Skip to main content
SpringerLink
Log in

Biological Responses to Seasonally Varying Fluxes of Organic Matter to the Ocean Floor: A Review

  • Published:
Journal of Oceanography Aims and scope Submit manuscript

Abstract

Deep-sea benthic ecosystems are sustained largely by organic matter settling from the euphotic zone. These fluxes usually have a more or less well-defined seasonal component, often with two peaks, one in spring/early summer, the other later in the year. Long time-series datasets suggest that inter-annual variability in the intensity, timing and composition of flux maxima is normal. The settling material may form a deposit of “phytodetritus” on the deep-seafloor. These deposits, which are most common in temperate and high latitude regions, particularly the North Atlantic, evoke a response by the benthic biota. Much of our knowledge of these responses comes from a few time-series programmes, which suggest that the nature of the response varies in different oceanographic settings. In particular, there are contrasts between seasonal processes in oligotrophic, central oceanic areas and those along eutrophic continental margins. In the former, it is mainly “small organisms” (bacteria and protozoans) that respond to pulsed inputs. Initial responses are biochemical (e.g. secretion of bacterial exoenzymes) and any biomass increases are time lagged. Increased metabolic activity of small organisms probably leads to seasonal fluctuations in sediment community oxygen consumption, reported mainly in the North Pacific. Metazoan meiofauna are generally less responsive than protozoans (foraminifera), although seasonal increases in abundance and body size have been reported. Measurable population responses by macrofauna and megafauna are less common and confined largely to continental margins. In addition, seasonally synchronised reproduction and larval settlement occur in some larger animals, again mainly in continental margin settings. Although seasonal benthic responses to pulsed food inputs are apparently widespread on the ocean floor, they are not ubiquitous. Most deep-sea species are not seasonal breeders and there are probably large areas, particularly at abyssal depths, where biological process rates are fairly uniform over time. As with other aspects of deep-sea ecology, temporal processes cannot be encapsulated by a single paradigm. Further long time-series studies are needed to understand better the nature and extent of seasonality in deep-sea benthic ecosystems.

This is a preview of subscription content, log in via an institution to check access.

Access this article

Log in via an institution

Price excludes VAT (USA)
Tax calculation will be finalised during checkout.

Instant access to the full article PDF.

Institutional subscriptions

Similar content being viewed by others

References

  • Altenbach, A. V. (1992): Short term processes and patterns in the foraminiferal response to organic flux rates. Mar. Micropaleo., 19, 119–129.

    Google Scholar 

  • Altenbach, A. V., U. Pflaumann, R. Schiebel, A. Thies, S. Timm and M. Trauth (1999): Scaling percentages and distributional patterns of benthic foraminifera with flux rates of organic carbon. J. Foram. Res., 29, 173–185.

    Google Scholar 

  • Ambrose, W. G. and P. E. Renaud (1995): Benthic response to water column productivity patterns: Evidence for benthopelagic coupling in the Northeast Water Polynya. J. Geophys. Res., 100, 4411–4421.

    Google Scholar 

  • Ambrose, W. G. and P. E. Renaud (1997): Does a pulsed food supply to the benthos affect polychaete recruitment patterns in the Northwest Water Polynya? J. Mar. Sys., 10, 483–495.

    Google Scholar 

  • Arntz, W. E., J. Tarazona, V. Gallardo, L. A. Flores and H. Salzwedel (1991): Benthos communities in oxygen deficient shelf and upper slope areas of the Peruvian and Chilean Pacific coast and changes caused by El Niño. p. 131–154. In Modern and Ancient Continental Shelf Anoxia, ed. by R. V. Tyson and T. H. Pearson, Geological Society, Tulsa Oklahoma.

    Google Scholar 

  • Asper, V. L., W. G. Deuser, G. A. Knauer and S. E. Lohrenz (1992): Rapid coupling of sinking particles between surface and deep ocean waters. Nature, 357, 670–672.

    Google Scholar 

  • Attrill, M. J., R. G. Hartnoll and A. L. Rice (1991): Aspects of the biology of the deep-sea crab Eryon trispinosus from the Porcupine Seabight. J. Mar. Biol. Ass., U.K., 71, 311–328.

    Google Scholar 

  • Auffret, G., A. Khripounoff and A. Vangriesheim (1994): Rapid post-bloom resuspension in the northeast Atlantic. Deep-Sea Res. I, 41, 925–939.

    Google Scholar 

  • Baldwin, R. J., R. C. Glatts and K. L. Smith (1998): Particulate matter fluxes into the benthic boundary layer at a long time-series station in the abyssal NE Pacific: composition and fluxes. Deep-Sea Res. II, 45, 643–665.

    Google Scholar 

  • Barnes, D. K. A. and A. Clarke (1995): Seasonality of feeding activity in Antarctic suspension feeders. Polar Biol., 15, 335–340.

    Google Scholar 

  • Barnett, P., B. Hardy and J. Watson (1982): Meiofaunal studies in the Rockall Trough. Deep-Sea Newslett., 6, 5.

    Google Scholar 

  • Bathmann, U. V. (1988): Mass occurrence of Salpa fusiformis in the spring of 1984 off Ireland: implications for sedimentation processes. Mar. Biol., 97, 127–135.

    Google Scholar 

  • Beaulieu, S. E. and K. L. Smith (1998): Phytodetritus entering the benthic boundary layer and aggregated on the seafloor in the abyssal NE Pacific: macro-and microscopic observations. Deep-Sea Res. II, 45, 781–815.

    Google Scholar 

  • Berger, W. H. and G. Wefer (1990): Export production: seasonality and intermittency, and paleoceanographic implications. Palaeogeo., Palaeoclim., Palaeoecol., 89, 245–254.

    Google Scholar 

  • Billett, D. S. M. and A. L. Rice (2001): The BENGAL programme: introduction and overview. Prog. Oceanogr., 50, 13–25.

    Google Scholar 

  • Billett, D. S. M., R. S. Lampitt, A. L. Rice and R. F. C. Mantoura (1983): Seasonal sedimentation of phytoplankton to the deep-sea benthos. Nature, 302, 520–522.

    Google Scholar 

  • Bishop, J. D. D. and S. H. Shalla (1994): Discrete seasonal reproduction in an abyssal peracarid crustacean. Deep-Sea Res. I, 41, 1789–1800.

    Google Scholar 

  • Blake, J. A. and L. Watling (1994): Life history studies of deep-sea benthic infauna: Polychaeta, Aplacophora, and Cumacea from the continental slope off Massachusetts. p. 244–260. In Reproduction, Larval Biology, and Recruitment of the Deep-Sea Benthos, ed. by C. M. Young and K. J. Eckelbarger, Columbia University Press, New York.

    Google Scholar 

  • Bodungen, B. Von, V. S. Smetacek, M. M. Tilzer and B. Zeitschel (1986): Primary production and sedimentation during spring in the Antarctic Peninsula region. Deep-Sea Res., 33, 177–194.

    Google Scholar 

  • Bodungen, B. Von, A. Antia, E. Bauerfiend, O. Haupt, W. Koeve, I. Peeken, R. Peinert, S. Reitmeier, C. Thomsen, M. Voss, M. Wunsch, U. Zeller and B. Zeitzschel (1995): Pelagic processes and vertical flux of particles: an overview over long-term comparative study in the Norwegian and Greenland Sea. Geol. Rundsch., 84, 11–27.

    Google Scholar 

  • Boetius, A. and K. Lochte (1994): Regulation of microbial enzymatic degradation of organic matter in deep-sea sediments. Mar. Ecol. Prog. Ser., 104, 299–307.

    Google Scholar 

  • Boetius, A. and K. Lochte (1996): Effects of organic enrichment on hydrolytic potentials and growth of bacteria in deep-sea sediments. Mar. Ecol. Prog. Ser., 140, 239–250.

    Google Scholar 

  • Boyd, P. and P. Newton (1995): Evidence of the potential influence of planktonic community structure on the interannual variability of particulate organic carbon flux. Deep-Sea Res. I, 42, 619–639.

    Google Scholar 

  • Brandt, A., J. Svavarsson and T. Brattegard (1994): Eurycope brevirostris (Isopoda: Asellota) from the deep Arctic Ocean: redescription, postmarsupial development, and reproductive pattern. Sarsia, 79, 127–143.

    Google Scholar 

  • Bronsdon, S. K., P. A. Tyler, A. L. Rive and J. D. Gage (1993): Reproductive biology of two epizoic anenomes from the deep north-east Atlantic Ocean. J. Mar. Biol. Ass., U.K., 73, 531–542.

    Google Scholar 

  • Campbell, J. W. and T. Aarup (1992): New production in the North Atlantic derived from seasonal patterns of surface chlorophyll. Deep-Sea Res., 39, 1669–1694.

    Google Scholar 

  • Campos-Creasey, L. S., P. A. Tyler, J. D. Gage and A. W. G. John (1994): Evidence for coupling the vertical flux of phytodetritus to the diet and seasonal life history of the deep-sea echinoid Echinus affinis. Deep-Sea Res. I, 41, 369–388.

    Google Scholar 

  • Cartes, J. E. (1998): Dynamics of the bathyal Benthic Boundary Layer in the northwestern Mediterranean: depth and temporal variations in macrofaunal-megafaunal communities and their possible connections with deep-sea trophic webs. Prog. Oceanogr., 41, 111–139.

    Google Scholar 

  • Cartes, J. E. and J. C. Sorbe (1996): Temporal population structure of deep-water cumaceans from the western Mediterranean slope. Deep-Sea Res. I, 43, 1423–1438.

    Google Scholar 

  • Cartes, J. E., J. B. Company and F. Maynou (1994): Deep-water decapod crustacean communities in the Northwest Mediterranean: influence of submarine canyons and season. Mar. Biol., 120, 221–229.

    Google Scholar 

  • Cartes, J. E., M. Elizalde and J. C. Sorbe (2000): Contrasting life histories and secondary production of populations of Munnopsurus atlanticus (Isopoda: Asellota) from two bathyal areas of the NE Atlantic and the NW Mediterranean. Mar. Biol., 136, 881–890.

    Google Scholar 

  • Coma, R., M. Ribes, J.-M. Gili and M. Zabala (2000): Seasonality in coastal benthic ecosytems. Trends Ecol. Evol., 15, 448–453.

    Google Scholar 

  • Danovaro, R., D. Dinet, G. Duinveld and A. Tselepides (1999): Benthic responses to particulate fluxes in different trophic environments: a comparison between the Gulf of Lions-Catalan Sea (western Mediterranean) and the Cretan Sea (eastern Mediterranean). Prog. Oceanogr., 44, 287–312.

    Google Scholar 

  • Danovaro, R., N. Della Croce, A. Dell'Anno, M. Fabiano, D. Marrale and D. Martorano (2000a): Seasonal changes and biochemical composition of the labile organic matter flux in the Cretan Sea. Prog. Oceanogr., 46, 259–278.

    Google Scholar 

  • Danovaro, R., D. Marrale, A. Dell'Anno, N. Della Croce, A. Tselepides and M. Fabiano (2000b): Bacterial response to seasonal changes in labile organic matter composition on the continental shelf and bathyal sediments of the Cretan Sea. Prog. Oceanogr., 46, 345–366.

    Google Scholar 

  • Danovaro, R., A. Tselepides, A. Otegui and N. Della Croce (2000c): Dynamics of meiofaunal assemblages on the continental shelf and deep-sea sediments of the Cretan Sea (NE Mediterranean): relationships with seasonal changes in food supply. Prog. Oceanogr., 46, 367–400.

    Google Scholar 

  • de Bovée, F., L. D. Guidi and J. Soyer (1990): Quantitative distribution of deep-sea meiobenthos in the northwestern Mediterranean (Gulf of Lions). Cont. Shelf Res., 10, 1123–1145.

    Google Scholar 

  • de Wilde, P. A. W. J., G. C. A. Duineveld, E. M. Berghuis, M. S. S. Lavaleye and A. Kok (1998): Late-summer mass deposition of gelatinous phytodetritus along the slope of the N.W. European Continental Margin. Prog. Oceanogr., 42, 165–187.

    Google Scholar 

  • Deuser, W. G. (1986): Seasonal and interannual variations in deep-water particle fluxes in the Sargasso Sea and their relation to surface hydrography. Deep-Sea Res. I, 33, 225–246.

    Google Scholar 

  • Deuser, W. G. and E. H. Ross (1980): Seasonal change in the flux of organic carbon to the deep Sargasso Sea. Nature, 283, 364–365.

    Google Scholar 

  • Deuser, W. G., T. D. Jickells, P. King and J. A. Commeau (1995): Decadal and annual changes in biogenic opal and carbonate fluxes to the deep Sargasso Sea. Deep-Sea Res., 42, 1923–1932.

    Google Scholar 

  • Dickson, R. R., W. J. Gould, P. A. Gurbutt and P. D. Killworth (1982): A seasonal signal in ocean currents to abyssal depths. Nature, 295, 193–198.

    Google Scholar 

  • Dickson, R. R., W. J. Gould, C. Griffiths, K. J. Medeler and E. M. Gmitrowicz (1986): Seasonality in currents of the Rockall Trough. Proc. Roy. Soc. Edin., 88B, 103–125.

    Google Scholar 

  • Drazen, J. E., R. J. Baldwin and K. L. Smith (1998): Sediment community response to a temporally varying food supply at an abyssal station in the NE Pacific Deep-Sea Res. II, 45, 893–913.

    Google Scholar 

  • Druffel, E. R. M. and K. L. Smith (eds.) (1998): Long time series monitoring of an abyssal site in the NE Pacific. Deep-Sea Res. II, 45, 569–913.

  • Ducklow, H. W. and R. P. Harris (eds.) (1993): JGOFS: The North Atlantic Bloom Experiment. Deep-Sea Res. II, 40, 641 pp.

  • Duco, A. and M. Roux (1981): Modalities particulière de croissance liées au milieu abyssal chez les Bathycrinidae (Echinodermes, Crinoides pédonculés). Oceanol. Acta, 4, 389–393.

    Google Scholar 

  • Duineveld, G. C. A., M. S. S. Lavaleye, E. M. Berghuis, P. A. W. J. de Wilde, J. van der Weele, A. Kok, S. D. Batten and J. W. De Leeuw (1997): Patterns of benthic fauna and benthic respiration on the Celtic continental margin in relation to the distribution of phytodetritus. Internationale Revue der gesampten Hydrobiologie, 82, 395–424.

    Google Scholar 

  • Duineveld, G. C. A., A. Tselepides, R. Witbaard, R. P. M. Bak, E. M. Berghuis, G. Nieuwland, J. van der Weele and A. Kok (2000): Benthopelagic coupling in the oligotrophic Cretan Sea. Prog. Oceanogr., 46, 457–480.

    Google Scholar 

  • Eckelbarger, K. J. and L. Watling (1995): Role of phylogenetic constraints in determining reproductive patterns in deep-sea invertebrates. Invert. Biol., 114, 256–269.

    Google Scholar 

  • Eglinton, G., H. Elderfield, M. Whitfield and P. J. Le B. Williams (1995): The role of the North Atlantic in the global carbon cycle. Phil. Trans. Roy. Soc. Ser. B, 348, 121–264.

    Google Scholar 

  • Elizalde, M., O. Weber, A. Pascual, J. C. Sorbe and H. Etcheber (1999): Benthic response of Munnopsurus atlanticus (Crustacea Isopoda) to the carbon content of the near-bottom sedimentary environment on the southern margin of the Cap-Ferret Canyon (Bay of Biscay, northeastern Atlantic Ocean). Deep-Sea Res. II, 46, 2331–2344.

    Google Scholar 

  • Féral, J.-P., J.-G. Ferrand and A. Guille (1990): Macrobenthic physiological responses to environmental fluctuations: the reproductive cycle and enzymatic polymorphism of a eurybathyal sea-urchin on the north-western Mediterranean continental shelf and slope. Cont. Shelf Res., 10, 1147–1155.

    Google Scholar 

  • Fischer, G. and G. Wefer (1996): Long-term observation of particle fluxes in the Eastern Atlantic: seasonality, changes in flux with depth and comparison with the sediment record. p. 325–344. In The South Atlantic: Present and Past Circulation, ed. by G. Wefer, W. H. Berger, G. Siedler and D. J. Webb, Springer-Verlag, Berlin, Heidelberg.

    Google Scholar 

  • Fischer, G., D. Fütterer, R. Gersonde, S. Honjo, D. Ostermann and G. Wefer (1988): Seasonal variability of particle flux in the Weddell Sea and its relation to ice cover. Nature, 335, 426–428.

    Google Scholar 

  • Flach, E. and C. Heip (1996): Seasonal variation in faunal distribution and activity across the continental slope of the Goban Spur area (NE Atlantic). J. Sea Res., 36, 203–215.

    Google Scholar 

  • Gage, J. D. (1985): The analysis of population dynamics in deep-sea benthos. p. 201–212. In Proceedings of the 19th European Marine Biology Symposium, ed. by P. E. Gibbs, Cambridge University Press, Cambridge.

    Google Scholar 

  • Gage, J. D. (1987): Growth of the deep-sea irregular sea urchins Echinosigra phiale and Hemiaster expergitus in the Rockall Trough (N.E. Atlantic Ocean). Mar. Biol., 96, 19–30.

    Google Scholar 

  • Gage, J. D. (1990): Skeletal growth markers in the deep-sea brittle stars Ophiura ljungmani and Ophiomusium lymani. Mar. Biol., 104, 427–435.

    Google Scholar 

  • Gage, J. D. (1991): Biological rates in the deep sea: a perspective from studies on processes in the benthic boundary layer. Reviews in Aquatic Biology, 5, 49–100.

    Google Scholar 

  • Gage, J. D. and P. A. Tyler (1981a): Non-viable seasonal settlement of larvae of the upper bathyal brittle star Ophiocten gracilis in the Rockall Trough abyssal. Mar. Biol., 64, 153–161.

    Google Scholar 

  • Gage, J. D. and P. A. Tyler (1981b): Re-appraisal of age composition, growth and survivorship of the deep-sea brittle star Ophiura ljungmani from size structure in a single sample time series from the Rockall Trough. Mar. Biol., 64, 163–172.

    Google Scholar 

  • Gage, J. D. and P. A. Tyler (1982): Growth and reproduction in the deep-sea brittle star Ophiomusium lymani Wyville Thomsen. Oceanol. Acta, 5, 73–83.

    Google Scholar 

  • Gage, J. D. and P. A. Tyler (1985): Growth and recruitment in the deep-sea urchin Echinus affinis. Mar. Biol., 90, 41–53.

    Google Scholar 

  • Gage, J. D. and P. A. Tyler (1991): Deep-Sea Biology. Cambridge University Press, Cambridge, 504 pp.

    Google Scholar 

  • Gage, J. D., R. H. Lightfoot, M. Pearson and P. A. Tyler (1980): An introduction to a sample time-series of abyssal macrobenthos: methods and principle sources of variability. Oceanol. Acta, 3, 169–176.

    Google Scholar 

  • Gage, J. D., P. A. Tyler and D. Nichols (1986): Reproduction and growth of Echinus acutus var. norvegicus Düben & Koren and E. elegans Düben & Koren on the continental slope off Scotland. J. Exp. Mar. Biol. Ecol., 101, 61–83.

    Google Scholar 

  • Gardener, W. D., L. G. Sullivan and E. M. Thorndike (1984): Long-term photographic, current, and nephelometer observations of manganese nodule environments in the Pacific. Earth Planet. Sci. Lett., 70, 95–100.

    Google Scholar 

  • Gehlen, M., C. Rabouille, U. Ezat and L. D. Guidi-Guilvard (1997): Drastic changes in deep-sea sediment porewater composition induced by episodic input of organic matter. Limnol. Oceanogr., 42, 980–986.

    Google Scholar 

  • Geistdoerfer, P. (1979): New data on the reproduction of macrourids (Teleostei, Gadiformes). Sarsia, 64, 109–112.

    Google Scholar 

  • Gooday, A. J. (1993): Deep-sea benthic foraminiferal species which exploit phytodetritus: characteristic features and controls on distribution. Mar. Micropaleo., 22, 187–205.

    Google Scholar 

  • Gooday, A. J. and A. E. Rathburn (1999): Temporal variability in living deep-sea foraminifera: a review. Earth Sci. Rev., 46, 187–212.

    Google Scholar 

  • Gooday, A. J. and C. M. Turley (1990): Responses by benthic organisms to inputs of organic material to the ocean floor: a review. Phil. Trans. Roy. Soc. London, Series A, 331, 119–138.

    Google Scholar 

  • Gooday, A. J., L. A. Levin, P. Linke and T. Heeger (1992): The role of benthic Foraminifera in deep-sea food webs and carbon cycling. p. 63–91. In Deep-Sea Food Chains and the Global Carbon Cycle, ed. by G. T. Rowe and V. Pariente, Proceedings of NATO Advanced Research Workshop, College Station, Texas. Dordrecht, Kluwer Academic Publishers.

    Google Scholar 

  • Gooday, A. J., B. J. Bett and D. N. Pratt (1993): Direct observation of episodic growth in an abyssal xenophyophore (Protista). Deep-Sea Res. I, 40, 2131–2143.

    Google Scholar 

  • Gooday, A. J., O. Pfannkuche and P. J. D. Lambshead (1996): An apparent lack of response by metazoan meiofauna to phytodetritus deposition in the bathyal northeast Atlantic. J. Mar. Biol. Ass., U.K., 76, 297–310.

    Google Scholar 

  • Gordon, J. D. M. (1979): Seasonal reproduction in deep-sea fish. p. 223–229. In Cyclic Phenomena in Marine Plants and Animals, ed. by E. Naylor and R. G. Hartnoll, Pergamon Press, Oxford.

    Google Scholar 

  • Graf, G. (1989): Bentho-pelagic coupling in a deep-sea benthic community. Nature, 341, 437–439.

    Google Scholar 

  • Graf, G. (1992): Benthopelagic coupling: a benthic view. Oceanogr. Mar. Biol. Ann. Rev., 30, 149–190.

    Google Scholar 

  • Graf, G., S. A. Gerlach, P. Linke, W. Queisser, W. Ritzrau, A. Scheltz, L. Thomsen and U. Witte (1995): Benthic-pelagic coupling in the Greenland-Norwegian Sea and its effect on the geological record. Geologische Rundschau, 84, 49–58.

    Google Scholar 

  • Grassle, J. F. and L. S. Morse-Porteous (1987): Macrofaunal colonization of disturbed deep-sea environments and the structure of deep-sea benthic communities. Deep-Sea Res., 34, 1911–1950.

    Google Scholar 

  • Green, A., P. A. Tyler, M. V. Angel and J. D. Gage (1994): Gametogenesis in deep-and surface-dwelling oceanic stalked barnacles from the NE Atlantic. J. Exp. Mar. Biol. Ecol., 184, 143–158.

    Google Scholar 

  • Guidi-Guilvard, L. (2002): DYFAMED-BENTHOS, a long time-series benthic survey at 2347 m depth in the NW Mediterranean: general introduction. Deep-Sea Res. II (in press).

  • Guidi-Guilvard, L., J.-C. Miquel and A. Kripounoff (2000): Metazoan meiobenthos responses to sedimentation events in the deep northwest Mediterranean. 9th Deep-Sea Biology Symposium, Abstracts, p. 26.

  • Gutt, J., A. Starmans and G. Diec kmann (1998): Phytodetritus deposition on the Antarctic shelf and upper slope: its relevance to the benthic system. J. Mar. Sys., 17, 435–444.

    Google Scholar 

  • Haake, B., V. Ittekkot, T. Rixen, V. Ramaswamy, R. R. Nair and W. B. Curry (1993): Seasonality and interannual variability of particle fluxes to the deep Arabian Sea. Deep-Sea Res., 40, 1323–1344.

    Google Scholar 

  • Harrison, K. (1988): Seasonal reproduction in deep-sea Crustacea (Isopoda: Asellota). Journal of Natural History, 22, 175–197.

    Google Scholar 

  • Hartnoll, R. G. and A. L. Rice (1985): Further studies on the biology of the deep-water spider crab Dorhynchus thomsoni: instar sequence and the annual cycle. p. 231–241. In Proceedings of the 19th European Marine Biology Symposium, ed. by P. E. Gibbs, Cambridge University Press, Cambridge.

    Google Scholar 

  • Hartnoll, R. G., A. L. Rice and D. I. Williamson (1987): Dorhynchus thomsoni: a deep-sea crab with a strange life history. Investigacion Pesquera, 51(Suppl. 1), 125–133.

    Google Scholar 

  • Hecker, B. (1990): Photographic evidence for the rapid flux of particles to the seafloor and their transport down the continental slope. Deep-Sea Res., 37, 1773–1782.

    Google Scholar 

  • Honjo, S. (1982): Seasonality of biogenic and lithogenic fluxes in the Panama Basin. Science, 218, 883–884.

    Google Scholar 

  • Honjo, S. and S. J. Manganini (1993): Annual biogenic particle fluxes to the interior of the North Atlantic Ocean: studied at 34°N 21°W and 48°N 21°W. Deep-Sea Res. II, 40, 587–607.

    Google Scholar 

  • Honjo, S., J. Dymond, R. Collier and S. J. Manganini (1995): Export production of particles to the interior of the equatorial Pacific Ocean during the 1992 EqPac experiment. Deep-Sea Res. II, 42, 831–870.

    Google Scholar 

  • Honjo, S., J. Dymond, W. Prell and V. Ittekot (1999): Monsoon-controlled export fluxes to the interior of the Arabian Sea. Deep-Sea Res. II, 46, 1859–1902.

    Google Scholar 

  • Hutchings, J. A. and R. L. Haedrich (1984): Growth and population structure in two species of bivalves (Nucularidae) from the deep sea. Mar. Ecol. Prog. Ser., 17, 135–142.

    Google Scholar 

  • Ittekkot, V., R. R. Nair, S. Honjo, V. Ramaswamy, M. Bartsch, S. Manganini and B. N. Desai (1991): Enhanced particle fluxes in Bay of Bengal induced by injection of fresh water. Nature, 351, 385–387.

    Google Scholar 

  • Jackson, J. B. C., M. X. Kirby, W. H. Berger, K. A. Bjorndal, L. W. Botsford, B. J. Bourque, R. H. Bradbury, R. Cooke, J. Erlandson, J. A. Estes, T. P. Hughes, S. Kidwell, C. B. Lange, H. S. Lenihan, J. M. Pandolfi, C. H. Peterson, R. S. Steneck, M. J. Tegner and R. R. Warner (2001): Historical overfishing and the recent collapse of coastal ecosystems. Science, 293, 629–638.

    Google Scholar 

  • Karl, D. M., J. R. Christiansen, J. E. Dore, D. V. Hebel, R. M. Letelier, L. M. Tupas and C. D. Winn (1996): Seasonal and interannual variability in primary production and particle flux at Station ALOHA. Deep-Sea Res. II, 43, 539–568.

    Google Scholar 

  • Kaufmann, R. S. and K. L. Smith (1997): Activity patterns of mobile epibenthic megafauna at an abyssal site in the eastern North Pacific: results from a 17–month time-lapse photographic study. Deep-Sea Res. I, 44, 559–579.

    Google Scholar 

  • Kitazato, H. and T. Ohga (1995): Seasonal changes in deep-sea benthic foraminiferal populations: results of long-term observations at Sagami Bay, Japan. p. 331–342. In Biogeochemical Processes and Ocean Flux in the Western Pacific, ed by H. Sakai and Y. Nozaki, Terra Scientific Pub., Tokyo.

    Google Scholar 

  • Kitazato, H., Y. Shirayama, T. Nakatsuka, S. Fujiwara, M. Shimanaga, Y. Kato, Y. Okada, J. Kanda, A. Yamaoka, T. Masuzawa and K. Suzuki (2000): Seasonal phytodetritus deposition and responses of bathyal benthic foraminiferal populations in Sagami Bay: preliminary results from “Project Sagami 1996–1999”. Mar. Micropaleo., 40, 135–149.

    Google Scholar 

  • Kurbjeweit, F., G. Schmiedl, R. Schiebl, Ch. Hemleben, O. Pfannkuche, K. Wallmann and P. Schäfer (2000): Distribution, biomass and diversity of benthic foraminifera in relation to sediment geochemistry in the Arabian Sea. Deep-Sea Res. II, 47, 2913–2955.

    Google Scholar 

  • Lampitt, R. S. (1985): Evidence for seasonal deposition of detritus to the deep-sea floor and its subsequent resuspension. Deep-Sea Res., 32, 885–897.

    Google Scholar 

  • Lampitt, R. S. (1990): Directly measured rapid growth of a deep-sea barnacle. Nature, 345, 805–807.

    Google Scholar 

  • Lampitt, R. S. and A. N. Antia (1997): Particle flux in deep seas: regional characteristics and temporal variability. Deep-Sea Res. I, 44, 1377–1403.

    Google Scholar 

  • Lampitt, R. S. and G. J. L. Paterson (1988): The feeding behaviour of an abyssal sea anemone from in situ photographs and trawl samples. Oceanol. Acta, 10, 455–461.

    Google Scholar 

  • Lampitt, R. S., N. R. Merrett and M. H. Thurston (1983): Interrelations of necrophagous amphipods, a fish predator, and tidal currents in the deep sea. Mar. Biol., 74, 73–78.

    Google Scholar 

  • Lampitt, R. S., R. C. Raine, D. M. S. Billett and A. L. Rice (1995): Material supply to the European continental slope: A budget based on benthic oxygen demand and organic supply. Deep-Sea Res. I, 42, 1865–1880.

    Google Scholar 

  • Lampitt, R. S., B. J. Bett, K. Kiriakoulakis, E. E. Popova, O. Ragueneau, A. Vangriesheim and G. A. Wolff (2001): Material supply to the abyssal seafloor in the Northeast Atlantic. Prog. Oceanogr., 50, 27–63.

    Google Scholar 

  • Lauerman, L. and R. S. Kaufmann (1998): Deep-sea epibenthic echinodermes and a temporally varying food supply: results from a one year rime series in the NE Pacific. Deep-Sea Res. II, 45, 817–842.

    Google Scholar 

  • Legendre, L. and J. Le Fèvre (1995): Microbial food webs and the export of biogenic carbon in oceans. Aquat. Microb. Ecol., 9, 66–77.

    Google Scholar 

  • Levin, L. A., N. E. Blair, C. M. Martin, D. J. DeMaster, G. Plaia and C. J. Thomas (1999): Macrofaunal processing of phytodetritus at two sites on the Carolina margin. Mar. Ecol. Prog. Ser., 182, 537–554.

    Google Scholar 

  • Lightfoot, R. H., P. A. Tyler and J. D. Gage (1979): Seasonal reproduction in deep-sea bivalves and brittle stars. Deep-Sea Res., 26, 967–973.

    Google Scholar 

  • Linke, P. (1992): Metabolic adaptations of deep-sea benthic foraminifera to seasonally varying food input. Mar. Ecol. Prog. Ser., 81, 51–63.

    Google Scholar 

  • Linke, P., A. V. Altenbach, G. Graf and T. Heeger (1995): Response of deep-sea benthic foraminifera to a simulated sedimentation event. J. Foram. Res., 25, 75–82.

    Google Scholar 

  • Lochte, K. (1992): Bacterial standing stock and consumption of organic carbon in the benthic boundary layer of the abyssal North Atlantic. p. 1–10. In Deep-Sea Food Chains and the Global Carbon Cycle, ed. by G. T. Rowe and V. Pariente, Kluwer Academic Publishers, Dordrecht.

    Google Scholar 

  • Lochte, K. and C. M. Turley (1988): Bacteria and cyanobacteria associated with phytodetritus in the deep sea. Nature, 333, 67–69.

    Google Scholar 

  • Longhurst, A. (1995): Seasonal cycles of pelagic production and consumption. Prog. Oceanogr., 36, 77–167.

    Google Scholar 

  • Longhurst, A. (1998): Ecological Geography of the Sea. Academic Press, San Diego, London, Boston, New York, Sydney, Tokyo, Toronto, 398 pp.

    Google Scholar 

  • Longhurst, A., S. Sathyendranath, T. Platt and C. Caverhill (1995): An estimate of global primary production in the ocean from satellite radiometer data. J. Plankton Res., 17, 1245–1271.

    Google Scholar 

  • Loubere, P. (1998): The impact of seasonality on the benthos as reflected in the assemblages of deep-sea foraminifera. Deep-Sea Res. I, 45, 409–432.

    Google Scholar 

  • Loubere, P. and M. Fariduddin (1999a): Quantitative estimation of global patterns of surface ocean biological productivity and its seasonal variation on timescales from centuries to millennia. Global Biogeochem. Cycles, 13, 115–133.

    Google Scholar 

  • Loubere, P. and M. Fariduddin (1999b): Benthic Foraminifera and the flux of organic carbon to the seabed. p. 181–199. In Modern Foraminifera, ed. by B. K. Sen Gupta, Kluwer Academic Publishers.

  • Martin, W. R. and M. L. Bender (1988): The variability of benthic fluxes and sedimentary remineralization rates in response to seasonally variable organic carbon rain rates in the deep sea: a modelling study. Am. J. Sci., 288, 561–574.

    Google Scholar 

  • Masson, D. G., N. H. Kenyon and P. P. E. Weaver (1996): Slides, debris flow, and turbidity currents. p. 136–151. In Oceanography: An Illustrated Guide, ed. by C. P. Summerhayes and S. A. Thorpe, Manson Publishing, London.

    Google Scholar 

  • Menzies, R. J. (1965): Conditions for the existence of life on the abyssal sea floor. Oceanogr. Mar. Biol. Ann. Rev., 3, 195–210.

    Google Scholar 

  • Menzies, R. J. and G. T. Rowe (1969): The distribution and significance of detrital turtle grass, Thalassia testudinata, on the deep-sea floor off North Carolina. Internationale Revue der gesampten Hydrobiologie und Hydrographie, 54, 217–222.

    Google Scholar 

  • Miquel, J. C., S. W. Fowler, J. La Rosa and P. Buat-Menard (1994): Dynamics of the downward flux of particles and carbon in the open northwestern Mediteranean Sea. Deep-Sea Res. I, 41, 243–261.

    Google Scholar 

  • Momma, H., R. Iwasw, K. Mitsuzawa, Y. Kaiho and Y. Fujiwara (1998): Preliminary results of a three-year continuous observation by a deep seafloor observatory in Sagami Bay, central Japan. Phys. Earth Planet. Inter., 108, 263–274.

    Google Scholar 

  • Monaco, A., P. E. Biscaye, J. Soyer, R. Pocklington and S. Heussner (1990): Particle fluxes and ecosystem responses on a continental margin: the 1985–1988 Mediterranean ECOMARGE experiment. Cont. Shelf Res., 10, 809–839.

    Google Scholar 

  • Monaco, A., P. E. Biscaye and P. Laborde (1999): The ECOFER (ECOsysteme du canyon du cap-FERret) experiment in the Bay of Biscay: introduction, objectives and major results. Deep-Sea Res. II, 46, 1967–1978.

    Google Scholar 

  • Moseley, H. N. (1880): Deep-sea dredging and life in the deep sea. Nature, 21, 591–593.

    Google Scholar 

  • Nair, R. R., V. Ittekkot, S. J. Manganini, V. Ramaswamy, B. Haake, E. T. Degens, B. N. Desai and S. Honjo (1989): Increased particle flux to the deep ocean related to monsoons. Nature, 338, 749–751.

    Google Scholar 

  • Newton, A. C. and G. T. Rowe (1995): The abundance of benthic calcareous Foraminifera and other meiofauna at a time series station in the Northeast Water Polynya, Greenland. J. Geophys. Res., 100, 4423–4438.

    Google Scholar 

  • Newton, P. P., R. S. Lampitt, T. D. Jickells, P. King and C. Boutle (1994): Temporal and spatial variability of biogenic particle fluxes during the JGOFS northeast Atlantic process studies at 47°N, 20°W. Deep-Sea Res. I, 41, 1617–1642.

    Google Scholar 

  • Nodder, S. D. and L. C. Northcote (2001): Episodic particulate fluxes at southern temperate mid-latitudes (42–45°S) in the Subtropical Front region, east of New Zealand. Deep-Sea Res. I, 48, 833–864.

    Google Scholar 

  • Ohga, T. and H. Kitazato (1997): Seasonal changes in bathyal foraminiferal populations in response to the flux of organic matter (Sagami Bay, Japan). Terra Nova, 9, 33–37.

    Google Scholar 

  • Patching, J. W., R. C. T. Raine, P. R. O. Barnett and J. Watson (1986): Abyssal benthic oxygen consumption in the Northeastern Atlantic: measurements using the suspended core technique. Oceanol. Acta, 9, 1–7.

    Google Scholar 

  • Pfannkuche, O. (1985): The deep-sea meiofauna of the Porcupine Seabight and abyssal plain (NE Atlantic): population structure, distribution, standing stocks. Oceanol. Acta, 8, 343–353.

    Google Scholar 

  • Pfannkuche, O. (1992): Organic carbon flux through the benthic community in the temperate Northeast Atlantic. p. 183–198. In Deep-Sea Food Chains and the Global Carbon Cycle, ed. by G. T. Rowe and V. Pariente, Kluwer Academic Publishers, Dordrecht.

    Google Scholar 

  • Pfannkuche, O. (1993): Benthic response to the sedimentation of particulate organic matter at the BIOTRANS station, 47°N, 20°W. Deep-Sea Res. II, 40, 135–149.

    Google Scholar 

  • Pfannkuche, O. and K. Lochte (1993): Open ocean pelagobenthic coupling: cyanobacteria as tracers of sedimenting salp faeces. Deep-Sea Res. I, 40, 727–737.

    Google Scholar 

  • Pfannkuche, O. and T. Soltwedel (1998): Small benthic size classes along the N.W. European continental margin: spatial and temporal variability in activity and biomass. Prog. Oceanogr., 42, 189–207.

    Google Scholar 

  • Pfannkuche, O. and H. Thiel (1987): Meiobenthic stocks and benthic activity on the NE-Svalbard shelf and in the Nansen Basin. Polar Biol., 7, 253–266.

    Google Scholar 

  • Pfannkuche, O., A. Boetius, K. Lochte, U. Lundgreen and H. Thiel (1999): Responses of deep-sea benthos to sedimentation patterns in the North-East Atlantic in 1992. Deep-Sea Res. I, 46, 573–596.

    Google Scholar 

  • Pfannkuche, O., S. Sommer and A. Kähler (2000): Coupling between phytodetritus deposition and the small-sized benthic biota in the deep Arabian Sea: analyses of biogenic sediment compounds. Deep-Sea Res. II, 47, 2805–2833.

    Google Scholar 

  • Poremba, K. and H.-G. Hoppe (1995): Spatial variation of benthic microbial production and hydrolytic enzyme activity down the continental slope of the Celtic Sea. Mar. Ecol. Prog. Ser., 118, 237–245.

    Google Scholar 

  • Priede, I. G. and P. M. Bagley (2000): In situ studies on deep-sea demersal fishes using autonomous unmanned lander platforms. Oceanogr. Mar. Biol. Ann. Rev., 38, 357–392.

    Google Scholar 

  • Priede, I. G., P. M. Bagley and K. L. Smith (1994): Seasonal changes in activity of abyssal demersal scavenging grenadiers Coryphaenoides (Nematonurus) armatus in the eastern North Pacific Ocean. Limnol. Oceanogr., 39, 279–285.

    Google Scholar 

  • Ramseier, R. O., C. Garrity, R. Bauerfeind and R. Peinert (1999): Sea-ice impact on long-term particle flux in the Greenland Sea's Is Odden-Nordbukta region. J. Geophys. Res., 104, 5329–5343.

    Google Scholar 

  • Rannou, M. (1976): Age et croissance d'un poisson bathyal: Nezumia sclerorhynchus (Macrouridae Gadiforme) de la Mer d'Alboran. Cahiers de Biologie Marine, 17, 413–421.

    Google Scholar 

  • Rice, A. L., D. S. M. Billett, J. Fry, A. W. G. John, R. S. Lampitt, R. F. C. Mantoura and R. J. Morris (1986): Seasonal deposition of phytodetritus to the deep-sea floor. Proc. Roy. Soc. Edin., 88B, 265–279.

    Google Scholar 

  • Rice, A. L., M. H. Thurston and B. J. Bett (1994): The IOS DEEPSEAS programme: introduction and photographic evidence for the presence and absence of a seasonal input of phytodetritus at contrasting abyssal sites in the northeastern Atlantic. Deep-Sea Res. I, 41, 1305–1320.

    Google Scholar 

  • Rice, A. L., J. D. Gage, R. S. Lampitt, O. Pfannkuche and M. Sibuet (1998): BENGAL: high resolution temporal and spatial study of the benthic biology and geochemistry of a northeast Atlantic abyssal locality. p. 273–287. In Third European Marine Science and Technology Conference, Lisbon 23–27 May 1998. Project Synopses, Vol. 1, Luxembourg Office of Official Publications of the European Communities.

  • Rixen, T., V. Ittekkot, B. Haake-Gaye and P. Schäfer (2000): The influence of the SW monsoon on the deep-sea organic carbon cycle in the Holocene. Deep-Sea Res. II, 47, 2629–2651.

    Google Scholar 

  • Rokop, F. J. (1974): Reproductive patterns in the deep-sea benthos. Science, 186, 743–745.

    Google Scholar 

  • Rokop, F. J. (1977a): Seasonal reproduction of the brachiopod Frieleia halli and the scaphopod Cadulus californicus at bathyal depths in the deep sea. Mar. Biol., 43, 237–246.

    Google Scholar 

  • Rokop, F. J. (1977b): Patterns of reproduction in the deep-sea benthic crustaceans: a re-evaluation. Deep-Sea Res., 24, 683–691.

    Google Scholar 

  • Sardà, F., J. E. Cartes and J. B. Company (1994): Spatio-temporal variations in megabenthos abundance in three different habitats of the Catalan deep-sea (Western Mediterranean). Mar. Biol., 120, 211–219.

    Google Scholar 

  • Sauter, E. M., M. Schlüter and E. Suess (2001): Organic carbon flux and mineralisation in surface sediments from the northern North Atlantic derived from pore-water oxygen microprofiles. Deep-Sea Res. I, 48, 529–553.

    Google Scholar 

  • Sayles, F. L., W. R. Martin and W. G. Deuser (1994): Response of benthic oxygen demand to particulate organic carbon supply in the deep sea near Bermuda. Nature, 371, 686–689.

    Google Scholar 

  • Scheltema, R. S. (1994): Adaptations for reproduction among deep-sea benthic molluscs: an appraisal of the existing evidence. p. 44–75. In Reproduction, Larval Biology, and Recruitment of the Deep-Sea Benthos, ed. by C. M Young and K. J. Eckelbarger, Columbia University Press, New York.

    Google Scholar 

  • Shimanaga, M. and Y. Shirayama (2000): Response of benthic organisms to seasonal change of organic matter deposition in the bathyal Sagami Bay, central Japan. Oceanol. Acta, 23, 91–107.

    Google Scholar 

  • Shimanaga, M., H. Kitazato and Y. Shirayama (2000): Seasonal patterns of vertical distribution between meiofaunal groups in relation to phytodetritus deposition in the bathyal Sagami Bay, central Japan. Oceanol. Acta, 56, 379–387.

    Google Scholar 

  • Smart, C. M., S. King, A. J. Gooday, J. W. Murray and E. Thomas (1994): A benthic foraminiferal proxy for pulsed organic matter palaeofluxes. Mar. Micropaleo., 23, 89–99.

    Google Scholar 

  • Smith, A., I. G. Priede, P. M. Bagley and S. W. Addison (1997): Interception and dispersal of artificial food falls by scavenging fishes in the abyssal Northeast Atlantic: early season observations prior to annual deposition of phytodetritus. Mar. Biol., 128, 329–336.

    Google Scholar 

  • Smith, C. R. (1986): Nekton falls, low-intensity disturbance and community structure in infaunal benthos in the deep sea. J. Mar. Res., 44, 567–600.

    Google Scholar 

  • Smith, C. R. (1994): Tempo and mode in deep-sea benthic ecology: punctuated equilibrium revisited. Palaios, 9, 3–13.

    Google Scholar 

  • Smith, C. R., H. Kukert, R. A. Wheatcroft, P. A. Jumars and J. W. Deming (1989): Vent faunas on whale remains. Nature, 341, 27–28.

    Google Scholar 

  • Smith, C. R., D. J. Hoover, S. E. Doan, R. H. Pope, D. J. Demaster, F. C. Dobbs and M. A. Altabet (1996): Phytodetritus at the abyssal seafloor across 10° of latitude in the central equatorial Pacific. Deep-Sea Res. II, 43, 1309–1338.

    Google Scholar 

  • Smith, K. L. (1987): Food energy supply and demand: a discrepancy between particulate organic carbon flux and sediment community oxygen consumption in the deep ocean. Limnol. Oceanogr., 32, 201–220.

    Google Scholar 

  • Smith, K. L. and R. J. Baldwin (1984): Seasonal fluctuations in deep-sea sediment community oxygen consumption: central and eastern North Pacific. Nature, 307, 624–626.

    Google Scholar 

  • Smith, K. L and E. R. M. Druffel (1998): Long time-series monitoring of an abyssal site in the NE Pacific: an introduction. Deep-Sea Res. II, 45, 573–586.

    Google Scholar 

  • Smith, K. L. and R. S. Kaufmann (1999): Long-term discrepancy between food supply and demand in the deep eastern North Pacific. Science, 284, 1174–1177.

    Google Scholar 

  • Smith, K. L., G. A. White and M. B. Laver (1979): Oxygen uptake and nutrient exchange measured in situ using a free vehicle grab respirometer. Deep-Sea Res., 26, 337–346.

    Google Scholar 

  • Smith, K. L., R. J. Baldwin and P. M. Williams (1992): Reconciling particulate organic carbon flux and sediment community oxygen consumption in the deep North Pacific. Nature, 359, 313–316.

    Google Scholar 

  • Smith, K. L., R. S. Kaufmann and R. J. Baldwin (1994): Coupling near-bottom pelagic and benthic processes at abyssal depths in the eastern North Pacific Ocean. Limnol. Oceanogr., 39, 1101–1118.

    Google Scholar 

  • Smith, K. L., R. J. Baldwin, R. C. Glatts, R. S. Kaufmann and E. C. Fisher (1998): Detrital aggregates on the sea floor: chemical composition and aerobic decomposition rates at a time-series station in the abyssal NE Pacific. Deep-Sea Res. II, 45, 843–880.

    Google Scholar 

  • Snelgrove, P. V. R., J. F. Grassle and R. F. Petrecca (1992): The role of food patches in maintaining high deep-sea diversity: Field experiments with hydrodynamically unbiased colonization trays. Limnol. Oceanogr., 37, 1543–1550.

    Google Scholar 

  • Snelgrove, P. V. R., J. F. Grassle and R. F. Petrecca (1994): Macrofaunal response to artificial enrichments and depressions in a deep-sea habitat. J. Mar. Res., 52, 345–369.

    Google Scholar 

  • Snelgrove, P. V. R., J. F. Grassle and R. F. Petrecca (1996): Experimental evidence for aging food patches as a factor contributing to high macrofaunal diversity. Limnol. Oceanogr., 41, 605–614.

    Google Scholar 

  • Soetaert, K., P. M. J. Herman and J. J. Middelburg (1996): Dynamic response of deep-sea sediments to seasonal variations: A model. Limnol. Oceanogr., 41, 1651–1668.

    Google Scholar 

  • Soltwedel, T. (1997): Temporal variabilities in benthic activity and biomass on the western European continental margin. Oceanol. Acta, 20, 871–879.

    Google Scholar 

  • Soltwedel, T. (2000): Metazoan meiobenthos along continental margins: a review. Prog. Oceanogr., 46, 59–84.

    Google Scholar 

  • Soltwedel, T. and H. Thiel (1995): Biogenic sediment compounds in relation to marine meiofauna abundances. Internationale Revue der gesampten Hydrobiologie, 80, 297–311.

    Google Scholar 

  • Soltwedel, T., O. Pfannkuche and H. Thiel (1996): The structure of deep-sea meiobenthos in the north-eastern Atlantic: nematode size spectra in relation to environmental variables. J. Mar. Biol. Ass., U.K., 76, 327–344.

    Google Scholar 

  • Sommer, S. and O. Pfannkuche (2000): Metazoan meiofauna of the deep Arabian Sea: standing stocks, size spectra and regional variability in relation to monsoonal induced enhanced sedimentation regimes of particulate organic matter. Deep-Sea Res. II, 47, 2957–2977.

    Google Scholar 

  • Sorbe, J. C. (1999): Deep-sea macrofaunal assemblages within the Benthic Boundary Layer of the Cap-Ferret Canyon (Bay of Biscay, NE Atlantic). Deep-Sea Res. II, 46, 2309–2329.

    Google Scholar 

  • Stavrakakis, S., G. Chronis, A. Tselepides, S. Heussner, A. Monaco and A. Abassi (2000): Downward fluxes of settling particles in the deep Cretan Sea (NE Mediterranean). Prog. Oceanogr., 46, 217–240.

    Google Scholar 

  • Stephens, M. P., D. C. Kadko, C. R. Smith and M. Latasa (1997): Chlorophyll-α and pheopigments as tracers of labile organic carbon at the central equatorial Pacific ocean floor. Geochim. Cosmochim. Acta, 61, 4605–4617.

    Google Scholar 

  • Sumida, P. Y. G., P. A. Tyler, R. S. Lampitt and J. D. Gage (2000): Reproduction, dispersal and settlement of the bathyal ophiuroid Ophiocten gracilis in the NE Atlantic Ocean. Mar. Biol., 137, 623–630.

    Google Scholar 

  • Takahashi, K., S. Honjo and S. Tabata (1989): Siliceous phytoplankton flux: interannual variability and response to hydrograophic changes in the northeastern Pacific. p. 151–160. In Aspects of Climate Variability in the Pacific and Western Americas, ed. by D. H. Peterson, AGU Geophysical Monograph, 55, American Geophysical Union, Washington, D.C.

    Google Scholar 

  • Takahashi, K., N. Fujitani, M. Yanada and Y. Maita (2000): Long-term biogenic particle fluxes in the Bering Sea and the central subarctic Pacific Ocean, 1990–1995. Deep-Sea Res. I, 47, 1723–1759.

    Google Scholar 

  • Thiel, H. (1983): Meiobenthos and nanobenthos of the deep sea. p. 167–230. In Deep-Sea Biology, ed. by G. T. Rowe, Wiley, New York.

    Google Scholar 

  • Thiel, H., O. Pfannkuche, G. Schriever, K. Lochte, A. J. Gooday, Ch. Hemleben, R. F. G. Mantoura, C. M. Turley, J. W. Patching and F. Riemann (1989): Phytodetritus on the deep-sea floor in a central oceanic region of the Northeast Atlantic. Biol. Oceanogr., 6, 203–239.

    Google Scholar 

  • Thistle, D., S. C. Ertman and C. Fauchald (1991): The fauna of the HEBBLE site: patterns in standing stock and sediment dynamic effects. Mar. Geol., 99, 413–422.

    Google Scholar 

  • Thomas, E. and A. J. Gooday (1996): Cenozoic deep-sea benthic foraminifers: tracers for changes in oceanic productivity? Geology, 24, 355–358.

    Google Scholar 

  • Tselepides, A. and T. Polychronaki (eds.) (2000): Pelagic-benthic coupling in the oligotrophic Cretan Sea. Prog. Oceanogr., 46, 85–480.

  • Turley, C. M. (2000): Bacteria in the cold deep-sea benthic boundary layer and sediment-water interface in the NE Atlantic. FEMS Microbiol. Ecol., 33, 89–99.

    Google Scholar 

  • Turley, C. M. and K. Lochte (1990): Microbial response to the input of fresh detritus to the deep-sea bed. Palaeogeo., Palaeoclim., Palaeoecol., 89, 3–23.

    Google Scholar 

  • Turley, C. M., K. Lochte and D. J. Patterson (1988): A barophilic flagellate isolated from 4500 m in the mid-North Atlantic. Deep-Sea Res., 35, 1079–1092.

    Google Scholar 

  • Turley, C. M., K. Lochte and R. S. Lampitt (1995): Transformations of biological particles during sedimentation in the northeastern Atlantic. Phil. Trans. Roy. Soc. London, Ser. B, 348, 179–189.

    Google Scholar 

  • Turner, R. D. (1973): Wood-boring bivalves, opportunistic species in the deep sea. Science, 180, 1377–1379.

    Google Scholar 

  • Tyler, P. A. (1986): Studies of a benthic time series: reproductive biology of benthic invertebrates in the Rockall Trough. Proc. Roy. Soc. Edin., 88B, 175–190.

    Google Scholar 

  • Tyler, P. A. (1988): Seasonality in the deep sea. Oceanogr. Mar. Biol. Ann. Rev., 26, 227–258.

    Google Scholar 

  • Tyler, P. A. (1996): Faunal responses to temporal disequilibrium in the deep sea. p. 261–270. In Deep-Sea and Extreme Shallow-Water Habitats: Affinities and Adaptations, ed. by F. Uiblein, J. Ott and M. Stachowitsch, Biosystematics and Ecology Series Vol. 11.

  • Tyler, P. A. and J. D. Gage (1980): Reproduction and growth of the deep-sea brittlestar Ophiura ljungmani (Lyman). Oceanol. Acta, 3, 177–185.

    Google Scholar 

  • Tyler, P. A. and J. D. Gage (1982): Ophiopluteus ramosus, the larval form of Ophiocten gracilis (Echinodermata: Ophiuroidea). J. Mar. Biol. Ass., U.K., 62, 485–486.

    Google Scholar 

  • Tyler, P. A. and J. A. Gage (1984): Seasonal reproduction in Echinus affinis (Echinodermata: Echinoidea) in the Rockall Trough, northeast Atlantic Ocean. Deep-Sea Res., 31, 387–402.

    Google Scholar 

  • Tyler, P. A. and S. L. Pain (1982): The reproductive biology of Plutonaster bifrons, Dytaster insignis and Philaster andromeda (Asteroidea: Astropectinidae) from the Rockall Trough. J. Mar. Biol. Ass., U.K., 62, 869–887.

    Google Scholar 

  • Tyler, P. A. and C. M. Young (1992): Reproduction in marine invertebrates in “stable” environments: the deep-sea model. Invertebrate Reproduction and Development, 22, 185–192.

    Google Scholar 

  • Tyler, P. A., A. Grant, S. L. Pain and J. D. Gage (1982): Is annual reproduction in deep-sea echinoderms a response to variability in their environment. Nature, 300, 747–749.

    Google Scholar 

  • Tyler, P. A., A. Muirhead and J. Coleman (1985): Observations on continuous reproduction in large deep-sea epibenthos. p. 223–230. In Proceedings of the 19th European Marine Biology Symposium, ed. by P. E. Gibbs, Cambridge University Press, Cambridge.

    Google Scholar 

  • Tyler, P. A., D. S. M. Billett and J. D. Gage (1990): Seasonal reproduction in the seastar Dytaster grandis from 4000 m in the North-East Atlantic Ocean. J. Mar. Biol. Ass., U.K., 70, 163–172.

    Google Scholar 

  • Tyler, P. A., J. D. Gage and D. S. M. Billett (1992a): Reproduction and recruitment in deep-sea invertebrate populations in the NE Atlantic Ocean: a review of the options. p. 257–262. In Proceeding of the 25th European Marine Biology Symposium, ed. by G. Colombo, Olsen & Olsen, Fredensborg.

    Google Scholar 

  • Tyler, P. A., R. Harvey, L. A. Giles and J. D. Gage (1992b): Reproductive strategies and diet in deep-sea nuculanid protobranchs (Bivalvia: Nuculoidea) from the Rockall Trough. Mar. Biol., 114, 571–580.

    Google Scholar 

  • Tyler, P. A., J. D. Gage, G. J. L. Paterson and A. L. Rice (1993): Dietary constraints on reproductive periodicity in two sympatric deep-sea astropectinid seastars. Mar. Biol., 115, 267–277.

    Google Scholar 

  • Tyler, P. A., L. S. Campos-Creasey and L. A. Giles (1994): Environmental control of quasi-continuous and seasonal reproduction in deep-sea invertebrates. p. 158–178. In Reproduction, Larval Biology, and Recruitment of the Deep-Sea Benthos, ed. by C. M. Young and K. J. Eckelbarger, Columbia University Press, New York.

    Google Scholar 

  • Van Dover, C. L. (2000): The Ecology of Deep-Sea Hydrothermal Vents. Princeton University Press, Princeton, NJ, 424 pp.

    Google Scholar 

  • Van Weering, Tj. C. E., I. N. McCave and I. B. Hall (eds.) (1998): Ocean Margin Exchange (OMEX I) benthic processes study. Prog. Oceanogr., 42, 1–257.

  • Van-Praet, M. (1985): Cycles biologiques de cnidaires abyssaux. Bull. Soc. Zool. Fr., 110, 461–463.

    Google Scholar 

  • Van-Praet, M. (1990): Gametogenesis and the reproductive cycle in the deep-sea anemone Paracalliactis stephensoni (Cnidaria: Actiniaria). J. Mar. Biol. Ass., U.K., 70, 163–172.

    Google Scholar 

  • Van-Praet, M., A. L. Rice and M.H. Thurston (1990): Reproduction in two deep-sea anemones (Actiniaria); Phelliactis hertwigi and P. robustus. Prog. Oceanogr., 24, 207–222.

    Google Scholar 

  • Vanreusel, A., N. Cosson-Sarradin, A. J. Gooday, G. J. L. Paterson, J. Galeron, M. Sibuet and M. Vincx (2001): Evidence for episodic recruitment in a small opheliid polychaete species from the abyssal NE Atlantic. Prog. Oceanogr., 50, 285–301.

    Google Scholar 

  • Wakeham, S. G., C. Lee, J. I. Hedges, P. J. Hernes and M. L. Peterson (1997): Molecular indicators of diagenetic status in marine organic matter. Geochim. Cosmochim. Acta, 61, 5363–5369.

    Google Scholar 

  • Wefer, G. and G. Fisher (1991): Annual production and export flux in the Southern Ocean from sediment trap data. Mar. Chem., 35, 597–613.

    Google Scholar 

  • Williams, R. and J. Moyse (1988): Occurrence, distribution and orientation of Poecilasma kaempferi Darwin (Cirripedia: Pedunculata) epizoic on Neolithodes grimaldi Milne-Edwards and Bouvier (Decapoda: Anomura) in the northeast Atlantic. J. Crust. Biol., 8, 177–186.

    Google Scholar 

  • Witbaard, R., G. C. A. Duineveld, J. A. Van der Weele, E. M. Berghuis and J. L. Reyss (2000): The benthic response to seasonal deposition of phytodetritus at the Porcupine Abyssal Plain in the North East Atlantic. J. Sea Res., 43, 15–31.

    Google Scholar 

  • Witte, U. (1996): Seasonal reproduction in deep-sea sponges—triggered by vertical particle flux. Mar. Biol., 124, 571–581.

    Google Scholar 

  • Wolff, T. (1979): Animals associated with seagrass in the deep sea. p. 199–224. In A Handbook of Seagrass Biology: An Ecosystem Perspective, ed. by R. C. Phillips and C. P. McRoy, Garland Publishing Company, New York.

    Google Scholar 

  • Wong, C. S., F. A. Whitney, D. W. Crawford, K. Iseki, R. J. Matear, W. K. Johnson, J. S. Page and D. Timothy (1999): Seasonal and interannual variability in particle fluxes of carbon, nitrogen and silicon from time series sediment traps at Ocean Station P, 1982–1993: relationship to changes in subarctic primary productivity. Deep-Sea Res. II, 46, 2735–2760.

    Google Scholar 

  • Yamaoka, A., H. Kitazato, R. Iwase and H. Momma (1998): Seasonally appeared dense clouds of suspended particles at benthic boundary layer which were recorded in video images at the Real-time Deep-sea floor Observatory off Hatsushima in Sagami Bay. JAMSTEC J. Deep-Sea Res., 14, 319–327.

    Google Scholar 

  • Young, C. M., P. A. Tyler, J. L. Cameron and S. G. Rumrill (1992): Seasonal breeding aggregations in low-density populations of a bathyal echinoid, Stylocidaris lineata. Mar. Biol., 113, 603–612.

    Google Scholar 

  • Zezina, O. N. (1975): On some deep-sea brachiopods from the Gay Head-Bermuda transect. Deep-Sea Res., 22, 903–912.

    Google Scholar 

Download references

Author information

Authors and Affiliations

  1. Empress Dock, European Way, Southampton Oceanography Centre, Southampton, SO14 3ZH, U.K.

    Andrew J. Gooday

Authors
  1. Andrew J. Gooday
    View author publications

    You can also search for this author in PubMed Google Scholar

Corresponding author

Correspondence to Andrew J. Gooday.

Rights and permissions

Reprints and permissions

About this article

Cite this article

Gooday, A.J. Biological Responses to Seasonally Varying Fluxes of Organic Matter to the Ocean Floor: A Review. Journal of Oceanography 58, 305–332 (2002). https://doi.org/10.1023/A:1015865826379

Download citation

  • Issue Date:

  • DOI: https://doi.org/10.1023/A:1015865826379

Search

Navigation