Jeffrey C. Cornwell

Influence of Eastern Oysters on Nitrogen and Phosphorus Regeneration in Chesapeake Bay, USA

Suspension-feeding bivalves couple pelagic and benthic processes because they consume seston from the water column, and their biodeposits (feces and pseudofeces) settle on the sediment surface. Abundant stocks of bivalves can exert grazer control on phytoplankton, and this results in some nitrogen and phosphorus being regenerated to the water column as excreta and via microbial decomposition of biodeposits. Bivalve biodeposition, however, enhances net ecosystem losses of N and P via sediment burial and bacterially mediated, coupled nitrification-denitrification. Bivalve feeding also reduces turbidity and thereby increases light available for microphytobenthos. Although microphytobenthos may compete with nitrifying bacteria for N, potentially reducing coupled nitrification-denitrification, they retain N and P within sediments, further reducing net regeneration to the water column.

Increased sediment accretion rates following invasion byPhragmites australis: The role of litter

Negative connotations of invasive plants worldwide have implicated them as the bearers of unfavorable ecosystem change. We contrasted 5-yr-old and 20-yr-oldPhragmites populations with pre-invasion areas occupied byTypha spp. andPanicum virgatum in an oligohaline tidal marsh of Chesapeake Bay. Peak live biomass was 3 times greater, while standing dead and litter was twice as great in the 20-yr-oldPhragmites. It is this abundance of concentrated litter on the marsh surface of maturePhragmites populations that we implicate as encouraging the trapping of organic and mineral matter. The rate of vertical accretion in 20-yr-oldPhragmites populations is 3–4 mm yr−1 above the adjacent populations. By integrating the constant initial concentration and constant rate of supply models on individual210Pb cores, we estimate thatPhragmites populations require a minimum of 7-yr post-colonization to enhance rates of accretion in this system. In ligh of the considerable loss of marsh habitat from relative sea-level rise, this finding contests the view that invasion creates strictly undesirable change at the ecosystem level.

Ecological Stoichiometry, Biogeochemical Cycling, Invasive Species, and Aquatic Food Webs: San Francisco Estuary and Comparative Systems

Eutrophication has altered food webs across aquatic systems, but effects of nutrient stoichiometry (varying nutrient ratios) on ecosystem structure and function have received less attention. A prevailing assumption has been that nutrients are not ecologically relevant unless concentrations are limiting to phytoplankton. However, changes in nutrient stoichiometry fundamentally affect food quality at all levels of the food web. Here, 30-year records of nitrogen and phosphorus concentrations and ratios, phytoplankton, zooplankton, macroinvertebrates, and fish in the San Francisco Estuary (Bay Delta) were examined to collectively interpret ecosystem changes within the framework of ecological stoichiometry. Changes in nutrient concentrations and nutrient ratios over time fundamentally affect biogeochemical nutrient dynamics that can lead to conditions conducive to invasions of rooted macrophytes and bivalve molluscs, and the harmful cyanobacterium Microcystis. Several other aquatic ecosystems considered here have exhibited similar changes in food webs linked to stoichiometric changes. Nutrient stoichiometry is thus suggested to be a significant driver of food webs in the Bay Delta by altering food quality and biogeochemical dynamics. Since nitrogen-to-phosphorus ratios have increased over time, an overall implication is that remediation of fish populations in the San Francisco Estuary will require significant nitrogen reductions to restore the historic ecological stoichiometric balance and the food web.

Respiratory succession and community succession of bacterioplankton in seasonally anoxic estuarine waters.

ABSTRACT Anoxia occurs in bottom waters of stratified estuaries when respiratory consumption of oxygen, primarily by bacteria, outpaces atmospheric and photosynthetic reoxygenation. Once water becomes anoxic, bacterioplankton must change their metabolism to some form of anaerobic respiration. Analysis of redox chemistry in water samples spanning the oxycline of Chesapeake Bay during the summer of 2004 suggested that there was a succession of respiratory metabolism following the loss of oxygen. Bacterial community doubling time, calculated from bacterial abundance (direct counts) and production (anaerobic leucine incorporation), ranged from 0.36 to 0.75 day and was always much shorter than estimates of the time that the bottom water was anoxic (18 to 44 days), indicating that there was adequate time for bacterial community composition to shift in response to changing redox conditions. However, community composition (as determined by PCR-denaturing gradient gel electrophoresis analysis of 16S rRNA genes) in anoxic waters was very similar to that in surface waters in June when nitrate respiration was apparent in the water column and only partially shifted away from the composition of the surface community after nitrate was depleted. Anoxic water communities did not change dramatically until August, when sulfate respiration appeared to dominate. Surface water populations that remained dominant in anoxic waters were Synechococcus sp., Gammaproteobacteria in the SAR86 clade, and Alphaproteobacteria relatives of Pelagibacter ubique, including a putative estuarine-specific Pelagibacter cluster. Populations that developed in anoxic water were most similar (<92% similarity) to uncultivated Firmicutes, uncultivated Bacteroidetes, Gammaproteobacteria in the genus Thioalcalovibrio, and the uncultivated SAR406 cluster. These results indicate that typical estuarine bacterioplankton switch to anaerobic metabolism under anoxic conditions but are ultimately replaced by different organisms under sulfidic conditions.

The Role of Oligohaline Marshes in Estuarine Nutrient Cycling

Oligohaline marshes, poised at the land-sea margin, often occur where the estuary is most enriched in inorganic particles and nutrients. Although light can limit the production of planktonic communities, high nutrient concentrations and regular tidal inundation results in highly productive macrophyte and algal communities. Despite potentially important water quality values, relatively few detailed studies of N and P cycling in oligohaline marshes are evident in the literature. Because of the temporal variability in marsh flux studies, the net annual retention of N and P is best assessed by measurement of N and P burial in the sediment. In the Chesapeake Bay and other estuaries and subestuaries, high rates of tidal marsh N and P burial indicate an important water quality function. A recent study shows the marshes of a Chesapeake Bay tributary retain a large portion of nitrogen and phosphorus entering the river from above the fall line. The marshes trap 35% of the nitrogen and 81% of the phosphorus which would otherwise be recycled, exported, or buried in the subtidal sediments of the estuary. Although there are few studies, high nitrate supply rates, potentially high nitrification rates, and high rates of sediment metabolism can result in high rates of denitrification. More complete studies of tidal marsh nutrient cycling, particularly nitrogen cycling, are needed for a better understanding of the importance of these tidal freshwater marshes to estuarine nutrient balances. Alternative methodologies for denitrification measurement are needed for more accurate measurements, and more attention needs to be paid to scaling individual measurements to whole marsh ecosystems. A new method for the measurement of net N2(g) exchange was applied to a Chesapeake Bay tributary to develop an annual estimate of net denitrification in the marsh sediments. Denitrification rates were ∼ 60 μmol m−2 h−1 with high seasonal variability. Annual calculations were made based on a loose correlation to annual ambient nitrate concentrations. This preliminary calculation suggests that an additional 10% of the fall line nitrogen may be removed by such marsh systems. More measurements of net N2(g) exchange and computer simulation models are required to determine the net removal of fall line nitrogen by the upper estuarine marshes.

Influence of Plant Communities on Denitrification in a Tidal Freshwater Marsh of the Potomac River, United States

We investigated whether marsh surface elevation, plant community composition (annuals vs. perennials), and organic matter quantity/quality were associated with differences in denitrification rates in an urban tidal freshwater marsh of the Potomac River, United States. We measured denitrification rates using both denitrification enzyme activity (DEA) with acetylene inhibition (June: n = 38, 3234 +/- 303; October: n = 38, 1557 +/- 368 ng N g dry soil(-1) h(-1)) and direct N(2) flux measurements with membrane inlet mass spectrometry (MIMS) (November: n = 6, 147 +/- 24 mumol m(-2) h(-1)). Organic carbon content and nitrate concentrations in soil, and plant community composition were correlated with elevation, but DEA rates did not differ across marsh surface elevation. Soil organic carbon was highest in plots dominated by perennial graminoids, but DEA rates did not differ across plant community types. The DEA rates increased with increasing soil ammonium concentrations and total N content, and DEA rates differed between summer and fall sampling. The MIMS rates did not differ across plant community types, but were correlated with soil organic N content. Denitrification rates suggest that potential N removal at the site could be substantial. In addition, denitrification rates measured in Dyke Marsh were higher than rates for sediments measured in the adjacent Potomac River. Tidal freshwater marshes can represent an important site for denitrification, and factors fostering denitrification should be considered when restoring urban tidal freshwater wetlands as they are faced with pressures from increasing land use change and sea level rise.

Determination of denitrification in the chesapeake bay from measurements of N2 accumulation in bottom water

This study demonstrates the feasibility of using direct N2 measurements in an estuary for determination of denitrification. High precision measurements of dinitrogen: argon ratios (N2∶Ar) were made by membrane inlet mass spectrometry on water samples taken along the length of the Chesapeake Bay in July and October 2004. The N2∶Ar ratio in low salinity surface water was elevated relative to air saturation by 0.3–0.5% with no systematic change along the length of the Bay. N2∶Ar in high salinity bottom water exhibited a linear increase in the landward direction along a 144-km longitudinal section. In this section of the Bay covering 20% of the main stem, the bottom water salinity was statistically uniform and the increase in N2∶Ar was in the direction of net residual current flow. The system was analyzed as a capped river with the assumption that N2 entered the water from the underlying sediment where denitrification is known to take place. The rate of denitrification needed to support the measured increase in N2 was calculated using an average residual current velocity and water column depth. The increase in N2 with distance (0.046μmol N l−1 km−1) equated to an average denitrification flux of 73 μmol N m−2 h−1. N2 fluxes determined on sediment cores taken from the source and terminus regions of the delineated water mass were 45±23 and 83±39 μmol N m−2 hr−1, respectively, which were not statistically different from the whole system estimate. The measured change in oxygen concentration within the bottom water was used to estimate nitrogen remineralization and the efficiency of denitrification. Denitrification efficiency (nitrogen denitrified/nitrogen remineralized) was estimated to be in the range of 22–28% for the bottom water sediment system and 30–37% considering the sediment zone alone.

Transitions in nirS-type denitrifier diversity, community composition, and biogeochemical activity along the Chesapeake Bay estuary

Chesapeake Bay, the largest estuary in North America, can be characterized as having steep and opposing gradients in salinity and dissolved inorganic nitrogen along the main axis of the Bay. In this study, the diversity of nirS gene fragments (encoding cytochrome cd1-type nitrite reductase), physical/chemical parameters, and benthic N2-fluxes were analyzed in order to determine how denitrifier communities and biogeochemical activity vary along the estuary salinity gradient. The nirS gene fragments were PCR-amplified, cloned, and sequenced from sediment cores collected at five stations. Sequence analysis of 96–123 nirS clones from each station revealed extensive overall diversity in this estuary, as well as distinct spatial structure in the nirS sequence distributions. Both nirS-based richness and community composition varied among stations, with the most dramatic shifts occurring between low-salinity (oligohaline) and moderate-salinity (mesohaline) sites. For four samples collected in April, the nirS-based richness, nitrate concentrations, and N2-fluxes all decreased in parallel along the salinity gradient from the oligohaline northernmost station to the highest salinity (polyhaline) station near the mouth of the Bay. The vast majority of the 550 nirS sequences were distinct from cultivated denitrifiers, although many were closely related to environmental clones from other coastal and estuarine systems. Interestingly, 8 of the 172 OTUs identified accounted for 42% of the total nirS clones, implying the presence of a few dominant and many rare genotypes, which were distributed in a non-random manner along the salinity gradient of Chesapeake Bay. These data, comprising the largest dataset to investigate nirS clone sequence diversity from an estuarine environment, also provided information that was required for the development of nirS microarrays to investigate the interaction of microbial diversity, environmental gradients, and biogeochemical activity.

Microtopography in Tidal Marshes : Ecosystem Engineering by Vegetation?

Hummock-hollow microtopography is characteristic of many freshwater wetland systems. It is comprised of elevated, vegetated hummocks and lower elevation hollows; the latter are usually unvegetated, with reducing conditions in sediments unfavorable for plant growth. This microtopography is also often found in interior regions of brackish marshes, where flood duration is high and salinity fluctuations are prominent. Previous investigation showed this spatial patterning to be relatively stable over time and suggested that these microenvironments are produced by the plants themselves. This study investigates the possible mechanisms and controlling factors of this microtopography and considers the effect of different salinity regimes. We examined microtopographic variability of vegetation and sediment biogeochemistry in two interior tidal marshes, a freshwater-oligohaline marsh and a mesohaline marsh, both of which exhibited fine-scale spatial variability. Within a 2-yr period, the freshwater-oligohaline site demonstrated a labile response of both vegetation and sediment chemistry to interannual variability in salinity and sulfide concentrations, whereas the microscale spatial variability of the mesohaline system persisted. Geochronological assessment of the mesohaline marsh, where microtopographic variability was relatively stable, supported the hypothesis that the formation of the hummock-hollow topography is driven by the plants, rather than developing as a result of underlying physical variability. We propose that brackish marsh vegetation alters the sedimentary environment in such a way as to maximize growth under high-stress, variable conditions. The adaptive advantage of this strategy was illustrated in the accretion rates measured at the higher salinity marsh, which were indistinguishable between the interior hummock sediments and those of an adjacent homogeneous bank marsh.

Metatranscriptomic analyses of plankton communities inhabiting surface and subpycnocline waters of the Chesapeake Bay during oxic-anoxic-oxic transitions.

ABSTRACT We used metatranscriptomics to study the gene transcription patterns of microbial plankton (0.2 to 64 μm) at a mesohaline station in the Chesapeake Bay under transitions from oxic to anoxic waters in spring and from anoxic to oxic waters in autumn. Samples were collected from surface (i.e., above pycnocline) waters (3 m) and from waters beneath the pycnocline (16 to 22 m) in both 2010 and 2011. Metatranscriptome profiles based on function and potential phylogeny were different between 2010 and 2011 and strongly variable in 2011. This difference in variability corresponded with a highly variable ratio of eukaryotic to bacterial sequences (0.3 to 5.5), reflecting transient algal blooms in 2011 that were absent in 2010. The similarity between metatranscriptomes changed at a lower rate during the transition from oxic to anoxic waters than after the return to oxic conditions. Transcripts related to photosynthesis and low-affinity cytochrome oxidases were significantly higher in shallow than in deep waters, while in deep water genes involved in anaerobic metabolism, particularly sulfate reduction, succinyl coenzyme A (succinyl-CoA)-to-propionyl-CoA conversion, and menaquinone synthesis, were enriched relative to in shallow waters. Expected transitions in metabolism between oxic and anoxic deep waters were reflected in elevated levels of anaerobic respiratory reductases and utilization of propenediol and acetoin. The percentage of archaeal transcripts increased in both years in late summer (from 0.1 to 4.4% of all transcripts in 2010 and from 0.1 to 6.2% in 2011). Denitrification-related genes were expressed in a predicted pattern during the oxic-anoxic transition. Overall, our data suggest that Chesapeake Bay microbial assemblages express gene suites differently in shallow and deep waters and that differences in deep waters reflect variable redox states.