Markus Huettel

Groundwater and pore water inputs to the coastal zone

Both terrestrial and marine forces drive underground fluid flows in the coastal zone. Hydraulic gradients on land result in groundwater seepage near shore and may contribute to flows further out on the shelf from confined aquifers. Marine processes such as tidal pumping and current-induced pressure gradients may induce interfacial fluid flow anywhere on the shelf where permeable sediments are present. The terrestrial and oceanic forces overlap spatially so measured fluid advection through coastal sediments may be a result of composite forcing. We thus define “submarine groundwater discharge” (SGD) as any and all flow of water on continental margins from the seabed to the coastal ocean, regardless of fluid composition or driving force. SGD is typically characterized by low specific flow rates that make detection and quantification difficult. However, because such flows occur over very large areas, the total flux is significant. Discharging fluids, whether derived from land or composed of re-circulated seawater, will react with sediment components. These reactions may increase substantially the concentrations of nutrients, carbon, and metals in the fluids. These fluids are thus a source of biogeochemically important constituents to the coastal ocean. Terrestrially-derived fluids represent a pathway for new material fluxes to the coastal zone. This may result in diffuse pollution in areas where contaminated groundwaters occur. This paper presents an historical context of SGD studies, defines the process in a form that is consistent with our current understanding of the driving forces as well as our assessment techniques, and reviews the estimated global fluxes and biogeochemical implications. We conclude that to fully characterize marine geochemical budgets, one must give due consideration to SGD. New methodologies, technologies, and modeling approaches are required to discriminate among the various forces that drive SGD and to evaluate these fluxes more precisely.

Hydrocarbon-Degrading Bacteria and the Bacterial Community Response in Gulf of Mexico Beach Sands Impacted by the Deepwater Horizon Oil Spill

ABSTRACT A significant portion of oil from the recent Deepwater Horizon (DH) oil spill in the Gulf of Mexico was transported to the shoreline, where it may have severe ecological and economic consequences. The objectives of this study were (i) to identify and characterize predominant oil-degrading taxa that may be used as model hydrocarbon degraders or as microbial indicators of contamination and (ii) to characterize the in situ response of indigenous bacterial communities to oil contamination in beach ecosystems. This study was conducted at municipal Pensacola Beach, FL, where chemical analysis revealed weathered oil petroleum hydrocarbon (C8 to C40) concentrations ranging from 3.1 to 4,500 mg kg−1 in beach sands. A total of 24 bacterial strains from 14 genera were isolated from oiled beach sands and confirmed as oil-degrading microorganisms. Isolated bacterial strains were primarily Gammaproteobacteria, including representatives of genera with known oil degraders (Alcanivorax, Marinobacter, Pseudomonas, and Acinetobacter). Sequence libraries generated from oiled sands revealed phylotypes that showed high sequence identity (up to 99%) to rRNA gene sequences from the oil-degrading bacterial isolates. The abundance of bacterial SSU rRNA gene sequences was ∼10-fold higher in oiled (0.44 × 107 to 10.2 × 107 copies g−1) versus clean (0.024 × 107 to 1.4 × 107 copies g−1) sand. Community analysis revealed a distinct response to oil contamination, and SSU rRNA gene abundance derived from the genus Alcanivorax showed the largest increase in relative abundance in contaminated samples. We conclude that oil contamination from the DH spill had a profound impact on the abundance and community composition of indigenous bacteria in Gulf beach sands, and our evidence points to members of the Gammaproteobacteria (Alcanivorax, Marinobacter) and Alphaproteobacteria (Rhodobacteraceae) as key players in oil degradation there.

Coral mucus functions as an energy carrier and particle trap in the reef ecosystem

Zooxanthellae, endosymbiotic algae of reef-building corals, substantially contribute to the high gross primary production of coral reefs, but corals exude up to half of the carbon assimilated by their zooxanthellae as mucus. Here we show that released coral mucus efficiently traps organic matter from the water column and rapidly carries energy and nutrients to the reef lagoon sediment, which acts as a biocatalytic mineralizing filter. In the Great Barrier Reef, the dominant genus of hard corals, Acropora, exudes up to 4.8 litres of mucus per square metre of reef area per day. Between 56% and 80% of this mucus dissolves in the reef water, which is filtered through the lagoon sands. Here, coral mucus is degraded at a turnover rate of at least 7% per hour. Detached undissolved mucus traps suspended particles, increasing its initial organic carbon and nitrogen content by three orders of magnitude within 2 h. Tidal currents concentrate these mucus aggregates into the lagoon, where they rapidly settle. Coral mucus provides light energy harvested by the zooxanthellae and trapped particles to the heterotrophic reef community, thereby establishing a recycling loop that supports benthic life, while reducing loss of energy and nutrients from the reef ecosystem.

Advective Transport Affecting Metal and Nutrient Distributions and Interfacial Fluxes in Permeable Sediments

Abstract Our laboratory flume experiments demonstrate that advective porewater flows produce biogeochemical reaction zones in permeable sediments, leading to specific and reproducible complex patterns of Fe, Mn, and nutrients. Oxygenated water, forced into the sediment when boundary flows were deflected by protruding sediment structures, generated distinct zones of nitrification and ferric iron precipitation. This inflow was balanced by ammonium-rich porewater ascending from deeper sediment layers, thereby creating an anoxic channel where dissolved Fe2+ and Mn2+ could reach the surface. Between the zones of ferric iron precipitation and Fe2+ upwelling, a layer with increased manganese oxide and solid phase Fe(II) concentrations formed, indicating redox reaction between these components. The establishment of topography on the previously smooth sediment surface reversed the net interfacial flux of solutes. While the smooth control core was found to be a sink for metals and nutrients, the sediment with mounds acted as a source for these substances. Our experiments show that in sandy sediment with an oxidised surface layer, reduced metal species can be released to the water column by flow-topography interactions. We conclude that advective transport processes constitute an important process controlling biogeochemical zonations and fluxes in permeable sea beds.

Hydrodynamical impact on biogeochemical processes in aquatic sediments

Boundary layer flow characteristics and sediment permeability control pathways and magnitude of material exchange in the surface layer of aquatic sediments. In fine-grained cohesive beds, bottom currents and sediment microtopography shape the diffusive boundary layer and locally produce areas where the interfacial solute fluxes are increased or reduced. Where sediment permeabilities exceed 10−12 m2, advective pore water flows driven by boundary flow–topography interaction dominate the sediment–water exchange of matter, with transport rates that exceed those of molecular diffusion by two orders of magnitude and more. The curved paths of the advective pore flows through the surface layers of such sandy beds generate complex three-dimensional biogeochemical patterns with extreme spatial and temporal variability ranging from millimeters to decimeters and seconds to seasons. High filtration rates, a bacterial community firmly attached to the mineral grains, rapidly changing biogeochemical zonations and winnowing of the sediment surface layers by frequent resuspension convert these beds into effective biocatalytical filter systems.

Benthic Exchange and Biogeochemical Cycling in Permeable Sediments

The sandy sediments that blanket the inner shelf are situated in a zone where nutrient input from land and strong mixing produce maximum primary production and tight coupling between water column and sedimentary processes. The high permeability of the shelf sands renders them susceptible to pressure gradients generated by hydrodynamic and biological forces that modulate spatial and temporal patterns of water circulation through these sediments. The resulting dynamic three-dimensional patterns of particle and solute distribution generate a broad spectrum of biogeochemical reaction zones that facilitate effective decomposition of the pelagic and benthic primary production products. The intricate coupling between the water column and sediment makes it challenging to quantify the production and decomposition processes and the resultant fluxes in permeable shelf sands. Recent technical developments have led to insights into the high biogeochemical and biological activity of these permeable sediments and their role in the global cycles of matter.

Activity and distribution of bacterial populations in Middle Atlantic Bight shelf sands

Abstract Spatiotemporal variation and metabolic activity of the microbial community were studied in coarse-grained Middle Atlantic Bight shelf sediments in relation to pools of dissolved and particulate carbon. Algal cells were present 8->70 mum) fraction of the sediment held the major share (61-98%) of benthic bacteria. Bacterial and algal cell abundances, exoenzymatic activity, and [DOC] generally showed higher values in May/July 2001 than in August/December 2000. Carbohydrates and proteins were hydrolyzed at potential rates of 1-12 nmol cm(-3) h(-1) (beta-glucosidase) and 3-70 nmol cm(-3) h(-1) (aminopeptidase), respectively. Fluorescence in situ hybridization analyses of the benthic microbes assigned 45-56% of DAPI-stained cells to Eubacteria and less than 2% to Eukarya. The prokaryotic community was dominated by planctomycetes and members of the Cytophaga/Flavobacterium cluster. Near the sediment surface, iodonitrotetrazolium violet reducing cells, that are considered actively respiring, amounted to 15-29% of total bacteria. Despite a low organic content (particulate organic carbon <0.03%) and relatively low bacterial abundances (<10(9) cm(-3)), the Middle Atlantic Bight shelf sediments showed organic matter turnover rates that are comparable to those found in organic-rich finer-grained deposits. Our findings suggest a high biocatalytic filtration activity in these coarse permeable sediments.

Bacteria, diatoms and detritus in an intertidal sandflat subject to advective transport across the water-sediment interface

This study focused on organic particles withrespect to their transport and sedimentarymineralisation in a North Sea intertidalsandflat previously characterised as stronglyinfluenced by advective transport across andbelow the water-sediment interface. Measuredpermeabilities of the sandy sediment rangedfrom 5.5 to 41⋅10−12 m2, andpermeabilities calculated from granulometricdata exceeded the measured values by a factorof 4.4 ∓ 2.8. Bacteria (2–9% of the POC)were highly variable in space and time. Theywere less mobile than interstitial fine (<70 µm) organic and inorganic particles, aspart of the population lived attached to large,heavy sand grains. The vertical distribution ofbacteria was closely related to the organiccarbon content of the fine-grained interstitialmaterial. In winter, bacterial numbers in theuppermost 5 cm amounted to 39–69% of thesummer ones. Carbon mineralisation rates rangedbetween 20 mg C m−2 d−1 in winter and580 mg C m−2 d−1 in summer, keepingstep with finer-grained sediments thatcontained an order of magnitude more organiccarbon. Sedimentary carbohydrates were mainlyintracellular or tightly bound to particles,and their concentrations were depth-invariantin winter, but exponentially decreasing withdepth in summer. Below 5 cm depth, the meanconcentration was (1590 ∓830) µg cm−3, without major downcoreor seasonal changes. Phytobenthos andphytodetritus were dominated by diatoms andcomprised merely minor amounts of other primaryproducers. Planktonic diatom depth profileswere related to weather and phytoplanktonconditions, and benthic diatoms showed similardepth distributions due to passive and activemotion. The penetration of relatively freshphytodetritus down to at least 5 cm, shown bychloropigment composition, emphasised the closecoupling between water column and sandysediment, facilitated by advective interfacialand subsurface flows.

Characterization of Nitrifying, Denitrifying, and Overall Bacterial Communities in Permeable Marine Sediments of the Northeastern Gulf of Mexico

ABSTRACT Sandy or permeable sediment deposits cover the majority of the shallow ocean seafloor, and yet the associated bacterial communities remain poorly described. The objective of this study was to expand the characterization of bacterial community diversity in permeable sediment impacted by advective pore water exchange and to assess effects of spatial, temporal, hydrodynamic, and geochemical gradients. Terminal restriction fragment length polymorphism (TRFLP) was used to analyze nearly 100 sediment samples collected from two northeastern Gulf of Mexico subtidal sites that primarily differed in their hydrodynamic conditions. Communities were described across multiple taxonomic levels using universal bacterial small subunit (SSU) rRNA targets (RNA- and DNA-based) and functional markers for nitrification (amoA) and denitrification (nosZ). Clonal analysis of SSU rRNA targets identified several taxa not previously detected in sandy sediments (i.e., Acidobacteria, Actinobacteria, Chloroflexi, Cyanobacteria, and Firmicutes). Sequence diversity was high among the overall bacterial and denitrifying communities, with members of the Alphaproteobacteria predominant in both. Diversity of bacterial nitrifiers (amoA) remained comparatively low and did not covary with the other gene targets. TRFLP fingerprinting revealed changes in sequence diversity from the family to species level across sediment depth and study site. The high diversity of facultative denitrifiers was consistent with the high permeability, deeper oxygen penetration, and high rates of aerobic respiration determined in these sediments. The high relative abundance of Gammaproteobacteria in RNA clone libraries suggests that this group may be poised to respond to short-term periodic pulses of growth substrates, and this observation warrants further investigation.

Microbial community successional patterns in beach sands impacted by the Deepwater Horizon oil spill

Although petroleum hydrocarbons discharged from the Deepwater Horizon (DWH) blowout were shown to have a pronounced impact on indigenous microbial communities in the Gulf of Mexico, effects on nearshore or coastal ecosystems remain understudied. This study investigated the successional patterns of functional and taxonomic diversity for over 1 year after the DWH oil was deposited on Pensacola Beach sands (FL, USA), using metagenomic and 16S rRNA gene amplicon techniques. Gamma- and Alphaproteobacteria were enriched in oiled sediments, in corroboration of previous studies. In contrast to previous studies, we observed an increase in the functional diversity of the community in response to oil contamination and a functional transition from generalist populations within 4 months after oil came ashore to specialists a year later, when oil was undetectable. At the latter time point, a typical beach community had reestablished that showed little to no evidence of oil hydrocarbon degradation potential, was enriched in archaeal taxa known to be sensitive to xenobiotics, but differed significantly from the community before the oil spill. Further, a clear succession pattern was observed, where early responders to oil contamination, likely degrading aliphatic hydrocarbons, were replaced after 3 months by populations capable of aromatic hydrocarbon decomposition. Collectively, our results advance the understanding of how natural benthic microbial communities respond to crude oil perturbation, supporting the specialization-disturbance hypothesis; that is, the expectation that disturbance favors generalists, while providing (microbial) indicator species and genes for the chemical evolution of oil hydrocarbons during degradation and weathering.