AOG Publications by Year


Title of Invention:  AQUATIC SAMPLE ANALYSIS SYSTEM, United States patent 14/808,757, awarded 07/24/2015 (co-inventor with Dr Ben Van Mooy)

In Progress/Submitted

Keil R.G. (in press 2016) Mechanisms of Organic Carbon and Calcium Carbonate Preservation in Marine Sediments.  Vol 9, Annual Review of Marine Science.

Fuchsman, C., Penn J., Keil R.G., Devol A.H., Deutsch C., and Rocap G., in preparation. Microbial ecological niche partitioning affects N2 gas production in the largest marine oxygen deficient zone. Global Biogeochemical Cycles

Ward N.D., Bianchi T.S., Sawakuchi H.S., Gagne-Maynard W., Cunha A.C., Brito D.C., Neu V., Krusche A.V.,  Keil R.G., and Richey J.E. (in press)  A comparison of plant litter leachate degradation rates and priming effects in the lower Amazon River. Geophysical Research Letters, submitted.


Keil RG, Neibauer J, Biladeau C., Van der Elst K., and Devol AH (2016).  Organic Geochemistry and Targeted Peptidomics of Sinking Particles in the Arabian Sea.  Biogeosciences, 13, 2077–2092, 2016,


Ward N.D., Bianchi T.S., Sawakuchi H.O., Gagne-Maynard W., Cunha A.C., Brito D.C., Neu V., de Matos A., da Silva R., Krusche A.V., Richey J.E., and Keil R.G. (in review) Reactivity of organic molecules and priming processes in the Amazon River.  JGR

Sollai M., Hopmans E.C., Schouten S., Keil R.G., and Sinninghe-Damsté J., (2015) Intact polar lipids of Thaumarchaeota and anammox bacteria as indicators of N-cycling in the Eastern Tropical North Pacific oxygen deficient zone. Biogeosciences.

Ward N.D., Krusche A.V., Sawakuchi H., Brito D.C., Cunha A.C., Sousa Moura J.M., da Silva R., Yager P.L., Keil R.G., and Richey J.E.  (2015)  The compositional evolution of dissolved and particulate organic matter along the lower Amazon River, In press, Marine Chemistry,

Keil R.G. (2015) Hoard of Fjord carbon. Nature Geosciences commentary, invited, March 2015.

Bridoux M.C., Neibauer J., Nunn B., Ingalls A.I., and Keil R.G.  (2015).  Suspended marine particulate proteins in coastal and oligotrophic waters.  Journal of Marine Systems


Babbin AR, Keil RG, Devol AH, Ward BB (in press) Organic matter stoichiometry, flux, and oxygen control nitrogen loss in the ocean. Science 25 April 2014: 344 (6182), 406-408. [DOI:10.1126/science.1248364]  (supplemental materials)

Keil R.G., and Mayer L.M. (2014) Mineral Matrices and Organic Matter. In: Holland H.D. and Turekian K.K. (eds.) Treatise on Geochemistry, Second Edition, vol. 12, pp. 337-359. Oxford: Elsevier.

Chang BX, Rich JR, Jayakumar A, Naik H, Prathihary A, Keil RG, Ward BB, Devol AH.  2014. The effect of organic carbon on fixed nitrogen loss in the eastern tropical South Pacific and Arabian Sea oxygen deficient zones.  Limnology and Oceanography  2014;59(4):1267-1274.


Ward ND, Keil RG, Medeiros PM, Brito DC, Cunha AC, Dittmar T, Yager PL, Krusche AV, Richey JE (2013b) Degradation of terrestrially derived macromolecules in the Amazon River. Nature Geosciences 6:530-533. doi:10.1038/ngeo1817

Ward ND, Finley RJ, Keil RG, Clay TG (2013a) Benefits and Limitations of iPads in the High School Science Classroom and a Trophic Cascade Lesson Plan. Journal of Geoscience Education 61:378-384. doi:10.5408/13-008.1

Martin EE, Ingalls AE, Richey JE, Keil RG, Santos GM, Truxal LT, Alin SR, Druffel ERM (2013) Age of riverine carbon suggests rapid export of terrestrial primary production in tropics. Geophysical Research Letters 40:5687-5691. doi:10.1002/2013gl057450


Hale R., C. Nittrouer, J. T. Liu, R. Keil, and A Ogston. 2012.  Effects of a Major Typhoon on Sediment Accumulation in Fangliao Submarine Canyon, SW Taiwan.  Marine Geology 326–328 (2012) 116–130,

Bridoux M., Annekov V., Menzel H, Keil, R., and Ingalls A. 2012. Widespread distribution and molecular diversity of diatom frustule bound aliphatic long chain polyamines (LCPAs) in Marine Sediments.  Organic Geochemistry 48, 9-20   DOI: 10.1016/j.orggeochem.2012.04.002

Bridoux M., Annekov V., Menzel H, Keil, R., and Ingalls A. 2012. Analysis of natural diatom communities reveals novel insights into the diversity of long chain polyamine (LCPA) structures involved in silica precipitation.  Organic Geochemistry 47 (2012) 9–21,

Ellis E.E., Ingalls A.E., Keil R.G. and Richey J.E. 2012.  Seasonal variability in the sources of particulate organic carbon of the Mekong River as discerned by elemental and lignin analyses.  Journal Of Geophysical Research, vol. 117, G01038, doi:10.1029/2011JG001816, 2012

Ward, N., Richey, J., and Keil. R 2012 Temporal variation in river nutrient and dissolved lignin phenol concentrations and the impact of storm events on nutrient loading to Hood Canal, Washington, USA.  Biogeochemistry. Biogeochemistry  DOI 10.1007/s10533-012-9700-9


Keil, R., Salemme, K., Forrest, B., Neibauer, J. and Logsdon, M., 2011. Differential presence of anthropogenic compounds dissolved in the marine waters of Puget Sound, WA and Barkley Sound, BC. Marine Pollution Bulletin, 62(11): 2404-2411.

Keil, R.G., 2011. Terrestrial influences on carbon burial at sea. Proceedings of the National Academy of Sciences, 108(24): 9729-9730.

Bridoux M., Annekov V., Menzel H, Keil, R., and Ingalls A. 2011. A new liquid chromatography/electrospray ionization mass spectrometry method for the analysis of underivatized aliphatic long chain polyamines.  Rapid Communications in Mass Spectrometry. 25(7) 877-888


Keil, R., Nuwer, J., and Strand S. 2010. Burial of Agricultural Byproducts in the Deep Sea as a form of Carbon Sequestration. Marine Chemistry122:91-95,  doi: 10.1016/j.marchem.2010.07.007

Ingalls, A., Ellis, E., Santos, G., McDuffee, K., Truxal, L., Keil, R., Druffel, E. 2010. HPLC Purification of higher plant-derived lignin phenols for compound specific radiocarbon analysis, Analytical Chemistry. Doi: 10.1021/ac1016584

Rezende, C., Pfeiffer W., Martinelli L., Tsamakis E., Hedges J and Keil R. 2010.  The diagenetic signature of lignin phenols delivered to Sepetiba Bay RJ, Brasil.  Estuarine, Coastal and Shelf Science 87, 479-486., doi:10.1016/j.ecss.2010.02.008.


Keil, R., and Neibauer, J. 2009.  Analysis of Cooking Spices in Natural Waters Limnology and Oceanography Methods 7: 848-855

Kimball, B., Myers-Pigg A., Clay T., Neibauer J., and Keil R.  2009.  SoundCitizen: Students and Citizens conducting environmental science together.  Proc NAAEE Symp 2009.


Walsh, E.M., Ingalls A.E. and Keil R.G. 2008. Sources and transport of terrestrial organic matter in Vancouver Island fjords and the Vancouver-Washington margin: A multiproxy approach using δ13Corg, lignin phenols and the ether lipid BIT index.  Limnology and Oceanography 53(3), 1054-1063.


Arnarson, T.S. and Keil, R.G. 2007. Importance of different physical protective mechanisms on organic matter preservation along a continuum of oxygen exposure time.  Geochimica et Cosmochimica Acta 71, 3545-3556.

Lau, W., Keil R.G. and Armbrust, E.V. 2007.  Succession and diel transcriptional response of the glycolate-utilizing component of the bacterial community during a spring phytoplankton bloom.  Applied and Environmental Microbiology Apr 2007, pp 2440-2450, doi:10.1128/AEM.01965-06


Nunn, B.A., and Keil, R.G. 2006. A comparison of non-hydrolytic methods for extracting amino acids and proteins from coastal marine sediments.  Marine Chemistry 98: 31-42. 


Nunn, B.A. and Keil, R.G. 2005. Size distribution and amino acid chemistry of base-extractable proteins from Washington coast sediments. Biogeochemistry 75: 177-200.

Nuwer, J. and Keil, R.G. 2005. Organic Carbon preservation in an intermittently anoxic fjord; Clayoquot Sound BC.  Limnology and Oceanography 50(4) 1119-1128.

Abell, J.T., Emerson, S. and Keil, R.G., 2005. Using preformed nitrate to infer decadal changes in DOM remineralization in the subtropical North Pacific. Global Biogeochemical Cycles, 19(GB1008, doi:10.1029/2004GB002285): 1-16.

Arnarson, T.S. and Keil, R.G., 2005. Influence of organic-mineral aggregates on microbial degradation of the dinoflagellate Scrippsiella trochoidea. Geochimica et Cosmochimica Acta, 69(8): 2111-2117.


Keil, R.G., Dickens, A.F., Arnarson, T., Nunn, B.L. and Devol, A.H., 2004. What is the oxygen exposure time of laterally transported organic matter along the Washington margin? Marine Chemistry, 92(1-4): 157-165.

Schnitzler Parker, M., Armbrust, E.V., Piovia-Scott, J. and Keil, R.G., 2004. Induction of photorespiration by light in the centric diatom Thallassiosira Weissflogii (Bacillariophyceae):  Molecular Characterization and Physiological Consequences. Journal of Phycology, 40(3): 557-567.

Van Mooy, B.A.S., Devol, A.H. and Keil, R.G., 2004a. Quantifying 3H-thymidine incorporation rates by a phylogenetically defined group of marine planktonic bacteria (Bacteriodetes phylum). Environmental Microbiology, 6(10): 1061–1069.

Van Mooy, B.A.S., Devol, A.H. and Keil, R.G., 2004b. Relationship between bacterial community structure, light, and carbon cycling in the eastern subarctic North Pacific. Limnology and Oceanography, 49(4): 1056–1062.


Nunn, B.L., Norbeck, A., and Keil R.G.  2003.  Hydrolysis patterns and the production of peptide intermediates during protein degradation in marine systems.  Marine Chemistry, 83 59-73.

Satterberg, J., T. Arnarson, E. Lessard and R.G. Keil.  2003.  Sorption of organic matter from four phytoplankton species to montmorillonite, chlorite and kaolinite in seawater.  Marine Chemistry 81 11-18.

Solomon, C., E. Lessard, R.G. Keil., and M. Foy.  2003.  Characterization of extracellular polymers of Phaeocystis globosa and P. antarctica  Marine Ecology Progress Series 250: 81-89


Van Mooy, B., R.G. Keil, and A.H. Devol.  2002  Impact of suboxia on sinking particulate organic carbon: enhanced flux and preferred degradation of amino acids via denitrification. Geochimica et cosmochimica acta, 66:457-465.

Van Mooy, B., and R.G. Keil.  2002.  Seasonal variations in sedimentary amino acids and the association of organic matter with mineral surfaces in a sandy eelgrass meadow. Marine Ecology Progress Series, 227: 275-280.


Arnarson T., and Keil R.G. 2001.  Organic-mineral interactions in marine sediments studied using density fractionation and X-ray photoelectron spectroscopy, Organic Geochemistry, 32:1401-1415.

Keil, R.G., and M.L. Fogel.  2001.  Stable carbon isotopic composition of amino acids along the Washington coast.  Limnology and Oceanography. 46, 14-23.

Holcombe, B., R.G. Keil, and A.H. Devol.  2001  Dissolved organic carbon fluxes from the suboxic sediment porewaters of the Mexican margin. Limnology and Oceanography, 46, 298-308

2000 and earlier

Arnarson T., and Keil R.G. 2000.  Mechanisms of natural organic matter sorption to montmorillonite surfaces in sea water. Marine Chemistry, 71, 309-320.

Keil, R.G., E. Tsamakis, and J.I. Hedges.  2000.  Early diagenesis of particulate amino acids in marine systems.  In: “Perspectives in Amino Acid and Protein Geochemistry” (G. A. Goodfriend, M. J. Collins, M. L. Fogel, S. A. Macko, and J. F. Wehmiller, eds.). Oxford University Press, NY.

Hedges, J.I., F.S. Hu, A.H. Devol, H.E. Hartnett, E. Tsamakis, and R.G. Keil.  1999  Sedimentary organic matter preservation: A test for selective degradation under oxic conditions. American Journal of Science, 299, 529-555.

Keil, R.G., and D.L. Kirchman.  1999  Utilization of dissolved protein and amino acids in the northern Sargasso sea. Aquatic Microbial Ecology.18, 293-300.

Keil, R.G., 1999  Early diagenesis of amino acids in high organic content marine sediments.  Geochemistry of the Earth’s Surface (Armannsson, H. ed) Balkema, Rotterdam.  259-262.

Arnarson, T.S. and Keil, R.G., 1999  Adsorption of marine pore water organic matter to montmorillonite.  Geochemistry of the Earth’s Surface (Armannsson, H. ed) Balkema, Rotterdam.  251-253.

Keil, R.G., and G.L. Cowie.  1999  Organic matter preservation through the oxygen minimum zone of the NE Arabian Sea as discerned by organic-mineral interactions. Marine Geology, 161, 13-22.

Hedges, J.I and R.G. Keil.  1999  Organic geochemical perspectives on estuarine processes: sorption reactions and consequences.   Marine Chemistry 65, 55-65.

Mayer, L., R.G. Keil, S.A. Macko, S.B. Joye, K.C. Ruttenberg and R.A. Aller.  1998  Importance of suspended particulates in riverine delivery of bioavailable nitrogen to coastal zones. Global Biogeochemical Cycles; 12, 573-579.

Keil, R.G., E. Tsamakis, J.C. Giddings and J.I. Hedges.  1998  Biochemical distributions (amino acids, neutral sugars and cupric oxide oxidation products) among size-classes of modern marine sediments.  Geochimica et Cosmochimica Acta, 62, 1347-1364.

Thimsen, C.A. and R.G. Keil.  1998  Potential interactions between sedimentary dissolved organic matter and mineral surfaces. Marine Chemistry, 62, 65-76.

Schmidt, J.L., J.W. Deming, P.A. Jumars and R.G. Keil,  1998  On the abundance and numeration of sedimentary bacteria; the need for new methods of normalization. Limnology and Oceanography,43, 976-982.

Hartnett, H. E., R.G. Keil, H.I. Hedges and A.H. Devol.  1998  Oxygen exposure time as a controlling factor in the presevation of organic carbon in continental margin sediments. Nature, 391, 572-574.

Keil, R.G., L. Mayer, P. Quay, J. Richey, and J.I. Hedges.  1997.  Loss of organic matter from riverine particles in deltas.  Geochimica et Cosmochimica Acta, 61;7  1507-1511.

Keil, R.G., E. Tsamakis, N. Wolf, J.I. Hedges, and M. Goni.  1997.  Relationships between Organic Carbon Preservation and Mineral Surface Area along the Amazon Fan (Sites 932A and 942A).  In:  Flood, R.D., Piper D.J.W., Klaus, A., and Peterson, L.C. (Eds). Proceedings of the Ocean Drilling Program , Scientific Results, Vol 155.  pp 531-538.

Ratanathanawongs Williams, S.K. and R.G. Keil.  1997.  The biological and physical reactivity of dextran carbohydrates in seawater incubations.  J. Liquid Chrom.  20(16&17): 2815-2833.

Hedges, J.I., R.G. Keil. and R. Benner.  1997.  What happens to land-derived organic matter in the ocean?  Organic Geochemistry,27, 195-212.

Cowie, G.L., Calvert, S., deLange, G., Keil, R., and J. Hedges.  1997.  The extent and implications of organic matter alteration at oxidation fronts in turbidites from the Madeira Abyssal Plain (Sites 951-952).  In:  Flood, R.D., Piper D.J.W., Klaus, A., and Peterson, L.C. (Eds). Proceedings of the Ocean Drilling Program , Scientific Results, Vol 157.

Bergamaschi, B.A., E. Tsamakis, R.G. Keil and J.I. Hedges.  1997.  The geochemistry of SPLITT-fractionated sediment from the Peru margin: effect of grain size and surface area on organic matter concentration and composition.  Geochimica et Cosmochimica Acta, 61:6, 1247-1260.

Keil, R.G., C.A Thimsen, E. Tsamakis, and J.I Hedges.  1995.  Evidence for Sorptive Preservation of Sedimentary Organic Matter. In: Organic Geochemistry: Developments and applications to energy, climate, environmental and human history. (Grimalt, J.O. and Dorron. C., eds) Donostia-San Sebastian, Spain. p850-853.

Hedges, J.I. and R.G. Keil.  1995.  Sedimentary organic matter preservation: an assessment and speculative synthesis. Mar. Chem. 49: 81-115.

Hedges, J.I. and R.G. Keil.  1995.  SOMP:  Authors closing comments. Mar. Chem. 49: 137-139.

Hedges, J.I., R.G. Keil, and G.L. Cowie.  Organic matter preservation in marine sediments: competition between sorptive protection and oxic degradation. In: Organic Geochemistry: Developments and applications to energy, climate, environmental and human history. (Grimalt, J.O. and Dorron. C., eds) Donostia-San Sebastian, Spain. p823-825.

Keil, R.G., D.B. Montluçon, F.G. Prahl, and J.I. Hedges.  1994.  Sorptive preservation of labile organic matter in marine sediments.  Nature; 370: 549-552.

Keil, R.G., F.S. Hu, E. Tsamakis, and J.I. Hedges.  1994.  Pollen in marine sediments trace oxygen-sensitive organic matter degradation.  Nature; 369: 639-641.

Keil, R.G., E. Tsamakis, C.B. Fuh, C.A. Giddings and J. Hedges.  1994.  Mineralogical and textural controls on the organic composition of coastal marine sediments: Hydrodynamic separation using SPLITT-fractionation. Geochimica et Cosmochimica Acta, 58(2): 879-893.

Keil, R.G., and D.L. Kirchman.  1994.  Abiotic transformation of dissolved protein to refractory protein in marine water.  Marine Chemistry, 45: 187-196.

Keil, R.G., and D.L. Kirchman.  1993.  Dissolved combined amino acids: chemical form and utilization by marine bacteria.  Limnology and Oceanography, 38(6): 1256-1270.

Keil, R.G., and J.I. Hedges.  1993.  Sorption of organic matter to mineral surfaces and the preservation of organic matter in coastal marine sediments.  Chemical Geology, 107: 385-388.

Hedges, J.I., R.G. Keil, and G.L. Cowie.  1993.  Sediment Diagenesis: organic perspectives with inorganic overlays.  Chemical Geology, 107: 487-492.

Kirchman, D.L., R.G. Keil, M. Simon, and N.A. Welschmeyer.  1993.  Biomass and production of heterotrophic bacterioplankton in the Oceanic Subarctic Pacific: SUPER 1987 and 1988. Deep Sea Research, 40: 967-988.

Keil, R.G., and D.L. Kirchman.  1992.  Bacterial utilization of protein and methylated protein and its implications for studies of protein degradation in aquatic systems.  Applied and Environmental Microbiology, 58(4) 1374-1375.

Kirchman, D.L., J. Moss, and R.G. Keil.  1992.  Evidence for nitrate uptake by heterotrophic bacteria in coastal waters of Delaware.  Arch. Hydrobiol. Beih,, 37:129-138

Keil, R.G., and D.L. Kirchman.  1991.  Contribution of amino acids and ammonium to the nitrogen requirements of heterotrophic bacterial assemblages in aquatic systems. Marine Ecology Progress Series 73:1-10.

Keil, R.G., and D.L. Kirchman.  1991.  Dissolved combined amino acids as estimated by a vapor phase hydrolysis method. Marine Chemistry, 33: 243-259.

Kirchman, D.L., R.G. Keil and P.A. Wheeler.  1990.  Carbon limitation of ammonium uptake by heterotrophic bacteria in the subarctic Pacific.  Limnology and Oceanography 35: 1258-1266.

Kirchman, D.L., R.G. Keil and P.A. Wheeler.  1989.  The effect of amino acids on ammonium utilization and regeneration by heterotrophic bacteria in the subarctic Pacific. Deep Sea Research, 36(11): 1763-1776.