Hidden biosphere in an oxygen-deficient Atlantic open-ocean eddy: Future implications of ocean deoxygenation on primary production in the eastern tropical North Atlantic

Biogeosciences

Volumn 12, Issue 24, 2015, Pages 7467-7482

  Loscher C R ^a,b   Fischer, M A ^a   Neulinger, S C ^a   Fiedler, B ^b   Philippi, M ^a   Schutte F ^b   Singh, A ^b,e   Hauss, H ^b   Karstensen, J ^b   Kortzinger A ^b,c   Kunzel S ^d   Schmitz, R A ^a  

^a UNIVERSITY HOSPITAL SCHLESWIG HOLSTEIN   (Germany)

^b GEOMAR HELMHOLTZ CENTRE FOR OCEAN RESEARCH KIEL   (Germany)

^c UNIVERSITY OF KIEL   (Germany)

^d MAX PLANCK INSTITUTE FOR EVOLUTIONARY BIOLOGY   (Germany)

^e PHYSICAL RESEARCH LABORATORY   (India)

Author keywords

[No Author keywords available]

Indexed keywords

ANOXIC CONDITIONS; BIOGEOCHEMISTRY; BIOSPHERE; CHEMICAL PROCESS; COMMUNITY STRUCTURE; ECOLOGICAL IMPACT; EDDY; FUTURE PROSPECT; MICROBIAL COMMUNITY; OPEN OCEAN; PRIMARY PRODUCTION; RESPIRATION; UPWELLING;

ATLANTIC OCEAN; ATLANTIC OCEAN (NORTH);

BACTERIA (MICROORGANISMS); PROCHLOROCOCCUS;

EID: 84951794780     PISSN: 17264170     EISSN: 17264189     Source Type: Journal    
DOI: 10.5194/bg-12-7467-2015     Document Type: Article

References (90)

1. Ahlgren, N., Rocap, G., and Chisholm, S.: Measurement of Prochlorococcus ecotypes using real-time polymerase chain reaction reveals different abundances of genotypes with similar light physiologies, Environ. Microbiol., 8, 441-454, 2006.
2. Altabet, M. A., Ryabenko, E., Stramma, L., Wallace, D. W. R., Frank, M., Grasse, P., and Lavik, G.: An eddy-stimulated hotspot for fixed nitrogen-loss from the Peru oxygen minimum zone, Biogeosciences, 9, 4897-4908, doi:10.5194/bg-9-4897-2012, 2012.
3. Angel, R., Matthies, D., and Conrad, R.: Activation of Methanogenesis in Arid Biological Soil Crusts Despite the Presence of Oxygen, PlosOne, 6, e20453, doi:10.1371/journal.pone.0020453, 2011.
4. Arévalo-Martínez, D. L., Kock, A., Löscher, C. R., Schmitz, R. A., Stramma, L., and Bange, H.W.: Influence of mesoscale eddies on the distribution of nitrous oxide in the eastern tropical South Pacific, Biogeosciences Discuss., 12, 9243-9273, doi:10.5194/bgd-12-9243-2015, 2015.
5. Baird, M. E., Suthers, I.M., Griffin, D. A., Hollings, B., Pattiaratchi, C., Everett, J. D., Roughan, M., Oubelkheir, K., and Doblin, M.: The effect of surface flooding on the physical-biogeochemical dynamics of a warm-core eddy off southeast Australia, Deep-Sea Res. Pt. II, 58, 592-605, 2011.
6. Bauer, M., Kube, M., Teeling, H., Richter, M., Lombardot, T., Allers, E., Würdemann, C. A., Quast, C., Kuhl, H., Knaust, F., Woebken, D., Bischof, K., Mussmann, M., Choudhuri, J. V., Meyer, F., Reinhardt, R., Amann, R. I., and Glöckner, F. O.: Whole genome analysis of the marine Bacteroidetes'Gramella forsetii' reveals adaptations to degradation of polymeric organic matter, Environ. Microbiol., 8, 2201-2213, 2006.
7. Beman, J. M. and Carolan, M. T.: Deoxygenation alters bacterial diversity and community composition in the ocean's largest oxygen minimum zone, Nat. Commun., 4, 2705, doi:10.1038/ncomms3705, 2013.
8. Beman, J. M., Chow, C. E., King, A. L., Feng, Y. Y., Fuhrman, J. A., Andersson, A., Bates, N. R., Popp, B. N., and Hutchins, D. A.: Global declines in oceanic nitrification rates as a consequence of ocean acidification, P. Natl. Acad. Sci. USA, 108, 208-213, doi:10.1073/pnas.1011053108, 2011.
9. Benjamini, Y. and Hochberg, Y.: Controlling the false discovery rate: A practical and powerful approach to multiple testing, J. Roy. Stat. Soc. B, 57, 289-300, 1995.
10. Bertillson, S., Berglund, O., Pullin, M. J., and Chisholm, S.W.: Release of dissolved organic matter by Prochlorococcus, ie et milieu - Life and environment, 55, 225-231, 2005.
11. Berube, P. M., Biller, S. J., Kent, A. G., Berta-Thompson, J. W., Roggensack, S. E., Roache-Johnson, K. H., Ackerman, M., Moore, L. R., Meisel, J. D., Sher, D., Thompson, L. R., Campbell, L., Martiny A., and Chisholm, S.W.: Physiology and evolution of nitrate acquisition in Prochlorococcus, ISME J., 9, 1195-1207, doi:10.1038/ismej.2014.211, 2014.
12. Biller, S. J., Berube, P. M., Lindell, D., and Chisholm, S. W.: Prochlorococcus: the structure and function of collective diversity, Nat. Rev. Microbiol., 13, 13-27, 2015.
13. Brown, M. V., Ostrowski, M., Grzymski, J. J., and Lauro, F. M.: A trait based perspective on the biogeography of common and abundant marine bacterioplankton clades, Marine Genomics, 15, 17-28, 2014.
14. Bryant, J. A., Stewart, F. J., Eppley, J. M., and DeLong, E. F.: Microbial community phylogenetic and trait diversity declines with depth in a marine oxygen minimum zone, Ecology, 93, 1659-1673, 2012.
15. Buchan, A., LeCleir, G. R., Gulvik, C. A., and González, J. M.: Master recyclers: features and functions of bacteria associated with phytoplankton blooms, Nat. Rev. Microbiol., 12, 686-698, doi:10.1038/nrmicro3326, 2014.
16. Carini, P., Steindler, L., Beszteri, S., and Giovannoni, S. J.: Nutrient requirements for growth of the extreme oligotroph "Candidatus Pelagibacter ubique" HTCC1062 on a defined medium, ISME J., 7, 592-602, 2013.
17. Chavez, F. P. and Messié, M.: A comparison of Eastern Boundary Upwelling Ecosystems, Prog. Oceanogr., 83, 80-96, 2009.
18. Chelton, D. B., Gaube, P., Schlax, M. G., Early, J. J., and Samelson, R. M.: The Influence of Nonlinear Mesoscale Eddies on Near-Surface Oceanic Chlorophyll, Science, 334, 328-332, 2011.
19. Chen, V. B., Davis, I. W., and Richardson, D. C.: KING (Kinemage, Next Generation): A versatile interactive molecular and scientific visualization program, Prot. Sci., 18, 2403-2409, doi:10.1002/pro.250, 2009.
20. Cole, J. R., Wang, Q., Fish, J. A., Chai, B., McGarrell, D. M., Sun, Y., Brown, C. T., Porras-Alfaro, A., Kuske, C. R., and Tiedje, J. M.: Ribosomal Database Project: data and tools for high throughput rRNA analysis, Nucleic Acids Res., 42, 633-642, doi:10.1093/nar/gkt1244, 2014.
21. De Cáceres, M. and Legendre, P.: Associations between species and groups of sites: indices and statistical inference, Ecology, 90, 3566-3574, doi:10.1890/08-1823.1, 2009.
22. DeLong, E. F.: The microbial ocean from genomes to biomes, Nature, 459, 200-206, doi:10.1038/nature08059, 2009.
23. DeSantis, T. Z., Hugenholtz, P., Larsen, N., Rojas, M., Brodie, E. L., Keller, K., Huber, T., Dalevi, D., Hu, P., and Andersen, G. L.: Greengenes, a chimera-checked 16S rRNA gene database and workbench compatible with ARB, Appl. Environ. Microb., 72, 5069-5072, doi:10.1128/aem.03006-05, 2006.
24. Deutsch, C., Sarmiento, J. L., Sigman, D. M., Gruber, N., and Dunne, J. P.: Spatial coupling of nitrogen inputs and losses in the ocean, Nature, 445, 163-167, doi:10.1038/nature05392, 2007.
25. Edgar, R. C., Haas, B. J., Clemente, J. C., Quince, C., and Knight, R.: UCHIME improves sensitivity and speed of chimera detection, Bioinformatics, 27, 2194-2200, doi:10.1093/bioinformatics/btr381, 2011.
26. Fernandez, C., Farias, L., and Ulloa, O.: Nitrogen Fixation in Denitrified Marine Waters, Plos One, 6, e20539, doi:10.1371/journal.pone.0020539, 2011.
27. Fiedler, B., Grundle, D., Hauss, H., Krahmann, G., Schütte, F., Monteiro, I., Silva, P., Vieira, N., and Körtzinger, A.: Biogeochemistry of oxygen depleted mesoscale Eddies in the open eastern tropical North Atlantic, Biogeosciences Discuss., in preparation, 2015.
28. Fierer, N., Hamady, M., Lauber, C. L., and Knight, R.: The influence of sex, handedness, and washing on the diversity of hand surface bacteria, P. Natl. Acad. Sci. USA, 105, 17994-17999, doi:10.1073/pnas.0807920105, 2008.
29. Fischer, G., Karstensen, J., Romero, O., Baumann, K.-H., Donner, B., Hefter, J., Mollenhauer, G., Iversen, M., Fiedler, B., Monteiro, I., and Körtzinger, A.: Bathypelagic particle flux signatures from a suboxic eddy in the oligotrophic tropical North Atlantic: production, sedimentation and preservation, Biogeosciences Discuss., 12, 18253-18313, doi:10.5194/bgd-12-18253-2015, 2015.
30. Flombaum, P., Gallegos, J. L., Gordillo, R. A; Rincon, J., Zabala, L. L., Jiao, N., Karld, D. M., Li, W. K. W., Lomas, M. W., Veneziano, D., Vera, C. S., Vrugt, J. A., and Martiny, A. C.: Present and future global distributions of the marine Cyanobacteria Prochlorococcus and Synechococcus, P. Natl. Acad. Sci. USA, 110, 9824-9829, 2013.
31. Forth, M., Liljebladh, B., Stigebrandt, A., Hall, P. O. J., and Treusch, A. H.: Effects of ecological engineered oxygenation on the bacterial community structure in an anoxic fjord in western Sweden, ISME J., 9, 656-669, doi:10.1038/ismej.2014.172, 2014.
32. Frank, D. N., St Amand, A. L., Feldman, R. A., Boedeker, E. C., Harpaz, N., and Pace, N. R.: Molecular-phylogenetic characterization of microbial community imbalances in human inflammatory bowel diseases, P. Natl. Acad. Sci. USA, 104, 13780-13785, doi:10.1073/pnas.0706625104, 2007.
33. Franz, J. M. S., Hauss, H., Sommer, U., Dittmar, T., and Riebesell, U.: Production, partitioning and stoichiometry of organic matter under variable nutrient supply during mesocosm experiments in the tropical Pacific and Atlantic Ocean, Biogeosciences, 9, 4629-4643, doi:10.5194/bg-9-4629-2012, 2012.
34. Ganesh, S., Parris, D. J., DeLong, E. F., and Stewart, F. J.: Metagenomic analysis of size-fractionated picoplankton in a marine oxygen minimum zone, ISME J., 8, 187-211, 2014.
35. Gao, H., Schreiber, F., Collins, G., Jensen, M. M., Kostka, J. E., Lavik, G., de Beer, D., Zhou, H.-Y., and Kuypers, M. M. M.: Aerobic denitrification in permeable Wadden Sea sediments, ISME J., 4, 417-426, 2010.
36. Giovannoni, S. J., Britschgi, T. B., Moyer, C. L., and Field, K. G.: Genetic diversity in Sargasso Sea bacterioplankton, Nature, 345, 60-63, doi:10.1038/345060a0, 1990.
37. Glöckner, F. O., Fuchs, B. M., and Amann, R.: Bacterioplankton compositions of lakes and oceans: a first comparison based on fluorescence in situ hybridization, Environ Microbiol., 65, 3721-3726, 1999.
38. Grasshoff, G., Kremling, K., and Erhardt, M.: Methods of seawater analysis, 3 Edn., Wiley VCH, Weinheim, 1999.
39. 2O activity in an oxygenated ocean, Nature Scientific Reports, submitted, 2015.
40. Hauss, H., Christiansen, S., Schütte, F., Kiko, R., Edvam Lima, M., Rodrigues, E., Karstensen, J., Löscher, C. R., Körtzinger, A., and Fiedler, B.: Dead zone or oasis in the open ocean? Zooplankton distribution and migration in low-oxygen modewater eddies, Biogeosciences Discuss., 12, 18315-18344, doi:10.5194/bgd-12-18315-2015, 2015.
41. Hood, E. M., Sabine, C. L., and Sloyan, B. M.: The GO-SHIP Repeat Hydrography Manual: A Collection of Expert Reports and Guidelines, IOCCP Report Number 14, ICPO Publication Series Number 134, available at: http://www.go-ship.org/HydroMan.html (last access: 19 March 2013), 2010.
42. Jickells, T. D., An, Z. S., Andersen, K. K., Baker, A. R., Bergametti, G., Brooks, N., Cao, J. J., Boyd, P. W., Duce, R. A., Hunter, K. A., Kawahata, H., Kubilay, N., laRoche, J., Liss, P. S., Mahowald, N., Prospero, J. M., Ridgwell, A. J., Tegen, I., and Torres, R.: Global iron connections between desert dust, ocean biogeochemistry, and climate, Science, 308, 67-71, 2005.
43. Jost, L.: Entropy and diversity, Oikos, 113, 363-375, doi:10.1111/j.2006.0030-1299.14714.x, 2006.
44. Jost, L.: Partitioning diversity into independent alpha and beta components, Ecology, 88, 2427-2439, doi:10.1890/06-1736.1, 2007.
45. Kalvelage, T., Lavik, G., Lam, P., Contreras, S., Artega, L., Löscher, C. R., Oschlies, A., Paulmier, A., Stramma, L., and Kuypers, M. M. M.: Nitrogen cycling driven by organic matter export in the South Pacific oxygen minimum zone, Nat. Geosci., 6, 228-234, 2013.
46. Karstensen, J., Stramma, L., and Visbeck, M.: Oxygen minimum zones in the eastern tropical Atlantic and Pacific oceans, Prog. Oceanogr., 77, 331-350, doi:10.1016/j.pocean.2007.05.009, 2008.
47. Karstensen, J., Fiedler, B., Schütte, F., Brandt, P., Körtzinger, A., Fischer, G., Zantopp, R., Hahn, J., Visbeck, M., and Wallace, D.: Open ocean dead zones in the tropical North Atlantic Ocean, Biogeosciences, 12, 2597-2605, doi:10.5194/bg-12-2597-2015, 2015a.
48. Karstensen, J., Schütte, F., Pietri, A., Krahmann, G., Fiedler, B., Körtzinger, A., Löscher, C. R., Grundle, D., and Hauss, H.: Anatomy of open ocean dead-zones based on high-resolution multidisciplinary glider data, Biogeosciences Discuss., in preparation, 2015b.
49. Kozich, J. J., Westcott, S. L., Baxter, N. T., Highlander, S. K., and Schloss, P. D.: Development of a dual-index sequencing strategy and curation pipeline for analyzing amplicon sequence data on the MiSeq Illumina sequencing platform, Appl. Environ. Microb., 79, 5112-5120, doi:10.1128/aem.01043-13, 2013.
50. Lam, P., Jensen, M. M., Lavik, G., McGinnis, D. F., Muller, B., Schubert, C. J., Amann, R., Thamdrup, B., and Kuypers, M. M. M.: Linking crenarchaeal and bacterial nitrification to anammox in the Black Sea, P. Natl. Acad. Sci. USA, 104, 7104-7109, doi:10.1073/pnas.0611081104, 2007.
51. Langfeldt, D., Neulinger, S. C., Heuer, W., Staufenbiel, I., Künzel, S., Baines, J. F., Eberhard, J., and Schmitz, R. A.: Composition of microbial oral biofilms during maturation in young healthy adults, PlosOne, 9, e87449, doi:10.1371/journal.pone.0087449, 2014.
52. Langlois, R. J., Hummer, D., and LaRoche, J.: Abundances and distributions of the dominant nifH phylotypes in the Northern Atlantic Ocean, Appl. Environ. Microb., 74, 1922-1931, doi:10.1128/aem.01720-07, 2008.
53. Lévy, M., Klein, P., and Treguier, A.-M.: Impacts of sub-mesoscale physics on phytoplankton production and subduction, J. Mar. Res., 59, 535-565, doi:10.1357/002224001762842181, 2001.
54. Lévy, M., Ferrari, R., Franks, P. J. S., Martin, A. P., and Riviere, P.: Bringing physics to life at the submesoscale, Geophys. Res. Lett., 39, L14602, doi:10.1029/2012gl052756, 2012.
55. Löscher, C. R., Kock, A., Könneke, M., LaRoche, J., Bange, H. W., and Schmitz, R. A.: Production of oceanic nitrous oxide by ammonia-oxidizing archaea, Biogeosciences, 9, 2419-2429, doi:10.5194/bg-9-2419-2012, 2012.
56. Löscher, C. R., Großkopf, T., Desai, F., Gill, D., Schunck, H., Croot, P., Schlosser, C., Neulinger, S. C., Lavik, G., Kuypers, M. M. M., LaRoche, J., and Schmitz, R. A.: Facets of diazotrophy in the oxygen minimum zone off Peru, ISME J., 8, 2180-2192, doi:10.1038/ismej.2014.71, 2014.
57. Löscher, C. R., Bange, H. W., Schmitz, R. A., Callbeck, C. M., Engel, A., Hauss, H., Kanzow, T., Kiko, R., Lavik, G., Loginova, A., Melzner, F., Neulinger, S. C., Pahlow, M., Riebesell, U., Schunck, H., Thomsen, S., and Wagner, H.: Water column biogeochemistry of oxygen minimum zones in the eastern tropical North Atlantic and eastern tropical South Pacific Oceans, Biogeosciences Discuss., 12, 4495-4556, doi:10.5194/bgd-12-4495-2015, 2015.
58. Mahadevan, A.: Ocean science: Eddy effects on biogeochemistry, Nat. Geosci., 506, 168-169, doi:10.1038/nature13048, 2014.
59. McGillicuddy, D. J., Anderson, L. A., Bates, N. R., Bibby, T., Buesseler, K. O., Carlson, C. A., Davis, C. S., Ewart, C., Falkowski, P. G., Goldthwait, S. A., Hansell, D. A., Jenkins, W. J., Johnson, R., Kosnyrev, V. K., Ledwell, J. R., Li, Q. P., Siegel, D. A., and Steinberg, D. K.: Eddy/wind interactions stimulate extraordinary mid-ocean plankton blooms, Science, 316, 1021-1026, doi:10.1126/science.1136256, 2007.
60. Morris, J. J., Kirkegaard, R., Szul, M. J., Johnson, Z. I., and Zinser, E. R.: Facilitation of robust growth of Prochlorococcus colonies and dilute liquid cultures by "helper" heterotrophic bacteria, Appl. Environ. Microb., 74, 4530-4534, 2008.
61. Morris, R. M., Rappé, M. S., Connon, S. A., Vergin, K. L., Siebold, W. A., Carlson, C. A., and Giovannoni, S. J.: SAR11 clade dominates ocean surface bacterioplankton communities, Nature, 420, 806-810, doi:10.1038/nature01240, 2002.
62. Oksanen, J., Blanchet, F. G., Kindt, R., Legendre, P., and Minchin, P. R.: vegan: Community ecology package, R package version 2.0-6., 2013.
63. Oschlies, A. and Garcon, V.: Eddy-induced enhancement of primary production in a model of the North Atlantic Ocean, Nature, 394, 266-269, 1998.
64. Parsons, T. R., Maita, Y., and Lalli, C. M.: A Manual of Chemical and Biological Methods for Seawater Analysis, Pergamon Press, Oxford, 173 pp., 1984.
65. Pinheiro, J. C., Bates, D. M., DebRoy, S., Sarkar, D., and R Core Team: nlme: Linear and nonlinear mixed effects models, R package version 3.1-120 ed., 2015.
66. Poretsky, R. S., Hewson, I., Sun, S. L., Allen, A. E., Zehr, J. P., and Moran, M. A.: Comparative day/night metatranscriptomic analysis of microbial communities in the North Pacific subtropical gyre, Environ. Microbiol., 11, 1358-1375, doi:10.1111/j.1462-2920.2008.01863.x, 2009.
67. Pruesse, E., Quast, C., Knittel, K., Fuchs, B. M., Ludwig, W., Peplies, J., and Glöckner, F. O.: SILVA: a comprehensive online resource for quality checked and aligned ribosomal RNA sequence data compatible with ARB, Nucleic Acids Res., 35, 7188-7196, doi:10.1093/nar/gkm864, 2007.
68. Rappé, M. S., Connon, S. A., Vergin, K. L., and Giovannoni, S. J.: Cultivation of the ubiquitous SAR11 marine bacterioplankton clade, Nature, 418, 630-633 doi:10.1038/nature00917, 2002.
69. R Core Team: R: A language and environment for statistical computing, R Foundation for Statistical Computing, Vienna, Austria, 2015.
70. Richardson, D. C. and Richardson, J. S.: The kinemage: A tool for scientific communication, Protein Sci., 1, 3-9, doi:10.1002/pro.5560010102, 1992.
71. Riebesell, U. and Gattuso, J.-P.: Lessons learned from ocean acidification research, Nature Climate Change, 5, 12-14, doi:10.1038/nclimate2456, 2015.
72. Ryabenko, E., Kock, A., Bange, H.W., Altabet, M. A., and Wallace, D.W. R.: Contrasting biogeochemistry of nitrogen in the Atlantic and Pacific Oxygen Minimum Zones, Biogeosciences, 9, 203-215, doi:10.5194/bg-9-203-2012, 2012.
73. Schloss, P. D., Westcott, S. L., Ryabin, T., Hall, J. R., Hartmann, M., Hollister, E. B., Lesniewski, R. A., Oakley, B. B., Parks, D. H., Robinson, C. J., Sahl, J. W., Stres, B., Thallinger, G. G., Van Horn, D. J., and Weber, C. F.: Introducing mothur: Open-source, platform-independent, community-supported software for describing and comparing microbial communities, Appl. Environ. Microb., 75, 7537-7541, doi:10.1128/aem.01541-09, 2009.
74. Schmid, M. C., Hooper, A. B., Klotz, M. G., Woebken, D., Lam, P., Kuypers, M. M. M., Pommerening-Roeser, A., op den Camp, H. J. M., and Jetten, M. S. M.: Environmental detection of octahaem cytochrome c hydroxylamine/hydrazine oxidoreductase genes of aerobic and anaerobic ammonium-oxidizing bacteria, Environ. Microbiol., 10, 3140-3149, doi:10.1111/j.1462-2920.2008.01732.x, 2008.
75. Schütte, F., Karstensen, J., Krahmann, G., Fiedler, B., Brandt, P., Visbeck, M., and Körtzinger, A.: Characterization of "dead-zone eddies" in the tropical North Atlantic Ocean, Biogeosciences Discuss., in preparation, 2015.
76. Stewart, F. J., Ulloa, O., and DeLong, E. F.: Microbial meta-transcriptomics in a permanent marine oxygen minimum zone, Environ. Microbiol., 14, 23-40, doi:10.1111/j.1462-2920.2010.02400.x, 2011.
77. Stramma, L., Johnson, G. C., Sprintall, J., and Mohrholz, V.: Expanding oxygen-minimum zones in the tropical oceans, Science, 320, 655-658, doi:10.1126/science.1153847, 2008.
78. Stramma, L., Bange, H. W., Czeschel, R., Lorenzo, A., and Frank, M.: On the role of mesoscale eddies for the biological productivity and biogeochemistry in the eastern tropical Pacific Ocean off Peru, Biogeosciences, 10, 7293-7306, doi:10.5194/bg-10-7293-2013, 2013.
79. Stratil, S. B., Neulinger, S. C., Knecht, H., Friedrichs, A. K., and Wahl, M.: Temperature-driven shifts in the epibiotic bacterial community composition of the brown macroalga Fucus vesiculosus, MicrobiologyOpen, 2, 338-349, doi:10.1002/mbo3.79, 2013.
80. Stratil, S. B., Neulinger, S. C., Knecht, H., Friedrichs, A. K., and Wahl, M.: Salinity affects compositional traits of epibacterial communities on the brown macroalga Fucus vesiculosus, FEMS Microbiol. Ecol., 88, 272-279, doi:10.1111/1574-6941.12292, 2014.
81. Suzuki, M., Preston, C., Chavez, F., and DeLong, E.: Quantitative mapping of bacterioplankton populations in seawater: field tests across an upwelling plume in Monterey Bay, Aquat. Microb. Ecol., 24, 117-127, 2001.
82. Swan, B. K., Martinez-Garcia, M., Preston, C. M., Sczyrba, A., Woyke, T., Lamy, D., Reinthaler, T., Poulton, N. J., Masland, E. D. P., Gomez, M. L., Sieracki, M. E., DeLong, E. F., Herndl, G. J., and Stepanauskas, R.: Potential for chemolithoautotrophy among ubiquitous bacteria lineages in the dark ocean, Science, 333, 1296-1300, doi:10.1126/science.1203690, 2011.
83. Ulloa, O., Canfield, D. E., DeLong, E. F., Letelier, R. M., and Stewart, F. J.: Microbial oceanography of anoxic oxygen minimum zones, P. Natl. Acad. Sci. USA, 109, 15996-16003, doi:10.1073/pnas.1205009109, 2012.
84. Vaquer-Sunyer, R. and Duarte, C. M.: Thresholds of hypoxia for marine biodiversity, P. Natl. Acad. Sci. USA, 105, 15452-15457, 2008.
85. Wang, Q., Garrity, G. M., Tiedje, J. M., and Cole, J. R.: Naïve Bayesian Classifier for Rapid Assignment of rRNA Sequences into the New Bacterial Taxonomy, Appl. Environ. Microb., 73, 5261-5267 doi:10.1128/AEM.00062-07, 2007.
86. West, N. J., Lebaron, P., Strutton, P. G., and Suzuki, M. T.: A novel clade of Prochlorococcus found in high nutrient low chlorophyll waters in the South and Equatorial Pacific Ocean, ISME J., 5, 933-944, 2011.
87. Wright, J. J., Konwar, K. M., and Hallam, S. J.: Microbial ecology of expanding oxygen minimum zones, Nat. Rev. Microbiol., 10, 381-394, doi:10.1038/nrmicro2778, 2012.
88. Yu, Y., Lee, C., Kim, J., and Hwang, S.: Group-specific primer and probe sets to detect methanogenic communities using quantitative real-time polymerase chain reaction, Biotechnol. Bioeng., 89, 670-679, doi:10.1002/bit.20347, 2005.
89. Zhu, M., and Ghodsi, A.: Automatic dimensionality selection from the scree plot via the use of profile likelihood, Comput. Stat. Data An., 51, 918-930, doi:10.1016/j.csda.2005.09.010, 2006.
90. Zuur, A. F., Ieno, E. N., Walker, N. J., Saveliev, A. A., and Smith, G. M.: Mixed effects models and extensions in ecology with R, Statistics for Biology and Health, edited by: Gail, M., Krickeberg, K., Samet, J. M., Tsiatis, A., and Wong, W., Springer, New York, USA, 2009.