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Progress in Oceanography 130 (2015) 205–248 Contents lists available at ScienceDirect Progress in Oceanography journal homepage: www.elsevier.com/locate/pocean Review Zooplankton fecal pellets, marine snow, phytodetritus and the ocean’s biological pump ⇑ Jefferson T. Turner Biology Department, University of Massachusetts Dartmouth, 285 Old Westport Road, North Dartmouth, MA 02747, USA School for Marine Science and Technology, University of Massachusetts Dartmouth, 706 South Rodney French Boulevard, New Bedford, MA 02744, USA article info abstract Article history: The ‘‘biological pump’’ is the process by which photosynthetically-produced organic matter in the ocean Received 17 August 2013 descends from the surface layer to depth by a combination of sinking particles, advection or vertical mix- Received in revised form 7 August 2014 ing of dissolved organic matter, and transport by animals. Particulate organic matter that is exported Accepted 8 August 2014 downward from the euphotic zone is composed of combinations of fecal pellets from zooplankton and Available online 27 August 2014 fish, organic aggregates known as ‘‘marine snow’’ and phytodetritus from sinking phytoplankton. Previ- ous 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 tech- niques 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) zoo- plankton 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, con- sumption and fragmentation of marine snow by animals, pathogens associated with marine snow; (4) phytodetritus, including pulsed export of phytodetritus, phytodetritus from Phaeocystis spp., picoplank- ton 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. Ó 2014 Elsevier Ltd. All rights reserved. Contents Introduction. ...................................................................................................... 206 The ocean’s biological pump ......................................................................................... 206 Recent studies of the export flux of particulate organic carbon (POC) . ...................................................... 208 Dissolved organic carbon (DOC). ......................................................................... 212 Zooplankton fecal pellets . ................................................................................... 213 Contribution of zooplankton fecal pellets to export flux . ...................................................... 213 Epipelagic retention of zooplankton fecal pellets due to zooplankton activities . ................................... 213 ⇑ Addresses: Biology Department, University of Massachusetts Dartmouth, 285 Old Westport Road, North Dartmouth, MA 02747, USA, School for Marine Science and Technology, University of Massachusetts Dartmouth, 706 South Rodney French Boulevard, New Bedford, MA 02744, USA. E-mail address: [email protected] http://dx.doi.org/10.1016/j.pocean.2014.08.005 0079-6611/Ó 2014 Elsevier Ltd. All rights reserved. 206 J.T. Turner / Progress in Oceanography 130 (2015) 205–248 Zooplankton vertical migration and fecal pellet repackaging. ............................................................ 215 Microbial ecology of fecal pellets. ............................................................................ 217 Sinking velocity of fecal pellets and aggregates, and ballasting by mineral contents. ......................................... 218 Phytoplankton cysts, intact cells and harmful algae toxins in fecal pellets and sediment traps . ...................... 219 Importance of fecal pellets from various types of zooplankton . ............................................................ 219 Zooplankton fecal pellets and picoplankton export . ............................................................ 220 Marine snow . ......................................................................................................... 221 Origins, abundance, and distributions of marine snow . ............................................................ 221 Particles and organisms associated with marine snow. ............................................................ 221 Consumption and fragmentation of marine snow by animals . ............................................................ 221 Pathogens associated with marine snow. ............................................................................ 221 Phytodetritus . ......................................................................................................... 222 Pulsed export of phytodetritus. ............................................................................ 222 Aggregation and disaggregation of phytodetritus and other particles. ......................................... 222 Transparent exopolymer particles (TEP) . ............................................................................ 222 Phytodetritus from Phaeocystis spp. ............................................................................ 223 Picoplankton in phytodetritus . ............................................................................ 224 The summer export pulse (SEP) of phytodetritus in the subtropical north pacific. ......................................... 224 Benthic community responses to phytodetritus . ............................................................ 224 Other components of the biological pump . ...................................................................... 225 Fish fecal pellets and fish-mediated export. ............................................................................ 225 Sinking carcasses of animals and macrophytes. ............................................................ 225 Feces from marine mammals . ............................................................................ 226 The biological pump and climate. ......................................................................................... 226 Origins of the biological pump. ............................................................................ 226 The biological pump and glacial/interglacial cycles . ............................................................ 227 The biological pump and contemporary climate variations . ............................................................ 228 The biological pump and anthropogenic climate change . ............................................................ 229 Conclusions . ......................................................................................................... 232 Acknowledgements . ......................................................................................... 232 References . ......................................................................................................... 232 Introduction polar waters (Buesseler, 1998). Usually only <3% of net primary production reaches bathypelagic depths (>1000 m) in the deep The ocean’s biological pump sea (De La Rocha and Passow, 2007, and references therein). Such variability in export to depth is partly due to variability in the The ‘‘biological pump’’ is the process through which photosyn- planktonic food webs in overlying epipelagic waters (Legendre thetically-produced organic matter in the ocean is exported from and Rivkin, 2002). The >97% of net primary production that does the surface layer to depth by a combination of sinking particles, not reach the deep sea is consumed or respired in the water advection or vertical mixing of dissolved organic matter, and trans- column, with approximately 15% being processed by bacteria port by animals. (Ducklow, 2000), 30–70% by microzooplankton 20–200 lm in size Together with the ‘‘solubility pump’’ (Volk and Hoffert, 1985), (Calbet and Landry, 2004) and 20–35% by the >200 lm mesozoo- whereby the solubility of CO2 increases with depth due to