Size-structured planktonic ecosystems: constraints, controls and assembly instructions

Here we present a nutrient-phytoplankton-zooplankton (NPZ) model that has arbitrary size-resolution within the phytoplankton- and zooplankton-state variables. The model assumes allometric scaling of biological parameters. This particular version of the model (herbivorous zooplankton only) has analytical solutions that allow efficient exploration of the effects of allometric dependencies of various biological processes on the model's equilibrium solutions. The model shows that there are constraints on the possible combinations of allometric scalings of the biological rates that will allow ecosystems to be structured as we observe (larger organisms added as the total biomass increases). The diversity (number of size classes occupied) of the ecosystem is the result of simultaneous bottom-up and top-down control: resources determine which classes can exist; predation determines which classes do exist. Thus, the simultaneous actions of bottom-up and top-down controls are essential for maintaining and structuring planktonic ecosystems. One important conclusion from this model is that there are multiple, independent ways of obtaining any given biomass spectrum, and that the spectral slope is not, in and of itself, very informative concerning the underlying dynamics. There is a clear need for improved size-resolved field measurements of biological rates; these will both elucidate biological processes in the field, and allow strong testing of size-structured models of planktonic ecosystems.