Glucuronidation in the polar bear (Ursus maritimus)

Polar bears bioaccumulate lipophilic pollutants, including polychlorinated biphenyls (PCBs), into their bodies from their exclusive diet of marine organisms. Hydroxylated PCB metabolites (OH-PCBs) have been found in plasma, presumably due to CYP-dependent biotransformation of PCBs in liver. Little is known about the phase 2 metabolism of hydroxylated xenobiotics in polar bears. The objective of this study was to examine UDP-glucuronosyltransferase (UGT) activity with OH-PCBs and a hydroxylated polycyclic aromatic hydrocarbon, 3-hydroxy-benzo(a)pyrene (3-OH-BaP), in polar bear liver. Samples of frozen polar bear liver were used to prepare microsomes. UGT activity with 3-OH-BaP in Brij-treated microsomes, measured by a fluorescence assay, was readily measurable with protein concentrations in assay tubes of up to 10 g/ml, but dropped off very sharply at higher protein concentrations. The apparent Km for 3-OH-BaP was 1.71 +/- 0.04 microM, and Vmax 1.26 +/- 0.16 nmol/min/mg protein (mean +/- SD, n=3). UGT activities with a model tetrachloro-OH-PCB (4'-OH-CB72) and a model hexachloro-OH-PCB (4'-OH-CB159) were assayed with [14-C]-UDPGA and separation of the [14-C]-glucuronide by ion-pair extraction and thin-layer chromatography. [14-C]-glucuronide conjugates were readily formed by polar bear liver microsomes in the absence of added substrate, apparently from contaminants present in liver. This phenomenon was not observed using hepatic microsomes from laboratory-held catfish. Glucuronidation efficiency was much higher with 4'-OH-CB72 (Km 7.3 microM; Vmax 1.55 nmol/min/mg) than 4'-OH-CB159 (Km 16.1 microM; Vmax 0.46 nmol/min/mg). The identities of the aglycones present in polar bear liver are not known, but could include OH-PCBs or hydroxylated metabolites of other persistent organic pollutants. This study demonstrates that UGT with high activity for 3-OH-BaP and other substrates is present in polar bear liver.     

Functional responses for zooplankton feeding on multiple resources: a review of assumptions and biological dynamics

Modelers often need to quantify the rates at which zooplankton consume a variety of species, size classes and trophic types. Implicit in the equations used to describe the multiple resource functional response (i.e. how nutritional intake varies with resource densities) are assumptions that are not often stated, let alone tested. This is problematic because models are sensitive to the details of these formulations. Here, we enable modelers to make more informed decisions by providing them with a new framework for considering zooplankton feeding on multiple resources. We define a new classification of multiple resource responses that is based on preference, selection and switching, and we develop a set of mathematical diagnostics that elucidate model assumptions. We use these tools to evaluate the assumptions and biological dynamics inherent in published multiple resource responses. These models are shown to simulate different resource preferences, implied single resource responses, changes in intake with changing resource densities, nutritional benefits of generalism, and nutritional costs of selection. Certain formulations are further shown to exhibit anomalous dynamics such as negative switching and sub-optimal feeding. Such varied responses can have vastly different ecological consequences for both zooplankton and their resources; inappropriate choices may incorrectly quantify biologically-mediated fluxes and predict spurious dynamics. We discuss how our classes and diagnostics can help constrain parameters, interpret behaviors, and identify limitations to a formulation's applicability for both regional (e.g. High-Nitrate-Low-Chlorophyll regions comprising large areas of the Pacific) and large-scale applications (e.g. global biogeochemical or climate change models). Strategies for assessing uncertainty and for using the mathematics to guide future experimental investigations are also discussed.     

Trace element diagenesis in pyrite-rich sediments of the Achterwasser lagoon, SW Baltic Sea

The early diagenesis of trace elements (V, Cr, Co, Cu, Zn, As, Cd, Ba, U) in anoxic sediments of the Achterwasser, a shallow lagoon in the non-tidal Oder estuary in the Baltic Sea, was investigated in the context of pyrite formation. The dissolved major redox parameters show a two-tier distribution with transient signals in the occasionally re-suspended fluid mud layer (FM) and a permanently established diagenetic sequence in the sediment below. Intense microbial respiration leads to rapid depletion of O₂ within the uppermost mm of the FM. The reduction zones of Mn, Fe and sulfate overlap in the FM and in the permanently anoxic sediment section which appears to be a typical feature of estuarine sediments, under low-sulfate conditions. Degrees of pyritization (DOP) range from 50% in the FM to remarkably high values > 90% at 50 cm depth. Pyrite formation at the sediment surface is attributed to the reaction of Fe-monosulfides with intermediate sulfur species via the polysulfide pathway. By contrast, intense pyritization in the permanently anoxic sediment below is attributed to mineral growth via adsorption of aqueous Fe-sulfide complexes onto pyrite crystals which had originally formed in the surface layer.The studied trace elements show differential behavior patterns which are closely coupled to the diagenetic processes described above: (i) Zn, Cu and Cd are liberated from organic matter in the thin oxic layer of the sediment and diffuse both upwards across the sediment/water boundary and downwards to be trapped as monosulfides, (ii) V, Cr, Co and As are released during reductive dissolution of Mn- and Fe-oxyhydroxides, (iii) U removal from pore water occurs concomitantly to Fe reduction in the FM and is attributed to reduction of U(VI) to U(IV), (iv) the Ba distribution is controlled by reductive dissolution of authigenic barite in the sulfate reduction zone coupled with upward diffusion and re-precipitation. The incorporation of trace elements into pyrite is most intense for Co, Mn and As, intermediate for Cu and Cr and little to negligible for U, Zn, Cd, V and Ba. The observed trend is largely in agreement with previous studies and may be explained with differing rates for ligand exchange. Slow and fast ligand exchange and thus precipitation kinetics are also displayed by downcore increasing (Mn, Cr, Co and As) or constantly low (Zn, Cu, Cd) pore water concentrations. The downward increasing degrees of trace metal pyritization (DTMP) for Co, Cu, Zn and As are, in analogy to pyrite growth, assigned to adsorption of sulfide complexes or As oxyanions onto preexisting pyrite minerals.     

Source regions for recruitment of Calanus finmarchicus to Georges Bank: evidence from molecular population genetic analysis of mtDNA

Molecular population genetic analysis has provided evidence that the copepod, Calanus finmarchicus, of the Labrador Current, Gulf of St Lawrence, Scotian Shelf, Gulf of Maine, and Georges Bank constitute a single, interbreeding population. The DNA sequence of a 350 base pair portion of the mitochondrial large subunit (16S) ribosomal RNA (rRNA) gene was determined for a total of 72 individuals collected in 1992, and 110 individuals collected in 1993 from these regions. There was significant heterogeneity in haplotype frequencies among the samples collected in 1992, but this heterogeneity did not resolve into regional patterns. The only regional differences seen were between pooled samples of the western N. Atlantic and those of the Norwegian Sea. There were no significant differences in haplotype frequencies among the samples collected in 1993, and fewer haplotypes were observed in these samples. Intraspecific molecular variation was typical of other marine species: there were 29 haplotypes among the 182 individuals sequenced. The frequency distribution of the haplotypes was highly skewed: 128 individuals shared one haplotype and 19 individuals were unique. There were 24 variable sites among the 350 bases sequenced; estimated nucleotide diversity was 0.0042. The genetic character of C. finmarchicus populations in the western N. Atlantic was stable over time in that three of the haplotypes (including the most abundant) occurred in both 1992 and 1993. However, haplotype frequencies differed significantly between the two years. The lack of regional structure in the 1992 samples and the genetic homogeneity of samples collected in 1993 across the domain from the Labrador Current to the Gulf of St Lawrence to Georges Bank and the Gulf of Maine indicated that there is significant gene flow across this region. The persistent genetic pattern suggests that the Gulf of St Lawrence may be an important source region for recruitment of C. finmarchicus to Georges Bank. Determination of zooplankton dispersal patterns within high gene flow species will provide information that may not be determined by conventional oceanographic analyses.     

The Bengal Fan: Some preliminary results from ODP drilling

Three deep holes, with a maximum penetration of 960 m below sea floor, were drilled into the distal Bengal Fan just south of the equator during ODP Leg 116. The entire section recovered is dominated by sandy silt and mud turbidites derived from the Ganges Delta and from the continental margin of the western Bay of Bengal, interbedded with thin pelagic clays and with biogenic turbidites probably from a local sea mount source. The effects of Hi-Malayan uplift, sea level fluctuations, local tectonics, and fan channel/lobe processes have closely interacted to produce the observed sedimentary record of the past 17 million years since the early Miocene.Coauthors include: James R. Cochran, Lamont-Doherty Geological Observatory (Co-chief scientist); Dorrik A. V. Stow, Nottingham University, England (Co-chief scientist); Christian A. Auroux, Texas A & M University, USA (ODP staff scientist); Kazou Amano, Ibaraki University, Japan; Peter S. Balson, British Geological Survey, Keyworth, Notts, England; Jacques Boulegue, Pierre & Marie Curie University, Paris, France; Garrett W. Brass, University of Miami, Florida, USA; Jeffrey Corrigan, University of Texas, Austin, USA; Stefan Gartner, Texas A & M University, USA; Stuart A. Hall, University of Houston, Texas, USA; Silvia Iaccarino, University of Parma, Italy; Toshio Ishizuka, University of Tokyo, Japan; trena Kacmarska, Mount Allison University, New Brunswick, Canada; Heidemarie Kassens, Kiel University, West Germany; Gregory Leger, Dalhousie University, Halifax, Nova Scotia, Canada; Franca Proto Decima, University of Padova, Italy; C. V. Raman, Andhra University, Visakhapatnam, India; William W. Sager, Texas A & M University, USA; Kozo Takahashi, Woods Hole Oceanographic Institution, USA; Thomas Thompson, 580 Euclid Ave, Boulder, Colo, USA; Jean-Jacques Tiercelin, University of Bretagne, Brest, France; Mark Townsend, University of Nottingham, England; Andreas Wetzel, Tubingen University, West Germany; N. P. Wijayananda, National Aquatatic Resource Agency, Colombo, Sri Lanka; and Colin Williams, Lamont-Doherty Geological Observatory.     

Copepod grazing during spring blooms: Can Pseudocalanus newmani induce trophic cascades?

During late winter and spring of 2002 and 2003, 24 two- to three-day cruises were conducted to Dabob Bay, Washington State, USA, to examine the grazing, egg production, and hatching success rates of adult female Calanus pacificus and Pseudocalanus newmani. Here, we discuss the results of our grazing experiments for P. newmani. Each week, we conducted traditional microzooplankton dilution experiments and “copepod dilution” experiments, each from two different layers. Grazing was measured by changes in chlorophyll concentration and direct cell counts. Clearance rates on individual prey species, as calculated by cell counts, showed that Pseudocalanus are highly selective in their feeding, and may have much higher grazing rates on individual taxa than calculated from bulk chlorophyll disappearance. The grazing rates of the copepods, however, are typically an order of magnitude lower than the grazing rates of the microzooplankton community, or the growth rates of the phytoplankton. P. newmani ingested diatoms, but, at certain times fed preferentially on microzooplankton, such as ciliates, tintinnids, and larger dinoflagellates. Removal of the microzooplankton may have released the other phytoplankton species from grazing pressure, allowing those species’ abundance to increase, which was measured as an apparent “negative” grazing on those phytoplankton species. The net result of grazing on some phytoplankton species, while simultaneously releasing others from grazing pressure resulted in bulk chlorophyll-derived estimates of grazing which were essentially zero or slightly negative; thus bulk chlorophyll disappearance is a poor indicator of copepod grazing. Whether copepods can significantly release phytoplankton from the grazing pressure by microzooplankton in situ, thus causing a trophic cascade, remains to be verified, but is suggested by our study.     

An ecosystem model with iron limitation of primary production in the equatorial Pacific at 140°W

We construct a one-dimensional ecosystem model (nitrate, ammonium, phytoplankton, zooplnakton and detritus) with simple physics and biology in order to focus on the structural relations and intrinsic properties of the food web that characterizes the biological regime in the central equatorial Pacific at 140°W. When possible, data collected during the EgPac and other cruises were used to calibrate model parameters for two simulations that differ in the limiting nutrient, i.e. nitrogen or iron. Both simulations show annual results in good agreement with the data, but phytoplankton biomass and primary production show a more pronounced annual variability when iron is used as the limiting nutrient. This more realistically reproduces the variability of biological production and illustrates the greater coupling between vertical physical processes and biological production when the limiting nutrient is iron rather than nitrogen. The iron simulation also illustrates how iron supply controls primary production variability, how grazingbalances primary production and controls phytoplankton biomass, and how both iron supply and grazingcontrol primary production. These results suggest that it is not possible to capture primary production variability in the central equatorial Pacific with biological models using nitrogen as the limiting nutrient. Other indirect results of this modeling study were: (1) partitioning of export production between dissolved and particulate matter is almost equal, suggesting that the importance of DOC export may have been previously overestimated; (2) lateral export of live biomass has to be taken into account in order to balance the nitrogen budget on the equator at 140°W; and (3) preferential uptake of ammonium (i.e. nitrate uptake inhibition by ammonium) associated with high regeneration of nitrogen (low f ratio as a consequence of the food web structure imposed by iron limitation) largely accounts for the surface build-up of upwelled nitrate.