title: Subsurface chlorophyll maximum layers: enduring enigma or mystery solved?
abstract: The phenomenon of subsurface chlorophyll maximum layers (SCMLs) is not a unique ecological response to environmental conditions; rather, a broad range of interacting processes can contribute to the formation of persistent layers of elevated chlorophyll a concentration (Chl) that are nearly ubiquitous in stratified surface waters. Mechanisms that contribute to the formation and maintenance of the SCMLs include a local maximum in phytoplankton growth rate near the nutricline, photoacclimation of pigment content that leads to elevated Chl relative to phytoplankton biomass at depth, and a range of physiologically influenced swimming behaviors in motile phytoplankton and buoyancy control in diatoms and cyanobacteria that can lead to aggregations of phytoplankton in layers, subject to grazing and physical control. A postulated typical stable water structure characterizes consistent patterns in vertical profiles of Chl, phytoplankton biomass, nutrients, and light across a trophic gradient structured by the vertical flux of nutrients and characterized by the average daily irradiance at the nutricline. Hypothetical predictions can be tested using a nascent biogeochemical global ocean observing system. Partial results to date are generally consistent with predictions based on current knowledge, which has strong roots in research from the twentieth century.
authors.affiliation: Department of Oceanography, Dalhousie University, Halifax B3H 4R2, Canada; email: john.cullen@dal.ca.
authors.forename: John J
authors.initials: JJ
authors.lastname: Cullen
mesh_terms: D000064:Acclimatization / Q000502:physiology; D018533:Biomass; D002734:Chlorophyll / Q000032:analysis*; D004784:Environmental Monitoring / Q000379:methods* / Q000706:statistics & numerical data; D008027:Light; D008962:Models, Theoretical; D009792:Oceans and Seas; D010839:Phytoplankton / Q000254:growth & development* / Q000378:metabolism / Q000502:physiology; D011156:Population Density; D012623:Seawater / Q000737:chemistry* / Q000382:microbiology; D013696:Temperature
chemical_list: D002734:Chlorophyll
keywords: acclimation; behavior; carbon-to-chlorophyll ratio; deep chlorophyll maximum; growth; phytoplankton; primary production; stratification; thin layers; turbulence