FOOD INPUTS, UTILIZATION, CARBON FLOW AND ENERGETICS
367
flux in the total estimate of net accumulation on the
bottom.
Epilogue
All these uncertainties, along with the need to model
a highly non-linear system, may prompt despair in
scaling models from particular sites up to the level of
the regional, or even global-scale ecosystem. As an
example of what might be possible, a comprehensive
modelling approach using a mass-balanced ecosystem
modelling package, Ecopath©, has been useful in
developing a model of trophic energy flow in the
Antarctic shelf ecosystem in the Weddell Sea (Schalk
et al., 1993; Jarre-Teichmann et al., 1996).
Clearly projection of variables to the larger scale will
involve large uncertanties. The studies reported above
at single sites represent huge effort. Their budgetary
deficiencies and the evidence for long-term changes
(e.g., K.L. Smith and Kaufmann, 1999) reveal how little
is really known about biological processes in the deep
sea and about variability in coupling to processes in
the upper ocean. But, despite their limitations, models
assembled on such data as are currently available have
proved at the very least useful and informative on what
is, or is not, important, and also on what parts of the
system are not being addressed.
ACKNOWLEDGEMENTS
The following have very kindly read parts of this
chapter and offered useful comments and criticism: Dr
Laurenz Thomsen, GEOMAR, Germany; Dr Thomas
Brey, Alfred Wegener Institute, Bremerhaven, Germany, and Dr Richard Lampitt, Southampton Oceanography Centre, Southampton, U.K.
REFERENCES
Agassiz, A., 1888. Three Cruises of the United States Coast and
Geodetic Survey Steamer ‘Blake’, 2 Vols. Sampson Low, Marston,
Searle and Rivington, London, 220 pp. and 314 pp.
Aldred, R.G., Riemann-Z¨ urneck, K., Thiel, H. and Rice, A.L., 1979.
Ecological observations on the deep-sea anemone Actinoscyphia
aurelia. Oceanol. Acta, 2: 389−395.
Alldredge, A.L. and Gotschalk, C.C., 1989. Direct observations
of mass flocculations of diatom blooms: characteristics, settling
velocities and formation of diatom aggregates. Deep-Sea Res.,
36A: 159−173.
Alldredge, A.L. and McGillivary, P., 1991. The attachment
probabilities of marine snow and their implications for particle
coagulation in the ocean. Deep-Sea Res., 38A: 431−443.
Alldredge, A.L. and Silver, M.W., 1988. Characteristics, dynamics
and significance of marine snow. Prog. Oceanogr., 20: 41−82.
Allen, J.A., 1978. Evolution of the deep-sea protobranch bivalves.
Philos. Trans. R. Soc. London Ser. B, 241: 387−401.
Allen, J.A. and Sanders, H.L., 1966. Adaptations to abyssal life
as shown by the bivalve Abra profundorum. Deep-Sea Res., 13:
1175−1184.
Allen, J.A. and Sanders, H.L., 1973. Studies on deep-sea
Protobranchia (Bivalvia); the families Siliculidae and Lametilidae.
Bull. Mus. Comp. Zool., 145: 263−310.
Aller, J.Y. and Aller, R.C., 1986. Evidence for localized enhancement
of biological activity associated with tube and burrow structures in
deep-sea sediments at the HEBBLE site, western North Atlantic.
Deep-Sea Res., 33A: 755−790.
Aller, R.C., 1982. The effects of macrobenthos on chemical properties
of marine sediment and overlying water. In: P.L. McCall and
M.J.S. Tevesz (Editors), Animal–Sediment Relations: the Biogenic
Alteration of Sediments. Plenum Press, London, pp. 53–102.
Alongi, D.M., 1987. The description and composition of deep-sea
microbenthos in a bathyal region of the western Coral Sea. DeepSea Res., 34A: 1245−1254.
Alongi, D.M., 1990. Bacterial growth rates, production and estimates
of detrital carbon utilization in deep-sea sediments of the Solomon
and Coral Seas. Deep-Sea Res., 37A: 731−746.
Angel, M.V., 1984. Detrital organic fluxes through pelagic
ecosystems. In: M.J. Fasham (Editor), Flows of Energy and
Materials in Marine Ecosystems. Plenum, London, pp. 475–516.
Angel, M.V., 1989. Vertical profiles of pelagic communities in the
vicinity of the Azores Front and their implications to deep ocean
ecology. Prog. Oceanogr., 22: 1−46.
Angel, M.V., 1990. Life in the benthic boundary layer: connections to
the mid-water and sea floor. Philos. Trans. R. Soc. London Ser. A,
331: 15−28.
Angel, M.V. and Baker, A. de C., 1982. Vertical distribution of the
standing crop of plankton and micronekton at three stations in the
northeast Atlantic. Biol. Oceanogr., 2: 1−29.
Archer, D. and Devol, A., 1992. Benthic oxygen fluxes on the
Washington shelf and slope: a comparison of in situ microelectrode
and chamber flux measurements. Limnol. Oceanogr., 37:
614−629.
Arillo, A., Bavestrello, G., Burlando, B. and Sara, M., 1993.
Metabolic integration between symbiotic cyanobacteria and
sponges: a possible mechanism. Mar. Biol., 117: 159−162.
Armstrong, J.D., Bagley, P.M. and Priede, I.G., 1992. Photographic
and acoustic tracking observations of the behaviour of the grenadier
Coryphaenoides (Nematonurus) armatus, the eel Synaphobranchus
bathybius, and other abyssal demersal fish in the North Atlantic
Ocean. Mar. Biol., 112: 535−544.
Asper, V.L., Deuser, W.G., Knauer, G.A. and Lohrenz, S.E., 1992.
Rapid coupling of sinking particle fluxes between surface and deep
ocean waters. Nature, 357: 670−672.
Auffret, G.A., Khripounoff, A. and Vangriesheim, A., 1994. Rapid
post-bloom resuspension in the northeastern Atlantic. Deep-Sea
Res. I, 41: 925−939.
367
flux in the total estimate of net accumulation on the
bottom.
Epilogue
All these uncertainties, along with the need to model
a highly non-linear system, may prompt despair in
scaling models from particular sites up to the level of
the regional, or even global-scale ecosystem. As an
example of what might be possible, a comprehensive
modelling approach using a mass-balanced ecosystem
modelling package, Ecopath©, has been useful in
developing a model of trophic energy flow in the
Antarctic shelf ecosystem in the Weddell Sea (Schalk
et al., 1993; Jarre-Teichmann et al., 1996).
Clearly projection of variables to the larger scale will
involve large uncertanties. The studies reported above
at single sites represent huge effort. Their budgetary
deficiencies and the evidence for long-term changes
(e.g., K.L. Smith and Kaufmann, 1999) reveal how little
is really known about biological processes in the deep
sea and about variability in coupling to processes in
the upper ocean. But, despite their limitations, models
assembled on such data as are currently available have
proved at the very least useful and informative on what
is, or is not, important, and also on what parts of the
system are not being addressed.
ACKNOWLEDGEMENTS
The following have very kindly read parts of this
chapter and offered useful comments and criticism: Dr
Laurenz Thomsen, GEOMAR, Germany; Dr Thomas
Brey, Alfred Wegener Institute, Bremerhaven, Germany, and Dr Richard Lampitt, Southampton Oceanography Centre, Southampton, U.K.
REFERENCES
Agassiz, A., 1888. Three Cruises of the United States Coast and
Geodetic Survey Steamer ‘Blake’, 2 Vols. Sampson Low, Marston,
Searle and Rivington, London, 220 pp. and 314 pp.
Aldred, R.G., Riemann-Z¨ urneck, K., Thiel, H. and Rice, A.L., 1979.
Ecological observations on the deep-sea anemone Actinoscyphia
aurelia. Oceanol. Acta, 2: 389−395.
Alldredge, A.L. and Gotschalk, C.C., 1989. Direct observations
of mass flocculations of diatom blooms: characteristics, settling
velocities and formation of diatom aggregates. Deep-Sea Res.,
36A: 159−173.
Alldredge, A.L. and McGillivary, P., 1991. The attachment
probabilities of marine snow and their implications for particle
coagulation in the ocean. Deep-Sea Res., 38A: 431−443.
Alldredge, A.L. and Silver, M.W., 1988. Characteristics, dynamics
and significance of marine snow. Prog. Oceanogr., 20: 41−82.
Allen, J.A., 1978. Evolution of the deep-sea protobranch bivalves.
Philos. Trans. R. Soc. London Ser. B, 241: 387−401.
Allen, J.A. and Sanders, H.L., 1966. Adaptations to abyssal life
as shown by the bivalve Abra profundorum. Deep-Sea Res., 13:
1175−1184.
Allen, J.A. and Sanders, H.L., 1973. Studies on deep-sea
Protobranchia (Bivalvia); the families Siliculidae and Lametilidae.
Bull. Mus. Comp. Zool., 145: 263−310.
Aller, J.Y. and Aller, R.C., 1986. Evidence for localized enhancement
of biological activity associated with tube and burrow structures in
deep-sea sediments at the HEBBLE site, western North Atlantic.
Deep-Sea Res., 33A: 755−790.
Aller, R.C., 1982. The effects of macrobenthos on chemical properties
of marine sediment and overlying water. In: P.L. McCall and
M.J.S. Tevesz (Editors), Animal–Sediment Relations: the Biogenic
Alteration of Sediments. Plenum Press, London, pp. 53–102.
Alongi, D.M., 1987. The description and composition of deep-sea
microbenthos in a bathyal region of the western Coral Sea. DeepSea Res., 34A: 1245−1254.
Alongi, D.M., 1990. Bacterial growth rates, production and estimates
of detrital carbon utilization in deep-sea sediments of the Solomon
and Coral Seas. Deep-Sea Res., 37A: 731−746.
Angel, M.V., 1984. Detrital organic fluxes through pelagic
ecosystems. In: M.J. Fasham (Editor), Flows of Energy and
Materials in Marine Ecosystems. Plenum, London, pp. 475–516.
Angel, M.V., 1989. Vertical profiles of pelagic communities in the
vicinity of the Azores Front and their implications to deep ocean
ecology. Prog. Oceanogr., 22: 1−46.
Angel, M.V., 1990. Life in the benthic boundary layer: connections to
the mid-water and sea floor. Philos. Trans. R. Soc. London Ser. A,
331: 15−28.
Angel, M.V. and Baker, A. de C., 1982. Vertical distribution of the
standing crop of plankton and micronekton at three stations in the
northeast Atlantic. Biol. Oceanogr., 2: 1−29.
Archer, D. and Devol, A., 1992. Benthic oxygen fluxes on the
Washington shelf and slope: a comparison of in situ microelectrode
and chamber flux measurements. Limnol. Oceanogr., 37:
614−629.
Arillo, A., Bavestrello, G., Burlando, B. and Sara, M., 1993.
Metabolic integration between symbiotic cyanobacteria and
sponges: a possible mechanism. Mar. Biol., 117: 159−162.
Armstrong, J.D., Bagley, P.M. and Priede, I.G., 1992. Photographic
and acoustic tracking observations of the behaviour of the grenadier
Coryphaenoides (Nematonurus) armatus, the eel Synaphobranchus
bathybius, and other abyssal demersal fish in the North Atlantic
Ocean. Mar. Biol., 112: 535−544.
Asper, V.L., Deuser, W.G., Knauer, G.A. and Lohrenz, S.E., 1992.
Rapid coupling of sinking particle fluxes between surface and deep
ocean waters. Nature, 357: 670−672.
Auffret, G.A., Khripounoff, A. and Vangriesheim, A., 1994. Rapid
post-bloom resuspension in the northeastern Atlantic. Deep-Sea
Res. I, 41: 925−939.