Physical drivers of benthic and pelagic algal biomass dynamics in lakes: a conceptual exploration with 'Lake2D'

Abstract

Size, depth and basin shape are important factors controlling the physics, chemistry and, ultimately, the productivity of lakes. To our knowledge, a comprehensive theoretical investigation of the physical determinants of lake primary production from a conceptual, process-based perspective has not been performed. To address this knowledge gap, we developed and analyzed Lake2D, a process-based, reaction-advection-diffusion model that adopts a 2-dimensional modeling approach by reducing lake bathymetry to its hypsographic depth distribution under the simplifying assumption of radial symmetry of the lake basin. In simpler terms, the model assumes the lake basin is perfectly circular and uses a cross-section to represent its depth and shape. We used Lake2D to explore and analyze the dynamics of pelagic and benthic algae on a whole-lake scale in response to six environmental drivers (lake area and depth, horizontal and vertical turbulent mixing, water transparency and nutrient status), which we varied over ranges that are representative of the vast majority of the worlds lakes. Numerical analyses reveal three distinct patterns across environmental parameter space. (1) Benthic algae dominate total biomass in shallow, clear lakes with low nutrient content. In these lakes, low horizontal and vertical mixing is beneficial to benthic but detrimental to pelagic algae, which experience high sinking losses, thus liberating nutrients and minimizing shading of benthic algae. (2) Increasing mean lake depth, abiotic turbidity and/or nutrient content strongly benefits pelagic algae, which increasingly shade out benthic algae and dominate total biomass. In these lakes, increased mixing affects both algal types positively due to increased nutrient transport to shallow, well-lit areas. (3) Finally, at high lake mean depth and/or abiotic turbidity, a large fraction of the lake volume is aphotic. In such lakes, high horizontal and vertical mixing is detrimental to pelagic algae but beneficial to benthic algae, because pelagic algae are mixed to aphotic depths, liberating nutrients and minimizing shading. Moving from shallow, clear lakes via deeper and/or nutrient-rich lakes to very deep and/or turbid lakes, benthic and pelagic algae therefore show opposite gradual biomass shifts in response to vertical and horizontal mixing. Specifically, benthic biomass is highest at low and lowest at high overall mixing in shallow, nutrient-poor lakes, but shows the opposite trend in all other lakes. Conversely, pelagic biomass is highest at low and lowest at high overall mixing in very deep and/or turbid lakes, but shows the opposite trend in all other lakes.