Zooplankton fecal pellets, marine snow, phytodetritus and the ocean’s biological pump

Abstract

The “biological pump” is the process by which photosynthetically-produced organic matter in the ocean descends from the surface layer to depth by a combination of sinking particles, advection or vertical mixing of dissolved organic matter, and transport by animals. Particulate organic matter that is exported downward from the euphotic zone is composed of combinations of fecal pellets from zooplankton and fish, organic aggregates known as “marine snow” and phytodetritus from sinking phytoplankton. Previous reviews by Turner and Ferrante (1979) and Turner (2002) focused on publications that appeared through late 2001. Since that time, studies of the biological pump have continued, and there have been >300 papers on vertical export flux using sediment traps, large-volume filtration systems and other techniques from throughout the global ocean. This review will focus primarily on recent studies that have appeared since 2001. Major topics covered in this review are (1) an overview of the biological pump, and its efficiency and variability, and the role of dissolved organic carbon in the biological pump; (2) zooplankton fecal pellets, including the contribution of zooplankton fecal pellets to export flux, epipelagic retention of zooplankton fecal pellets due to zooplankton activities, zooplankton vertical migration and fecal pellet repackaging, microbial ecology of fecal pellets, sinking velocities of fecal pellets and aggregates, ballasting of sinking particles by mineral contents, phytoplankton cysts, intact cells and harmful algae toxins in fecal pellets, importance of fecal pellets from various types of zooplankton, and the role of zooplankton fecal pellets in picoplankton export; (3) marine snow, including the origins, abundance, and distributions of marine snow, particles and organisms associated with marine snow, consumption and fragmentation of marine snow by animals, pathogens associated with marine snow; (4) phytodetritus, including pulsed export of phytodetritus, phytodetritus from Phaeocystis spp., picoplankton in phytodetritus, the summer export pulse (SEP) of phytodetritus in the subtropical North Pacific, benthic community responses to phytodetritus; (5) other components of the biological pump, including fish fecal pellets and fish-mediated export, sinking carcasses of animals and macrophytes, feces from marine mammals, transparent exopolymer particles (TEP); (6) the biological pump and climate, including origins of the biological pump, the biological pump and glacial/interglacial cycles, the biological pump and contemporary climate variations, and the biological pump and anthropogenic climate change. The review concludes with potential future modifications in the biological pump due to climate change.