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Seeps
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Marsh and Estuarine Biogeochemistry
Research
conducted by Joye's group examines the biogeochemical cycling of
nutrients (nitrogen and phosphorus), trace metals (iron and manganese),
carbon, sulfur and oxygen in a variety of systems, ranging from saline
lakes to coastal environments to deep ocean sediments. Several projects
include parallel studies of biogeochemical and molecular ecological
dynamics with the aim of identifying fundamental links between environmental
variables, bacterial community composition and bacterial activity.
Areas of focus include coastal ecology and the study of microbial
metabolism in and adaptation to "extreme" environments (a.k.a. astrobiology)
Coastal
Ecology
We strive to identify and understand
the environmental and physiological controls on elemental cycling
(e. g., N, P, C, O, S, Fe, Mn ...) and ecosystem metabolism in coastal
regions. A primary goal of this work is to identify how coastal
ecosystems respond to global change and various natural and anthropogenic
forcing functions.
We utilize a broad, cross-disciplinary
approach to address biogeochemical research questions. We value innovative
technologies and incorporate new methodologies and approaches into
our research program as they become available. Our work
exploits traditional biogeochemical (e.g., radio- and stable- isotopic
tracers, physiological inhibitor-based studies), microbiological (e.g.,
enrichment culture work, microbial isolations), molecular biological
(e.g., PCR, cloning and sequencing), and geochemical (e.g., rate measurements,
stable isotopes, and constituent concentration profiles) to examine
rates of individual processes at local and system-wide scales. Significant
effort is put towards developing novel approaches and techniques to
address research questions.
We have worked or are working in a variety
of coastal environments, including salt marshes along the coast of
Georgia, South Carolina and Massachusetts, California, North Carolina,
Georgia and Texas, and mangrove forests in Belize and Panama. Current
projects are summarized briefly below. If you have questions, comments
or would like additional information, please contact Mandy Joye at mjoye@uga.edu.
Our coastal ecology research is supported
by the National Science Foundation's Georgia Coastal Ecosystems
LTER program and the Biocomplexity in the Environment program,
by the Environmental Protection Agency's Climate Change program and
by NOAA Sea Grant programs in Georgia and South Carolina.
 

Salt Marsh and Estuarine Ecology The role of groundwater in coastal nutrient
budgets {Learn More}
Groundwater
Biogeochemistry
As human population has grown and agricultural, commercial, and industrial activities
have expanded in coastal watersheds, loading of anthropogenic wastes from localized
(e.g., sewage, industrial effluent) and diffuse (e.g., agricultural run-off)
sources to surface waters and groundwater has increased. Ultimately, these materials
end up in coastal waters, where they cause eutrophication, the increase in labile
organic matter supply to an ecosystem. Today, increases in the frequency of
harmful algal blooms, water column hypoxia/anoxia, fish kills, and reduced water
quality are clear signs of eutrophication in the coastal zone. Understanding
the response of the coastal ecosystem to eutrophication is one of the most important
research challenges facing marine scientists today.
Many factors, including land use (natural
vs. developed, e.g., residential, agricultural or commercial), soil
type and erosion rate, influence water quality. To date, the scientific
effort has focused largely on documenting the composition and flow
of surface waters (e.g., rivers, streams), the most conspicuous component
of the hydrologic cycle. However, predicting the response of coastal
ecosystems to land use change and eutrophication requires robust models
that include all relevant sources of nutrients and organic materials.
Among these, groundwater isan important, but poorly understood, source of nutrients and organics
to coastal waters. Groundwater-derived fluxes need to be quantified,
particularly in Southeastern U.S., coastal systems where such data
are rare. The lack of information regarding groundwater flux and groundwater
quality in coastal systems makes it impossible to predict how the
quality (chemistry) or quantity (freshwater flux) of this "input" or "source
term" will respond to increased development pressures in coastal regions.
We are assessing the flux of groundwater
to coastal systems using a variety of approaches, including seepage
chambers (see figures to the left and below; seepage meters are the
brown circles beneath the water surface). We collaborate with Dr. Carolyn
Ruppel at Georgia Tech, who uses indirect geophysical techniques
to estimate groundwater fluxes, and researchers at the University
of South Carolina (Drs. Marj Aelion and Billy
Moore), who use geochemical and microbiological techniques to
evaluate the fate of groundwater derived materials. An integrated
effort is being made to quantify the impact of coastal development
and land use change on ecological, chemical, and physical processes
through the study of both groundwater and surface water components
in Georgia and South Carolina. In the course of this project, we will
quantify the flux and chemical signature of shallow coastal groundwater
and contrast the chemical composition of groundwater and surface waters.
The impact of groundwater flux on a suite of biological processes
will be evaluated in the marsh and in estuarine waters (tidal creeks).
These data will be integrated into a GIS-based modeling framework,
with the aim of predicting the response of groundwater-derived fluxes
to land use change.

Sediment Biogeochemistry To evalaute the role of groundwater-derived nutrient loading
in an ecosystem-level context, we characterize sediment biogeochemical
regimes, determine rates processes like denitrification, and quantify
benthic fluxes. We are examining sites where surface and/or
groundwater quality has been altered to varying degrees due to development
in the upper (or adjacent) watershed. Over the past 50 years,
increased population density and industrial and agricultural activity
in coastal watersheds has elevated the loading rates of nutrients,
particularly nitrogen (N) and phosphorus (P), to coastal waters resulting
in the eutrophication of many nearshore environments. Identifying
ways to improve coastalenvironmental quality and formulating a suite of systematic
environmental quality indicators are primary objectives of coastal
managers around the globe.
Sound management decisions require a
basic understanding of the structure and function of coastal ecosystems. However,
such basic information is lacking for many systems and scientists
are unable to predict how they will respond to future perturbations.
Examining the biogeochemical response of coastal sediments to increased
nutrient loading provides a mechanism to assess how these critical
habitats may respond to future perturbation and may allow us to identify
important feedback mechanisms necessary for improving coastal environmental
quality in the future.
Salt marsh and mudflat habitats, the
interface between terrestrial and oceanic ecosystems, serve as filters
of materials exchanged between these two end-members. These regions
may act as sources, sinks and/or transformers of organic materials
and nutrients. Determining which role they play and elucidating the
factors that control net nutrient exchanges have important implications
for understanding nutrient flux and production dynamics in both estuarine
and coastal oceanic environments. In this project, we are assessing
spatio-temporal patterns in sediment metabolism and nutrient dynamics
at sites within two intertidal marsh-mudflat complexes along the Georgia
coast. We have chosen to study sediment processes because they determine
net fluxes of materials between the water column and sediment and
as a group are capable of significantly altering net metabolism on
the ecosystem scale in shallow coastal regions.
Our hypotheses concern linkages between
benthic primary production, net sediment metabolism, and the cycling
of nitrogen and phosphorus. We will investigate interactions among
processes on various time and space scales, but will focus our efforts
on documenting diel (day-night) and seasonal trends in carbon (C),
N and P dynamics. With the recent emergence of ephemeral microalgal
blooms in the pristine Satilla River, understanding the mechanisms
leading to the proliferation of benthic microalgae and on how their
activity influences N and P processing has never been more critical.
We will quantify rates of benthic metabolism and N and P fluxes as
well as rates of "indicator" processes, specifically, benthic primary
production and denitrification, in marshes at the mouth of Satilla
River and adjacent to Sapelo Island.
Impacts of Climate Change on Coastal Ecosystem
Dynamics {Learn More}
Conceptual models will be used to develop
a framework for examining how ecosystem services of tidal marshes
vary along the salinity gradient and how climate change will alter
the delivery of ecosystem services. We hypothesize that tidal
freshwater marshes provide higher levels of regulation, habitat and
(plant) production functions relative to salt marshes. Salt
marshes though provide higher levels of disturbance regulation and
fisheries habitat. Accelerated sea level rise is predicted to
reduce the area of tidal marsh through submergence and conversion
of tidal freshwater marsh to brackish and salt marsh. The result
is a reduction in ecosystem services of salt and brackish marshes
along with a complete loss of services of tidal freshwater marshes. Predicted
greater inter-annual variability of climate in the future will lead
to greater frequency of drought that reduces delivery of ecosystem
services and episodic freshwater pulsing which we predict will enhance
delivery of ecosystem services.
We will test the effects of rising sea
level and greater inter-annual variability of climate on alteration
of area and ecosystem services of tidal marshes in three watersheds
(Altamaha, Satilla and Savannah Rivers, Georgia) of the South Atlantic
coast. Ecosystem services related to disturbance regulation
(shoreline protection), gas regulation (CO2 & CH4 flux),
soil formation (C sequestration), nutrient regulation (N, P retention)
and waste treatment (sediment deposition, denitrification), refugium
and food (production & diversity of macrophytes & marsh nekton)
will be measured in replicate (n=2) salt, brackish and tidal freshwater
marshes of each watershed. GIS in conjunction the SLAMM model
will be employed to predict changes in marsh area resulting from submergence
and habitat conversion. Overlay of ecosystem-scale measurements will
be used to predict how the cumulative delivery of ecosystem services
in each watershed is altered in response to rising sea level. SLAMM
also will be used to predict changes in shoreline protection potential
of tidal marshes, commercial shrimp yields and the effects of alternate
management scenarios (maintenance of existing dikes, construction
of new dikes) on delivery of ecosystem services.
This work will provide a basis to understand
how ecosystem services vary among salt-, brackish- and tidal fresh-water
marshes, predicts how sea level rise alters marsh area and delivery
of ecosystem services, compares the effects of diking (to protect
marshes) on delivery of ecoystem services and elucidates how climate
variability affects temporal patterns of primary production & diversity,
sediment deposition and marsh accretion.

Mangrove Biogeochemistry {Learn More}
Microbial and nutrient controls in mangrove ecosystems
[this Biocomplexity project is directed by Dr.
Candy Feller at the Smithsonian Environmental Research Center]
Mangrove forests, the intertidal, tree-dominated
communities characteristic of tropical coasts, are often described
as "simple systems." This is a relative description, however, perhaps
due to the apparent greater complexity of terrestrial systems with
larger numbers of plant species. Nonetheless, these tidal wetlands
are systems with complex trophic structures and sharp gradients in
nutrient dynamics. Furthermore, mangroveorganisms exhibit intricate
physiological and structural adaptations to stresses and environmental
variables associated with the zone of overlap between marine and terrestrial
ecosystems. Ecological interactions and physiological processes in
mangrove forests are complex and controlled by interactions among
the microbial community structure, function, and interaction of higher
trophic groups, and feedback among different trophic levels. Hydrology,
physico-chemical factors, and the physiological tolerances of mangrove
ecosystem species mediate these interactions. We propose to elucidate
the linkages and feedback loops among mangrove community components
and place them in the context of forest structure and trophic (grazing
and detrital) connections in mangrove ecosystems. Our major goal is
to determine how the interactions controlling structure and function
at a number of trophic levels act to produce the emergent properties
that characterize mangrove forests.
Our Biocomplexity program seeks to answer
the question,"What are the functional relationships among microorganisms,
geochemical processes, hydrology, and nutrient availability in mangrove
forests, and how do these relationships interact to generate biological
complexity in this ecosystem?" We will qualify and quantify
a set of potentially interactive ecological and physiological processes
that span multiple levels of organization and scale, i.e., microbial
community structure, microbially-mediated nutrient transformations,
soil physico-chemical properties, primary productivity, plant physiology
and stress adaptations, within-plant and ecosystem-level nutrient
cycling, and primary and detrital consumption. Using an established
fertilization experiment in Belize and a multidisciplinary approach,
our objectives are to determine how these processes are interrelated
by examining the effects of increased availability of limiting nutrients
on feedback relationships among hydro-edaphic conditions; microbial,
plant, and animal communities: primary production and decomposition;
and peat formation. The proposed study site is a range of mangrove-dominated
islands located in an oligotrophic setting, remote from anthropogenic
influences. This system is nitrogen (N) limited at the mangrove fringe,
phosphorus (P) limited in the interior, and both N and P limited in
a transition zone.
To unify our study, we will use both
diagenetic modeling of sediment processes and Network Analysis. Diagenetic
model information will feed into the Network Analysis to add robustness
to nutrient cycling components. The latter provides a powerful set
of mathematical tools to determine direct and indirect interactions
of one ecosystem compartment with another, to produce a synopsis of
the trophic structure, to enumerate and quantify routes via which
nutrients are recycled, and to define the overall status of system
activity.

Extreme
Environments
Examining the interrelationships between
organisms and the environments within which they exist is a basic
goal of biogeochemistry. When such studies focus on life-supporting
environments that exist near the extremes of planetary conditions,
new insight can be obtained regarding the evolution of life on Earth
and the ability of microorganisms to adapt to a variety of physiological
extremes, e.g., temperature, salinity, pressure, etc. The study
of extreme environments is often referred to as "Astrobiology".
Our research in extreme environments
examines chemoautotrophic processes, mainly methane oxidation, nitrification
and nitrification. We use traditional biogeochemical approaches and
molecular biological and organic geochemical methods to quantify rates
of specific processes, access microbial diversity and genetic potential,
and to identify novel microorganisms with unique physiologies.
This research is funded by the National
Science Foundation's Microbial Observatory and Life in Extreme Environments
programs, by NOAA's National Undersea Research Program, and by the
American Chemical Society's Petroleum Research Fund. 

Cold
Seeps {Learn More}
Molecular ecology and biogeochemistry of methane
hydrates and brine
pools in the Gulf of Mexico.
Low
molecular weight hydrocarbons (methane through pentane) and hydrogen
sulfide form ice-like clathrates with water called "gas hydrates".
Frequently, methane is the dominant gas, so these structures are referred
to as "methane hydrates" or "methane clathrates". Submarine methane
hydrates comprise a large reservoir of labile organic carbon. Methane
hydrate forms under conditions of methane saturation, high pressure
and low temperature by the inclusion of gas molecules, primarily methane
but also higher hydrocarbon gases up to pentane, into a lattice of
water molecules. The resulting solid is stable at temperatures below
about 7 ºC and pressures greater than about 50 bar. The
widespread occurrence of gas hydrates became apparent when deeply
buried layers, known as bottom simulation reflectors (BSR), were detected
by seismic and deep-sea drilling studies. BSR features are widely
distributed at the base of continental margins at sub-bottom depths
of about 300m. In the Gulf of Mexico, hydrates occur as shallow layers
and vein-filling plugs that breach the sediment interface and slowly
decompose in contact with seawater. The "extreme" nature of the hydrate
niche includes low temperature and high pressure, limited carbon sources
(C1-C5), and desiccation (little available free water). Thus microbes
living in this environment require special adaptations.
Little is known about the types of microorganisms
dwelling in hydrate environments, their phylogenetic diversity, taxonomy,
ecology or ecophysiology. This project will test the hypotheses that
hydrate microbes inhabit distinct microbial niches requiring specialized
adaptation, that these niches have predictable temporal and spatial
characteristics, and that there is dynamic interaction between microbial
activity and the geochemistry of the system. We will use molecular
biological and organic geochemical techniques to assess the microbial
community composition and identify 'new' microorganisms. Rates of
processes will be determined using classic biogeochemical methods.

Saline
Lakes {Learn
More} Mono Lake Microbial Observatory
What might the microbial ecology of Mono
Lake have to do with microbial life on Mars? Present day alkaline
(or soda), hypersaline lakes on Earth are believed to be good
analogs to the ancient seas of Earth and Mars (shown to the right).
Studying the microbial ecology of soda lakes like Mono Lake may provide
insight to the types and physiology of microbes that inhabited early
Earth and Martian oceans.
Available data suggests that early in
the history of Mars, climatic conditions could have supported development
of a microbial biosphere. Degassing of volatiles from the Martian
interior would have generated shallow oceans (from depths of 50 m
to 0.5 km on the surface) and a dense atmosphere rich in carbon dioxide.
Martian "oceans" were probably standing water bodies that filled impact
craters and thus were quite different from Earth's oceans. Nonetheless,
liquid water was present on the surface (as evidence by the canyon
shown to the left). A carbon dioxide-rich atmosphere would have generated
a greenhouse sufficient to maintain liquid water for millions to hundreds
of millions of years. A carbon dioxide rich atmosphere would have
also resulted in surface waters rich in carbonate and bicarbonate,
important anions in present-day soda lakes. During this period, microbial
life could have evolved on Mars. Geological evidence suggests that
Earth's microbial biosphere developed in about a hundred billion years.
When conditions on the Martian surface deteriorated, the microbial
biosphere would have become extinct.
So, the Archean seas of Earth and Mars were probably
saline, highly alkaline soda lakes and present day soda lakes, like
Mono Lake (shown below) are thus reasonable analogs for Archean aquatic
systems. Our work in MonoLake, California, a soda lake in the
Eastern Sierra Nevada region, investigates aspects of the microbial
ecology and biogeochemistry of lake waters and sediment. The Mono Lake
environment represents many environmental extremes, including elevated
pH (~ 10), alkalinity (>> 10 times average sea water) and salinity
(> 80 ppt). Because of the high salinity and depth-dependent salt
gradients, lakes like Mono Lake are often meromictic (i.e., they do
not mix completely each year), and with prolonged meromixis, anoxic
bottom waters can attain high concentrations of reduced compounds like
dissolved sulfide and ammonia. Despite this, microbial life thrives
in Mono Lake. Understanding the diversity and physiology of Mono Lake
microbes is a primary goal of our study.
We thank the National Science Foundation,
the EPA, NOAA Sea Grant LU-CES programs in GA and SC, NOAA National Undersea
Research Program, and the ACS-Petroleum Research Fund for supporting this
work.
**Disclaimer** Any
opinions, findings, and conclusions or recommendations expressed
here are those of the author (Mandy
Joye) and
do not necessarily reflect the views of the funding agencies.

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