Biological Oceanography

(ff) #1

quantifying oxygen concentration most often now are μmol kg−1 seawater; in an era


not very long back they were ml O2, (^) STP liter−1, referring to the gas volume at
“standard temperature and pressure” of 0°C and 1 atmosphere. Both units (and also
mass units, parts/million, partial pressures, ... ) appear in useful literature. Oxygen
concentration can be measured with good precision and to low values by the Winkler
titration, a series of redox reactions that are relatively simple to carry through to a
sharp end-point. It can also be measured with Clark electrodes and recently with
optodes, in which luminescence from molecules embedded in an oxygen-permeable
plastic is variably quenched depending upon adjacent oxygen concentration, and is
measured by a light meter. Optodes have fairly long response times (many seconds),
depending upon the permeability of the plastic, but the non-linear calibrations are
stable over long periods and can cover the seawater concentration range.
(^) If oxygen-depleted water is brought into the surface mixing layer and churns against
the interface with the atmosphere, its oxygen content will rise toward saturation.
However, the equilibration occurs relatively slowly, mostly because the mixing layer
can be thick, but also because the oxygen itself has no particularly great “drive”
around reaching saturation. The subjects of gas exchange coefficients and “piston
velocities” are left to texts on chemical oceanography (e.g. Pilson 1998). Despite the
slow oxygen-exchange rate, most deep water is “formed” at high latitudes where the
water is very cold. Thus, the initial charge of oxygen for much of the subsurface
ocean is close to the coldest possible saturation level.
(^) Once oxygen is in solution, consumption by respiration of everything from bacteria
to tiger sharks begins. Not all life depends upon oxygen (there are anaerobic
microbes), but larger organisms with high activity levels and high metabolic rates
depend upon oxidative respiration, a function primarily carried out in eukaryotic
organisms by mitochondria – cell organelles specialized for this function. Down in the
stratified layers well below vertical mixing from the surface, oxygen used is only
replaced by the horizontal flow of water that left the surface some distance, often a
very great distance, away. Thus, as organic matter sinks into these layers and is eaten
and respired, the oxygen levels decrease. Additional decrease comes from animals
that move between feeding near the surface and resting or hiding at depth, where
those fish, squid, and plankton continue to use oxygen. Depletion generates oxygen-
minimum zones in intermediate levels of the water column, particularly in the
Arabian Sea and Pacific and most reduced from saturation in the Pacific toward the
north (Plate 1.1).
(^) There are also some nearly or completely anoxic layers beneath upwelling areas,
particularly parts of the Peru Current and the Arabian Gulf. In the latter sites,
dissolved oxygen as O 2 can be completely exhausted. Microbes in the anoxic layers
continue to respire, at least to a point, using the oxygen atoms in nitrate and sulfate

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