Many natural streams are net heterotrophic, so I propose that trophic state be divided into autotrophic and heterotrophic state. This division allows consideration of the influence of external carbon sources as well as nutrients such as nitrogen and phosphorus. Empirical results suggest that phosphorus and nitrogen are the most important nutrients regulating autotrophic state in flowing waters and that benthic algal biomass is positively correlated to gross primary production in streams. Reference (minimally influenced by human activities) nutrient concentrations and correlations of nutrients with algal biomass are used to characterize reference distributions of stream autotrophic state. Only when reference nutrient concentrations are in the upper one third of those expected in the United States, is maximum benthic chlorophyll projected to exceed 100 mg m ÏȘ2 (a concentration commonly used to indicate nuisance levels) ÏŸ30% of the time. Average reference nutrient concentrations lead to sestonic chlorophyll concentrations above those considered typical of eutrophic lakes (ÏŸ8 mg m ÏȘ3 ) less than half the time. Preliminary analysis suggests that autotrophic state is variable in small pristine streams because it is influenced by canopy cover (light), but heterotrophic state is less variable because it can be based on allochthonous or autochthonous production. Nitrogen and phosphorus enrichment can influence both heterotrophic and autotrophic state, and these effects could cascade to animal communities. Stoichiometry should be considered because carbon, nitrogen, and phosphorus are all involved in trophic state. The proposed definition of trophic state offers a starting conceptual framework for such considerations.
The evolution of concepts regarding enrichment in streamsIn its course from the source to the sea, the progressive eutrophication of a river water by drainage from cultivated and inhabited districts is an almost inevitable natural process.-Butcher 1947Although current concerns about stream eutrophication mainly focus on nitrogen (N) and phosphorus (P) enrichment (e.g., Smith 2003), early water quality and nutrient enrichment studies in lotic systems focused on carbon (C) enrichment from untreated sewage. Excessive loading of biochemical oxygen demand (BOD) made rivers completely anoxic downstream of sewage treatment plants. Hynes (1960) considered the physical, biological, and chemical effects of sewage loading to create a general conceptual model on the basis of the research of Butcher (1946) and others. The conceptual model of Hynes in part considered the influence of increased organic C on dissolved oxygen (O 2 ) and subsequently on hypoxia-and anoxia-sensitive animals. He noted that most animals immediately downstream from a sewage outfall disappear under anoxic conditions and that, as O 2 enters the stream via aeration, high densities of pollution-tolerant fauna 1 Corresponding author (wkdodds@ksu.edu).