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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.  相似文献   
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The equatorial Pacific is an HNLC (High-Nitrate Low-Chlorophyll) region. Modeling and in-situ process studies have confirmed the importance of microzooplankton grazing in this ecosystem. Unfortunately, both the parameters and functions representing microzooplankton grazing within current ecosystem models are poorly constrained. We used a simple 4-component food web model to test the assumption that a lower grazing threshold, which is common in many models, is necessary to achieve the HNLC condition. Without the grazing threshold, the model did not reproduce the HNLC condition. However, by raising the half-saturation constant within the microzooplankton functional response with no threshold, it was possible to reproduce the critical dynamics of the HNLC condition under both steady and moderate seasonal variability in nutrient input. It was also possible to reproduce the HNLC system using a sigmoidal functional response for the microzooplankton, with results somewhere between the other two forms of the model, although this version had the highest sensitivity to changes in its parameters. The three models predicted similar phytoplankton biomass and primary productivity under steady nutrient input, but diverge in these metrics as the amplitude of nutrient input variability increases. These three functional responses also imply certain important differences in the microzooplankton community. Whereas the threshold model had the least sensitivity to parameter choice, the high half-saturation constant, no-threshold model may actually be a better approximation when modeling a community of grazers. Ecosystem models that predict carbon production and export in HNLC regions can be very sensitive to assumptions concerning microzooplankton grazing; future studies need to concentrate on the functional responses of microzooplankton before these models can be used for predicting fluxes in times or regions where forcing is beyond that used to constrain the original model.  相似文献   
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