Why are diverse relationships observed between phytoplankton biomass and transport time?

Transport time scales such as flushing time and residence time are often used to explain variability in phytoplankton biomass. In many cases, empirical data are consistent with a positive phytoplankton-transport time relationship (i.e., phytoplankton biomass increases as transport time increases). However, negative relationships, varying relationships, or no significant relationship may also be observed. We present a simple conceptual model, in both mathematical and graphical form, to help explain why phytoplankton may have a range of relationships with transport time, and we apply it to several real systems. The phytoplankton growth-loss balance determines whether phytoplankton biomass increases with, decreases with, or is insensitive to transport time. If algal growth is faster than loss (e.g., grazing, sedimentation), then phytoplankton biomass increases with increasing transport time. If loss is faster than growth, phytoplankton biomass decreases with increasing transport time. If growth and loss are approximately balanced, then phytoplankton biomass is relatively insensitive to transport time. In analyses of several systems, portions of an individual system, or time periods, apparent insensitivity of phytoplankton biomass to changes in transport time could arise due to the superposition of cases with different phytoplankton-transport time relationships. Thus, in order to understand or predict responses of phytoplankton biomass to changes in transport time, the relative rates of algal growth and loss must be known.