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.     

Mussels document loss of bioavailable polycyclic aromatic hydrocarbons and the return to baseline conditions for oiled shorelines in Prince William Sound, Alaska

Polycyclic aromatic hydrocarbons (PAH) were measured in mussels (Mytilus trossulus) collected between 1990 and 2002 from 11 sites on the shores of Prince William Sound (PWS), Alaska, that were heavily oiled by the 1989 Exxon Valdez oil spill (EVOS). This study, utilizing the methods of the NOAA Status and Trends Mussel Watch Program, found that concentrations of PAH released from spill remnants have decreased dramatically with time and by 2002 were at or near the range of total PAH (TPAH) of 3-355 ng/g dry weight obtained for mussels from unoiled reference sites in PWS. Time-series TPAH data indicate a mean TPAH half-life in mussel tissues of 2.4 years with a range from 1.4 to 5.3, yielding an annual mean loss of bioaccumulated TPAH of 25%. The petroleum-derived TPAH fraction in mussel tissues has decreased with time, reflecting the decreasing release of EVOS residues in shoreline sediments. These results show that PAH from EVOS residues that remain buried in shoreline sediments after the early 1990s are in a form and at locations that have a low accessibility to mussels living in the intertidal zone.     

A permanent seismic station beneath the Ocean Bottom

The Hawaii Institute of Geophysics began development of the Ocean Subbottom Seisometer (OSS) system in 1978, and OSS systems were installed in four locations between 1979 and 1982. The OSS system is a permanent, deep ocean borehole seismic recording system composed of a borehole sensor package (tool), an electromechanical cable, recorder package, and recovery system. Installed near the bottom of a borehole (drilled by the D/V Glomar Challenger), the tool contains three orthogonal, 4.5-Hz geophones, two orthogonal tilt meters; and a temperature sensor. Signals from these sensors are multiplexed, digitized (with a floating point technique), and telemetered through approximately 10 km of electromechanical cable to a recorder package located near the ocean bottom. Electrical power for the tool is supplied from the recorder package. The digital seismic signals are demultiplexed, converted back to analog form, processed through an automatic gain control (AGC) circuit, and recorded along with a time code on magnetic tape cassettes in the recorder package. Data may be recorded continuously for up to two months in the self-contained recorder package. Data may also be recorded in real time (digital formal) during the installation and subsequent recorder package servicing. The recorder package is connected to a submerged recovery buoy by a length of bouyant polypropylene rope. The anchor on the recovery buoy is released by activating either of the acoustical command releases. The polypropylene rope may also be seized with a grappling hook to effect recovery. The recorder package may be repeatedly serviced as long as the tool remains functionalA wide range of data has been recovered from the OSS system. Recovered analog records include signals from natural seismic sources such as earthquakes (teleseismic and local), man-made seismic sources such as refraction seismic shooting (explosives and air cannons), and nuclear tests. Lengthy continuous recording has permitted analysis of wideband noise levels, and the slowly varying parameters, temperature and tilt.Hawaii Institute of Geophysics Contribution 1909.     

A summary of the geology and oil habitat of the Eastern Flank Hydrocarbon Province of South Oman

The Eastern Flank Hydrocarbon Province borders the southeastern edge of the South Oman Salt Basin in the southern part of Oman. An intensive exploration programme by PDO over the past seven years has led to the discovery of almost 2 × 10⁹ m³ of oil with current production of some 15000 m³/d from six fields.In stark contrast to other oil habitats of the Arabian Peninsula, the main play involves rock units and geological events ranging in age from Late Precambrian to Quaternary Times, while trap development and charging are closely related to syn-tectonic salt dissolution.The principal reservoirs and seals are poorly consolidated, Palaeozoic clastics which drape, due to salt dissolution, over residual ‘cores’ principally composed of either Lower Palaeozoic sandstones (Haima Group), or Late Proterozoic carbonates (Huqf Group), or some combination of these. Oils have been geochemically correlated with algal source rocks of the Huqf Eastern Flank. Structures are considered to have been ‘charged’ from Late Mesozoic times onwards in conjunction with progressive subsurface salt removal and the onset of maturation of local source rocks. The effectiveness of ‘charging’ along the retreating salt edge is indicated by the small percentage of dry wells.Major oil zones are found in both anticlinal and truncation traps which are commonly developed around sandstone (Haima Group) cores. Oils show a wide range of characteristics but are typically heavy, early expulsion crudes, high sulphur with little associated gas. Bacterial transformation is not uncommon.Producing reservoirs mainly occur at relatively shallow depths (600–1200 m). Primary recovery factors of around 7% reflect the high density and viscosity of the Eastern Flank crudes and have initiated interest in Enhanced Oil Recovery (EOR) techniques. Two such pilot projects, involving steam and polymer flooding, are due to commence in 1984 and, if successful, will substantially increase the reserve base of these fields.     

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.     

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.     

Chalk shore platform erosion in the vicinity of sea defence structures and the impact of construction methods

Erosion of the intertidal chalk platform in the vicinity of groynes and seawalls is evident to the naked eye along many stretches of the engineered coastline of southeast England, leading to undermining and eventually failure of these structures. However, quantification of the magnitude and spatial extent of the erosion has been difficult to date because of a lack of data about the past elevation of the platform. The application of softcopy photogrammetry makes it possible to recreate past platform elevations from historic air photographs and to compare these with elevations from modern air photographs. Coastal sea defence structures have been installed along the chalk coast east of Brighton at various dates over the past 70 years. During this period, the construction methods have changed from predominantly manual labour to a reliance on heavy machines. The analysis of erosion patterns around structures built since the 1970s using heavy machinery show that surface lowering is 4 to 25 times greater in the vicinity of these structures than across the platform as a whole. In contrast, there is no similar pattern of increased erosion around structures built using predominantly manual labour in the 1930s. A four fold increase in average surface lowering is found also along a vehicle trackway that crosses the mid platform. Depressions developed by enhanced lowering in the front of seawalls generate their own dynamic of increased erosion by trapping pebbles and cobbles that enhance the abrasion of the chalk through bedload transport under standing waves in front of the walls.     

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.