upper water column, northward flowing warm and saline Atlantic Waters in the east, and southward flowing cold and fresh Polar Surface Water in the west. Here, we assess how physical processes control nutrient dynamics in the Fram Strait using nitrogen isotope data collected during 2016 and 2018. In Atlantic Waters, a weakly stratified water column and a shallow nitracline reduce nitrogen limitation. To the west, in Polar Surface Water, nitrogen limitation is greater because stronger stratification inhibits nutrient resupply from deeper water and lateral nitrate supply from central Arctic waters is low. A historical hindcast simulation of ocean biogeochemistry from 1970 to 2019 corroborates these findings and highlights a strong link between nitrate supply to Atlantic Waters and the depth of winter mixing, which shoaled during the simulation in response to a local reduction in sea‐ice formation. Overall, we find that while the eastern Fram Strait currently experiences seasonal nutrient replenishment and high primary production, the loss of winter sea ice and continued atmospheric warming has the potential to inhibit deep winter mixing and limit primary production in the future. Plain Language Summary: The Fram Strait is the main gateway of the Arctic Ocean. In the east, warm, salty waters from the Atlantic flow north into the Arctic basin, and in the west, cold, fresh waters flow south from the central Arctic into the North Atlantic. We examined how changes to the availability of nutrients (which are essential for algae to grow) may limit algae growth in the Fram Strait, both as a result of changes to their source and also how easily the upper ocean mixes nutrients from depth. In the eastern Fram Strait, there is a high availability of nitrate, one of the main nutrients to support algae growth, and winter mixing sustains nutrient supply and biological production in recent decades. However, in the western Fram Strait, the outflowing surface waters do not easily mix with deeper waters and are depleted in nitrate, and nutrient supply from the central Arctic has been declining in recent decades. Our work suggests that although the eastern Fram Strait is sustaining higher levels of algae growth, which supports fisheries and higher trophic levels, warming over the coming decades could shoal winter mixed layers enough to decrease summertime nutrients and limit biological production. Key Points: Nitrate isotope signatures in the western Arctic outflow of Polar Surface Water and eastern inflow of Atlantic Water are characterizedWestern Fram Strait is strongly stratified and nitrate deplete compared to the east where winter mixing sustains nutrient supplyFuture warming may shoal the winter mixed layer in the east, decreasing nitrate supply, and reducing primary production below current rates [ABSTRACT FROM AUTHOR]

The thecosome pteropods Limacina helicina and L. retroversa are important contributors to the zooplankton community in high-latitude environments but little is known about their distribution and life cycle under polar conditions. We collected the early life stages (< 1 mm) of the thecosome population in 2012 and 2013 at a bi-weekly to monthly resolution in fjord highly influenced by Arctic waters as well as Atlantic inflows (Adventfjorden, Svalbard, 78°N), together with environmental parameters. L. retroversa only occurred episodically, in association with the inflow of Atlantic water, with low numbers and random size distributions. This suggests that this boreal species does not fulfill its life cycle in Adventfjorden. In contrast, young specimens of L. helicina were present during the entire study. Veligers hatched in late summer/autumn and measured 0.14 mm on average. They grew with rates of 0.0006 mm day−1 over the 10–11 months of development. Only thereafter, growth accelerated by one order of magnitude and maximal rates were reached in autumn (0.0077 mm day−1). Our results indicate that L. helicina reaches a size of 1 mm after approximately 1.5 years in Adventfjorden. We therefore suggest that L. helicina overwinters the first year as a small juvenile and that it needs at least 2 years to reach an adult size of 5 mm in Adventfjorden. This reveals an complex and delicate aspect of the life-cycle of L. helicina and further research is needed to determine if it makes the population especially vulnerable towards climate changes. [ABSTRACT FROM AUTHOR]

Submesoscale eddies and fronts are important components of oceanic mixing and energy fluxes. These phenomena occur in the surface ocean for a period of several days, on scales between a few hundred meters and few tens of kilometers. Remote sensing and modeling suggest that eddies and fronts may influence marine ecosystem dynamics, but their limited temporal and spatial scales make them challenging for observation and in situ sampling. Here, the study of a submesoscale filament in summerly Arctic waters (depth 0–400 m) revealed enhanced mixing of Polar and Atlantic water masses, resulting in a ca. 4 km wide and ca. 50 km long filament with distinct physical and biogeochemical characteristics. Compared to the surrounding waters, the filament was characterized by a distinct phytoplankton bloom, associated with depleted inorganic nutrients, elevated chlorophyll a concentrations, as well as twofold higher phyto‐ and bacterioplankton cell abundances. High‐throughput 16S rRNA gene sequencing of bacterioplankton communities revealed enrichment of typical phytoplankton bloom‐associated taxonomic groups (e.g., Flavobacteriales) inside the filament. Furthermore, linked to the strong water subduction, the vertical export of organic matter to 400 m depth inside the filament was twofold higher compared to the surrounding waters. Altogether, our results show that physical submesoscale mixing can shape distinct biogeochemical conditions and microbial communities within a few kilometers of the ocean. Hence, the role of submesoscale features in polar waters for surface ocean biodiversity and biogeochemical processes need further investigation, especially with regard to the fate of sea ice in the warming Arctic Ocean. [ABSTRACT FROM AUTHOR]

The study revealed species- and stage-specific differences in lipid accumulation of the dominant Antarctic copepods, the primarily herbivorous Calanoides acutus (copepodite stage V (CV), females) and the more omnivorous Calanus propinquus (females) storing wax esters and triacylglycerols, respectively, which were collected in summer (end of December). Feeding carbon-labelled diatoms to these copepods, 13C elucidated assimilation and turnover rates of copepod total lipids as well as specific fatty acids and alcohols. The 13C incorporation was monitored by compound-specific stable isotope analysis (CSIA). CV stages of C. acutus exhibited an intense total lipid turnover and 55% of total lipidswere labelled after 9 days of feeding. By contrast, total lipid assimilation of female C. acutus and C. propinquus was lower with 29% and 32%, respectively. The major dietary fatty acids 16:0, 16:1(n - 7) and 20:5(n - 3) had high turnover rates in all specimens. In C. acutus CV, the high rates of the de novo synthesized long-chain monounsaturated fatty acids and alcohols 20:1(n - 9) and 22:1(n - 11) indicate intense lipid deposition, whereas these rates were low in females. The differences in lipid assimilation and turnover clearly show that the copepod species exhibit a high variability and plasticity to adapt their lipid production to their various life phases. [ABSTRACT FROM AUTHOR]

Mitochondrial respiration is a multi-step pathway that involves matrix and membrane-associated enzymes and plays a key role in acclimation to variable environmental conditions, but until now it has not been clear which of these steps would be most important in acclimation to changing temperatures and CO2 levels. Considering scenarios of ocean warming and acidification we assessed the role and limitation to phenotypic plasticity in the hearts of two Gadoid species adapted to different thermal ranges: the polar cod (Boreogadus saida), an Arctic stenotherm, and the Northeast Arctic population of Atlantic cod (NEAC, Gadus morhua), a cold eurytherm. We analysed the capacity of single enzymes involved in mitochondrial respiration [citrate synthase (CS), succinate dehydrogenase (SDH), cytochrome c oxidase (CCO)], the capacity of the electron transport system and the lipid class composition of the cellular membranes. Juveniles of the two species were held for four months at four temperatures (0, 3, 6, 8 °C for polar cod and 3, 8, 12, 16 °C for NEAC), at both ambient and elevated P CO 2 (400 µatm and 1170 µatm, respectively). Polar cod showed no changes in mitochondrial enzyme capacities and in the relative lipid class composition in response to altered temperature or elevated P CO 2. The lack of cardiac cellular plasticity together with evidence at the whole-animal level coming from other studies is indicative of little or no ability to overcome stenothermy, in particular during acclimation to 8 °C. In contrast, eurythermal NEAC exhibited modifications of membrane composition towards a more rigid structure and altered enzyme capacities to preserve functionality at higher temperatures. Furthermore, in NEAC, the capacities of SDH, CCO and CS were increased by high levels of CO2 if combined with high temperatures (12 and 16 °C), suggesting the compensation of an inhibitory effect. These results indicate that the cold eurythermal species (NEAC) is able to alter its mitochondrial function to a far greater extent than the Arctic stenotherm (polar cod), indicating greater resilience to variable environmental conditions. This difference in plasticity may underpin differences in the resilience to climate change and affect future species distributions and, eventually, survival. [ABSTRACT FROM AUTHOR]

Fatty acid (FA) compositions provide insights about storage and feeding modes of marine organisms, characterizing trophic relationships in the marine food web. Such compositional data, which are normalized to sum to 1, have values—and thus derived statistics as well—that depend on the particular mix of components that constitute the composition. In FA studies, if the set of FAs under investigation is different in two separate studies, all the summary statistics and relationships between the FAs that are common to the two studies are artificially changed due to the normalization, and thus incomparable. Ratios of FAs, however, are invariant to the particular choice of FAs under consideration—they are said to be subcompositionally coherent. Here, we document the collaboration between a biochemist (M.G.) and a statistician (M.J.G.) to determine a suitable small set of FA ratios that effectively replaces the original data set for the purposes of univariate and multivariate analysis. This strategy is applied to two FA data sets, on copepods and amphipods, respectively, and is widely applicable in other contexts. The selection of ratios is performed in such a way as to satisfy substantive requirements in the context of the respective data set, namely to explain phenomena of interest relevant to the particular species, as well as the statistical requirement to explain as much variance in the FA data set as possible. Benefits of this new approach are (1) univariate statistics that can be validly compared between different studies, and (2) a simplified multivariate analysis of the reduced set of ratios, giving practically the same results as the analysis of the full FA data set. [ABSTRACT FROM AUTHOR]

Increasing sea surface temperatures (SST) and blooms of lipid‐poor, filamentous cyanobacteria can change mesozooplankton metabolism and foraging strategies in marine systems. Lipid shortage and imbalanced diet may challenge the build‐up of energy pools of lipids and proteins, and access to essential fatty acids (FAs) and amino acids (AAs) by copepods. The impact of cyanobacterial blooms on individual energy pools was assessed for key species temperate Temora longicornis and boreal Pseudo‐/Paracalanus spp. that dominated field mesozooplankton communities isolated by seasonal stratification in the central Baltic Sea during the hot and the cold summer. We looked at (a) total lipid and protein levels, (b) FA trophic markers and AA composition, and (c) compound‐specific stable carbon isotopes (δ13C) in bulk mesozooplankton and in a subset of parameters in particulate organic matter. Despite lipid‐poor cyanobacterial blooms, the key species were largely able to cover both energy pools, yet a tendency of lipid reduction was observed in surface animals. Omni‐ and carnivory feeding modes, FA trophic makers, and δ13C patterns in essential compounds emphasized that cyanobacterial FAs and AAs have been incorporated into mesozooplankton mainly via feeding on mixo‐ and heterotrophic (dino‐) flagellates and detrital complexes during summer. Foraging for essential highly unsaturated FAs from (dino‐) flagellates may have caused night migration of Pseudo‐/Paracalanus spp. from the deep subhalocline waters into the upper waters. Only in the hot summer (SST>19.0°C) was T. longicornis submerged in the colder subthermocline water (~4°C). Thus, the continuous warming trend and simultaneous feeding can eventually lead to competition on the preferred diet by key copepod species below the thermocline in stratified systems. A comparison of δ13C patterns of essential AAs in surface mesozooplankton across sub‐basins of low and high cyanobacterial biomasses revealed the potential of δ13C‐AA isoscapes for studies of commercial fish feeding trails across the Baltic Sea food webs. [ABSTRACT FROM AUTHOR]

The meroplanktonic larvae of benthic organisms are an important seasonal component of the zooplankton in temperate coastal waters. The larvae of the reef-building polychaete Lanice conchilega contribute up to 15% of the summer zooplankton biomass in the North Sea. Despite their importance for reef maintenance (which positively affects the benthic community), little is known about the trophic ecology of this meroplanktonic larva. Qualitative and quantitative estimates of carbon (C) transfer between trophic levels and of fatty acid (FA)—specific assimilation, biosynthesis, and bioconversion can be obtained by compound-specific stable isotope analysis of FA. The present work tested the hypothesis that the concept of fatty acid trophic markers (FATM), widely used for studies on holoplankton with intermediate to high lipid contents, is also applicable to lipid-poor organisms such as meroplanktonic larvae. The incorporation of isotopically-enriched dietary C by L. conchilega larvae was traced, and lipid assimilation did not follow FA-specific relative availabilities in the diet. Furthermore, FAs that were unavailable in the diet, such as 22:5(n-3), were recorded in L. conchilega, suggesting their bioconversion by the larvae. The results indicate that L. conchilega larvae preferentially assimilate certain FAs and regulate their FA composition (lipid homeostasis) independently of that of their diet. Their quasi-homeostatic response to dietary FA availability could imply that the concept of FATM has limited application in lipid-poor organisms such as L. conchilega larvae. [ABSTRACT FROM AUTHOR]

Abstract: How the abundant pelagic life of the Southern Ocean survives winter darkness, when the sea is covered by pack ice and phytoplankton production is nearly zero, is poorly understood. Ice‐associated (“sympagic”) microalgae could serve as a high‐quality carbon source during winter, but their significance in the food web is so far unquantified. To better understand the importance of ice algae‐produced carbon for the overwintering of Antarctic organisms, we investigated fatty acid (FA) and stable isotope compositions of 10 zooplankton species, and their potential sympagic and pelagic carbon sources. FA‐specific carbon stable isotope compositions were used in stable isotope mixing models to quantify the contribution of ice algae‐produced carbon (αIce) to the body carbon of each species. Mean αIce estimates ranged from 4% to 67%, with large variations between species and depending on the FA used for the modelling. Integrating the αIce estimates from all models, the sympagic amphipod Eusirus laticarpus was the most dependent on ice algal carbon (αIce: 54%–67%), and the salp Salpa thompsoni showed the least dependency on ice algal carbon (αIce: 8%–40%). Differences in αIce estimates between FAs associated with short‐term vs. long‐term lipid pools suggested an increasing importance of ice algal carbon for many species as the winter season progressed. In the abundant winter‐active copepod Calanus propinquus, mean αIce reached more than 50% in late winter. The trophic carbon flux from ice algae into this copepod was between 3 and 5 mg C m−2 day−1. This indicates that copepods and other ice‐dependent zooplankton species transfer significant amounts of carbon from ice algae into the pelagic system, where it fuels the food web, the biological carbon pump and elemental cycling. Understanding the role of ice algae‐produced carbon in these processes will be the key to predictions of the impact of future sea ice decline on Antarctic ecosystem functioning. [ABSTRACT FROM AUTHOR]

The Arctic represents an extreme habitat for phototrophic algae due to long periods of darkness caused by the polar night (~4 months darkness). Benthic diatoms, which dominate microphytobenthic communities in shallow water regions, can survive this dark period, but the underlying physiological and biochemical mechanisms are not well understood. One of the potential mechanisms for long-term dark survival is the utilisation of stored energy products in combination with a reduced basic metabolism. In recent years, water temperatures in the Arctic increased due to an ongoing global warming. Higher temperatures could enhance the cellular energy requirements for the maintenance metabolism during darkness and, therefore, accelerate the consumption of lipid reserves. In this study, we investigated the macromolecular ratios and the lipid content and composition of Navicula cf. perminuta Grunow, an Arctic benthic diatom isolated from the microphytobenthos of Adventfjorden (Svalbard, Norway), over a dark period of 8 weeks at two different temperatures (0 and 7 °C). The results demonstrate that N. perminuta uses the stored lipid compound triacylglycerol (TAG) during prolonged dark periods, but also the pool of free fatty acids (FFA). Under the enhanced temperature of 7 °C, the lipid resources were used significantly faster than at 0 °C, which could consequently lead to a depletion of this energy reserves before the end of the polar night. On the other hand, the membrane building phospho- and glycolipids remained unchanged during the 8 weeks darkness, indicating still intact thylakoid membranes. These results explain the shorter survival times of polar diatoms with increasing water temperatures during prolonged dark periods. [ABSTRACT FROM AUTHOR]

Knowledge on the capability of zooplankton to adapt to the rapidly changing environmental conditions in the Arctic is crucial to predict future ecosystem processes. The key species on the Arctic shelf, the calanoid copepod Calanus glacialis, grows and accumulates lipid reserves in spring and summer in surface waters. The winter is spent in dormancy in deeper water layers with low metabolic activity. As timing and intensity of metabolic changes have been poorly investigated, our study aims to characterize the physiology of C. glacialis over an entire year, from July 2012 to July 2013. We followed anabolic and catabolic enzyme activities and the biochemical composition of this species, taking depth-stratified samples once a month in Billefjorden, a high-Arctic sill fjord. A large part of the population had migrated to depths >100 m by July 2012. Only thereafter, anabolic activities decreased slowly, suggesting that low metabolism is related to ceased feeding rather than to endogenous regulation. During overwintering, anabolic enzyme activities were reduced by half as compared to peak activities in spring. The biochemical composition of the copepods changed little from July to December. Then, the lipid catabolic activity increased and the lipid content decreased, likely fuelling moulting and gonad maturation. The protein content did not change significantly during winter, suggesting that proteins are not much catabolized during that time. The relatively high metabolic activity in C. glacialis in winter suggests that this species is not entering a true diapause and should thus be able to respond flexible to changing environmental conditions. [ABSTRACT FROM AUTHOR]

To better predict ecological consequences of changing Arctic sea ice environments, we aimed to quantify the contribution of ice algae-produced carbon ( αIce) to pelagic food webs in the central Arctic Ocean. Eight abundant under-ice fauna species were submitted to fatty acid (FA) analysis, bulk stable isotope analysis (BSIA) of nitrogen ( δ15N) and carbon ( δ13C) isotopic ratios, and compound-specific stable isotope analysis (CSIA) of δ13C in trophic marker FAs. A high mean contribution αIce was found in Apherusa glacialis and other sympagic (ice-associated) amphipods (BSIA: 87% to 91%, CSIA: 58% to 92%). The pelagic copepods Calanus glacialis and C. hyperboreus, and the pelagic amphipod Themisto libellula showed substantial, but varying αIce values (BSIA: 39% to 55%, CSIA: 23% to 48%). Lowest αIce mean values were found in the pteropod Clione limacina (BSIA: 30%, CSIA: 14% to 18%). Intra-specific differences in FA compositions related to two different environmental regimes were more pronounced in pelagic than in sympagic species. A comparison of mixing models using different isotopic approaches indicated that a model using δ13C signatures from both diatom-specific and dinoflagellate-specific marker FAs provided the most conservative estimate of αIce. Our results imply that ecological key species of the central Arctic Ocean thrive significantly on carbon synthesized by ice algae. Due to the close connectivity between sea ice and the pelagic food web, changes in sea ice coverage and ice algal production will likely have important consequences for food web functioning and carbon dynamics of the pelagic system. [ABSTRACT FROM AUTHOR]

This study aimed at understanding how life-cycle strategies of the primarily herbivorous Pseudocalanus minutus and the omnivorous Oithona similis are reflected by their lipid carbon turnover capacities. The copepods were collected in Billefjorden, Svalbard, and fed with 13C labeled flagellates and diatoms during 3 weeks. Fatty acid (FA) and fatty alcohol compositions were determined by gas chromatography, 13C incorporation was monitored using isotope ratio mass spectrometry. Maximum lipid turnover occurred in P. minutus, which exchanged 54.4% of total lipid, whereas 9.4% were exchanged in O. similis. In P. minutus, the diatom markers 16:1(n-7), 16:2(n-4) and 16:3(n-4) were almost completely renewed from the diet within 21 days, while 15% of the flagellate markers 18:2(n-6), 18:3(n-3) and 18:4(n-3) were exchanged. In O. similis, 15% of both flagellate and diatom markers were renewed. P. minutus exhibited typical physiological adaptations of herbivorous copepod species, with a very high lipid turnover rate and the ability to integrate FAs more rapidly from diatoms than from flagellates. O. similis depended much less on lipid reserves and had a lower lipid turnover rate, but was able to ingest and/or assimilate lipids with the same intensity from various food sources, to sustain shorter periods of food shortage. [ABSTRACT FROM AUTHOR]

Decreasing Arctic sea ice cover and increasing stratification of ocean surface waters make the exposure of pelagic microalgae to high irradiances more likely. Apart from light being a necessary prerequisite for photosynthesis, rapidly changing and/or high irradiances are potentially detrimental. An in situ study was performed in the high Arctic (79°N) to determine the effect of high irradiances in general, and ultraviolet radiation (UVR, 280-400 nm) in particular, on cell concentrations, fatty acid composition, and photoprotective pigments of three diatom species isolated from seawater around Svalbard. Unialgal cultures were exposed in situ at 0.5- and 8 m-depth. After 40 h, cell concentrations of Synedropsis hyperborea and Thalassiosira sp., were lower at 0.5 than at 8 m, and the content of the photoprotective xanthophyll-cycle pigment diatoxanthin in all species ( S. hyperborea, Thalassiosira sp., Porosira glacialis) was higher in the 0.5 m exposure compared to 8 m. In S. hyperborea, growth was additionally inhibited by UVR at 0.5-m depth. In situ radiation conditions led, furthermore, to a significant decrease in polyunsaturated fatty acids (PUFAs) in all three species, but UVR had no additional effect. Hence, we conclude that natural radiation conditions close to the surface could reduce growth and PUFA concentrations, but the effects are species specific. The diatoms' potential to acclimate to these conditions over time has to be evaluated. [ABSTRACT FROM AUTHOR]

Our study deals with the lipid biochemistry of the krill community in the ecosystem of the high Arctic Kongsfjord (Svalbard). During the last decades, Kongsfjord experienced a change in krill species composition due to recent increased advection of Atlantic water masses carrying characteristic boreal as well as subtropical-boreal euphausiids into the ecosystem. The lipid biochemistry and trophic relationships of the species recently inhabiting the Arctic water masses are scarcely known, although a change in a krill population may have a significant impact on the ecosystem. A comparison of nutrition and energy storage strategies, stable isotopes, lipid profiles and fatty acid compositions showed remarkable differences between the krill species. These reflected the diverse feeding behaviours and specific adaptations to the environments of their origin: the boreal Meganyctiphanes norvegica and subtropical Nematoscelis megalops appear more carnivorous and have significantly lower mean lipid contents (29 and 10 %, respectively) and a different energy storage pattern (triacylglycerols and polar lipids, respectively) than the arcto-boreal Thysanoessa inermis, which consists of up to 54 % of lipids mainly stored as wax esters (>40 %). These differences may have significant implications for the rapidly changing marine food web of Kongsfjord-especially for higher trophic levels relying on the nutritional input of animal lipids. [ABSTRACT FROM AUTHOR]

It is commonly assumed that the most accurate data on fatty acid (FA) contents are obtained when samples are analyzed immediately after collection. For logistical reasons, however, this is not always feasible and samples are often kept on ice or frozen at various temperatures and for diverse time periods. We quantified temporal changes of selected FA (μg FAME per mg tissue dry weight) from 6 fish species subjected to 2 handling and 3 storage methods and compared them to FA contents from muscle tissue samples that were processed immediately. The following species were investigated: Common Carp (Cyprinus carpio), Freshwater Drum (Aplodinotus grunniens), Channel Catfish (Ictalurus punctatus), Antarctic Eelpout (Pachycara brachycephalum), Rainbow Trout (Oncorhynchus mykiss) and Arctic Charr (Salvelinus alpinus). The impact of storage method and duration of storage on FA contents were species-specific, but not FA-specific. There was no advantage in using nitrogen gas for tissue samples held on ice for 1 week; however, holding tissue samples on ice for 1 week resulted in a loss of FA in Charr. In addition, most FA in Trout and Charr decreased in quantity after being stored between 3 and 6 hours on ice. Freezer storage temperature (-80 or -20°C) also had a significant effect on FA contents in some species. Generally, we recommend that species with high total lipid content (e.g. Charr and Trout) should be treated with extra caution to avoid changes in FA contents, with time on ice and time spent in a freezer emerging as significant factors that changed FA contents. [ABSTRACT FROM AUTHOR]

The giant kelp, Macrocystis pyrifera, is exposed to highly variable irradiance and temperature regimes across its geographic and vertical depth gradients. The objective of this study was to extend our understanding of algal acclimation strategies on different temporal scales to those varying abiotic conditions at various water depths. Different acclimation strategies to various water depths (0.2 and 4 m) between different sampling times (Jan/Feb and Aug/Sept 2012; long-term acclimation) and more rapid adjustments to different depths (0.2, 2 and 4 m; short-term acclimation) during 14 d of transplantation were found. Adjustments of variable Chl a fluorescence, pigment composition (Chl c, fucoxanthin), and the de-epoxidation state of the xanthophyll cycle pigments were responsible for the development of different physiological states with respect to various solar radiation and temperature climates. Interestingly, the results indicated that phlorotannins are important during long-term acclimation while antioxidants have a crucial role during short-term acclimation. Furthermore, the results suggested that modifications in total lipids and fatty acid compositions apparently also might play a role in depth acclimation. In Aug/Sept (austral winter), M. pyrifera responded to the transplantation from 4 m to 0.2 m depth with a rise in the degree of saturation and a switch from shorter- to longer-chain fatty acids. These changes seem to be essential for the readjustment of thylakoid membranes and might, thus, facilitate efficient photosynthesis under changing irradiances and temperatures. Further experiments are needed to disentangle the relative contribution of solar radiation, temperature and also other abiotic parameters in the observed physiological changes. [ABSTRACT FROM AUTHOR]

Gut content examination and trophic markers (fatty acids, stable isotopes of C and N) were combined to delineate the diet of the dominant species of amphipods from Mediterranean Posidonia oceanica seagrass meadows and to highlight trophic diversity among this community. Our results indicate that, although all dominant species heavily relied on macroalgal epiphytes, considerable interspecific dietary differences existed. Carbon stable isotope ratios notably showed that some of the amphipod species favored grazing on epiphytes from leaves or litter fragments (Apherusa chiereghinii, Aora spinicornis, Gammarus aequicauda), while others such as Dexamine spiniventris preferred epiphytes from rhizomes. The remaining amphipods (Caprella acanthifera, Ampithoe helleri and Gammarella fucicola) readily consumed both groups. In addition, SIAR modeling suggested that most species had a mixed diet, and relied on several food items. Fatty acid analysis and gut contents revealed that contributions of microepiphytic diatoms and of benthic and suspended particulate organic matter to the diet of amphipods were anecdotal. None of the examined species seemed to graze on their seagrass host [low 18:2(n-6) and 18:3(n-3) fatty acids contents], but Gammarus aequicauda partly relied on seagrass leaf detritus, as demonstrated by the lesser 13C-depletion of their tissues. Overall, our findings suggest that amphipods, because of their importance in the transfer of organic matter from primary producers and detritus to higher rank consumers, are key items in P. oceanica-associated food webs. [ABSTRACT FROM AUTHOR]

Polar regions are characterized by a strong seasonality in primary production and distinct fluctuations in food supply. The storage of energy in the form of lipids is expected to follow a seasonal pattern; however, year-round observations are lacking. The seasonal variations of lipid classes and fatty acid compositions of four pelagic amphipods have been investigated. Individuals were collected at various depths during summer and winter expeditions to the Fram Strait and Svalbard archipelago (78°-81°N). Our results show a year-round dominance of wax esters and triacylglycerols over phospholipids for Themisto abyssorum, Themisto libellula, Themisto compressa and Cyclocaris guilelmi. High levels of the Calanus-marker fatty acids 20:1 and 22:1 (both isomers) during summer and winter indicated that all four species are part of the Calanus-based food web. Specific fatty acid trophic markers for diatoms and flagellates indicated that the lipid-based energy transfer could be traced back to the respective algal sources.We found a distinct difference between animals collected from very deep and shallow waters, which indicates that the lipid reservoir of these amphipods and their biochemical modifications seem to buffer seasonal variations in available prey and to allow an active, predatory life-style year-round in Arctic waters. [ABSTRACT FROM AUTHOR]

Plankton fractions from a saline lake in Argentina were studied using a combined trophic marker approach. A strong seasonality of biomarkers was characteristic for the different fractions, particularly the variations in the 18:4(n - 3) and 20:4(n - 3) fatty acids and the δ13C values. The primary production in the lake was mainly driven by diatoms, reflected by the close relation of δ13C, chlorophyll a and diatom fatty acid markers. The combined approach of δ13C and 20:4(n - 3) enabled processes in the lipid metabolism of the copepod Boeckella poopoensis to be inferred. The polyunsaturated fatty acid 22:6(n - 3) and the δ15N separated the trophic levels in this food web with copepods at higher trophic level. Nutritional stress and omnivory of B. poopoensis partially explained the δ15N variations in mesozooplankton. The δ15N signature was probably driven by cyanobacteria in the microplankton and by microbial processes in the nanoplankton fraction. Warmer temperatures may favour the saturation of microalgae fatty acids and the abundance of plankton groups richer in saturated fatty acids. The tendency to unsaturation in mesozooplankton at colder temperatures was probably influenced by diet and metabolic requirements. Future temperature increase and eutrophication-like processes may increase the importance of cyanobacterial and bacterial markers under climate change scenarios. [ABSTRACT FROM AUTHOR]

Lessonia nigrescens used to be an abundant kelp species along the Chilean coast, but recent molecular studies revealed the existence of a L. nigrescens species complex, which includes the two cryptic species Lessonia berteroana and Lessonia spicata. Since these species have different distributions (16°S-30°S for L. berteroana and 29°S-42°S for L. spicata), they experience differences in environmental conditions, such as solar irradiance, seawater temperature and air exposure during low tide. This study tested to what extent the genetic distinctness of each of the two species [identified by a mitochondrial marker ( atp8/trnS)] is reflected by ecophysiological traits (total lipids, fatty acid composition, phlorotannins, pigments and variable chlorophyll a fluorescence of PSII) in response to the respective environmental conditions, prevailing along the latitudinal gradient. We studied algal individuals from eight populations (27°S-32°S, including the species overlapping zone). Phlorotannins, pigments and Chl a fluorescence of PSII were most crucial for species-specific adaptations at the respective growth sites, whereas changes in total lipids and fatty acid compositions were negligible. Hence, species differentiation within the L. nigrescens complex is also manifested at the ecophysiological level. These findings may help to predict kelp responses towards future environmental changes. [ABSTRACT FROM AUTHOR]

Few studies have been made on planktonic food webs of temperate ecosystems, especially those from the Southern Atlantic Ocean, using molecular biomarkers. The fatty acid compositions of suspended particulate matter (SPM), microplankton and mesozooplankton were studied during summer and winter at a sewage-impacted and a control site in the Bahía Blanca Estuary (Argentina). The aim was to identify trophic relationships on a spatial and seasonal scale and to detect allochthonous inputs to the food web. Fatty acid trends were consistent with the seasonal succession of the plankton community structure supporting our underlying hypothesis that regional seasonality is mostly responsible for changes in fatty acid composition. Sewage had no clear impact on the fatty acids and may not be a significant source of SPM in the estuary. However, at the sewage site the composition of the SPM was more related to terrestrially derived compounds, diatoms and bacteria, and mesozooplankton fatty acids suggested grazing on terrestrial components and on diatoms over flagellates. Saltmarshes likely have a crucial role as the main contributors to the organic fraction of SPM followed by plankton. The seasonal fatty acid pattern of the mesozooplankton indicated different feeding strategies suggesting an active feeding mode during summer and a more terrestrially associated diet in winter. The fatty acid trophic marker approach provided relevant information to clarify planktonic trophic interactions and to trace the origin of organic matter in this highly dynamic temperate coastal system. [ABSTRACT FROM AUTHOR]

Antarctic notothenioid fish are characterized by their evolutionary adaptation to the cold, thermostable Southern Ocean, which is associated with unique physiological adaptations to withstand the cold and reduce energetic requirements but also entails limited compensation capacities to environmental change. This study compares the capacities of mitochondrial acclimation to ocean warming and acidification between the Antarctic nototheniid Notothenia rossii and the sub-Antarctic Lepidonotothen squamifrons, which share a similar ecology, but different habitat temperatures. After acclimation of L. squamifrons to 9°C and N. rossii to 7°C (normocapnic/hypercapnic, 0.2 kPa CO2/2000 ppm CO2) for 4–6 weeks, we compared the capacities of their mitochondrial respiratory complexes I (CI) and II (CII), their P/O ratios (phosphorylation efficiency), proton leak capacities and mitochondrial membrane fatty acid compositions. Our results reveal reduced CII respiration rates in warm-acclimated L. squamifrons and cold hypercapnia-acclimated N. rossii. Generally, L. squamifrons displayed a greater ability to increase CI contribution during acute warming and after warm-acclimation than N. rossii. Membrane unsaturation was not altered by warm or hypercapnia-acclimation in both species, but membrane fatty acids of warm-acclimated L. squamifrons were less saturated than in warm normocapnia−/hypercapnia-acclimated N. rossii. Proton leak capacities were not affected by warm or hypercapnia-acclimation of N. rossii. We conclude that an acclimatory response of mitochondrial capacities may include higher thermal plasticity of CI supported by enhanced utilization of anaplerotic substrates (via oxidative decarboxylation reactions) feeding into the citrate cycle. L. squamifrons possesses higher relative CI plasticities than N. rossii, which may facilitate the usage of energy efficient NADH-related substrates under conditions of elevated energy demand, possibly induced by ocean warming and acidification. The observed adjustments of electron transport system complexes with a higher flux through CI under warming and acidification suggest a metabolic acclimation potential of the sub-Antarctic L. squamifrons, but only limited acclimation capacities for N. rossii. [ABSTRACT FROM AUTHOR]

Radiation damage can inter alia result in lipid peroxidation of macroalgal cell membranes. To prevent photo-oxidation within the cells, photoprotective substances such as phlorotannins are synthesized. In the present study, changes in total fatty acids (FA), FA composition and intra/extracellular phlorotannin contents were determined by gas chromatography and the Folin-Ciocalteu method to investigate the photoprotective potential of phlorotannins to prevent lipid peroxidation. Alaria esculenta juveniles (Phaeophyceae) were exposed over 20 days to high/low photosynthetically active radiation (PAR) in combination with UV radiation (UVR) in the treatments: PAB (low/high PAR + UV-B + UV-A), PA (low/high PAR + UV-A) or low/high PAR only. While extracellular phlorotannins increased after 10 days, intracellular phlorotannins increased with exposure time and PA and decreased under PAB. Interactive effects of time:radiation wavebands, time:PAR dose as well as radiation wavebands:PAR dose were observed. Low FA contents were detected in the PA and PAB treatments; interactive effects were observed between time:high PAR and PAB:high PAR. Total FA contents were correlated to extra/intracellular phlorotannin contents. Our results suggest that phlorotannins might play a role in intra/extracellular protection by absorption and oxidation processes. Changes in FA content/composition upon UVR and high PAR might be considered as an adaptive mechanism of the A. esculenta juveniles subjected to variations in solar irradiance. [ABSTRACT FROM AUTHOR]

The copepod Calanus glacialis plays a key role in the lipid-based energy flux in Arctic shelf seas. By utilizing both ice algae and phytoplankton, this species is able to extend its growth season considerably in these seasonally ice-covered seas. This study investigated the impacts of the variability in timing and extent of the ice algal bloom on the reproduction and population success of C. glacialis. The vertical distribution, reproduction, amount of storage lipids, stable isotopes, fatty acid and fatty alcohol composition of C. glacialis were assessed during the Circumpolar Flaw Lead System Study. Data were collected in the Amundsen Gulf, south-eastern Beaufort Sea, from January to July 2008 with the core-sampling from March to April. The reduction in sea ice thickness and coverage observed in the Amundsen Gulf in 2007 and 2008 affected the life strategy and reproduction of C. glacialis. Developmental stages CIII and CIV dominated the overwintering population, which resulted in the presence of very few CV and females during spring 2008. Spawning began at the peak of the ice algal bloom that preceded the precocious May ice break-up. Although the main recruitment may have occurred later in the season, low abundance of females combined with a potential mismatch between egg production/development to the first feeding stage and phytoplankton bloom resulted in low recruitment of C. glacialis in the early summer of 2008. [ABSTRACT FROM AUTHOR]

We have studied the seasonal dynamics of abundance and feeding characteristics of three species of calanoid copepods ( Acartia spp ., Centropages hamatus and Temora longicornis) in the White Sea from the surface water layer (0-10 m), in order to assess their role in the pelagic food web and to determine the major factors governing their population dynamics during the productive season. These species dominated in the upper water layer (0-10 m) from June through September, producing up to 3 generations per year. Data on the food spectra revealed all species to be omnivorous; but some inter- and intraspecific differences were observed. Generally, copepods consumed diatoms, dinoflagellates and microzooplankton. The omnivory index 'UC' (i.e., fatty acid unsaturation coefficient) varied from 0.2 to 0.6, which implied ingestion of phytoplankton. The different degree of selectivity on the same food items by the studied species was observed, and therefore, successful surviving strategy with minimal overlapping could be assumed. In total, the populations of the three studied copepod species grazed up to 2.15 g C m day and released up to 0.68 g C m day in faecal pellets. They consumed up to 50% of particulate organic carbon, or up to 85% of phytoplankton standing stock (in terms of Chl. a), and thus played a significant role in the transformation of particulate organic matter. Seasonal changes in abundance of the studied species depended mostly on water temperature in the early summer, but were also affected by food availability (Chl. a concentration) during the productive season. [ABSTRACT FROM AUTHOR]

Phlorotannins are considered inter alia to act protective to a variety of stressors, while lipids in spores are known to fuel various metabolic processes during spore release and settlement. Here, phlorotannin production in zoospores/juvenile gametophytes in relation to lipid metabolism was investigated for the first time in order to study-related metabolic costs. The experiment was carried out in Ny-Ålesund (Svalbard, Arctic) within the development from spores to juvenile gametophytes of the brown alga Saccharina latissima over 20 days. In the release stage, the total phlorotannin content of single zoospores was 1.5 × 10 μg and 1.9 × 10 μg in the surrounding medium. Upon release, the total amount of lipids was 1.76 × 10 μg lipid zoospore containing the major fatty acids 16:0 and 18:0, 18:1(n-9), 18:2 (n-6), 18:3(n-3), 20:4(n-6), and 20:5(n-3). During development from spores to gametophytes, a decrease in fatty acids was observed via electron microscopy and a decrease in the fatty acid 18:1(n-9) from 45 to 30% was measured by gas chromatography in particular. While phlorotannin content within the spores remained stable, phlorotannins accumulated in the surrounding medium of gametophytes to 4.0 × 10 μg phlorotannins spore indicating exudation processes. Results obtained support the key and multifunctional role of lipids in zoospore/gametophyte metabolism and may indicate that gametophytes of S. latissima need approximately 10 days to switch to photo-autotrophic strategies, becoming independent of storage lipids. In addition, fatty acids might represent an essential energy source to fuel adaptive responses. [ABSTRACT FROM AUTHOR]

The Arctic bloom consists of two distinct categories of primary producers, ice algae growing within and on the underside of the sea ice, and phytoplankton growing in open waters. Long chain omega-3 fatty acids, a subgroup of polyunsaturated fatty acids (PUFAs) produced exclusively by these algae, are essential to all marine organisms for successful reproduction, growth, and development. During an extensive field study in the Arctic shelf seas, we followed the seasonal biomass development of ice algae and phytoplankton and their food quality in terms of their relative PUFA content. The first PUFA-peak occurred in late April during solid ice cover at the onset of the ice algal bloom, and the second PUFA-peak occurred in early July just after the ice break-up at the onset of the phytoplankton bloom. The reproduction and growth of the key Arctic grazer Calanus glacialis perfectly coincided with these two bloom events. Females of C. glacialis utilized the high-quality ice algal bloom to fuel early maturation and reproduction, whereas the resulting offspring had access to ample high-quality food during the phytoplankton bloom 2 months later. Reduction in sea ice thickness and coverage area will alter the current primary production regime due to earlier ice break-up and onset of the phytoplankton bloom. A potential mismatch between the two primary production peaks of high-quality food and the reproductive cycle of key Arctic grazers may have negative consequences for the entire lipid-driven Arctic marine ecosystem. [ABSTRACT FROM AUTHOR]

We present an accurate, fast, simple and non-destructive photographic method to estimate wax ester and lipid content in single individuals of the calanoid copepod genus Calanus and test this method against gas-chromatographic lipid measurements. [ABSTRACT FROM PUBLISHER]

In coastal waters, Antarctic rhodophytes are exposed to harsh environmental conditions throughout the year, like low water temperatures ranging from −1.8°C to 2°C and high light during the summer season. Photosynthetic performance under these conditions may be affected by slowed down enzymatic reactions and the increased generation of reactive oxygen species. The consequence might be a chronic photoinhibition of photosynthetic primary reactions related to increased fragmentation of the D1 reaction centre protein in photosystem II. It is hypothesized that changes in lipid composition of biomembranes may represent an adaptive trait to maintain D1 turnover in response to temperature variation. The interactive effects of high light and low temperature were studied on an endemic Antarctic red alga, Palmaria decipiens, sampled from two shore levels, intertidal and subtidal, and exposed to mesocosm experiments using two levels of natural solar radiation and two different temperature regimes (2–5°C and 5–10°C). During the experimental period of 23 days, maximum quantum yield of photosynthesis decreased in all treatments, with the intertidal specimens exposed at 5–10°C being most affected. On the pigment level, a decreasing ratio of phycobiliproteins to chlorophyll a was found in all treatments. A pronounced decrease in D1 protein concentration occurred in subtidal specimens exposed at 2–5°C. Marked changes in lipid composition, i.e. the ratio of saturated to unsaturated fatty acids, indicated an effective response of specimens to temperature change. Results provide new insights into mechanisms of stress adaptation in this key species of shallow Antarctic benthic communities. [ABSTRACT FROM AUTHOR]

Lipid content, fatty acid composition, and feeding activity of the dominant Antarctic copepods, Calanoides acutus, Calanus propinquus, and Metridia gerlachei, were studied at a quasi-permanent station in the eastern Weddell Sea in December 2003. During 3 weeks of the spring phytoplankton development, total lipid levels of females and copepodite stages V (CVs) of C. acutus were almost doubled. Meanwhile, only a slight increase in total lipid content occurred in M. gerlachei, and no clear trend was observed in lipids of C. propinquus females. The pronounced increase of lipids in C. acutus was due to an accumulation of wax esters. The proportion of wax esters in the lipids of M. gerlachei was clearly lower, while triacylglycerols played a more important role. In C. propinquus, triacylglycerols were the only neutral lipid class. There were no pronounced changes in the feeding activity of M. gerlachei, whereas the feeding activity of C. acutus had rapidly increased with the development of the phytoplankton bloom in December, which explains its rapid lipid accumulation. The combination of gut content and fatty acid trophic marker analyses showed that C. acutus was feeding predominantly on diatoms. The typical diatom fatty acid marker, 16:1(n-7), slightly decreased and the tracer for flagellates, 18:4(n-3), increased in females and CVs of C. acutus. This shift indicates the time, when the significance of flagellates started to increase. The three copepod species exhibited different patterns of lipid accumulation in relation to their trophic niches and different duration of their active phases. The investigations filled a crucial data gap in the seasonal lipid dynamics of dominant calanoid copepods in the Weddell Sea in December and support earlier hypotheses on their energetic adaptations and life cycle strategies. [ABSTRACT FROM AUTHOR]

Seasonal dynamics of major biochemical features were studied for three abundant egg-diapausing copepods Acartia bifilosa, Centropages hamatus and Temora longicornis, in the White Sea (66°N), between June 2002 and September 2002. Dry weight (DW) and prosome length varied from 0.54 μg ind−1 and 0.163 ± 0.012 mm ( A. bifilosa, CI) to 9.58 ± 0.72 μg ind−1 and 1.135 ± 0.167 mm ( C. hamatus, females). Corg and Norg content reached up to 5.91 ± 0.44 and 1.23 ± 0.09 μg ind−1 ( C. hamatus, females). Protein and lipid content varied greatly from 31.8 to 67.3% DW and from 8.7 to 42.6% DW, respectively. These species show somewhat different biology compared to species at lower latitudes. The copepods use lipid stores to survive during short-term food shortage (e.g. in autumn) and successfully complete their life cycle. In the isolated White Sea during last post-glacial period, species probably evolved some special biochemical features (especially wax esters presence). Food quality demands and long ice coverage are possible factors limiting early development of species in spring. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

The Arctic pteropod Clione limacina was collected in Kongsfjorden, Svalbard, in mid June 2004, to study the lipid metabolism within the sites of lipid storage structures during long-term starvation. Animals survived in an aquarium without any food for nearly 1 year (356 days). Size, number of lipid droplets, dry and lipid mass, lipid class and fatty acid compositions of C. limacina were determined and separately analysed for the digestive gland and the remaining integument. During the starvation period, animals shrunk from 22.4 to 12 mm in length on average, and the number of lipid droplets decreased from 1,600 to 1,000 per animal. Dry mass (DM) and total lipid mass both dropped by about 80% from day 200 to the end. The lipid content as percentage DM of the total organism did not decrease significantly ranging from 43.8 to 32.3%DM. The lipid content of the trunk was moderate with about 20%DM. The digestive gland was very rich in lipids with more than 70%DM throughout the experiment and is the major site of lipid metabolism and storage. Triacylglycerols (TAG) decreased, in the total organism, from high initial levels of 62.6 to 43% of total lipid at the end. In contrast, the proportions of 1- O-alkyldiacylglycerols [diacylglycerol ethers (DAGE)] remained almost constant, varying between 20.4 and 28.4%. In the digestive gland, TAG ranged from 60.3 to 64.8% and DAGE from 23.6 to 32.2% from day 200 to the end of the experiment. TAG and DAGE of the trunk were most likely located in the lipid droplets and were almost depleted at the end of starvation. Besides their function as lipid deposit DAGE may also act as protecting substance against bacterial and fungal infections. During the first 200 days of starvation, the fatty acid compositions showed only small variations. Thereafter, fatty acids typical for storage lipids decreased in all body compartments. In adaptation to long periods of food scarcity, C. limacina has evolved various strategies as body shrinkage, utilisation of body constituents not essential for survival, a very low metabolism and slow lipid consumption. [ABSTRACT FROM AUTHOR]

Calorie restriction (CR) without undernutrition has been found to enhance stress resistance and life span in endotherms and ectotherms. We investigated the effect of 30% reduction in food offering on growth, aerobic capacities and oxidative stress parameters in young turbot ( Scophthalmus maximus, L.). No differences in body weight, length and hepatosomatic index between the ad libitum (AL)-fed and the CR group occurred after 55 days of diet application. Of the mitochondrial marker enzymes, only citrate synthase (CS) activity in the liver was reduced under CR, whereas muscle CS activity and cytochrome oxidase (CCO) activity in both tissues remained the same in both feeding groups. The concentration of reduced glutathione increased significantly during feeding in muscle of CR fish, resulting in a more reduced glutathione redox ratio (GSH/GSSG) compared with AL fish muscle. Thiobarbituric acid-reactive lipid hydroperoxides (lipid peroxidation) but not protein carbonyl content (protein oxidation) was significantly reduced in CR fish muscle. Liver oxidative stress parameters did not vary significantly between experimental feeding groups. We conclude that 30% CR over 8 weeks has no adverse effect on young turbot. On the contrary, CR supports a reduced tissue oxidation state and reduces accumulation of lipid peroxidation products in muscle at sustained muscular aerobic capacity. [ABSTRACT FROM AUTHOR]

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

The sea butterfly Limacina helicina was collected from May to September 2001 in Kongsfjorden, Spitsbergen, to investigate population structure and body and lipid composition with regard to life cycle and reproductive strategy. Veligers and juveniles were only found in late autumn and spring, whereas females occurred from July to September. The size of the females increased until mid-August and decreased in September. Dry and lipid mass were closely related to size; dry mass increased exponentially and lipids linearly with size. The lipid content was highest in veligers (31.5% of dry mass) and juveniles (23.6%) but low in females (<10%). Phospholipids were the dominating lipid class followed by triacylglycerols. Females, veligers, and egg ribbons, all from September, were richest in phospholipids. Juveniles contained the highest amounts of triacylglycerols, whereas females had low levels in July and the beginning of August. In mid-August, levels of triacylglycerols were variable and higher. This suggests that females were in their main spawning period and the high variability in triacylglycerols points to different stages within the spawning cycle. Enhanced amounts of free fatty acids in females from July may be related to gonad development. The 16:1(n-7) fatty acid was more dominant in spring whereas 18:4(n-3) increased in summer and autumn, which reflects a change in diet from diatom-dominated food items in spring to dinoflagellates in summer/autumn. Small amounts of long-chain monounsaturated fatty acids suggest ingestion of copepods, and the fatty acid composition of veligers feeding on particulate matter. L. helicina has a one-year life cycle with peak spawning in August and over-winters as veligers that may grow to juveniles during the winter period. They metamorphose into juveniles during spring, develop to males in early summer, and mature into females in July and August. [ABSTRACT FROM AUTHOR]

Limnocalanus macrurus is an important member of the zooplankton communities of the Siberian shelf seas. During the cruise, Boris Petrov 1999, in August/September to the southern Kara Sea and the Ob and Yenisej estuaries, its abundance and vertical distribution were investigated. In adults, salinity tolerance, egg production, feeding and lipid composition were studied. L. macrurus occurred in water with salinities ranging from 1.7 to >33 without clear preference, as revealed from salinity-tolerance experiments. The dominance of adults and their high wax-ester content, as well as the lack of egg production and feeding activity, suggest that the population was in the pre-overwintering condition. Wax esters allow L. macrurus to survive long starvation periods and to reproduce in times of little food availability, but through its potential carnivory, it should be able to replenish its diet by preying on other zooplankton. Morphology and swimming behaviour of L. macrurus resemble the omnivorous copepod Metridia longa, which, however, is mainly found in the open ocean. The overall lipid composition and the mode of lipid storage also point to an omnivorous feeding behaviour. However, the high proportion of the marker fatty acid 16:1(n-7) suggests that L. macrurus strongly exploited the existent phytoplankton bloom, consisting mainly of diatoms. A striking characteristic of its lipids is the high level of the 20:1(n-7) fatty alcohol in addition to the 18:1(n-7) fatty acid and alcohol. It is the first copepod species known to produce such high amounts of 20:1(n-7) alcohol. Since this alcohol and the corresponding fatty acid are not abundant in any prey, this long-chain monounsaturated wax-ester moiety has to be produced de novo. Owing to these particular lipid characteristics in its distribution, feeding, and life-cycle strategy, L. macrurus can be described as a very versatile and opportunistic copepod. [ABSTRACT FROM AUTHOR]

Climate change has led to a ~ 40% reduction in summer Arctic sea-ice cover extent since the 1970s. Resultant increases in light availability may enhance phytoplankton production. Direct evidence for factors currently constraining summertime phytoplankton growth in the Arctic region is however lacking. GEOTRACES cruise GN05 conducted a Fram Strait transect from Svalbard to the NE Greenland Shelf in summer 2016, sampling for bioessential trace metals (Fe, Co, Zn, Mn) and macronutrients (N, Si, P) at ~ 79°N. Five bioassay experiments were conducted to establish phytoplankton responses to additions of Fe, N, Fe + N and volcanic dust. Ambient nutrient concentrations suggested N and Fe were deficient in surface seawater relative to typical phytoplankton requirements. A west-to-east trend in the relative deficiency of N and Fe was apparent, with N becoming more deficient towards Greenland and Fe more deficient towards Svalbard. This aligned with phytoplankton responses in bioassay experiments, which showed greatest chlorophyll-a increases in + N treatment near Greenland and + N + Fe near Svalbard. Collectively these results suggest primary N limitation of phytoplankton growth throughout the study region, with conditions potentially approaching secondary Fe limitation in the eastern Fram Strait. We suggest that the supply of Atlantic-derived N and Arctic-derived Fe exerts a strong control on summertime nutrient stoichiometry and resultant limitation patterns across the Fram Strait region. [ABSTRACT FROM AUTHOR]

Assessing the relative importance of sea ice algal-based production is often vital for studies about climate change impacts on Arctic marine ecosystems. Several types of lipid biomarkers and stable isotope ratios are widely used for tracing sea ic-associated (sympagic) vs. pelagic particulate organic matter (POM) in marine food webs. However, there has been limited understanding about the plasticity of these compounds in space and time, which constrains the robustness of some of those approaches. Furthermore, some of the markers are compromised by not being unambiguously specific for sea ice algae, whereas others might only be produced by a small sub-group of species. We analyzed fatty acids, highly branched isoprenoids (HBIs), stable isotope ratios of particulate organic carbon (POC) (δ13C), as well as δ13C of selected fatty acid markers during an Arctic sea ice algal bloom, focusing on spatial and temporal variability. We found remarkable differences between these approaches and show that inferences about bloom characteristics might even be contradictory between markers. The impact of environmental factors as causes of this considerable variability is highlighted and explained. We emphasize that awareness and, in some cases, caution is required when using lipid and stable isotope markers as tracers in food web studies and offer recommendations for the proper application of these valuable approaches. [ABSTRACT FROM AUTHOR]

Sea ice algae can constitute an important carbon source for high-Antarctic euphausiids during winter. To quantify the importance of this 'sympagic carbon' during summer, the three most abundant Antarctic euphausiids, Euphausia superba, E. crystallorophias, and Thysanoessa macrura, collected off the Filchner Ice Shelf, were analyzed regarding their fatty acid (FA) and stable isotope compositions. Fingerprints of diatom- and dinoflagellate-associated FAs in the euphausiids indicated a mixed carbon source composition for all three species. Bulk and FA-specific carbon stable isotope compositions (δ13C) were used to quantify the contribution of sympagic carbon versus phytoplankton-produced carbon to the euphausiids' carbon budget, suggesting a lower proportional contribution of sympagic carbon in E. superba (5–18%) compared to E. crystallorophias (16–36%) and T. macrura (15–36%). The latter two species probably received sympagic carbon through heterotrophic prey, a hitherto overlooked source of sympagic carbon for pelagic species. Euphausiids collected close to the surface indicated a higher importance of sympagic carbon to their carbon budget compared to individuals caught at greater depths. Our results imply that, in the southern Weddell Sea, ice algae play a significant, but possibly not critical role as a carbon source for the three euphausiids during summer. Their ability to utilize carbon of different origins implies a certain resilience to environmental change during summer. The winter period, however, remains the critical bottle neck of survival when Antarctic sea ice declines, because during this season of minimal pelagic productivity, ice algae standing stocks constitute the only dependable carbon source. [ABSTRACT FROM AUTHOR]

Maternal investment (MI), the energy allocated by mothers to offspring, has important effects on the life-history traits of marine organisms. Variation in such traits shows strong correlation with latitude for several marine taxa (Thorson’s rule). Large-scale latitudinal variation in MI within a single species suggests population genetic divergence, while temporal changes in MI, rather, reflect plasticity. At higher latitudes (i.e., colder waters), traits associated with MI (brood weight, fecundity, egg volume, and energy content) increase. To identify phenotypic plasticity along a latitudinal gradient in MI traits (brood weight, egg volume, density number, and egg lipid composition), five populations of the kelp crab Taliepus dentatus along the coast of Chile (30°S-42°S) were investigated during the summer (December-February) and winter months (June-August) of 2015-2016. Despite this wide latitudinal range, the sea surface temperature (SST) difference between the northernmost and the southernmost sites was only approximately 2.0 °C in winter and 5.5 °C in summer. In summer, when latitudinal variation in SST was highest, brood weight, egg density, fecundity, and egg lipids increased with latitude, while egg volume decreased. No trends in MI were observed in winter when the SST gradient was almost non-existent. These results suggest that the relationship between MI and latitude is shaped by temperature rather than being site-specific. The seasonality of latitudinal MI traits also suggests a trade-off between the costs of female maintenance and/or brooding behaviours and MI. When investigating latitudinal and temporal variation in marine brooder MI, the effect of temperature on life-history traits and the associated costs of female brooding should be quantified. [ABSTRACT FROM AUTHOR]

The rhodophytes Mastocarpus stellatus and Chondrus crispus occupy the lower intertidal zone of rocky shores along North Atlantic coastlines, with C. crispus generally occurring slightly deeper. Consequently, M. stellatus is exposed to more variable environmental conditions, related to a generally higher stress tolerance of this species. In order to extend our understanding of seasonal modulation of stress tolerance, we subjected local populations of M. stellatus and C. crispus from Helgoland, North Sea, to short-term high-light stress experiments over the course of a year (October 2011, March, May and August 2012). Biochemical analyses (pigments, antioxidants, total lipids, fatty acid compositions) allowed to reveal mechanisms behind modulated high-light tolerances. Overall, C. crispus was particularly more susceptible to high-light at higher water temperatures (October 2011 and August 2012). Furthermore, species-specific differences in antioxidants, total lipid levels and the shorter-chain/longer-chain fatty acid ratio (C14 + C16/C18 + C20) were detected, which may enhance the tolerance to high-light and other abiotic stress factors in M. stellatus, so that this species is more competitive in the highly variable upper intertidal zone compared to C. crispus. Since the high-light tolerance in C. crispus seemed to be affected by water temperature, interactions between both species may be impacted in the future by rising mean annual sea surface temperature around the island of Helgoland. [ABSTRACT FROM AUTHOR]