High diatom production and export in stratified waters – A potential negative feedback to global warming

It is widely held that increased stratification and reduced vertical mixing in the ocean driven by global warming will promote the replacement of diatoms by smaller phytoplankton and lead to an overall decrease in productivity and carbon export. Here we present contrary evidence from a synergy of modern observations and palaeo-records that reveal high diatom production and export from stratified waters. Diatom adaptations to stratified waters include the ability to grow in low light conditions in deep chlorophyll maxima; vertical migrations between nutricline depths and the surface, and symbioses with N2-fixing cyanobacteria in diatom–diazotroph associations (DDA). These strategies foster the maintenance of seed populations that may then exploit mixing events induced by storms or eddies, but may also inherently promote blooms. Recent oceanographic observations in the subtropical gyres, at increasingly high temporal and spatial resolutions, have monitored short-lived but often substantial blooms and export of stratified-adapted diatoms including rhizosolenids and the diazotroph-associated Hemiaulus hauckii. Aggregate formation by such diatoms is common and promotes rapid settling thereby minimizing water column remineralization and optimizing carbon flux. Convergence zones associated with oceanic fronts or mesoscale features may also generate substantial flux of stratified-adapted diatom species. Conventional oceanographic observing strategies and sampling techniques under-represent such activity due to the lack of adequate capability to sample the large sized diatoms and colonies involved, the subsurface location of many of these blooms, their common development in thin <3 m layers and their episodic nature. Superbly preserved “palaeo-sediment trap” records in laminated sediments reveal enhanced production and export from stratified waters mediated by the same taxa. The organic carbon-rich Mediterranean sapropels were the result of production within deep chlorophyll maxima in highly stratified waters dominated by rhizosolenid diatoms but also including H. hauckii. In the Cretaceous when CO2 levels were higher than present, laminated sediments reveal dominant flux also including Hemiaulus and rhizosolenid species. Nitrogen isotope and biomarker studies suggest that, analogous to modern DDA blooms within the subtropical gyres, Hemiaulus blooms in the ancient Mediterranean and within the Cretaceous seas were aided by nitrogen-fixing cyanobacterial symbionts. These lines of evidence suggest that diatom production and associated export of organic carbon, may not decrease, as is widely predicted, but may actually increase with greater ocean stratification, and potentially act as a negative feedback to global warming. However, the key genera involved in such potential feedbacks are underrepresented in both laboratory and field studies and are poorly represented in models. Our findings suggest that a reappraisal is necessary of the way diatoms are represented as plankton functional types (PFTs) in ocean biogeochemical models and that new observing and sampling strategies are also required to study these processes