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.     

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.     

The seasonal cycle of arsenic in estuarine and nearshore waters of the South Atlantic Bight

Arsenic concentrations vary with season on the continental shelf of the South Atlantic Bight. During periods of high winds in the winter and early spring, inorganic arsenic concentrations are reduced to as little as 20% of typical open ocean concentrations by sorption onto resuspended sediments or incorporation into phytoplankton. In the early fall, arsenic sequestered in sediments in the spring is regenerated and returned to the water column, creating elevated arsenic concentrations in the nearshore zone that are up to 50% greater than open ocean concentrations. Arsenic in the adjacent estuaries is nearly conservative over the seasonal cycle, although its distribution in the estuaries is greatly affected by the seasonal changes in arsenic concentrations in the nearshore waters.     

On the sulphur chemistry of a super-anoxic fjord, Framvaren, South Norway

The concentrations of total carbonate (C_t), sulphate, sulphide, thiols and oxygen, the ratio between the stable sulphur isotopes ³⁴S and ³²S in sulphate and sulphide, and the density (used to calculate salinity) were determined on samples from the water column of Framvaren, a superanoxic fjord in southern Norway. From a depth of 18m (the oxic-anoxic boundary) the initial sulphate concentration, ([SO₄]_init), as calculated from salinity, is significantly higher than the sum of the measured sulphur species. This is attributed to a loss of sulphur from the water column. The amount of total carbonate produced, corrected for the initial concentration (C_t - 2.4 Sal/35) is found to be proportional to the amount of sulphate consumed, ([SO₄]_init - [SO₄]), according to the following relation C_t- 2.4 Sal/35 = 1.84 ([SO₄]_init - [SO₄]). Isotopic fractionation caused by bacterial sulphate reduction in the anoxic part of the water column produces sulphide with a δ³⁴S 40‰ lower than the δ³⁴S for sulphate at corresponding depths. The isotopic fractionation also results in δ³⁴S value for the remaining sulphate at depths below 80 m being considerably higher than the mean value for ocean water, which is close to + 20‰. The δ³⁴S values for sulphate at depths between 10 and 50 m were lower than + 20‰ which indicates oxidation of sulphide, which follows upon diffusion of sulphide from deeper parts of the water column and inflow of oxygenated seawater over the sill into the anoxic water of the fjord. A conclusive scenario of the Framvaren sulphur chemistry is presented.     

A community-wide intercomparison exercise for the determination of dissolved iron in seawater

The first large-scale international intercomparison of analytical methods for the determination of dissolved iron in seawater was carried out between October 2000 and December 2002. The exercise was conducted as a rigorously “blind” comparison of 7 analytical techniques by 24 international laboratories. The comparison was based on a large volume (700 L), filtered surface seawater sample collected from the South Atlantic Ocean (the “IRONAGES” sample), which was acidified, mixed and bottled at sea. Two 1-L sample bottles were sent to each participant. Integrity and blindness were achieved by having the experiment designed and carried out by a small team, and overseen by an independent data manager. Storage, homogeneity and time-series stability experiments conducted over 2.5 years showed that inter-bottle variability of the IRONAGES sample was good (< 7%), although there was a decrease in iron concentration in the bottles over time (0.8–0.5 nM) before a stable value was observed. This raises questions over the suitability of sample acidification and storage.     

Environmental overview of Eckernförde Bay,northern Germany

This environmental overview of Eckernförde Bay (northern Germany) summarizes the results of previous studies relevant to the Office of Naval Research's Coastal Benthic Boundary Layer (CBBL) Baltic field exercise conducted during 1993–1994. Significant environmental characteristics include the following: (1) surface sediment distribution is related to water depth, dictated primarily by hydrodynamic reworking of older glacial deposits; (2) the origin and characteristics of small-scale sedimentary structures depend on storm-generated waves and currents; (3) the proximity of the sea surface and sediment —water interface results in a pelagic—benthic coupling that drives biogeochemical processes and produces organicrich, acoustically turbid sediments; and (4) the bay floor is complicated topographically by pockmarks and manmade sedimentary structures.