Water column contributions to the metabolism and nutrient dynamics of mid-sized rivers

Lotic and lentic ecosystems are traditionally viewed as dominated by either benthic or water column processes. However, mid-sized rivers represent a transition zone where both benthic and water column processes may both contribute substantially to ecosystem dynamics. Ecosystem processes such as gross primary production (GPP), ecosystem respiration (ER), or nutrient uptake, and the relative contribution of the water column to these processes at the reach scale, are poorly understood in non-wadeable, mid-sized rivers. To clarify the role of the water column at the reach-scale, and to quantify controls on water column processes, we measured GPP, ER, and uptake of nitrate (NO3-), ammonium (NH4+), and soluble reactive phosphorus (SRP) in the water columns of 15 mid-sized rivers (discharge: 13.5-83.3 m(3) s(-1)) spanning nutrient and total suspended solids gradients. We compared water column metabolic and nutrient uptake rates to reach-scale rates to estimate the contribution of the water column to the entire river. Water column metabolism was autotrophic on the day when measured, GPP increased with nutrient availability, and the water column contributed more to whole river GPP than to ER. Water column nutrient uptake increased with GPP across solutes, and there was a positive relationship between human land use and water column uptake of NO3--N and SRP. The water column accounted for a substantial proportion of reach-scale metabolism and nutrient uptake, but this contribution depended on suspended material and nutrient availability. Integrating the water column into theory describing lotic ecosystem function should clarify mechanisms controlling metabolism and nutrient processing and enhance management of non-wadeable rivers.