Macrophyte Complexity Controls Nutrient Uptake in Lowland Streams

Macrophytes act as ecosystem engineers in lowland stream ecosystems, enhancing habitat complexity and physical structure. Studies have demonstrated that macrophyte abundance and growth form can dictate the degree to which physical and biological stream characteristics are altered. However, few studies have investigated the influence of macrophytes and their speciesspecific variation in morphological complexity on functional processes, such as nutrient uptake. We injected ¹⁵N-labeled ammonium (¹⁵N-NH₄⁺) into four macrophyte-rich lowland streams in Denmark to quantify the uptake of NH₄⁺ by macrophytes, epiphytic biofilms, benthic biofilms, and suspended particulate organic matter in the water column. Overall, macrophytes and their epiphytic biofilms accounted for 71-98% of the reach-weighted uptake across the study streams. While macrophytes had the highest rates of NH₄⁺ uptake among the compartments we measured, the epiphytic biofilms had the highest uptake efficiency, ranging from 0.06 to 0.6 mg N mg $N_{biomass}^{ - 1}{d^{ - 1}}$. Among all compartments, the uptake efficiency was inversely related to the carbon-to-nitrogen ratio. Macrophyte complexity, expressed as leaf perimeter-to-area ratio (P:A), varied among the five species found in the study streams. The uptake rates by macrophyte species with high leaf P:A were, on average, an order of magnitude higher than the rates for species with simple leaf morphology (430 vs. 49 mg N m⁻²d⁻¹). In summary, our results indicate that macrophytes regulate stream function both via direct uptake of NH₄⁺ from the water column and by providing a substrate for epiphytic biofilms. Furthermore, the effect of leaf architecture on nutrient uptake rates provides evidence that physical complexity can enhance ecosystem function.