Gerhard Kattner

Fatty acid trophic markers in the pelagic marine environment

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 worlds 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.

Microbial production of recalcitrant dissolved organic matter: long-term carbon storage in the global ocean

The biological pump is a process whereby CO2 in the upper ocean is fixed by primary producers and transported to the deep ocean as sinking biogenic particles or as dissolved organic matter. The fate of most of this exported material is remineralization to CO2, which accumulates in deep waters until it is eventually ventilated again at the sea surface. However, a proportion of the fixed carbon is not mineralized but is instead stored for millennia as recalcitrant dissolved organic matter. The processes and mechanisms involved in the generation of this large carbon reservoir are poorly understood. Here, we propose the microbial carbon pump as a conceptual framework to address this important, multifaceted biogeochemical problem.

The biogeochemistry of the river and shelf ecosystem of the Arctic Ocean: a review

The Arctic Ocean is, on a volume basis, the ocean with the highest terrestrial input in terms of freshwater and organic matter. The drainage areas of the Arctic contain more than half of the organic carbon stored globally in soils and are extremely sensitive to climate change. These changes may considerably influence the huge continental flux of water and organic and inorganic constituents to the Arctic Ocean. Because of the immediate global concerns we here review the current knowledge about the biogeochemistry of the Arctic river and shelf ecosystem. Organic matter concentrations in the Arctic rivers are among the highest reported in worlds rivers. Dissolved organic carbon (DOC) reaches concentrations of up to 1000 µM C. The total amount of DOC discharged by rivers into the Arctic Ocean is 18 to 26·1012g C·yr-1 and similar to that of the Amazon. The discharge of particulate organic carbon is much lower with 4 to 6·1012g C·yr-1. Nitrogen and phosphorus are principally discharged as organic compounds. The concentrations of inorganic nutrients are among the lowest worldwide (inorganic nitrogen: 0 to 20 µM; phosphate: 0 to 0.8 µM), with the exception of silicate in some rivers (0.5 to 110 µM).Freshly produced organic matter is labile and its turnover rates are high in the Arctic Ocean. Riverine organic matter, in contrast, is soil-derived and refractory. It seems to behave biogeochemically stable in the estuaries and shelves and therefore does not substantially support the productivity of the Arctic Ocean. Suspended organic matter from the rivers principally settles in the estuaries and on the shelves, hence the terrigenous signature in the sediment decreases with distance from the coast. However, a fraction of terrigenous suspended matter escapes the shelves and is present in considerable amounts even in sediments of the central Arctic Ocean. Terrigenous dissolved organic matter, on the other hand, behaves primarily conservatively in the Arctic Ocean. There are practically no removal mechanisms in the estuaries and shelves. The molecular composition of dissolved organic matter can largely be explained as a mixture of refractory marine and terrigenous compounds. Therefore, the Arctic river discharge plays an important role as a contemporary sink in the global carbon cycle. The few available data on the biogeochemistry of the Russian rivers indicate that the proportion of taiga and tundra in the drainage areas has no considerable influence on the concentration and chemical composition of dissolved organic matter, with the exception of lignin-derived phenols, which can be used as chemotaxonomic tracers. It can therefore be speculated that changes in vegetation due to climate warming may not considerably influence the composition of dissolved organic matter discharged to the Arctic Ocean. The discharge of inorganic nutrients, however, may already has increased in the last decades, as indicated by long-term increases in winter water discharge and the seasonality of nutrient concentrations. For a reliable assessment of future changes long-term and seasonal data of nutrient and organic matter discharge, as well as more detailed biogeochemical information is urgently needed.

Biogeochemical characteristics of dissolved and particulate organic matter in Russian rivers entering the Arctic Ocean

Abstract The biogeochemical signature of riverine matter in the Russian Arctic was investigated to establish a background for tracing terrestrial organic material in the Arctic Ocean. Elemental and lignin compositions of particulate and dissolved organic matter (POM, DOM), stable carbon isotope ratios of POM and nutrient concentrations are reported for 12 Russian rivers along 4000 km of coastline. The 12 rivers account for about 43% of the freshwater supply to the Arctic Ocean. Nine rivers drain both tundra and taiga areas and three rivers only tundra. Concentrations of nitrogenous nutrients and phosphate were low, whereas silicate values were generally high with only few exceptions. The concentrations of particulate organic carbon (POC) varied between 25.5 and 291 μmol/L C, contributing 0.4–2.1% to the total suspended sediment (TSS). Dissolved organic carbon (DOC) ranging from 230 to 1006 μmol/L C was on average eight times higher than POC. The concentrations of particulate and dissolved organic nitrogen were similar (ca. 11 μmol/L N) resulting in four times higher C/N ratios in the dissolved fraction (48) compared to the particulate fraction (11). The δ13C ratios were uniform (−25.6 to −27.4‰) and similar in taiga and tundra draining rivers. The exclusively terrestrial component lignin, determined as lignin phenols after cupric oxide oxidation, ranged from 5.6 to 37.6 nmol/L in the particulate fraction and from 34 to 319 nmol/L in the dissolved fraction. The syringyl/vanillyl (S/V) and cinnamyl/vanillyl (C/V) ratios of the particulate and dissolved lignin phenols were significantly correlated with the proportion of tundra and taiga in the drainage areas. This is true despite different formation processes and diagenetic degree of POM and DOM, as evident from acid/aldehyde ratios of vanillyl phenols [(Ad/Al)v]. Export rates were calculated from the carbon and lignin data. The 12 rivers studied transport about 10 × 1012 g of total organic carbon per year into the Arctic Ocean. The largest proportion is discharged by the Yenisey River with about 5 × 1012 g/yr. Total lignin discharge was about 42 × 109 g/yr. The contribution of lignin to the overall carbon flux (0.26% of carbon export) was almost negligible, which does not limit its quality as a quantitative tracer for organic matter fluxes from tundra and taiga regions.

Diet-induced changes in the fatty acid composition of Arctic herbivorous copepods: Experimental evidence of trophic markers

To verify the potential of fatty acids as trophic markers, feeding experiments were carried out with the dominant herbivorous copepods Calanus finmarchicus, C. hyperboreus and C. glacialis from the Greenland Sea during two Arctic expeditions in June/July 1991. Depending on the fatty acid composition of these copepods, the diatom Thalassiosira antarctica or the dinoflagellate Amphidinium carterae were offered as food to induce deviating fatty acid compositions. Since the copepodite Stages CV of C. finmarchicus had very low amounts of the 16:1(n−7) fatty acid but high amounts of 18:4(n−3), the specimens were fed on T. antarctica rich in 16:1(n−7) over a period of up to 42 days. At the end of the feeding experiment the portion of the 16:l(n-7) fatty acid had strongly increased by 11%, while the 18:4(n−3) fatty acid was almost depleted. In contrast, high amounts of the 16:1(n−7) fatty acid in C. hyperboreus (CV) suggested feeding on diatoms, therefore its diet was changed to A. carterae dominated by high amounts of the 18:4(n−3) fatty acid. After 47 days the portion of 18:4(n−3) increased by 8%, whereas 16:1(n−7) decreased by 3 %. In female C. glacialis the changes in the fatty acid composition after feeding with A. carterae were less pronounced as compared to the other species, due to a severe lipid loss during the experiment. The feeding experiments document the incorporation and turnover of dietary fatty acids under controlled laboratory conditions and provide clear evidence for the potential of specific fatty acids as trophic marker lipids.

Lipids and life strategy of Arctic Calanus

Abstract The three Arctic Calanus species, C. finmarchicus (Gunnerus, 1765), C. glacialis (Jaschov, 1955), and C. hyperboreus, are the most important herbivores in Arctic seas in terms of species biomass. They play a key role in the lipid-based energy flux in the Arctic, converting low-energy carbohydrates and proteins in ice algae and phytoplankton into high-energy wax esters. In this paper we review the over-wintering strategy, seasonal migration, stage development, life span, feeding strategy, body size, lipid biochemistry and the geographic distribution of the three dominant Calanus species in Arctic waters. We then relate these parameters to other biotic and abiotic factors, such as the timing of the Arctic phytoplankton and ice algae bloom, sea ice cover and climate variability. We also present new data on fatty acid and fatty alcohol content in the three Calanus species in addition to reviewing the available literature on these topics. These data are analysed for species homogeneity and geographic grouping. The dominance of diatom fatty acid trophic markers in the lipids of Calanus underpins the importance of diatoms as Arctic primary producers, even if dinoflagellates and Phaeocystis pouchetii can also be important food sources for the calanoid copepods. We conclude that the Arctic Calanus species are herbivores, engineered to feed on the Arctic bloom, and that the timing of the bloom is the most important factor in determining the life strategies of the individual species.

Lipids in Arctic benthos: does the fatty acid and alcohol composition reflect feeding and trophic interactions?

Arctic benthic organisms of various taxa (Anthozoa, Polychaeta, Pantopoda, Crustacea, Echinodermata) were collected on the shelves off northeast Greenland, Spitsbergen and the western Barents Sea. Their fatty acid compositions were generally characterised by the predominance of the polyunsaturated fatty acids 20:5(n-3) and 22:6(n-3) together with the saturated fatty acid 16:0, which reflect the dominance of phospholipids. The fatty acid compositions of most benthic specimens were influenced by fatty acids of dietary origin. High amounts of the fatty acid 16:1(n-7), typical of diatoms, were found in different taxa from the northeast Greenland shelf. The 18:4(n-3) fatty acid, often typical of non-diatom input, was only dominant in Ophiopholis aculeata from the Spitsbergen shelf. In some taxa small amounts of wax esters were detected with alcohol moieties similar to those of the dominant Arctic copepods. The occurrence of intact wax esters, as well as the wax ester typical fatty acids 20:1(n-9) and 22:1(n-11), also suggested ingestion of large herbivorous copepods. An unusual fatty acid composition was found for most brittle stars, due to a ratio of the 18:1(n-9) and (n-7) fatty acid isomers below 1 with lowest ratios of 0.1. A similar low ratio was also detected in the polychaete Onuphis conchylega. The extremely low portions of the 18:1(n-9) fatty acid are striking, since carnivores are generally characterised by high levels of this fatty acid. A clear gradient from low 18:1(n-9) to (n-7) ratios in suspension feeders, via predatory decapods, to higher ratios in the scavenging amphipods was a major characteristic of the benthic species. Our investigations showed that lipid analyses can give important hints on trophic relationships of benthic species and may serve as means to establish the intensity of pelagic-benthic coupling.

Water masses and circulation in the Eurasian basin. Results from the Oden-91 expedition

The Oden 91 North Pole expedition obtained Oceanographic measurements on four sections in the Nansen and Amundsen basins of the Eurasian Basin and in the Makarov Basin of the Canadian Basin, thereby proving the feasibility of carrying out a typical Oceanographic program using an icebreaker in the Arctic Ocean. The data show greater spatial variability in water structure and circulation than was apparent from previous data. The results show that a clear front exists between the Eurasian and Canadian basins such that upper halocline water in the Canadian Basin is almost absent from the Eurasian Basin. The lower halocline water produced in the Barents-Kara Sea region permeates much of the Eurasian Basin and flows along the continental slope into the Canadian Basin. The deeper circulation is strongly influenced by topography. Three return flows of the Atlantic layer are identified, one over the Nansen-Gakkel Ridge, one over the Lomonosov Ridge, and a third flowing from the Canadian Basin. The slight differences observed in salinity and temperature characteristics of the deeper waters of the Nansen and Amundsen basins do not lead to an obvious explanation of their origin or flow pattern.

River or mangrove? Tracing major organic matter sources in tropical Brazilian coastal waters

The influence of mangrove-fringed tropical estuaries on coastal carbon budgets has been widely recognised. However, a quantitative differentiation between riverine and mangrove-derived inputs to the dissolved (DOM) and microparticulate organic matter (POM) pool of these environments has been hitherto not possible. Based on lignin-derived phenols and stable carbon isotopes a chemical signature for mangrove, terrestrial and marine-derived organic matter was established for a mangrove estuary in North Brazil. A mixing model was applied to calculate the contribution of each of the three sources to the DOM and POM pool in the estuary throughout 18 tidal cycles in the course of one year. Best source assignment for POM was reached with the yield of lignin phenols and d13C as paired indicators, while the origin of DOM was best identified by the yield of lignin phenols and the acid to aldehyde ratio of vanillyl phenols. Although only about 6 % of the fluvial catchment area is covered by mangroves, their contribution to the estuarine DOM and POM pool generally exceeded several times the terrigenic input from the hinterland. This outwelling of mangrove-derived organic matter was enhanced during the rainy season. DOM and POM were exported from the mangrove to the estuary in similar proportions. Most mangrove-POM was rapidly removed from the water column, while mangrove-DOM behaved conservatively. In contrast, terrestrial DOM was almost entirely removed in the outer part of the estuary, which was accompanied by a concomitant increase in terrestrial POM. This seems to be the result of a geochemical barrier zone for this type of DOM in the estuary. Generally a high proportion of mangrove-DOM was present in the outer part of the estuary, even at high tide. This indicates DOM outwelling from mangroves in adjacent bays or estuaries and points to similar driving forces controlling this process on a regional scale. Mangroves probably play a more important role than rivers for marine carbon budgets along the North Brazilian coast south of the Amazon estuary.

Herbivorous or omnivorous? On the significance of lipid compositions as trophic markers in Antarctic copepods

Three dominant Antarctic copepods, Calanoides acutus, Rhincalanus gigas and Metridia gerlachei (copepodite stages V and females), were collected during summer (January/ February) in the southern Weddell Sea south of 70°S. Detailed analyses of their lipid and fatty acid/ alcohol compositions were carried out. The trophic positions of these copepods were elucidated by means of the lipid compositions (“marker lipids”). High amounts of wax esters were found in C. acutus (92% of total lipids) and in R. gigas (84–86%). The level of wax esters in M. gerlachei was relatively low (27–42%), while the accumulation of triacylglycerols tended to be higher (19–22%). Characteristic lipid components of C. acutus were the long-chain monounsaturated fatty acids and fatty alcohols 20:1 (n-9) and 22:1 (n-11). These components together with elevated amounts of the 18:4 (n-3) and, to a lesser degree, of the 16:! (n-7) fatty acids, typical of phytoplankton lipids, indicate herbivorous feeding for C. acutus. Other abundant fatty acids were 20:5 (n-3) and 22:6 (n-3). The fatty acid composition of M. gerlachei was characterized by very high amounts of these 22:6 and 20:5 acids. Other important fatty acids were 18:1 (n-9) and 16:0, but only small amounts of 16:1 (n-7) and 18:4 (n-3) occurred. In contrast to C. acutus the fatty alcohols of M. gerlachei consisted almost exclusively of the short-chain components 14:0 and 16:0 M. gerlachei is known as an omnivorous species, which was clearly reflected by its lipid and fatty acid/alcohol pattern. Few data are available on the feeding of R. gigas, but it is usuaally described as an herbivorous small-particle feeder. R. gigas showed fatty acid/alcohol characteristics typical of either C. acutus or M. gerlachei. Higher amounts of the 16:1 (n-7) and 18:4 (n-3) fatty acids suggest herbivorous feeding, whereas the dominance of short-chain alcohols (14:0 and 16:0) resembled the lipid pattern found in the omnivorous M. gerlachei. Hence, the lipid composition of R. gigas showed an intermediate pattern, which implies a tendency towards an opportunistic feeding mode, positioned somewhere between the other two species.