Phytoplankton patchiness: the role of lateral stirring and mixing

Explanations for the patchy distribution of marine phytoplankton are critically reviewed with the focus on the role played by lateral advection and mixing. Generating mechanisms for what is increasingly seen as a ubiquitous feature of the oceans have been sought in biology, behaviour, population dynamics, physics and various combinations thereof. As the mesoscale and sub-mesoscale (1-500 km) contain the "weather" of the ocean--eddies and fronts capable of strong stirring and mixing of any tracers advected by them--the physical circulation in this regime may have a strong role in the generation of patchiness. As a consequence, the focus here is on the ways the turbulent currents at these scales can interact with other processes to produce the complex spatial structures in phytoplankton distributions seen in countless satellite images and cruise data. The mechanisms reviewed include patch formation (KiSS theory), filamentation, Turing and related instabilities, diffusion waves, shear effects and turbulent stirring. Parallel to this is a discussion of observations and of the techniques that have been used to analyse them, particularly spectral analysis. It will be seen that many of the limitations in extant theories stem from a paucity of data and ambiguity in its interpretation. A synthesis of recent developments in sampling, instrumentation, image analysis and turbulence theory is used to suggest alternatives to conventional approaches, to highlight important work in related fields and to motivate new lines of research.