Sv. Smith et Jt. Hollibaugh, ANNUAL CYCLE AND INTERANNUAL VARIABILITY OF ECOSYSTEM METABOLISM IN ATEMPERATE CLIMATE EMBAYMENT, Ecological monographs, 67(4), 1997, pp. 509-533
We have studied the net and gross metabolism of Tomales Bay, a tempera
te climate estuary in northern California. Tomales Bay has proved to b
e heterotrophic, implying that the bay oxidizes a subsidy of organic c
arbon from outside the system, in excess of inorganic nutrients suppli
ed to it from outside and in addition to material cycling within it. N
et organic oxidation releases dissolved inorganic nutrients, and the s
ystem exports these dissolved inorganic products. Dissolved inorganic
phosphorus is exported to the ocean via mixing and constitutes the mos
t direct record of net ecosystem production (NEP). Excess dissolved in
organic nitrogen is lost to denitrification. Excess dissolved inorgani
c carbon largely results in alkalinity elevation and hydrographic expo
rt of alkalinity due to sulfate reduction. The negative NEP of this sy
stem results in little release of CO2 to the atmosphere, because of th
is alkalinity elevation. A major purpose of the study was to ascertain
the relative importance of various sources of organic material suppli
ed to the system from outside its boundaries and undergoing net reacti
ons within it. In order to address the question, we used stoichiometri
cally linked whole-system budgets of carbon, nitrogen, and phosphorus.
The difference between dissolved inorganic phosphorus (DIP) fluxes to
and from the bay is a measure of net internal sources or sinks of DIP
and is used as a quantitative index of NEP, with the assumption that
the C:P ratio of organic matter is constant (similar to 106:1). The sy
stem is thus defined in terms of water column dissolved material compo
sition; this definition includes time, as well as space. Net changes i
n the standing stocks of dissolved materials can originate from (spati
al) transport to or from the system or from internal (temporal) transf
ormations between the dissolved and particulate materials (i.e., chang
es in organic storage). Over the 8-yr study, the system respired 12 mm
ol.m(-2).d(-1) more organic C than the internal system primary product
ion of similar to 100 mmol.m(-2)d(-1). The system is thus heterotrophi
c by similar to 10%, with substantial seasonality in the extent of het
erotrophy. By deconvoluting the time series of NEP into a seasonal cyc
le and interannual variation, we infer that terrestrial and marine sou
rces each account for about half of the carbon required to support neg
ative NEP in this system, but with quite different turnover time scale
s. Temporal response of NEP to terrigenous input appears to be extreme
ly modulated, so that there is no obvious immediate (same year) respon
se to extreme interannual variation in terrigenous organic loading. In
contrast, NEP responds both interannually and seasonally to marine or
ganic inputs. We interpret the differences in response to loading of t
errestrial vs. marine organic matter as reflecting differences in the
reactivity of these carbon reservoirs.