Characterization of novel pathways in the phosphorus cycle of lakes

Abstract

Phosphorus (P) is a limiting nutrient regulating productivity in both freshwater and marine ecosystems. A full knowledge of the sources and pathways of the P cycle is essential for understanding aquatic ecosystem function and for managing eutrophication. However, two significant pathways are poorly understood or remain uncharacterized. First, aquatic metazoans represent a significant internal regenerative pathway of P through the mineralization, translocation (i.e., benthic pelagic coupling) and excretion of nutrients. Rates of P excreted are expected to vary across taxa (i.e., zooplankton vs. mussels vs. benthic macroinvertebrates vs. fish), yet the significance of any one group of taxa in supplying P to bacteria and algae is unknown. Therefore, I developed the first comprehensive set of empirical models of nutrient release for aquatic metazoans (zooplankton, mussels, “other” benthic macroinvertebrates, and detritivorous and non-detritivorous fish) and compared inter-taxonomic differences in P excretion. I demonstrated that detritivorous fish excrete P at rates greater than all other taxa (as a function of individual organism mass); whereas, mussels generally excreted P at rates less than other taxa. Significant differences in the rate of P excretion between zooplankton and non-detritivorous fish were not observed [i.e., the allometry of P excretion was similar between zooplankton and non-detritivorous fish (as a function of individual body mass)]. I subsequently applied the models to assemblage biomass and abundance data to examine and compare the relative contribution of each taxa to the internal supply of P, and to examine the turnover time of P bound in metazoan biomass. I clearly demonstrated a hierarchy in the contribution by different metazoan assemblages to P cycling (zooplankton > benthic macroinvertebrates > mussels > fish) and clarified the significance of different metazoan taxa in P cycling. Moreover, I demonstrated that the slow turnover time of P bound in fish biomass (relative to other metazoans) indicates that fish are important as sinks rather than sources of P. A second potentially significant P pathway is through the influence of ultraviolet radiation (UVR) on P cycling. UVR may alter P cycling abiotically through changes in P availability and biotically through changes in the acquisition and regeneration of dissolved P by plankton. However, the significance of P released from the photodecomposition of dissolved organic P compounds (DOP), and the effect of UVR on the uptake and regeneration of dissolved P, the turnover of particulate P, and on ambient phosphate (PO43-) concentration has not been investigated and remains unknown. Therefore, my initial experiments applied the novel use of radiophosphate uptake assays to quantify the significance of the photodecomposition of DOP to PO43-. I concluded that the liberation of PO43- through the photodecomposition of DOP is not a significant pathway. However, the photochemical liberation of PO43- from suspended sediments was evident and should be an important pathway supplying PO43- to plankton in shallow polymictic lakes. This represents the first study to identify this P pathway in lakes. The turnover time of the PO43- pool increased under UVR irradiance (i.e., uptake of P by plankton decreased), while the regeneration rate of dissolved P and turnover rate of planktonic P were generally not affected. The net effect of UVR was an increase in steady state PO43- concentration (ssPO43-). Alkaline phosphatase activity (APA) in the dissolved and particulate fractions was significantly reduced in UVR treatments, but unrelated to changes in P uptake as proposed in the literature. This is the first study to comprehensively investigate the biotic effects of UVR on P cycling and represents a major advancement in the field of photobiology. In summary, I have characterized several poorly understood pathways in the P cycle of lakes. With the models I have developed, aquatic metazoans can now be integrated into the P cycle of lakes, for example, with other internal and external sources of P (e.g., from inlets, lake sediments and the atmosphere). This will advance our knowledge of P cycling, and will provide researchers with a better understanding of the nutrient pathways supporting primary production.