Fatty acids have been used as qualitative markers to trace or confirm predator-prey relationships in the marine environment for more than thirty years. More recently, they have also been used to identify key processes impacting the dynamics of some of the world's major ecosystems. The fatty acid trophic marker (FATM) concept is based on the observation that marine primary producers lay down certain fatty acid patterns that may be transferred conservatively to, and hence can be recognized in, primary consumers. To identify these fatty acid patterns the literature was surveyed and a partial least squares (PLS) regression analysis of the data was performed, validating the specificity of particular microalgal FATM. Microalgal group specific FATM have been traced in various primary consumers, particularly in herbivorous calanoid copepods, which accumulate large lipid reserves, and which dominate the zooplankton biomass in high latitude ecosystems. At higher trophic levels these markers of herbivory are obscured as the degree of carnivory increases, and as the fatty acids originate from a variety of dietary sources. Such differences are highlighted in a PLS regression analysis of fatty acid and fatty alcohol compositional data (the components of wax esters accumulated by many marine organisms) of key Arctic and Antarctic herbivorous, omnivorous and carnivorous copepod species. The analysis emphasizes how calanoid copepods separate from other copepods not only by their content of microalgal group specific FATM, but also by their large content of long-chain monounsaturated fatty acids and alcohols. These monounsaturates have been used to trace and resolve food web relationships in, for example, hyperiid amphipods, euphausiids and fish, which may consume large numbers of calanoid copepods. Results like these are extremely valuable for enabling the discrimination of specific prey species utilized by higher trophic level omnivores and carnivores without the employment of invasive techniques, and thereby for identifying the sources of energetic reserves. A conceptual model of the spatial and temporal dominance of group-specific primary producers, and hence the basic fatty acid patterns available to higher trophic levels is presented. The model is based on stratification, which acts on phytoplankton group dominance through the availability of light and nutrients. It predicts the seasonal and ecosystem specific contribution of diatom and flagellate/microbial loop FATM to food webs as a function of water column stability. Future prospects for the application of FATM in resolving dynamic ecosystem processes are assessed.

Fatty acid trophic markers in the pelagic marine environment.

Zooplankton storage lipids play an important role during reproduction, food scarcity, ontogeny and diapause, as shown by studies in various oceanic regions. While triacylglycerols, the primary storage lipid of terrestrial animals, are found in almost all zooplankton species, wax esters are the dominant storage lipid in many deep-living and polar zooplankton taxa. Phospholipids and diacylglycerol ethers are the unique storage lipids used by polar euphausiids and pteropods, respec- tively. In zooplankton with large stores of wax esters, triacylglycerols are more rapidly turned over and used for short-term energy needs, while wax esters serve as long-term energy deposits. Zooplankton groups found in polar, westerlies, upwelling and coastal biomes are characterized by accumulation of large lipid stores. In contrast, zooplankton from the trades/tropical biomes is mainly composed of omnivorous species with only small lipid reserves. Diapausing copepods, which enter deep water after feeding on phytoplankton during spring/summer blooms or at the end of upwelling periods, are characterized by large oil sacs filled with wax esters. The thermal expansion and com- pressibility of wax esters may allow diapausing copepods and other deep-water zooplankton to be neutrally buoyant in cold deep waters, and they can thus avoid spending energy to remain at these depths. Lipid droplets are often noted in zooplankton ovaries, and a portion of these droplets can be transferred to developing oocytes. In addition to lipid droplets, zooplankton eggs have yolks with lipovitellin, a lipoprotein with approximately equal amounts of protein and lipid. The lipovitellin lipid is predominantly phosphatidylcholine, so during reproduction females must convert a portion of their storage lipid into this phospholipid. Developing embryos use their lipovitellin and lipid droplets for energy and materials until feeding begins. The various functions storage lipids serve during the dif- ferent life history stages of zooplankton are very complex and still not fully understood and hence offer a multitude of fascinating research perspectives.

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Lipid storage in marine zooplankton

Kongsfjorden is a glacial fjord in the Arctic (Svalbard) that is influenced by both Atlantic and Arctic water masses and harbours a mixture of boreal and Arctic flora and fauna. Inputs from large tidal glaciers create steep environmental gradients in sedimentation and salinity along the length of this fjord. The glacial inputs cause reduced biomass and diversity in the benthic community in the inner fjord. Zooplankton suffers direct mortality from the glacial outflow and primary production is reduced because of limited light levels in the turbid, mixed inner waters. The magnitude of the glacial effects diminishes towards the outer fjord. Kongsfjorden is an important feeding ground for marine mammals and seabirds. Even though the fjord contains some boreal fauna, the prey consumed by upper trophic levels is mainly Arctic organisms. Marine mammals constitute the largest top-predator biomass, but seabirds have the largest energy intake and also export nutrients and energy out of the marine environment. Kongsfjorden has received a lot of research attention in the recent past. The current interest in the fjord is primarily based on the fact that Kongsfjorden is particularly suitable as a site for exploring the impacts of possible climate changes, with Atlantic water influx and melting of tidal glaciers both being linked to climate variability. The pelagic ecosystem is likely to be most sensitive to the Atlantic versus Arctic influence, whereas the benthic ecosystem is more affected by long-term changes in hydrography as well as changes in glacial runoff and sedimentation. Kongsfjorden will be an important Arctic monitoring site over the coming decades and a review of the current knowledge, and a gap analysis, are therefore warranted. Important knowledge gaps include a lack of quantitative data on production, abundance of key prey species, and the role of advection on the biological communities in the fjord.

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The marine ecosystem of Kongsfjorden, Svalbard

Higher plants resist the forces of gravity and powerful lateral forces through the cumulative strength of the walls that surround individual cells. These walls consist mainly of cellulose, noncellulosic polysaccharides and lignin, in proportions that depend upon the specific functions of the cell and its stage of development. Spatially and temporally controlled heterogeneity in the physicochemical properties of wall polysaccharides is observed at the tissue and individual cell levels, and emerging in situ technologies are providing evidence that this heterogeneity also occurs across a single cell wall. We consider the origins of cell wall heterogeneity and identify contributing factors that are inherent in the molecular mechanisms of polysaccharide biosynthesis and are crucial for the changing biological functions of the wall during growth and development. We propose several key questions to be addressed in cell wall biology, together with an alternative two-phase model for the assembly of noncellulosic polysaccharides in plants.

Heterogeneity in the chemistry, structure and function of plant cell walls

The North Water Polynya is an area of high biological activity that supports large numbers of higher trophic-level organisms such as seabirds and marine mammals. An overall objective of the Upper Trophic-Level Group of the International North Water Polynya Study (NOW) was to evaluate carbon and contaminant flux through these high trophic-level (TL) consumers. Crucial to an evaluation of the role of such consumers, however, was the establishment of primary trophic linkages within the North Water food web. We used δ15N values of food web components from particulate organic matter (POM) through polar bears (Ursus maritimus) to create a trophic-level model based on the assumptions that Calanus hyperboreus occupies TL 2.0 and there is a 2.4‰ trophic enrichment in 15N between birds and their diets, and a 3.8‰ trophic enrichment for all other components. This model placed the planktivorous dovekie (Alle alle) at TL 3.3, ringed seal (Phoca hispida) at TL 4.5, and polar bear at TL 5.5. The copepods C. hyperboreus, Chiridius glacialis and Euchaeta glacialis formed a trophic continuum (TL 2.0–3.0) from primary herbivore through omnivore to primary carnivore. Invertebrates were generally sorted according to planktonic, benthic and epibenthic feeding groups. Seabirds formed three trophic groups, with dovekie occupying the lowest, black-legged kittiwake (Rissa tridactyla), northern fulmar (Fulmarus glacialis), thick-billed murre (Uria aalge), and ivory gull (Pagophilia eburnea) intermediate (TL 3.9–4.0), and glaucous gull (Larus hyperboreus) the highest (TL 4.6) trophic positions. Among marine mammals, walrus (Odobenus rosmarus) occupied the lowest (TL 3.2) and bearded seal (Erignathus barbatus), ringed seal, beluga whale (Delphinapterus leucas), and narwhal (Monodon monoceros) intermediate positions (TL 4.1–4.6). In addition to arctic cod (Boreogadus saida), we suggest that lower trophic-level prey, in particular the amphipod Themisto libellula, contribute fundamentally in transferring energy and carbon flux to higher trophic-level seabirds and marine mammals. We measured PCB 153 among selected organisms to investigate the behavior of bioaccumulating contaminants within the food web. Our isotopic model confirmed the trophic magnification of PCB 153 in this high-Arctic food web due to a strong correlation between contaminant concentration and organism δ15N values, demonstrating the utility of combining isotopic and contaminant approaches to food-web studies. Stable-carbon isotope analysis confirmed an enrichment in 13C between POM and ice algae (–22.3 vs. –17.7‰). Benthic organisms were generally enriched in 13C compared to pelagic species. We discuss individual species isotopic data and the general utility of our stable isotope model for defining carbon flux and contaminant flow through the North Water food web.

http://web4.uwindsor.ca/users/f/fisk/main.nsf/0/48abbaa50ed52cc08525724a006839ac/$FILE/DSR%202002%20Hobson%20NOW%20SI%20food%20web.pdf

A stable isotope (δ13C, δ15N) model for the North Water food web: implications for evaluating trophodynamics and the flow of energy and contaminants

Stage IV and V copepodites were the dominant forms of Calanus finmarchicus, C. glacialis and C. hyperboreus in Kongsfjorden in late September 1997. Stage IV and V copepodites of C. glacialis and C. hyperboreus were rich in lipid, largely wax esters, and were well fitted to overwinter. Stage IV copepodites of C. finmarchicus were also rich in wax esters, but stage V copepodites of C. finmarchicus were less wax ester-rich. Large size increments between stage IV and V copepodites and between stage V copepodites and females were noted in C. finmarchicus. A very large increment between stage IV and V copepodites was noted for C. glacialis but the size difference between stage V copepodites and females was very small in this species. Particularly large increments were noted between stage IV and V copepodites of C. hyperboreus and also between stage V copepodites and females of this species. The very large, wax ester-rich C. hyperboreus is well adapted to survive the most extreme variations in the Arctic, in Arctic basin waters, whereas the smaller, wax ester-rich C. glacialis is adapted to survive less extreme Arctic variations, as in Arctic shelf waters. The smallest of the three, C. finmarchicus, is best adapted to survive the more predictable waters of the North Atlantic and the Barents Sea.

Lipids and life strategies of Calanus finmarchicus, Calanus glacialis and Calanus hyperboreus in late autumn, Kongsfjorden, Svalbard

Gammarus wilkitzkii, Apherusa glacialis, Onismus nanseni, Onismus glacialis, Boreogadus saida, Parathemisto libellula and Calanus hyperboreus, collected in late June in the Barents Sea marginal ice zone, contained substantial levels (28–51% of the dry mass) of total lipid, the highest levels (51% and 41% respectively) being in A. glacialis and C. hyperboreus. Neutral lipids were present in greater amounts than polar lipids in all species. Triacylglycerols were major neutral lipids in A. glacialis, G. wilkitzkii and O. nanseni; triacylglycerols and wax esters were present in similar amounts in O. glacialis; higher levels of wax esters than triacylglycerols occurred in P. libellula; wax esters greatly exceeded triacylglycerols in C. hyperboreus, the opposite being true for B. saida. Diatom fatty acid markers were prominent in the triacylglycerols of G. wilkitzkii, O. nanseni, O. glacialis and, particularly, of A. glacialis; 20:1(n-9) and 22:1(n-11) moieties were abundant in wax esters of G. wilkitzkii, O. nanseni, O. glacialis, P. libellula and C. hyperboreus, and in triacylglycerols of B. saida. We deduce that A. glacialis feeds mainly on ice algae and phytodetritus, G. wilkitzkii and the Onismus spp. feed on calanoid copepods as well as ice algae, whereas P. libellula and especially B. saida feed extensively on calanoid copepods.

Lipids and trophic interactions of ice fauna and pelagic zooplankton in the marginal ice zone of the Barents Sea

The percentage (%) fatty alcohol and fatty acid compositions of the wax esters of large numbers of Stage V and females of Calanus hyperboreus, C. glacialis and C. finmarchicus taken in late August to late September from Arctic waters (Kongsfjord in Svalbard, 78° 57' N, 11° 50' E) are presented. The data reveal that these stages of development of the 3 species can be discriminated on the basis of the % of 22:1n-11 fatty alcohol in their large levels of wax esters, with C. hyperboreus having the highest % followed by C. finmarchicus and then C. glacialis. Equally, C. hyperboreus has the lowest % of 20:1n-9 fatty alcohol in its wax esters with C. finmarchicus having a higher % and C. glacialis the highest %. Relatively minor differences occur in the fatty acid compositions of the wax esters of the 3 species, which consisted principally of 20:1n-9 (15 to 18%) and 22:1n-11 (10 to 15%), together with the diatom-derived fatty acids 16:1n-7 (20 to 23%) and 20:5n-3 (11 to 13%). The flagellate-derived fatty acids, 18:4n-3 (3 to 6%) and 22:6n-3 (1 to 3%), were minor constituents. The fatty acid compositions of the small amounts of polar lipid in the 3 species were indistinguishable with 22:6n-3 (41 to 46%) and 20:5n-3 (22 to 24%) being the major components. We conclude that Stage V and females of the species can be distinguished in autumn on the basis of the different % of 22:1n-11 and 20:1n-9 fatty alcohols in their wax esters and that de novo lipid biosynthetic activity in the cope- pods increases in the order C. finmarchicus < C. glacialis < C. hyperboreus. We discuss the results in terms of the contributions of fatty acids and fatty alcohols biosynthesised de novo and fatty acids derived from the diet to the copepods' lipids, the role of 20:1 and 22:1 fatty alcohols and fatty acids as energy sources, and the possible role of 22:6n-3 in the copepods' physiology.

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Species differences, origins and functions of fatty alcohols and fatty acids in the wax esters and phospholipids of Calanus hyperboreus, C. glacialis and C. finmarchicus from Arctic waters

The lipid class compositions of the ctenophores Mertensia ovum and Beroe cucumis were very similar, with polar lipid and wax esters each accounting for 35 to 40% of the total. The fatty acid compositions of the polar lipids in the 2 species were essentially the same, with 22:6n3 and 20:5n3 in a ratio of 2:1 accounting for more than 40% of the total. The fatty acid and fatty alcohol com- positions of the wax esters of both species were also essentially the same, with 20:1n9 and 22:1n11 fatty alcohols present in equal amounts accounting for 60% of the total fatty alcohol composition. The fatty acid and fatty alcohol compositions of the wax esters of M. ovum and B. cucumis were averaged and compared, using principal component analyses, to averages derived from published data for the potential prey species: Calanus finmarchicus, C. glacialis, C. hyperboreus, Pseudocalanus acuspes, Acartia longiremis and Metridia longa. The results were consistent with Calanus spp., especially C. glacialis, being the major prey of M. ovum and with M. ovum being the major prey of B. cucumis.

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Lipid biomarkers and trophic linkages between ctenophores and copepods in Svalbard waters

Galactomannan deposition was investigated in developing endosperms of three leguminous species representative of taxonomic groups which have galactomannans with high, medium and low galactose content. These were fenugreek (Trigonella foenum-graecum L.; mannose/galactose (Man/Gal) = 1.1), guar (Cyamopsis tetragonoloba (L.) Taub.; Man/Gal = 1.6) and Senna occidentalis (L.) Link. (Man/Gal = 3.3), respectively. Endosperms were analysed at different stages of seed development for galactomannan content and the levels, in cell-free extracts, of a mannosyltransferase and a galactosyltransferase which have been shown to catalyse galactomannan biosynthesis in vitro (M. Edwards et al., 1989, Planta 178, 41-51). There was a close correlation in each case between the levels of the biosynthetic mannosyl- and galactosyltransferases and the deposition of galactomannan. The relative in vitro activities of the mannosyl- and galactosyltransferases in fenugreek and guar were similar, and almost constant throughout the period of galactomannan deposition. In Senna the ratio mannosyltransferase/galactosyltransferase was always higher than in the other two species, and it increased substantially throughout the period of galactomannan deposition. In fenugreek and guar the galactomannans present in the endosperms of seeds at different stages of development had the Man/Gal ratios characteristic of the mature seeds. By contrast the galactomannan present in Senna endosperms at the earliest stages of deposition had a Man/Gal ratio of about 2.3. During late deposition this ratio increased rapidly, stabilising at about 3.3, the ratio characteristic of the mature seed. The levels of α-galactosidase in the developing endosperms of fenugreek and guar were low and remained fairly constant throughout the deposition of the galactomannan. In Senna, α-galactosidase activity in the endosperm was low during early galactomannan deposition, but increased subsequently, peaking during late galactomannan deposition. The developmental patterns of the α-galactosidase activity and of the increase in Man/Gal ratio of the Senna galactomannan were closely similar, indicating a cause-and-effect relationship. The endosperm α-galactosidase activity in Senna was capable, in vitro, of removing galactose from guar galactomannan without prior depolymerisation of the molecule. In fenugreek and in guar the genetic control of the Man/Gal ratio in galactomannan is not the result of a post-depositional modification, and must reside in the biosynthetic process. In Senna, the Man/Gal ratio of the primary biosynthetic galactomannan product is controlled by the biosynthetic process. Yet the final Man/Gal ratio of the galactomannan in the mature seed is, to an appreciable extent, the result of galactose removal from the primary biosynthetic product by an α-galactosidase activity which is present in the endosperm during late galactomannan deposition.

Catherine L. Scott

Papers

Lipids and life strategies of Calanus finmarchicus, Calanus glacialis and Calanus hyperboreus in late autumn, Kongsfjorden, Svalbard

Lipids and trophic interactions of ice fauna and pelagic zooplankton in the marginal ice zone of the Barents Sea

Species differences, origins and functions of fatty alcohols and fatty acids in the wax esters and phospholipids of Calanus hyperboreus, C. glacialis and C. finmarchicus from Arctic waters

Lipid biomarkers and trophic linkages between ctenophores and copepods in Svalbard waters

Control of mannose/galactose ratio during galactomannan formation in developing legume seeds.