ALBERT

Beschreibung: Deep‐water nitrate is a major reservoir of oceanic combined nitrogen and has long been considered to be the major source of new nitrogen supporting primary production in the oligotrophic ocean. 15N:14N ratios in plankton provide an integrative record of the nitrogen cycle processes at work in the ocean, and near‐surface organic matter in oligotrophic waters like the Sargasso Sea is characterized by an unusually low 15N content relative to average deep‐water nitrate. Herein we show that the low dΔ15N of suspended particles and zooplankton from the tropical North Atlantic cannot arise through isotopic fractionation associated with nutrient uptake and food web processes but are instead consistent with a significant input of new nitrogen to the upper water column by N2 fixation. These results provide direct, integrative evidence that N2 fixation makes a major contribution to the nitrogen budget of the oligotrophic North Atlantic Ocean.

Beschreibung: In many small aquatic ecosystems, watershed loading of organic C exceeds autochthonous primary production. Although this allochthonous organic C has long been thought of as refractory, multiple lines of evidence indicate that substantial portions are respired in the receiving aquatic ecosystem. To what extent does this terrestrial C support secondary production of invertebrates and fish? Do current models adequately trace the pathways of allochthonous and autochthonous C through the food web? We evaluated the roles of allochthonous and autochthonous organic C by manipulating 13C content of dissolved inorganic C in a small, softwater, humic lake, thereby labeling autochthonous primary production for about 20 d. To ensure rapid and sufficient uptake of inorganic 13C, we enriched the lake with modest amounts of N and P. We constructed a carbon flow model based on the ambient and manipulated levels of 13C in C compartments in the lake, along with information on key rate processes. Despite the short nature of this experiment, several results emerged. (1) Fractionation of photosynthetically assimilated 13C‐CO2 by phytoplankton (ɛ) is lower (~6‰) than physiologic models would estimate (~20‰). (2) Bacteria respire, but do not assimilate, a large amount of terrestrially derived dissolved organic C (DOC) and pass little of this C to higher trophic levels. (3) The oxidation of terrestrial DOC is the major source of dissolved inorganic C in the lake. (4) Zooplankton production, a major food of young‐of‐year fishes, is predominantly derived from current autochthonous carbon sources under the conditions of this experiment.

Beschreibung: Lake Tanganyika, East Africa, has a simple pelagic food chain, and trophic relationships have been established previously from gut‐content analysis. Instead of expected isotopic enrichment from phytoplankton to upper level consumers, there was a depletion of 15N in August 1999. The isotope signatures of the lower trophic levels were an indicator of a recent upwelling event, identified by wind speed and nitrate concentration data, that occurred over a 4‐d period several days prior to sampling. The isotope structure of the food web suggests that upwelled nitrate is a nutrient source rapidly consumed by phytoplankton, but the distinctive signature of this nitrate is diluted by time averaging in the upper trophic levels. This time averaging is a consequence of the fact that the isotopic signature of an organism is related to variable nitrogen sources used throughout the life of the organism. This study illustrates the importance of recognizing differences in time averaging among trophic levels.

Beschreibung: Gross and net O2 production between May 1996 and February 1999 was determined in bottle incubation experiments with H218O spike and from the change in O2 concentration. Carbon fixation rates were obtained from 14C incubations. In general, production rates determined using the H218O‐spike were about twice the primary production determined by the 14C method, where the latter was close to net oxygen evolution. These relationships are similar to results for the open ocean. During the spring bloom, when the dinoflagellate Peridinium was abundant, the ratio of gross O2 production to carbon fixation was about 7.5, and net O2 production was greater than carbon fixation. The difference between O2 gross production and carbon fixation results, at least in part, from uptake by Mehler reaction and from recycling of the 14C tracer by dark respiration and the alternative oxidase (AOX). We used the difference in isotopic discrimination against 18O, occurring during O2 consumption by various biological pathways, to place constraints on the relative engagement of these pathways. We estimated the overall discrimination against 18O in the lake from O2 isotopic mass balance as 20.5—29‰. The only mechanism that can explain the strong overall fractionation in the lake is AOX, which strongly discriminates against 18O (~31‰). Our results show, for the first time, that uptake by AOX is widespread and quantitatively important to oxygen consumption in aquatic systems.

Beschreibung: High‐latitude rivers supply the Arctic Ocean with a disproportionately large share of global riverine discharge and terrigenous dissolved organic matter (DOM). We used the abundance of lignin, a macromolecule unique to vascular plants, and stable carbon isotope ratios (δ13C) to trace the high molecular weight fraction of terrigenous DOM in major water masses of the Arctic Ocean. Lignin oxidation products in ultrafiltered DOM (UDOM; 〉1,000 Da) from Arctic rivers were depleted in syringyl relative to vanillyl phenols (S/V = 0.3–0.5) compared to UDOM in temperate and tropical rivers (S/V = 0.5–1.2), indicating that gymnosperm vegetation is a major source of terrigenous UDOM to the Arctic Ocean. High concentrations of lignin oxidation products (83–320 ng L−1) and a depletion of 13C (δ13C = −23.0 to −21.9) in UDOM throughout the surface Arctic Ocean indicate that terrigenous UDOM accounts for a much greater fraction of the UDOM in the surface Arctic (5–33%) than in the Pacific and Atlantic oceans (0.7–2.4%). In contrast, UDOM in deep water from the Arctic Ocean as well as waters from throughout the Greenland Gyre had relatively low concentrations of lignin oxidation products (24–45 ng L−1 ) and was enriched in 13C (δ13C = −21.0 to −20.8). Terrigenous UDOM has a relatively short residence (∼1–6 yr) in surface polar waters prior to export to the north Atlantic Ocean. Assuming that the bulk of Arctic‐derived DOM is compositionally similar to the UDOM fraction, we estimate that 12–41% of terrigenous DOM (2.9–10.3 Tg C yr−1 ) discharged by rivers to the Arctic Ocean is exported to the North Atlantic via surface waters of the East Greenland Current. It appears very little terrigenous DOM from the Arctic is incorporated into North Atlantic Deep Water and distributed globally via deep thermohaline circulation.

Beschreibung: Estimates of carbon sources, as determined from ratios of stable isotopes, were used in conjunction with estimates of secondary production to determine the relative contribution of algal carbon to macroinvertebrate production across a gradient of elevation in a Rocky Mountain stream (North St. Vrain Creek, Colorado). The relative contribution of algal carbon to macroinvertebrate production was then compared to the relative availability of algal carbon. Although algal production accounted for less than 2–40% of the combined sources of organic matter to North St. Vrain Creek, the relative contribution of algal carbon to annual macroinvertebrate production ranged from approximately 40% at a subalpine site to nearly 80% at a more open site in the foothills. Thus, the proportional contribution of algal carbon to consumer production greatly exceeded the relative availability of algal carbon in North St. Vrain Creek. Despite the disproportionate importance of algal carbon to consumers, most macroinvertebrates in North St. Vrain Creek used some vascular plant carbon.

Beschreibung: We report measurements of seasonal variability in the C‐N stable isotope ratios of plants collected across the habitat mosaic of San Francisco Bay, its marshes, and its tributary river system. Analyses of 868 plant samples were binned into 10 groups (e.g., terrestrial riparian, freshwater phytoplankton, salt marsh) to determine whether C‐N isotopes can be used as biomarkers for tracing the origins of organic matter in this river‐marsh‐estuary complex. Variability of δ13C and δ15N was high (~5–10‰) within each plant group, and we identified three modes of variability: (1) between species and their microhabitats, (2) over annual cycles of plant growth and senescence, and (3) between living and decomposing biomass. These modes of within‐group variability obscure any source specific isotopic signatures, confounding the application of C‐N isotopes for identifying the origins of organic matter. A second confounding factor was large dissimilarity between the δ13C‐δ15N of primary producers and the organic matter pools in the seston and sediments. Both confounding factors impede the application of C‐N isotopes to reveal the food supply to primary consumers in ecosystems supporting diverse autotrophs and where the isotopic composition of organic matter has been transformed and become distinct from that of its parent plant sources. Our results support the advice of others: variability of C‐N stable isotopes within all organic‐matter pools is high and must be considered in applications of these isotopes to trace trophic linkages from primary producers to primary consumers. Isotope‐based approaches are perhaps most powerful when used to complement other tools, such as molecular biomarkers, bioassays, direct measures of production, and compilations of organic‐matter budgets.

Beschreibung: We used variation in algal d13C between river habitats to study the spatial scale of energy flow through river food webs. We found a strong negative relationship between herbivore d13C (which reflects algal d13C) and water velocity in three productive Northern California rivers but not in unproductive streams. The contrast among habitats suggests that water velocity affects algal d13C most strongly when CO2 availability is low relative to photosynthetic rates. Our results help explain the wide variation in published river biota d13C and show that past studies using carbon isotope analyses may have significantly underestimated the importance of algal‐derived carbon to river food webs. While flow‐related variation in d13C complicates this common application of carbon isotope analysis, we show that it provides a natural tracer of the flux of algal production derived from different habitats within rivers to higher trophic levels. Measurements of consumer d13C showed that most invertebrate and vertebrate consumers relied on local production, except for filter‐feeding insects and steelhead trout, which relied on production derived from multiple sources. Stable carbon isotopes may thus be used to spatially delineate the habitats that support river food webs, providing previously unavailable information for understanding and managing river ecosystems.

Beschreibung: Marine diatoms generally form large blooms during periods of cool temperature (〈20°C), high NO3− fluxes (〉25 μM‐N), and turbulent mixing, but the adaptations that allow diatoms to bloom under these conditions are not well understood. We have conducted both NO3− uptake kinetics and direct short‐term temperature manipulation studies on field diatom‐dominated populations from Chesapeake and Delaware Bays during both spring and fall blooms. Absolute rates of NO3− uptake by a Rhizoseleni‐dominated population did not appear to saturate even at concentrations as high as 180 µM‐N. We observed contrasting patterns of NO2−, NH+, and urea utilization as a function of experimental temperature (ambient ± 9°C). Over the temperature range of 7–25°C, absolute uptake rates of NO3− (ρNO3−) decreased an average of 46% with increasing temperature from 7 to 25°C (nine individual experiments), while ρNH4+ and ρUREA increased with increasing temperature by an average of 179 and 86% (eight individual experiments), respectively. Based on these observations and the nature of the physical environment, we hypothesize that these diatom‐dominated populations were taking up NO3− in excess of nutritional requirements, the reduction of which may serve as a sink for electrons during transient periods of imbalance between light energy harvesting and utilization. We suggest that the increase in non‐nutritional NO3− uptake increases proportionately with the magnitude of the imbalance between light energy harvesting and imbalance. This hypothesis reconciles previous observations of low C:N uptake ratios, high release rates of dissolved organic nitrogen or NO2− by diatom‐dominated assemblages, other observations of nonsaturating NO3− kinetics in field populations, and the apparent "preference" for NO3− by the netplankton size fraction. The two phenomena described here, nonsaturable kinetics and a negative relationship between NO3− uptake and short‐term temperature shifts, have important ecological implications. The hypothesized ability of these diatom‐dominated populations to better modulate the flow of photosynthetic electron energy, via NO3− reduction, in variable environments may provide a competitive advantage to diatoms and could potentially explain why diatoms frequently dominate in regions of cool temperature, high NO3− flux, and turbulent mixing. Also, models of new production may need to incorporate terms for temperature dependence of NO3− uptake. Finally, if a significant fraction of NO3− uptake is regulated by non‐nutritional mechanisms in the cell, and if some fraction of nitrogen reduced by this mechanism is subsequently released in the form of NO2−, NH+, or dissolved organic nitrogen (DON), then estimates of new production based solely on NO3− uptake could be seriously biased.