Nutrient Limitation and Physiology Mediate the Fine-Scale (De)coupling of Biogeochemical Cycles

Nutrients in the environment are coupled over broad timescales (days to seasons) when organisms add or withdraw multiple nutrients simultaneously and in ratios that are roughly constant. But at finer timescales (seconds to days), nutrients become decoupled if physiological traits such as nutrient storage limits, circadian rhythms, or enzyme kinetics cause one nutrient to be processed faster than another. To explore the interactions among these coupling and decoupling mechanisms, we introduce a model in which organisms process resources via uptake, excretion, growth, respiration, and mortality according to adjustable trait parameters. The model predicts that uptake can couple the input of one nutrient to the export of another in a ratio reflecting biological demand stoichiometry, but coupling occurs only when the input nutrient is limiting. Temporal nutrient coupling may, therefore, be a useful indicator of ecosystem limitation status. Fine-scale patterns of nutrient coupling are further modulated by, and potentially diagnostic of, physiological traits governing growth, uptake, and internal nutrient storage. Together, limitation status and physiological traits create a complex and informative relationship between nutrient inputs and exports. Understanding the mechanisms behind that relationship could enrich interpretations of fine-scale time-series data such as those now emerging from in situ solute sensors.