Jeremy M. Testa

The metabolism of aquatic ecosystems: history, applications, and future challenges

Measurements of the production and consumption of organic material have been a focus of aquatic science for more than 80xa0years. Over the last century, a variety of approaches have been developed and employed for measuring rates of gross primary production (Pg), respiration (R), and net ecosystem production (Pnxa0=xa0Pgxa0−xa0R) within aquatic ecosystems. Here, we reconsider the range of approaches and applications for ecosystem metabolism measurements, and suggest ways by which such studies can continue to contribute to aquatic ecology. This paper reviews past and contemporary studies of aquatic ecosystem-level metabolism to identify their role in understanding and managing aquatic systems. We identify four broad research objectives that have motivated ecosystem metabolism studies: (1) quantifying magnitude and variability of metabolic rates for cross-system comparison, (2) estimating organic matter transfer between adjacent systems or subsystems, (3) measuring ecosystem-scale responses to perturbation, both natural and anthropogenic, and (4) quantifying and calibrating models of biogeochemical processes and trophic networks. The magnitudes of whole-system gross primary production, respiration and net ecosystem production rates vary among aquatic environments and are partly constrained by the chosen methodology. We argue that measurements of ecosystem metabolism should be a vital component of routine monitoring at larger scales in the aquatic environment using existing flexible, precise, and durable sensor technologies. Current and future aquatic ecosystem studies will benefit from application of new methods for metabolism measurements, which facilitate integration of process measurements and calibration of models for addressing fundamental questions involving ecosystem-scale processes.

Spatial and Temporal Patterns of Winter–Spring Oxygen Depletion in Chesapeake Bay Bottom Water

Although seasonal hypoxia is a well-studied phenomenon in many coastal systems, most previous studies have only focused on variability and controls on low-oxygen water masses during warm months when hypoxia is most extensive. Surprisingly, little attention has been given to investigations of what controls the development of hypoxic water in the months leading up to seasonal oxygen minima in temperate ecosystems. Thus, we investigated aspects of winter–spring oxygen depletion using a 25-year time series (1985–2009) by computing rates of water column O2 depletion and the timing of hypoxia onset for bottom waters of Chesapeake Bay. On average, hypoxia (O2 <62.5xa0μM) initiated in the northernmost region of the deep, central channel in early May and extended southward over ensuing months; however, the range of hypoxia onset dates spanned >50xa0days (April 6 to May 31 in the upper Bay). O2 depletion rates were consistently highest in the upper Bay, and elevated Susquehanna River flow resulted in more rapid O2 depletion and earlier hypoxia onset. Winter–spring chlorophyll a concentration in the bottom water was highly correlated with interannual variability in hypoxia onset dates and water column O2 depletion rates in the upper and middle Bay, while stratification strength was a more significant driver in the timing of lower Bay hypoxia onset. Hypoxia started earlier in 2012 (April 6) than previously recorded, which may be related to unique climatic and biological conditions in the winter–spring of 2012, including the potential carryover of organic matter delivered to the system during a tropical storm in September 2011. In general, mid-to-late summer hypoxic volumes were not correlated to winter–spring O2 depletion rates and onset, suggesting that the maintenance of summer hypoxia is controlled more by summer algal production and physical forcing than winter-spring processes. This study provides a novel synthesis of O2 depletion rates and hypoxia onset dates for Chesapeake Bay, revealing controls on the phenology of hypoxia development in this estuary.

What drives interannual variability of hypoxia in Chesapeake Bay: Climate forcing versus nutrient loading?

Oxygen depletion in estuaries is a worldwide problem with detrimental effects on many organisms. Although nutrient loading has been stabilized for a number of these systems, seasonal hypoxia persists and displays large year-to-year variations, with larger hypoxic volumes in wetter years and smaller hypoxic volumes in drier years. Data analysis points to climate as a driver of interannual hypoxia variability, but nutrient inputs covary with freshwater flow. Here we report an oxygen budget analysis of Chesapeake Bay to quantify relative contributions of physical and biogeochemical processes. Vertical diffusive flux declines with river discharge, whereas longitudinal advective flux increases with river discharge, such that their total supply of oxygen to bottom water is relatively unchanged. However, water column respiration exhibits large interannual fluctuations and is correlated with primary production and hypoxic volume. Hence, the model results suggest that nutrient loading is the main mechanism driving interannual hypoxia variability in Chesapeake Bay.

Progress and Challenges in Coupled Hydrodynamic-Ecological Estuarine Modeling

Numerical modeling has emerged over the last several decades as a widely accepted tool for investigations in environmental sciences. In estuarine research, hydrodynamic and ecological models have moved along parallel tracks with regard to complexity, refinement, computational power, and incorporation of uncertainty. Coupled hydrodynamic-ecological models have been used to assess ecosystem processes and interactions, simulate future scenarios, and evaluate remedial actions in response to eutrophication, habitat loss, and freshwater diversion. The need to couple hydrodynamic and ecological models to address research and management questions is clear because dynamic feedbacks between biotic and physical processes are critical interactions within ecosystems. In this review, we present historical and modern perspectives on estuarine hydrodynamic and ecological modeling, consider model limitations, and address aspects of model linkage, skill assessment, and complexity. We discuss the balance between spatial and temporal resolution and present examples using different spatiotemporal scales. Finally, we recommend future lines of inquiry, approaches to balance complexity and uncertainty, and model transparency and utility. It is idealistic to think we can pursue a “theory of everything” for estuarine models, but recent advances suggest that models for both scientific investigations and management applications will continue to improve in terms of realism, precision, and accuracy.

Decadal Changes in Water Quality and Net Productivity of a Shallow Danish Estuary Following Significant Nutrient Reductions

We utilized an extensive data set (1977–2013) from a water quality monitoring program to investigate the recovery of a Danish estuary following large reductions in total phosphorus (TP) and total nitrogen (TN) loading. Monthly rates of net transport and biogeochemical transformation of dissolved inorganic nitrogen (DIN) and phosphorus (DIP) were computed in two basins of the estuary using a box model approach, and oxygen-based rates of net ecosystem production (NEP) were determined. Since 1990, nutrient loading was reduced by 58xa0% for nitrogen and 80xa0% for phosphorus, causing significant decreases in DIN (60xa0%) and DIP (85xa0%) concentrations. Reductions in nutrient loadings and concentrations reduced annual chlorophyll levels by 50xa0% in the inner estuary and improved Secchi depth by approximately 1xa0m during the same period, particularly in the summer period. In the outer, deeper region of the estuary trends in water quality was less evident. Improvements in the inner estuary were strongly coupled to declines in DIN. Thresholds of DIN and DIP concentrations limiting phytoplankton growth indicated that both regions of the estuary were nitrogen limited. NEP rates indicated the development of more net autotrophic conditions over time that were likely associated with higher benthic primary production stimulated by improved light conditions. Box model computations revealed a modest reduction in summer net production of DIP over time, despite the persistence of elevated fluxes for several years after external loads were reduced. Since the mid-1990s, nutrient loading and transformation were stable while nutrient concentrations continued to decline and water quality improved in the inner estuary. The oligotrophication trajectory involved an initial fast transformation and modest retention of nutrients followed by a gradual decline in the rate of improvement towards a new stable condition.

Redox reactions and weak buffering capacity lead to acidification in the Chesapeake Bay

The combined effects of anthropogenic and biological CO2 inputs may lead to more rapid acidification in coastal waters compared to the open ocean. It is less clear, however, how redox reactions would contribute to acidification. Here we report estuarine acidification dynamics based on oxygen, hydrogen sulfide (H2S), pH, dissolved inorganic carbon and total alkalinity data from the Chesapeake Bay, where anthropogenic nutrient inputs have led to eutrophication, hypoxia and anoxia, and low pH. We show that a pH minimum occurs in mid-depths where acids are generated as a result of H2S oxidation in waters mixed upward from the anoxic depths. Our analyses also suggest a large synergistic effect from river–ocean mixing, global and local atmospheric CO2 uptake, and CO2 and acid production from respiration and other redox reactions. Together they lead to a poor acid buffering capacity, severe acidification and increased carbonate mineral dissolution in the USA’s largest estuary.The potential contribution of redox reactions to acidification in coastal waters is unclear. Here, using measurements from the Chesapeake Bay, the authors show that pH minimum occurs at mid-depths where acids are produced via hydrogen sulfide oxidation in waters mixed upward from anoxic depths.

Challenges and Directions for the Advancement of Estuarine Ecosystem Science

Estuarine ecosystem ecology is a dynamic field of study that has historically focused on a spectrum of compelling research topics, and here we present a series of perspectives on the major challenges to be overcome and key research questions to be addressed toward making progress over the coming decades. The challenges we identify include (1) maintaining and improving spatially distributed time-series datasets, (2) maximizing innovation by harnessing new technologies, (3) resuscitating experimental ecosystem research for estuaries, (4) integrating diagnostic ecological models into ecosystem research, and (5) improving basic science by linking it to applied research. We also raise a number of key research questions for the field, including (1) how does food web function respond to changing climate and nutrients, (2) what are likely trajectories of ecosystem recovery in response to restoration, (3) how does climate alter seasonality of estuarine ecosystem processes, (4) how do estuaries affect the global carbon budget and what are key feedbacks, and (5) how will tidal wetland ecosystems respond to sea level rise and climate change? Looking ahead, we envision that the field of estuarine ecosystem ecology will continue to build upon its rich tradition to address fundamental research questions with an expanded toolkit and enlightened perspective to focus basic science on the knowledge needs of society.

Submersed aquatic vegetation in Chesapeake Bay: Sentinel species in a changing world

Abstract Chesapeake Bay has undergone profound changes since European settlement. Increases in human and livestock populations, associated changes in land use, increases in nutrient loadings, shoreline armoring, and depletion of fish stocks have altered the important habitats within the Bay. Submersed aquatic vegetation (SAV) is a critical foundational habitat and provides numerous benefits and services to society. In Chesapeake Bay, SAV species are also indicators of environmental change because of their sensitivity to water quality and shoreline development. As such, SAV has been deeply integrated into regional regulations and annual assessments of management outcomes, restoration efforts, the scientific literature, and popular media coverage. Even so, SAV in Chesapeake Bay faces many historical and emerging challenges. The future of Chesapeake Bay is indicated by and contingent on the success of SAV. Its persistence will require continued action, coupled with new practices, to promote a healthy and sustainable ecosystem.

Long-term nutrient reductions lead to the unprecedented recovery of a temperate coastal region

Significance Human actions, including nutrient pollution, are causing the widespread degradation of coastal habitats, and efforts to restore these valuable ecosystems have been largely unsuccessful or of limited scope. We provide an example of successful restoration linking effective management of nutrients to the successful recovery of submersed aquatic vegetation along thousands of kilometers of coastline in Chesapeake Bay, United States. We also show that biodiversity conservation can be an effective path toward recovery of coastal systems. Our study validates 30 years of environmental policy and provides a road map for future ecological restoration. Humans strongly impact the dynamics of coastal systems, yet surprisingly few studies mechanistically link management of anthropogenic stressors and successful restoration of nearshore habitats over large spatial and temporal scales. Such examples are sorely needed to ensure the success of ecosystem restoration efforts worldwide. Here, we unite 30 consecutive years of watershed modeling, biogeochemical data, and comprehensive aerial surveys of Chesapeake Bay, United States to quantify the cascading effects of anthropogenic impacts on submersed aquatic vegetation (SAV), an ecologically and economically valuable habitat. We employ structural equation models to link land use change to higher nutrient loads, which in turn reduce SAV cover through multiple, independent pathways. We also show through our models that high biodiversity of SAV consistently promotes cover, an unexpected finding that corroborates emerging evidence from other terrestrial and marine systems. Due to sustained management actions that have reduced nitrogen concentrations in Chesapeake Bay by 23% since 1984, SAV has regained 17,000 ha to achieve its highest cover in almost half a century. Our study empirically demonstrates that nutrient reductions and biodiversity conservation are effective strategies to aid the successful recovery of degraded systems at regional scales, a finding which is highly relevant to the utility of environmental management programs worldwide.

Modeling the influence of deep water application of dispersants on the surface expression of oil: A sensitivity study

Although the effects of chemical dispersants on oil droplet sizes and ascent speeds are well-known, the fate and transport of dispersed oil droplets of different sizes under varying hydrodynamic conditions can be difficult to assess with observations alone. We used a particle tracking model to evaluate the effect of changes in droplet sizes due to dispersant application on the short-term transport and surface expression of oil released under conditions similar to those following the 3 June 2010 riser cutting during the Deepwater Horizon event. We used simulated injections of oil droplets of varying size and number under conditions associated with no dispersant application and with dispersant application at 50% and 100% efficiency. Due to larger droplet sizes in the no-dispersant scenario, all of the simulated oil reached the surface within 7 h, while only 61% and 28% of the oil reached the surface after 12 h in the 50% and 100% dispersant efficiency cases, respectively. The length of the surface slick after 6 h was 2 km in the no-dispersant case whereas there was no surface slick after 6 h in the 100% dispersant case, because the smaller oil droplets which resulted from dispersant application had not yet reached the surface. Model results suggest that the application of dispersants at the well head had the following effects: (1) less oil reached the surface in the 6-12 h after application, (2) oil had a longer residence time in the water-column, and (3) oil was more highly influenced by subsurface transport.