Mulholland
Margaret R.
Mulholland
Margaret R.
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PreprintOcean urea fertilization for carbon credits poses high ecological risks( 2008) Glibert, Patricia M. ; Azanza, Rhodora ; Burford, Michele ; Furuya, Ken ; Abal, Eva ; Al-Azri, Adnan ; Al-Yamani, Faiza ; Andersen, Per ; Anderson, Donald M. ; Beardall, John ; Berg, Gry M. ; Brand, Larry E. ; Bronk, Deborah ; Brookes, Justin ; Burkholder, JoAnn M. ; Cembella, Allan D. ; Cochlan, William P. ; Collier, Jackie L. ; Collos, Yves ; Diaz, Robert ; Doblin, Martina ; Drennen, Thomas ; Dyhrman, Sonya T. ; Fukuyo, Yasuwo ; Furnas, Miles ; Galloway, James ; Graneli, Edna ; Ha, Dao Viet ; Hallegraeff, Gustaaf M. ; Harrison, John A. ; Harrison, Paul J. ; Heil, Cynthia A. ; Heimann, Kirsten ; Howarth, Robert W. ; Jauzein, Cecile ; Kana, Austin A. ; Kana, Todd M. ; Kim, Hakgyoon ; Kudela, Raphael M. ; Legrand, Catherine ; Mallin, Michael ; Mulholland, Margaret R. ; Murray, Shauna A. ; O’Neil, Judith ; Pitcher, Grant C. ; Qi, Yuzao ; Rabalais, Nancy ; Raine, Robin ; Seitzinger, Sybil P. ; Salomon, Paulo S. ; Solomon, Caroline ; Stoecker, Diane K. ; Usup, Gires ; Wilson, Joanne ; Yin, Kedong ; Zhou, Mingjiang ; Zhu, MingyuanThe proposed plan for enrichment of the Sulu Sea, Philippines, a region of rich marine biodiversity, with thousands of tonnes of urea in order to stimulate algal blooms and sequester carbon is flawed for multiple reasons. Urea is preferentially used as a nitrogen source by some cyanobacteria and dinoflagellates, many of which are neutrally or positively buoyant. Biological pumps to the deep sea are classically leaky, and the inefficient burial of new biomass makes the estimation of a net loss of carbon from the atmosphere questionable at best. The potential for growth of toxic dinoflagellates is also high, as many grow well on urea and some even increase their toxicity when grown on urea. Many toxic dinoflagellates form cysts which can settle to the sediment and germinate in subsequent years, forming new blooms even without further fertilization. If large-scale blooms do occur, it is likely that they will contribute to hypoxia in the bottom waters upon decomposition. Lastly, urea production requires fossil fuel usage, further limiting the potential for net carbon sequestration. The environmental and economic impacts are potentially great and need to be rigorously assessed.
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ArticleHigh rates of N-2 fixation in temperate, western North Atlantic coastal waters expand the realm of marine diazotrophy(American Geophysical Union, 2019-06-10) Mulholland, Margaret R. ; Bernhardt, Peter W. ; Widner, Brittany ; Selden, Corday ; Chappell, Phoebe Dreux ; Clayton, Sophie A. ; Mannino, Antonio ; Hyde, Kimberly J. W.Dinitrogen (N2) fixation can alleviate N limitation of primary productivity by introducing fixed nitrogen (N) to the world's oceans. Although measurements of pelagic marine N2 fixation are predominantly from oligotrophic oceanic regions, where N limitation is thought to favor growth of diazotrophic microbes, here we report high rates of N2 fixation from seven cruises spanning four seasons in temperate, western North Atlantic coastal waters along the North American continental shelf between Cape Hatteras and Nova Scotia, an area representing 6.4% of the North Atlantic continental shelf area. Integrating average areal rates of N2 fixation during each season and for each domain in the study area, the estimated N input from N2 fixation to this temperate shelf system is 0.02 Tmol N/year, an amount equivalent to that previously estimated for the entire North Atlantic continental shelf. Unicellular group A cyanobacteria (UCYN‐A) were most often the dominant diazotrophic group expressing nifH, a gene encoding the nitrogenase enzyme, throughout the study area during all seasons. This expands the domain of these diazotrophs to include coastal waters where dissolved N concentrations are not always depleted. Further, the high rates of N2 fixation and diazotroph diversity along the western North Atlantic continental shelf underscore the need to reexamine the biogeography and the activity of diazotrophs along continental margins. Accounting for this substantial but previously overlooked source of new N to marine systems necessitates revisions to global marine N budgets.
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ArticlePotential virus-mediated nitrogen cycling in oxygen-depleted oceanic waters(Springer Nature, 2020-11-16) Gazitúa, M. Consuelo ; Vik, Dean R. ; Roux, Simon ; Gregory, Ann C. ; Bolduc, Benjamin ; Widner, Brittany ; Mulholland, Margaret R. ; Hallam, Steven J. ; Ulloa, Osvaldo ; Sullivan, Matthew B.Viruses play an important role in the ecology and biogeochemistry of marine ecosystems. Beyond mortality and gene transfer, viruses can reprogram microbial metabolism during infection by expressing auxiliary metabolic genes (AMGs) involved in photosynthesis, central carbon metabolism, and nutrient cycling. While previous studies have focused on AMG diversity in the sunlit and dark ocean, less is known about the role of viruses in shaping metabolic networks along redox gradients associated with marine oxygen minimum zones (OMZs). Here, we analyzed relatively quantitative viral metagenomic datasets that profiled the oxygen gradient across Eastern Tropical South Pacific (ETSP) OMZ waters, assessing whether OMZ viruses might impact nitrogen (N) cycling via AMGs. Identified viral genomes encoded six N-cycle AMGs associated with denitrification, nitrification, assimilatory nitrate reduction, and nitrite transport. The majority of these AMGs (80%) were identified in T4-like Myoviridae phages, predicted to infect Cyanobacteria and Proteobacteria, or in unclassified archaeal viruses predicted to infect Thaumarchaeota. Four AMGs were exclusive to anoxic waters and had distributions that paralleled homologous microbial genes. Together, these findings suggest viruses modulate N-cycling processes within the ETSP OMZ and may contribute to nitrogen loss throughout the global oceans thus providing a baseline for their inclusion in the ecosystem and geochemical models.
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ArticleToward resolving disparate accounts of the extent and magnitude of nitrogen fixation in the Eastern Tropical South Pacific oxygen deficient zone(Association for the Sciences of Limnology and Oceanography, 2021-04-07) Selden, Corday ; Mulholland, Margaret R. ; Widner, Brittany ; Bernhardt, Peter W. ; Jayakumar, AmalExamination of dinitrogen (N2) fixation in the Eastern Tropical South Pacific oxygen deficient zone has raised questions about the range of diazotrophs in the deep sea and their quantitative importance as a source of new nitrogen globally. However, technical considerations in the deployment of stable isotopes in quantifying N2 fixation rates have complicated interpretation of this research. Here, we report the findings of a comprehensive survey of N2 fixation within, above and below the Eastern Tropical South Pacific oxygen deficient zone. N2 fixation rates were measured using a robust 15N tracer method (bubble removal) that accounts for the slow dissolution of N2 gas and calculated using a conservative approach. N2 fixation was only detected in a subset of samples (8 of 125 replicated measurements) collected within suboxic waters (< 20 μmol O2 kg−1) or at the oxycline. Most of these detectable rates were measured at nearshore stations, or where surface productivity was high. These findings support the hypothesis that low oxygen/high organic carbon conditions favor non-cyanobacterial diazotrophs. Nevertheless, this study indicates that N2 fixation is neither widespread nor quantitatively important throughout this region.
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ArticleCarbon budget of tidal wetlands, estuaries, and shelf waters of eastern North America(John Wiley & Sons, 2018-04-04) Najjar, Raymond G. ; Herrmann, Maria ; Alexander, Richard ; Boyer, Elizabeth W. ; Burdige, David J. ; Butman, David ; Cai, Wei-Jun ; Canuel, Elizabeth A. ; Chen, Robert F. ; Friedrichs, Marjorie A. M. ; Feagin, Russell A. ; Griffith, Peter C. ; Hinson, Audra L. ; Holmquist, James R. ; Hu, Xinping ; Kemp, William M. ; Kroeger, Kevin D. ; Mannino, Antonio ; McCallister, S. Leigh ; McGillis, Wade R. ; Mulholland, Margaret R. ; Pilskaln, Cynthia H. ; Salisbury, Joseph E. ; Signorini, Sergio R. ; St-Laurent, Pierre ; Tian, Hanqin ; Tzortziou, Maria ; Vlahos, Penny ; Wang, Zhaohui Aleck ; Zimmerman, Richard C.Carbon cycling in the coastal zone affects global carbon budgets and is critical for understanding the urgent issues of hypoxia, acidification, and tidal wetland loss. However, there are no regional carbon budgets spanning the three main ecosystems in coastal waters: tidal wetlands, estuaries, and shelf waters. Here we construct such a budget for eastern North America using historical data, empirical models, remote sensing algorithms, and process‐based models. Considering the net fluxes of total carbon at the domain boundaries, 59 ± 12% (± 2 standard errors) of the carbon entering is from rivers and 41 ± 12% is from the atmosphere, while 80 ± 9% of the carbon leaving is exported to the open ocean and 20 ± 9% is buried. Net lateral carbon transfers between the three main ecosystem types are comparable to fluxes at the domain boundaries. Each ecosystem type contributes substantially to exchange with the atmosphere, with CO2 uptake split evenly between tidal wetlands and shelf waters, and estuarine CO2 outgassing offsetting half of the uptake. Similarly, burial is about equal in tidal wetlands and shelf waters, while estuaries play a smaller but still substantial role. The importance of tidal wetlands and estuaries in the overall budget is remarkable given that they, respectively, make up only 2.4 and 8.9% of the study domain area. This study shows that coastal carbon budgets should explicitly include tidal wetlands, estuaries, shelf waters, and the linkages between them; ignoring any of them may produce a biased picture of coastal carbon cycling.
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ArticleAssessing phytoplankton nutritional status and potential impact of wet deposition in seasonally oligotrophic waters of the Mid‐Atlantic Bight(John Wiley & Sons, 2018-04-06) Sedwick, Peter N. ; Bernhardt, Peter W. ; Mulholland, Margaret R. ; Najjar, Raymond G. ; Blumen, L. M. ; Sohst, Bettina M. ; Sookhdeo, C. ; Widner, BrittanyTo assess phytoplankton nutritional status in seasonally oligotrophic waters of the southern Mid‐Atlantic Bight, and the potential for rain to stimulate primary production in this region during summer, shipboard bioassay experiments were performed using natural seawater and phytoplankton collected north and south of the Gulf Stream. Bioassay treatments comprised iron, nitrate, iron + nitrate, iron + nitrate + phosphate, and rainwater. Phytoplankton growth was inferred from changes in chlorophyll a, inorganic nitrogen, and carbon‐13 uptake, relative to unamended control treatments. Results indicated the greatest growth stimulation by iron + nitrate + phosphate, intermediate growth stimulation by rainwater, modest growth stimulation by nitrate and iron + nitrate, and no growth stimulation by iron. Based on these data and analysis of seawater and atmospheric samples, nitrogen was the proximate limiting nutrient, with a secondary limitation imposed by phosphorus. Our results imply that summer rain events increase new production in these waters by contributing nitrogen and phosphorus, with the availability of the latter setting the upper limit on rain‐stimulated new production.
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ArticleDinitrogen fixation across physico-chemical gradients of the Eastern Tropical North Pacific oxygen deficient zone(American Geophysical Union, 2019-09-04) Selden, Corday ; Mulholland, Margaret R. ; Bernhardt, Peter W. ; Widner, Brittany ; Macias-Tapia, Alfonso ; Ji, Qixing ; Jayakumar, D. A.The Eastern Tropical North Pacific Ocean hosts one of the world's largest oceanic oxygen deficient zones (ODZs). Hot spots for reactive nitrogen (Nr) removal processes, ODZs generate conditions proposed to promote Nr inputs via dinitrogen (N2) fixation. In this study, we quantified N2 fixation rates by 15N tracer bioassay across oxygen, nutrient, and light gradients within and adjacent to the ODZ. Within subeuphotic oxygen‐deplete waters, N2 fixation was largely undetectable; however, addition of dissolved organic carbon stimulated N2 fixation in suboxic (<20 μmol/kg O2) waters, suggesting that diazotroph communities are likely energy limited or carbon limited and able to fix N2 despite high ambient concentrations of dissolved inorganic nitrogen. Elevated rates (>9 nmol N·L−1·day−1) were also observed in suboxic waters near volcanic islands where N2 fixation was quantifiable to 3,000 m. Within the overlying euphotic waters, N2 fixation rates were highest near the continent, exceeding 500 μmol N·m−2·day−1 at one third of inshore stations. These findings support the expansion of the known range of diazotrophs to deep, cold, and dissolved inorganic nitrogen‐replete waters. Additionally, this work bolsters calls for the reconsideration of ocean margins as important sources of Nr. Despite high rates at some inshore stations, regional N2 fixation appears insufficient to compensate for Nr loss locally as observed previously in the Eastern Tropical South Pacific ODZ.
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ArticleNitrogen fixation at the Mid‐Atlantic Bight Shelfbreak and transport of newly fixed nitrogen to the Slope Sea(American Geophysical Union, 2024-04-05) Selden, Corday R. ; Mulholland, Margaret R. ; Crider, Katie E. ; Clayton, Sophie A. ; Macias-Tapia, Alfonso ; Bernhardt, Peter W. ; McGillicuddy, Dennis J. ; Zhang, Weifeng Gordon ; Chappell, Phoebe DreuxContinental shelves contribute a large fraction of the ocean's new nitrogen (N) via N2 fixation; yet, we know little about how physical processes at the ocean's margins shape diazotroph biogeography and activity. Here, we test the hypothesis that frontal mixing favors N2 fixation at the Mid-Atlantic Bight shelfbreak. Using the 15N2 bubble release method, we measured N2 fixation rates on repeat cross-frontal transects in July 2019. N2 fixation rates in shelf waters (median = 5.42 nmol N L−1 d−1) were higher than offshore (2.48 nmol N L−1 d−1) but did not significantly differ front frontal waters (8.42 nmol N L−1 d−1). However, specific N2 uptake rates, indicative of the relative contribution of diazotroph-derived N to particulate N turnover, were significantly higher in frontal waters, suggesting that diazotroph-derived N is of greater importance in supporting productivity there. This study furthered captured an ephemeral shelf-water streamer, which resulted from the impingement of a warm core ring on the shelf. The streamer transported shelf-water diazotrophs (including UCYN-A and Richelia spp., as assessed by qPCR) offshore with sustained high N2 fixation rates. This feature injected >50 metric tons d−1 of newly fixed N to the Slope Sea—a rate equivalent to ∼4% of the total N flux estimated for the entire Mid-Atlantic Bight. As intrusions of Gulf Stream meanders and eddies onto the shelf are increasing in frequency due to climate change, episodic lateral fluxes of new N into the Slope Sea may become increasingly important to regional budgets and ecosystem productivity.