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.     

Geoacoustic and physical properties of carbonate sediments of the Lower Florida Keys

 Near-surface sediment geoacoustic and physical properties were measured from a variety of unconsolidated carbonate sediments in the Lower Florida Keys. Surficial values of compressional and shear speed correlate with sediment physical properties and near-surface acoustic reflectivity. Highest speeds (shear 125–150 m s^-1; compressional 1670–1725 m s^-1) are from sandy sediments near Rebecca Shoal and lowest speeds (shear 40–65 m s^-1; compressional 1520–1570 m s^-1) are found in soft, silty sediments which collect in sediment ponds in the Southeast Channel of the Dry Tortugas. High compressional wave attenuation is attributed to scattering of acoustic waves from heterogeneity caused by accumulation of abundant shell material and other impedance discontinuities rather than high intrinsic attenuation. Compared to siliciclastic sediments, carbonate sediment shear wave speed is high for comparable values of sediment physical properties. Sediment fabric, rather than changes due to the effects of biogeochemical processes, is responsible for these differences.     

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.     

JGOFS studies in the equatorial Pacific

This special issue is the third and final volume containing results from the JGOFS Process Study in the equatorial Pacific. Most of the Study in the equational Pacific. Most of the contributions evolved either from the US JGOFS workshop in 1994 on the equatorial Pacific in Scottsdale, AZ or from the NATO Advanced Research Workshop on the Carbon Cycle of the Equatorial Pacific in 1995 in Noumea, New Caledonia.     

Feeding of benthic foraminifera on diatoms and sewage-derived organic matter: an experimental application of lipid biomarker techniques

Foraminiferal ecology at sewage outfalls has been investigated in numerous field studies over the last 30 years. Foraminifera have been frequently used as biomonitors of sewage pollution since they are both abundant and ubiquitous. Sewage outfalls have been demonstrated to have both positive and negative effects on adjacent foraminiferal populations, but it has never been shown conclusively why sewage affects foraminifera in these ways. Such information on the impact mechanisms of sewage pollution is essential if foraminifera are to be used as sewage pollution biomonitors, and also to understand the ecology of these important protists. One possible cause of a positive effect is the direct consumption of sewage-derived particulate organic matter (POM) by the foraminifera themselves. However this hypothesis has never been tested experimentally. Here, lipid (fatty acid and sterol) biomarker techniques were applied to study the ingestion of two potential food items by the foraminiferan Haynesina germanica in the laboratory. An experiment was conducted to confirm that the laboratory conditions were conducive to the survival and feeding of the foraminifera. In this experiment, foraminifera were provided with the pennate diatom Phaeodactylum tricornutum, which was considered to be a suitable food source. After 2 weeks, a four-fold increase in the levels of the diatom fatty acid biomarker, 20:5(n-3), in the foraminifera suggested that they had fed actively on the diatoms and survived under the experimental conditions. These experimental conditions were used in the main experiment, where foraminifera were fed the POM from sewage. Lipid biomarker analysis indicated that H. germanica did not consume secondary treated sewage-derived POM. Neither fatty acid profiles in the sewage nor coprostanol, the diagnostic human faecal sterol, were detected in foraminifera after exposure to the potential sewage food source. However, foraminifera may have consumed bacteria associated with the sewage in the experiment. The findings are discussed in terms of current EU legislation on sewage treatment that has affected the composition of sewage discharges, and therefore possibly reduced the nutritive value of sewage to the marine benthos.     

Multi-scale variability of beach profiles at Duck: A wavelet analysis

Beach profiles have been observed to change over a range of spatial and temporal scales; however techniques for quantifying this variability have not been fully established. In this paper, a wavelet technique is introduced as a method to study the multi-scale variability of beach profiles. The beach profile data comprising a 22-year time series surveyed at the US Army Corps of Civil Engineers Field Research Facility (FRF) at Duck are analysed using the adapted maximal overlap discrete wavelet transform (AMODWT). The analysis successfully identifies strong local features in the variability of beach profiles in time and space separately that cannot be isolated by traditional statistical methods. The analysis of spatial wavelet variances provides a new means of investigating the depth of closure. Analysis of variances by temporal scales shows that the combined effects of several temporal scales with one or two dominant scales can be seen at particular points across profiles whilst the dominant temporal scales are different at different portions of the profiles. The method allows for the extremely nonstationary behaviour of beach profile to be analysed into separate frequency bands that can facilitate the interpretation of morphological changes in terms of physical processes.     

Uptake of plutonium from seawater and sediment by a marine polychaete worm

The results of the uptake of plutonium 239 from sediment by a marine polychaete wormNereis diversicolor is reported in the present paper. A comparison is made of the relative importance of two possible uptake pathways, sediment and seawater by which this worm may obtain its plutonium body-burden. Under the laboratory conditions used for the comparison of those two routes it would appear thatNereis obtains greater than 98% of its body activity from seawater.