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Direct measurements of the partial pressure of CO 2 (pCO 2 ) and dissolved inorganic carbon (DIC) were made over a 2-yr period in surface waters of the York River estuary in Virginia. The pCO 2 in surface waters exceeded that in the overlying atmosphere, indicating that the estuary was a net source of CO 2 to the atmosphere at most times and locations. Salinity-based DIC mixing curves indicate there was also an internal source of both DIC and alkalinity, implying net alkalinity generation within the estuary. The DIC and alkalinity source displayed seasonal patterns similar to that of pCO 2 and were reproducible over a 2-yr study period.We propose that the source of inorganic carbon necessary for both the sustained CO 2 evasion to the atmosphere and the advective export of DIC is respiration in excess of primary production (e.g., net heterotrophy). The rates of CO 2 evasion and DIC export were estimated to provide an annual rate of net heterotrophy of ϳ100 g C m Ϫ2 yrϪ1 . Approximately 40% of this excess inorganic carbon production was exported as DIC to the coastal ocean, whereas 60% was lost as CO 2 evasion to the atmosphere. The alkalinity generation needed to sustain the export of inorganic carbon, as HCO 3 Ϫ , is most likely provided by net sulfate reduction in sediments. Accumulation of sulfide in the sediments of a representative site directly adjacent to the York River estuary is sufficient to account for the net export of alkalinity. The seasonality of net heterotrophy causes large variations in annual CO 2 and DIC concentrations, and it stresses the need for comprehensive temporal data sets when reporting annual rates of CO 2 evasion, DIC advection, and net heterotrophy.
Direct measurements of the partial pressure of CO 2 (pCO 2 ) and dissolved inorganic carbon (DIC) were made over a 2-yr period in surface waters of the York River estuary in Virginia. The pCO 2 in surface waters exceeded that in the overlying atmosphere, indicating that the estuary was a net source of CO 2 to the atmosphere at most times and locations. Salinity-based DIC mixing curves indicate there was also an internal source of both DIC and alkalinity, implying net alkalinity generation within the estuary. The DIC and alkalinity source displayed seasonal patterns similar to that of pCO 2 and were reproducible over a 2-yr study period.We propose that the source of inorganic carbon necessary for both the sustained CO 2 evasion to the atmosphere and the advective export of DIC is respiration in excess of primary production (e.g., net heterotrophy). The rates of CO 2 evasion and DIC export were estimated to provide an annual rate of net heterotrophy of ϳ100 g C m Ϫ2 yrϪ1 . Approximately 40% of this excess inorganic carbon production was exported as DIC to the coastal ocean, whereas 60% was lost as CO 2 evasion to the atmosphere. The alkalinity generation needed to sustain the export of inorganic carbon, as HCO 3 Ϫ , is most likely provided by net sulfate reduction in sediments. Accumulation of sulfide in the sediments of a representative site directly adjacent to the York River estuary is sufficient to account for the net export of alkalinity. The seasonality of net heterotrophy causes large variations in annual CO 2 and DIC concentrations, and it stresses the need for comprehensive temporal data sets when reporting annual rates of CO 2 evasion, DIC advection, and net heterotrophy.
We measured dissolved organic carbon (DOC), dissolved inorganic carbon (DIC), and their corresponding ⌬ 14 C and ␦ 13C values in order to study the sources and fates of DOC in the York River Estuary (Virginia, U.S.A.). The ⌬ 14 C and ␦ 13C values of DOC and DIC at the freshwater end-member indicate that during periods of moderate to high flow, riverine DOC entering the York was composed of decadal-aged terrestrially organic matter. In nearly all cases, DOC concentrations exceeded conservative mixing lines and were therefore indicative of a net DOC input flux from within the estuary that averaged 1.. The nonconservative behavior of DOC in the York River Estuary was also apparent in carbon isotopic mixing curves and the application of an isotopic mixing model. The model predicted that 20-38% of the DOC at the mouth of the estuary was of riverine (terrestrial ϩ freshwater) origin, while 38-56% was added internally, depending on the isotopic values assigned to the internally added DOC. Measurements of ⌬ 14 C and ␦ 13 C of DOC and DIC and marsh organic matter suggest that the internal sources originated from estuarine phytoplankton and marshes. The isotopic mixing model also indicates a significant concomitant loss (27-45%) of riverine DOC within the estuary.Changes in DOC concentration, ⌬ 14 C-DOC, and ␦ 13 C-DOC were also measured during incubation experiments designed to quantify the amounts, sources, and ages of DOC supporting the carbon demands of estuarine bacteria. Results of these experiments were consistent with an estuarine source of phytoplankton and marsh DOC and the preferential utilization of young ( 14 C-enriched) DOC in the low-salinity reaches of the York. However, the average removal of riverine DOC by bacteria accounts for only ϳ4-19% of the riverine pool; therefore, other significant sinks for DOC exist within the estuary.
The cycling of dissolved inorganic carbon (DIC) and the role of tidal marshes in estuarine DIC dynamics were studied in a Virginia tidal freshwater marsh and adjacent estuary. DIC was measured over diurnal cycles in different seasons in a marsh tidal creek and at the junction of the creek with the adjacent Pamunkey River. In the creek, DIC concentrations around high tide were controlled by the same processes affecting whole-estuary DIC gradients. Near low tide, DIC concentrations were 1.5-5-fold enriched relative to high tide concentrations, indicating an input of DIC from the marsh. Similar patterns (although dampened in magnitude) were observed at the creek mouth and indicated that DIC was exported from the marsh. Marsh pore-water DIC concentrations were up to 5 mmol L Ϫ1 greater than those in the creek and suggested a significant input of sediment pore water to the creek. A model of tidal marsh DIC export showed that, on a seasonal basis, DIC export rates were influenced by water temperature. The composition of exported DIC averaged 19% dissolved CO 2 and 81% HCO and CO . Although CO 2 can belost to the atmosphere during transit through the estuary, DIC in the form of carbonate alkalinity is subject to export from the estuary to the coastal ocean. When extrapolated to an estuarywide scale, the export of marsh-derived DIC to the York River estuary explained a significant portion (47 Ϯ 23%) of excess DIC production (i.e., DIC in excess of that expected from conservative mixing between seawater and freshwater and equilibrium with the atmosphere) in this system. Therefore, CO 2 supersaturation, by itself, does not indicate that an estuary is net heterotrophic.One approach to understanding the cycling of organic carbon within ecosystems is through measurements of total system metabolism, given that the production and removal of organic matter are intimately linked to total dissolved inorganic carbon (DIC, or ⌺CO 2 ) and O 2 cycling. Recent studies of estuarine CO 2 and DIC dynamics have shown that estuaries are generally supersaturated with respect to CO 2 and exhibit high rates of net heterotrophy (i.e., respiration Ͼ photosynthesis; Smith and Hollibaugh 1993;Frankignoulle et al. 1998; Gatusso et al. 1998), although the main stem of Chesapeake Bay is net autotrophic (Kemp et al. 1997). Sources of CO 2 and DIC to estuarine waters include watercolumn and benthic respiration, riverine and groundwater inputs, photodegradation of dissolved organic matter, and inputs from intertidal marshes (Hopkinson and Vallino 1995;Kemp et al. 1997;Cai and Wang 1998). Accurate quantification of rates of net heterotrophy requires that one account 1 Corresponding author. Present address:
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