Aquatic microbiology for ecosystem scientists: New and recycled paradigms in ecological microbiology

In all ecosystems, bacteria are the most numerous organisms and through them flows a large fraction of annual primary production. In the past decade we have learned a great deal about some of the factors that regulate bacteria and their activities, and how these activities, in turn, alter ecosystem-level processes. Here I review three areas in which recent progress has been made with particular reference to pelagic ecosystems: the problem of bacterial cell dormancy; the effect of solar radiation on organic matter lability; and, the maintenance of net heterotrophy. In a system in which grazing is the major source of mortality for bacteria, bacterial cell dormancy is something of a paradox. I argue that the degree to which bacteria are grazed by flagellates (highly selective grazers) versus other grazers (cladocerans, bivalves) may explain the degree and variation in the proportion of active cells which recent evidence shows to be large. Another factor affecting bacterial activity that has come to the fore in recent years is solar radiation. Irradiation, especially in the ultra-violet range has long been thought of as simply deleterious to some bacteria. A wealth of newer evidence shows that refractory dissolved organic compounds may be converted into microbially labile compounds by solar radiation in several wavebands. This interaction between irradiation and organic matter (photolysis) may explain, in part, how dissolved organic carbon (C) may be refractory in the dark environment of the soil but become labile in the illumunated surface waters of lakes or rivers. The newer evidence shows that aquatic ecosystems, at least oligotrophic ones, are significantly subsidized by terrestrially-produced organic matter. I review here multiple lines of evidence that suggest that freshwater ecosystems are predominantly systems which respire more organic C than they produce by photosynthesis, and are therefore net heterotrophic. While net heterotrophy is an interesting exception for terrestrial ecosystems, it appears to be commonplace for aquatic systems and represents an important linkage between terrestrial and aquatic ecosystems.