A seasonal study of extracellular enzyme activities (EEA) was conducted in the coastal, sandy aquifer system located in the greater fluvial-littoral ecosystem of Doñana (SW, Spain). Glucosidase, leucine aminopeptidase, alkaline phosphatase, and phenol oxidase activities were determined over a 2-year period in 30 piezometers spread in an area of approximately 100 km(2). Taking into account all enzymes, piezometers, and seasons, EEA ranged over several orders of magnitude, from 1.01 × 10(-5) ± 2.92 × 10(-6) to 1.37 ± 0.13 nmol (methylumbelliferyl, amido-4-methylcoumarin, or dihydroxyphenylalanine) mL(-1) h(-1). The quality, much more than the quantity, of organic matter and nutrients seemed to be the major variables that controlled the spatiotemporal patterns showed by EEA. EEA patterns obtained in this study agree with several functional models of microbial communities, such as optimal resource allocation and nutrient co-limitation. This study probably represents the first one in which these functional models have been tested in subsurface systems. Results obtained in this study seem to suggest that microbial communities inhabiting groundwaters in Doñana are not dead or compromised cells. By contrast, these communities play relevant roles in carbon and nutrient cycling, continue with the decomposition process that begins in the sediments of the shallow lakes located in the area, provide remineralized carbon and nutrients to producers of these surface aquatic systems, and close energy and matter cycles. This study proposes that groundwater systems should be considered dynamic systems, comparable in functional complexity to surface systems.