Nitrogen cycling in oxygen deficient zones
The major marine oxygen deficient zones (ODZs), namely, Eastern Tropical North Pacific (ETNP), Eastern Tropical South Pacific (ETSP) and the Arabian Sea, comprise only 0.1 - 0.2% of the total volume of the world ocean, yet have long been recognized to be important in global nitrogen (N) cycling. At least a quarter of marine fixed N is lost (i.e. converted to gaseous end products) in these regions and the impact of this is observed in both the fixed N mass and stable isotope budgets. The major ODZs also contribute disproportionately to the atmospheric nitrous oxide budget, a greenhouse gas. Despite intense study, there is much uncertainty about the rates and pathways of N transformations in these regions. The main fixed N loss processes are denitrification and anammox (anaerobic ammonium oxidation). The substrates that support these processes are ultimately derived from the rain of organic matter produced in surface waters as well as the diffusion of inorganic N. Thus denitrification and anammox are linked to other processes in the N cycle to supply the N substrates that they convert to N2. (Figure 1) Most of these processes are the subject of research currently underway in the Ward laboratory.
Figure 1. Marine nitrogen cycle: Production of organic matter by assimilation of inorganic nitrogen occurs in the surface water, including N2 fixation. This material is mineralized at depth, leading to the release of NH4. Ammonia oxidation, followed by nitrite oxidation produced the nitrate that is the main DIN component in deep water. In oxygen minimum zones, the nitrate is denitrified to N2, via transformations including dissimilatory reduction to ammonium, anammox and canonical denitrification. The net input term for fixed N is N2 fixation, and the main loss terms are denitrification and anammox.
Phytoplankton N utilization
Photosynthesis by marine phytoplankton is thought to account for half of the oxygen in our atmosphere. Phytoplankton form the basis of the marine food web and thus are a crucial element in the biological pump whereby CO2 from the atmosphere is sequestered in the deep ocean. Biologically available nitrogen (N) is essential for photosynthesis and phytoplankton growth, and yet much of the global ocean appears to be N limited. Thus, N availability has the potential to not only limit marine productivity, but may also determine the composition/diversity of the phytoplankton community.
For decades, biological oceanography focused on the eukaryotic phytoplankton. In the 1970-80’s, it was discovered that the single celled picocyanobacteria are numerically dominant in the oceans and are responsible for a large fraction of ocean photosynthesis, leading to a new paradigm in which the picocyanobacteria dominate upper ocean biology and biogeochemistry. In fact, new data support the classic view that the eukaryotic phytoplankton are disproportionately important in both N and C cycling, even in regions where very small cells dominate the biomass and where the eukaryotes themselves are in the “pico” size fraction. On the basis of still quite limited molecular surveys, we now recognize that the diversity of both large and small eukaryotic phytoplankton is greater than previously thought and that the most abundant and widespread eukaryotes are probably not in culture and may not be closely related to known cultivated organisms.
The structure of phytoplankton communities is a major determinant in the magnitude and fate of primary production. We are interested in identifying the sources of N supporting phytoplankton growth with a view to understanding how N availability affects phytoplankton community diversity, both in the N-poor subtropical ocean (Figure 2) and in productive coastal upwelling regions. Blooms, in which highest productivity rates are observed, are usually dominated by a small number of cell types. Diatoms contribute disproportionately to new production by virtue of their dominance in upwelling systems, where they exhibit episodic blooms in response to nitrate supply. Although temporary escape from predation may be the ultimate reason that bloom biomass accumulates, the ability of diatoms to exploit episodic nitrate supply must also be a factor.
We are investigating the taxonomic, genetic and functional diversity of eukaryotic phytoplankton in many ocean environments and in the context of several different experiments.
Figure 2. Sampling the Sargasso Sea from the Atlantic Explorer at the BATS station.
Nitrogen cycling in the coastal marine environment
The coastal environment, including marshes and estuaries, plays an important role as a buffer and a link between terrestrial and marine environments. On the east coast of the U.S., many coastal systems are subject to anthropogenically enhanced nitrogen loading (Figure 3), which has the potential to alter natural ecosystem processes by influencing microbial activities. We are interested in the biogeochemistry of shallow sediments and coastal waters, using a suite of analytical, experimental and modeling approaches to investigate the magnitudes and distributions of N transformation rates and the structure of microbial assemblages responsible for those transformations.
Figure 3. Great Sippewissett Marsh. The stakes outline one of the 10-m diameter plots that has received high levels of nutrient additions for the past 40 years.