Deep ocean metagenomes provide insight into the metabolic architecture of bathypelagic microbial communities.
Journal
Communications biology
ISSN: 2399-3642
Titre abrégé: Commun Biol
Pays: England
ID NLM: 101719179
Informations de publication
Date de publication:
21 05 2021
21 05 2021
Historique:
received:
25
09
2020
accepted:
16
04
2021
entrez:
22
5
2021
pubmed:
23
5
2021
medline:
10
8
2021
Statut:
epublish
Résumé
The deep sea, the largest ocean's compartment, drives planetary-scale biogeochemical cycling. Yet, the functional exploration of its microbial communities lags far behind other environments. Here we analyze 58 metagenomes from tropical and subtropical deep oceans to generate the Malaspina Gene Database. Free-living or particle-attached lifestyles drive functional differences in bathypelagic prokaryotic communities, regardless of their biogeography. Ammonia and CO oxidation pathways are enriched in the free-living microbial communities and dissimilatory nitrate reduction to ammonium and H
Identifiants
pubmed: 34021239
doi: 10.1038/s42003-021-02112-2
pii: 10.1038/s42003-021-02112-2
pmc: PMC8139981
doi:
Substances chimiques
DNA, Bacterial
0
Types de publication
Journal Article
Research Support, Non-U.S. Gov't
Research Support, U.S. Gov't, Non-P.H.S.
Langues
eng
Sous-ensembles de citation
IM
Pagination
604Références
Cho, B. C. & Azam, F. major role of bacteria in biogeochemical fluxes in the ocean´s interior. Nature 332, 441–443 (1988).
doi: 10.1038/332441a0
Bar-On, Y. M., Phillips, R. & Milo, R. The biomass distribution on Earth. Proc. Natl Acad. Sci. USA 115, 6506–6511 (2018).
Aristegui, J., Gasol, J. M., Duarte, C. M. & Herndl, G. J. Microbial oceanography of the dark ocean’s pelagic realm. Limnol. Oceanogr. 54, 1501–1529 (2009).
doi: 10.4319/lo.2009.54.5.1501
Baltar, F., Arístegui, J., Gasol, J. M., Lekunberri, I. & Herndl, G. J. Mesoscale eddies: hotspots of prokaryotic activity and differential community structure in the ocean. ISME J. 4, 975–988 (2010).
pubmed: 20357833
doi: 10.1038/ismej.2010.33
Del Giorgio, P. A. & Duarte, C. M. Respiration in the open ocean. Nature 420, 379–384 (2002).
pubmed: 12459775
doi: 10.1038/nature01165
Arístegui, J. et al. Oceanography: dissolved organic carbon support of respiration in the dark ocean. Science 298, 1967 (2002).
pubmed: 12471250
doi: 10.1126/science.1076746
Herndl, G. J. & Reinthaler, T. Microbial control of the dark end of the biological pump. Nat. Geosci. 6, 718–724 (2013).
pubmed: 24707320
pmcid: 3972885
doi: 10.1038/ngeo1921
Baltar, F. et al. Significance of non-sinking particulate organic carbon and dark CO
doi: 10.1029/2010GL043105
Boyd, P. W., Claustre, H., Levy, M., Siegel, D. A. & Weber, T. Multi-faceted particle pumps drive carbon sequestration in the ocean. Nature 568, 327–335 (2019).
pubmed: 30996317
doi: 10.1038/s41586-019-1098-2
Stukel, M. R., Song, H., Goericke, R. & Miller, A. J. The role of subduction and gravitational sinking in particle export, carbon sequestration, and the remineralization length scale in the California Current Ecosystem. Limnol. Oceanogr. 63, 363–383 (2018).
doi: 10.1002/lno.10636
Omand, M. M. et al. Eddy-driven subduction exports particulate organic carbon from the spring bloom. Science 348, 222–225 (2015).
pubmed: 25814062
doi: 10.1126/science.1260062
Jónasdóttir, S. H., Visser, A. W., Richardson, K. & Heath, M. R. Seasonal copepod lipid pump promotes carbon sequestration in the deep North Atlantic. Proc. Natl Acad. Sci. USA 112, 12122–12126 (2015).
pubmed: 26338976
doi: 10.1073/pnas.1512110112
pmcid: 4593097
Dall’Olmo, G., Dingle, J., Polimene, L., Brewin, R. J. W. & Claustre, H. Substantial energy input to the mesopelagic ecosystem from the seasonal mixed-layer pump. Nat. Geosci. 9, 820–823 (2016).
pubmed: 27857779
pmcid: 5108409
doi: 10.1038/ngeo2818
Herndl, G. J. et al. Contribution of archaea to total prokaryotic production in the deep Atlantic Ocean. Appl. Environ. Microbiol. 71, 2303–2309 (2005).
pubmed: 15870315
pmcid: 1087563
doi: 10.1128/AEM.71.5.2303-2309.2005
Wuchter, C. et al. Archaeal nitrification in the ocean. Proc. Natl Acad. Sci. USA 103, 12317–12322 (2006).
pubmed: 16894176
doi: 10.1073/pnas.0600756103
pmcid: 1533803
Reinthaler, T., van Aken, H. M. & Herndl, G. J. Major contribution of autotrophy to microbial carbon cycling in the deep North Atlantic’s interior. Deep Res. Part II Top. Stud. Oceanogr. 57, 1572–1580 (2010).
doi: 10.1016/j.dsr2.2010.02.023
Swan, B. K. et al. Potential for chemolithoautotrophy among ubiquitous bacteria lineages in the dark ocean. Science 333, 1296–1300 (2011).
pubmed: 21885783
doi: 10.1126/science.1203690
Pachiadaki, M. G. et al. Major role of nitrite-oxidizing bacteria in dark ocean carbon fixation. Science 358, 1046–1051 (2017).
pubmed: 29170234
doi: 10.1126/science.aan8260
Hügler, M. & Sievert, S. M. Beyond the Calvin Cycle: autotrophic carbon fixation in the ocean. Ann. Rev. Mar. Sci. 3, 261–289 (2011).
pubmed: 21329206
doi: 10.1146/annurev-marine-120709-142712
Sorokin, D. Y. Oxidation of inorganic sulfur compounds by obligately organotrophic bacteria. Microbiology 72, 641–653 (2003).
doi: 10.1023/B:MICI.0000008363.24128.e5
Alonso-Sáez, L., Galand, P. E., Casamayor, E. O., Pedrós-Alió, C. & Bertilsson, S. High bicarbonate assimilation in the dark by Arctic bacteria. ISME J. 4, 1581–1590 (2010).
pubmed: 20555365
doi: 10.1038/ismej.2010.69
Turner, J. T. Zooplankton fecal pellets, marine snow and sinking phytoplankton blooms. Aquat. Microb. Ecol. 27, 57–102 (2002).
doi: 10.3354/ame027057
Ploug, H., Iversen, M. H. & Fischer, G. Ballast, sinking velocity, and apparent diffusivity within marine snow and zooplankton fecal pellets: implications for substrate turnover by attached bacteria. Limnol. Oceanogr. 53, 1878–1886 (2008).
doi: 10.4319/lo.2008.53.5.1878
Agusti, S. et al. Ubiquitous healthy diatoms in the deep sea confirm deep carbon injection by the biological pump. Nat. Commun. 6, 7608 (2015).
pubmed: 26158221
doi: 10.1038/ncomms8608
Smith, K. L., Ruhl, H. A., Huffard, C. L., Messié, M. & Kahru, M. Episodic organic carbon fluxes from surface ocean to abyssal depths during long-term monitoring in NE Pacific. Proc. Natl Acad. Sci. USA 115, 12235–12240 (2018).
pubmed: 30429327
doi: 10.1073/pnas.1814559115
pmcid: 6275536
Salazar, G. et al. Global diversity and biogeography of deep-sea pelagic prokaryotes. ISME J. 10, 596–608 (2016).
pubmed: 26251871
doi: 10.1038/ismej.2015.137
Mestre, M. et al. Sinking particles promote vertical connectivity in the ocean microbiome. Proc. Natl Acad. Sci. USA 115, E6799–E6807 (2018).
pubmed: 29967136
doi: 10.1073/pnas.1802470115
pmcid: 6055141
Salazar, G. et al. Particle-association lifestyle is a phylogenetically conserved trait in bathypelagic prokaryotes. Mol. Ecol. 24, 5692–5706 (2015).
pubmed: 26462173
doi: 10.1111/mec.13419
DeLong, E. F. et al. Community genomics among stratified microbial assemblages in the ocean’s interior. Science 311, 496–503 (2006).
pubmed: 16439655
doi: 10.1126/science.1120250
Martín-Cuadrado, A.-B. et al. Metagenomics of the deep mediterranean, a warm bathypelagic habitat. PLoS ONE 2, e914 (2007).
pubmed: 17878949
pmcid: 1976395
doi: 10.1371/journal.pone.0000914
Boeuf, D. et al. Biological composition and microbial dynamics of sinking particulate organic matter at abyssal depths in the oligotrophic open ocean. Proc. Natl Acad. Sci. USA 116, 11824–11832 (2019).
pubmed: 31127042
pmcid: 6575173
doi: 10.1073/pnas.1903080116
Ganesh, S. et al. Size-fraction partitioning of community gene transcription and nitrogen metabolism in a marine oxygen minimum zone. ISME J. 9, 2682–2696 (2015).
pubmed: 25848875
pmcid: 4817638
doi: 10.1038/ismej.2015.44
Rusch, D. B. et al. The Sorcerer II global ocean sampling expedition: northwest Atlantic through eastern tropical pacific. PLoS Biol. 5, e77 (2007).
pubmed: 17355176
pmcid: 1821060
doi: 10.1371/journal.pbio.0050077
Sunagawa, S. et al. Structure and function of the global ocean microbiome. Science 348, 1261359 (2015).
pubmed: 25999513
doi: 10.1126/science.1261359
Louca, S., Parfrey, L. W. & Doebeli, M. Decoupling function and taxonomy in the global ocean microbiome. Science 353, 1272–1277 (2016).
pubmed: 27634532
doi: 10.1126/science.aaf4507
Duarte, C. M. Seafaring in the 21St Century: the Malaspina 2010 circumnavigation expedition. Limnol. Oceanogr. Bull. 24, 11–14 (2015).
doi: 10.1002/lob.10008
Salazar, G. et al. Gene expression changes and community turnover differentially shape the global ocean metatranscriptome. Cell 179, 1068–1083.e21 (2019).
pubmed: 31730850
pmcid: 6912165
doi: 10.1016/j.cell.2019.10.014
Baltar, F. et al. Prokaryotic extracellular enzymatic activity in relation to biomass production and respiration in the meso- and bathypelagic waters of the (sub)tropical Atlantic. Environ. Microbiol 11, 1998–2014 (2009).
pubmed: 19508555
doi: 10.1111/j.1462-2920.2009.01922.x
Bergauer, K. et al. Organic matter processing by microbial communities throughout the Atlantic water column as revealed by metaproteomics. Proc. Natl Acad. Sci. USA 115, E400–E408 (2018).
pubmed: 29255014
doi: 10.1073/pnas.1708779115
Zhao, Z., Baltar, F. & Herndl, G. J. Linking extracellular enzymes to phylogeny indicates a predominantly particle-associated lifestyle of deep-sea prokaryotes. Sci. Adv. 6, 1–11 (2020).
doi: 10.1126/sciadv.aaz4354
Ruiz‐González, C. et al. Major imprint of surface plankton on deep ocean prokaryotic structure and activity. Mol. Ecol. 29, 1820–1838 (2020).
Pernice, M. C. et al. Large variability of bathypelagic microbial eukaryotic communities across the world’s oceans. ISME J. 10, 945–958 (2016).
pubmed: 26451501
doi: 10.1038/ismej.2015.170
Hingamp, P. et al. Exploring nucleo-cytoplasmic large DNA viruses in Tara Oceans microbial metagenomes. ISME J. 7, 1678–1695 (2013).
pubmed: 23575371
pmcid: 3749498
doi: 10.1038/ismej.2013.59
Kanehisa, M. & Goto, S. KEGG: Kyoto Encyclopedia of genes and genomes. Nucleic Acids Res. 28, 27–30 (2000).
pubmed: 10592173
pmcid: 102409
doi: 10.1093/nar/28.1.27
Galperin, M. Y., Makarova, K. S., Wolf, Y. I. & Koonin, E. V. Expanded Microbial genome coverage and improved protein family annotation in the COG database. Nucleic Acids Res. 43, D261–D269 (2015).
pubmed: 25428365
doi: 10.1093/nar/gku1223
El-Gebali, S. et al. The Pfam protein families database in 2019. Nucleic Acids Res. 47, D427–D432 (2019).
pubmed: 30357350
doi: 10.1093/nar/gky995
Allen, L. Z. et al. Influence of nutrients and currents on the genomic composition of microbes across an upwelling mosaic. ISME J. 6, 1403–1414 (2012).
pubmed: 22278668
pmcid: 3379637
doi: 10.1038/ismej.2011.201
López-Pérez, M., Kimes, N. E., Haro-Moreno, J. M. & Rodríguez-Valera, F. Not all particles are equal: the selective enrichment of particle-associated bacteria from the mediterranean sea. Front. Microbiol. 7, 996 (2016).
pubmed: 27446036
pmcid: 4916215
doi: 10.3389/fmicb.2016.00996
Smith, M. W., Zeigler Allen, L., Allen, A. E., Herfort, L. & Simon, H. M. Contrasting genomic properties of free-living and particle-attached microbial assemblages within a coastal ecosystem. Front. Microbiol. 4, 120 (2013).
pubmed: 23750156
pmcid: 3668451
doi: 10.3389/fmicb.2013.00120
Könneke, M. et al. Isolation of an autotrophic ammonia-oxidizing marine archaeon. Nature 437, 543–546 (2005).
pubmed: 16177789
doi: 10.1038/nature03911
Alonso-Saez, L. et al. Role for urea in nitrification by polar marine Archaea. Proc. Natl Acad. Sci. USA 109, 17989–17994 (2012).
pubmed: 23027926
doi: 10.1073/pnas.1201914109
pmcid: 3497816
Cordero, P. R. F. et al. Atmospheric carbon monoxide oxidation is a widespread mechanism supporting microbial survival. ISME J. 13, 2868–2881 (2019).
pubmed: 31358912
pmcid: 6794299
doi: 10.1038/s41396-019-0479-8
Anantharaman, K., Breier, J. A., Sheik, C. S. & Dick, G. J. Evidence for hydrogen oxidation and metabolic plasticity in widespread deep-sea sulfur-oxidizing bacteria. Proc. Natl Acad. Sci. USA 110, 330–335 (2013).
pubmed: 23263870
doi: 10.1073/pnas.1215340110
Brazelton, W. J., Nelson, B. & Schrenk, M. O. Metagenomic evidence for H
pubmed: 22232619
pmcid: 3252642
doi: 10.3389/fmicb.2011.00268
Ragsdale, S. W. Life with carbon monoxide. Crit. Rev. Biochem. Mol. Biol. 39, 165–195 (2004).
pubmed: 15596550
doi: 10.1080/10409230490496577
Weber, C. F. & King, G. M. Physiological, ecological, and phylogenetic characterization of Stappia, a marine CO-oxidizing bacterial genus. Appl. Environ. Microbiol. 73, 1266–1276 (2007).
pubmed: 17142374
doi: 10.1128/AEM.01724-06
Martín-Cuadrado, A. B., Ghai, R., Gonzaga, A. & Rodríguez-Valera, F. CO dehydrogenase genes found in metagenomic fosmid clones from the deep Mediterranean Sea. Appl. Environ. Microbiol. 75, 7436–7444 (2009).
pubmed: 19801465
pmcid: 2786428
doi: 10.1128/AEM.01283-09
Einsle, O. et al. Structure of cytochrome c nitrite reductase. Nature 400, 476–480 (1999).
pubmed: 10440380
doi: 10.1038/22802
Harborne, N. R., Griffiths, L., Busby, S. J. W. & Cole, J. A. Transcriptional control, translation and function of the products of the five open reading frames of the Escherichia coli nir operon. Mol. Microbiol. 6, 2805–2813 (1992).
pubmed: 1435259
doi: 10.1111/j.1365-2958.1992.tb01460.x
Bianchi, D., Weber, T. S., Kiko, R. & Deutsch, C. Global niche of marine anaerobic metabolisms expanded by particle microenvironments. Nat. Geosci. 11, 263–268 (2018).
doi: 10.1038/s41561-018-0081-0
Bowers, R. M. et al. Minimum information about a single amplified genome (MISAG) and a metagenome-assembled genome (MIMAG) of bacteria and archaea. Nat. Biotechnol. 35, 725–731 (2017).
pubmed: 28787424
pmcid: 6436528
doi: 10.1038/nbt.3893
Delmont, T. O. et al. Nitrogen-fixing populations of Planctomycetes and Proteobacteria are abundant in surface ocean metagenomes. Nat. Microbiol. 3, 804–813 (2018).
pubmed: 29891866
pmcid: 6792437
doi: 10.1038/s41564-018-0176-9
Tully, B. J., Sachdeva, R., Graham, E. D. & Heidelberg, J. F. 290 metagenome-assembled genomes from the Mediterranean Sea: a resource for marine microbiology. PeerJ 5, e3558 (2017).
pubmed: 28713657
pmcid: 5507172
doi: 10.7717/peerj.3558
Parks, D. H., Imelfort, M., Skennerton, C. T., Hugenholtz, P. & Tyson, G. W. CheckM: assessing the quality of microbial genomes recovered from isolates, single cells, and metagenomes. Genome Res. 25, 1043–1055 (2015).
pubmed: 25977477
pmcid: 4484387
doi: 10.1101/gr.186072.114
Huber, H. et al. A new phylum of Archaea represented by a nanosized hyperthermophilic symbiont. Nature 417, 63–67 (2002).
pubmed: 11986665
doi: 10.1038/417063a
Sharma, G., Khatri, I. & Subramanian, S. Complete genome of the starch-degrading myxobacteria Sandaracinus amylolyticus DSM 53668
pubmed: 27358428
pmcid: 5010890
doi: 10.1093/gbe/evw151
Mohr, K. Diversity of myxobacteria—we only see the tip of the iceberg. Microorganisms 6, 84 (2018).
pmcid: 6164225
doi: 10.3390/microorganisms6030084
Farnelid, H. et al. Nitrogenase gene amplicons from global marine surface waters are dominated by genes of non-cyanobacteria. PLoS ONE 6, e19223 (2011).
pubmed: 21559425
pmcid: 3084785
doi: 10.1371/journal.pone.0019223
Moisander, P. H. et al. Chasing after non-cyanobacterial nitrogen fixation in marine pelagic environments. Front. Microbiol. 8, 1736 (2017).
Zehr, J. P., Weitz, J. S. & Joint, I. How microbes survive in the open ocean. Science 357, 646–647 (2017).
pubmed: 28818930
doi: 10.1126/science.aan5764
Zehr, J. P. & Capone, D. G. Changing perspectives in marine nitrogen fixation. Science 368, eaay9514 (2020).
pubmed: 32409447
doi: 10.1126/science.aay9514
Hewson, I. et al. Characteristics of diazotrophs in surface to abyssopelagic waters of the Sargasso Sea. Aquat. Microb. Ecol. 46, 15–30 (2007).
doi: 10.3354/ame046015
Hamersley, M. R. et al. Nitrogen fixation within the water column associated with two hypoxic basins in the Southern California Bight. Aquat. Microb. Ecol. 63, 193–205 (2011).
doi: 10.3354/ame01494
Farnelid, H. et al. Diverse diazotrophs are present on sinking particles in the North Pacific Subtropical Gyre. ISME J. 13, 170–182 (2019).
pubmed: 30116043
doi: 10.1038/s41396-018-0259-x
Sorokin, D. Y., Tourova, T. P. & Muyzer, G. Citreicella thiooxidans gen. nov., sp. nov., a novel lithoheterotrophic sulfur-oxidizing bacterium from the Black Sea. Syst. Appl. Microbiol. 28, 679–687 (2005).
pubmed: 16261857
doi: 10.1016/j.syapm.2005.05.006
Tiirola, M. A., Männistö, M. K., Puhakka, J. A. & Kulomaa, M. S. Isolation and characterization of Novosphingobium sp. strain MT1, a dominant polychlorophenol-degrading strain in a groundwater bioremediation system. Appl. Environ. Microbiol. 68, 173–180 (2002).
pubmed: 11772624
pmcid: 126562
doi: 10.1128/AEM.68.1.173-180.2002
Yuan, J., Lai, Q., Zheng, T. & Shao, Z. Novosphingobium indicum sp. nov., a polycyclic aromatic hydrocarbon-degrading bacterium isolated from a deep-sea environment. Int. J. Syst. Evol. Microbiol. 59, 2084–2088 (2009).
pubmed: 19605709
doi: 10.1099/ijs.0.002873-0
Addison, S. L., Foote, S. M., Reid, N. M. & Lloyd-Jones, G. Novosphingobium nitrogenifigens sp. nov., a polyhydroxyalkanoate-accumulating diazotroph isolated from a New Zealand pulp and paper wastewater. Int. J. Syst. Evol. Microbiol 57, 2467–2471 (2007).
pubmed: 17978201
doi: 10.1099/ijs.0.64627-0
Kim, S. H. et al. Ketobacter alkanivorans gen. nov., sp. nov., an n-alkane-degrading bacterium isolated from seawater. Int. J. Syst. Evol. Microbiol. 68, 2258–2264 (2018).
pubmed: 29809120
doi: 10.1099/ijsem.0.002823
Tully, B. J., Graham, E. D. & Heidelberg, J. F. The reconstruction of 2,631 draft metagenome-assembled genomes from the global oceans. Sci. Data 5, 170203 (2018).
pubmed: 29337314
pmcid: 5769542
doi: 10.1038/sdata.2017.203
Teira, E., Lebaron, P., Van Aken, H. & Herndl, G. J. Distribution and activity of bacteria and archaea in the deep water masses of the North Atlantic. Limnol. Oceanogr. 51, 2131–2144 (2006).
doi: 10.4319/lo.2006.51.5.2131
Yakimov, M. M. et al. Contribution of crenarchaeal autotrophic ammonia oxidizers to the dark primary production in Tyrrhenian deep waters (Central Mediterranean Sea). ISME J. 5, 945–961 (2011).
pubmed: 21209665
pmcid: 3131861
doi: 10.1038/ismej.2010.197
La Cono, V. et al. Contribution of bicarbonate assimilation to carbon pool dynamics in the deep Mediterranean Sea and cultivation of actively nitrifying and CO
pubmed: 29403458
pmcid: 5780414
doi: 10.3389/fmicb.2018.00003
Zarzycki, J., Brecht, V., Müller, M. & Fuchs, G. Identifying the missing steps of the autotrophic 3-hydroxypropionate CO
pubmed: 19955419
doi: 10.1073/pnas.0908356106
pmcid: 2795484
Landry, Z., Swan, B. K., Herndl, G. J., Stepanauskas, R. & Giovannoni, S. J. SAR202 genomes from the dark ocean predict pathways for the oxidation of recalcitrant dissolved organic matter. mBio 8, 1e00413-17–19e00413-17 (2017).
doi: 10.1128/mBio.00413-17
Mehrshad, M., Rodríguez-Valera, F., Amoozegar, M. A., López-García, P. & Ghai, R. The enigmatic SAR202 cluster up close: shedding light on a globally distributed dark ocean lineage involved in sulfur cycling. ISME J. 12, 655–668 (2018).
pubmed: 29208946
doi: 10.1038/s41396-017-0009-5
Tabita, F. R., Satagopan, S., Hanson, T. E., Kreel, N. E. & Scott, S. S. Distinct form I, II, III, and IV Rubisco proteins from the three kingdoms of life provide clues about Rubisco evolution and structure/function relationships. J. Exp. Bot. 59, 1515–1524 (2008).
pubmed: 18281717
doi: 10.1093/jxb/erm361
Carter, M. S. et al. Functional assignment of multiple catabolic pathways for D-apiose. Nat. Chem. Biol. 14, 696–705 (2018).
pubmed: 29867142
pmcid: 6435334
doi: 10.1038/s41589-018-0067-7
Yelton, A. P. et al. Global genetic capacity for mixotrophy in marine picocyanobacteria. ISME J. 10, 2946–2957 (2016).
pubmed: 27137127
pmcid: 5148188
doi: 10.1038/ismej.2016.64
Buesseler, K. O. et al. An assessment of the use of sediment traps for estimating upper ocean particle fluxes. J. Mar. Res. 65, 345–416 (2007).
doi: 10.1357/002224007781567621
Crump, B. C., Armbrust, E. V. & Baross, J. A. Phylogenetic analysis of particle-attached and free-living bacterial communities in the Columbia River, Its Estuary, and the Adjacent Coastal Ocean. Appl. Environ. Microbiol. 65, 3192–3204 (1999).
pubmed: 10388721
pmcid: 91474
doi: 10.1128/AEM.65.7.3192-3204.1999
Ghiglione, J. F., Conan, P. & Pujo-Pay, M. Diversity of total and active free-living vs. particle-attached bacteria in the euphotic zone of the NW Mediterranean Sea. FEMS Microbiol. Lett. 299, 9–21 (2009).
pubmed: 19686348
doi: 10.1111/j.1574-6968.2009.01694.x
Logares, R. et al. Metagenomic 16S rDNA Illumina tags are a powerful alternative to amplicon sequencing to explore diversity and structure of microbial communities. Environ. Microbiol. 16, 2659–2671 (2014).
pubmed: 24102695
doi: 10.1111/1462-2920.12250
Huntemann, M. et al. The standard operating procedure of the DOE-JGI Metagenome Annotation Pipeline (MAP v.4). Stand. Genom. Sci. 11, 1–5 (2016).
Oksanen, J. et al. vegan: community ecology package. https://cran.r-project.org/package=vegan (2019).
R Core Team. R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. https://www.R-project.org/ (2020).
Li, W. & Godzik, A. Cd-hit: a fast program for clustering and comparing large sets of protein or nucleotide sequences. Bioinformatics 22, 1658–1659 (2006).
doi: 10.1093/bioinformatics/btl158
pubmed: 16731699
Suzek, B. E., Wang, Y., Huang, H., McGarvey, P. B. & Wu, C. H. UniRef clusters: a comprehensive and scalable alternative for improving sequence similarity searches. Bioinformatics 31, 926–932 (2015).
pubmed: 25398609
doi: 10.1093/bioinformatics/btu739
Buchfink, B., Xie, C. & Huson, D. H. Fast and sensitive protein alignment using DIAMOND. Nat. Methods 12, 59–60 (2014).
pubmed: 25402007
doi: 10.1038/nmeth.3176
Federhen, S. The NCBI Taxonomy database. Nucleic Acids Res. 40, 136–143 (2012).
doi: 10.1093/nar/gkr1178
Edgar, R. C. Search and clustering orders of magnitude faster than BLAST. Bioinformatics 26, 2460–2461 (2010).
pubmed: 20709691
doi: 10.1093/bioinformatics/btq461
Guillou, L. et al. The Protist Ribosomal Reference database (PR2): a catalog of unicellular eukaryote small sub-unit rRNA sequences with curated taxonomy. Nucleic Acids Res. 41, 597–604 (2013).
doi: 10.1093/nar/gks1160
Yutin, N., Wolf, Y. I., Raoult, D. & Koonin, E. V. Eukaryotic large nucleo-cytoplasmic DNA viruses: clusters of orthologous genes and reconstruction of viral genome evolution. Virol. J. 6, 223 (2009).
pubmed: 20017929
pmcid: 2806869
doi: 10.1186/1743-422X-6-223
Altschul, S. F. et al. Gapped BLAST and PSI-BLAST: a new generation of protein database search programs. Nucleic Acids Res. 25, 3389–3402 (1997).
pubmed: 9254694
pmcid: 146917
doi: 10.1093/nar/25.17.3389
Brum, J. R. et al. Patterns and ecological drivers of ocean viral communities. Science 348, 1261498 (2015).
pubmed: 25999515
doi: 10.1126/science.1261498
Roux, S. et al. Ecogenomics and potential biogeochemical impacts of globally abundant ocean viruses. Nature 537, 689–693 (2016).
pubmed: 27654921
doi: 10.1038/nature19366
Anderson, M. J. A new method for non-parametric multivariate analysis of variance. Austral Ecol. 26, 32–46 (2008).
doi: 10.1046/j.1442-9993.2001.01070.x
Li, D. et al. MEGAHIT v1.0: a fast and scalable metagenome assembler driven by advanced methodologies and community practices. Methods 102, 3–11 (2016).
pubmed: 27012178
doi: 10.1016/j.ymeth.2016.02.020
Langmead, B. & Salzberg, S. L. Fast gapped-read alignment with Bowtie 2. Nat. Methods 9, 357–359 (2012).
pubmed: 22388286
pmcid: 3322381
doi: 10.1038/nmeth.1923
Li, H. et al. The Sequence Alignment/Map format and SAMtools. Bioinformatics 25, 2078–2079 (2009).
pubmed: 19505943
pmcid: 2723002
doi: 10.1093/bioinformatics/btp352
Kang, D. D., Froula, J., Egan, R. & Wang, Z. MetaBAT, an efficient tool for accurately reconstructing single genomes from complex microbial communities. PeerJ 3, e1165 (2015).
pubmed: 26336640
pmcid: 4556158
doi: 10.7717/peerj.1165
Huang, X. & Madan, A. CAP3: a DNA sequence assembly program resource 868 genome research. Genome Res. 9, 868–877 (1999).
Letunic, I. & Bork, P. Interactive tree of life (iTOL) v4: recent updates and new developments. Nucleic Acids Res. 47, W256–W259 (2019).
pubmed: 30931475
pmcid: 6602468
doi: 10.1093/nar/gkz239
Chaumeil, P.-A., Mussig, A. J., Hugenholtz, P. & Parks, D. H. GTDB-Tk: a toolkit to classify genomes with the Genome Taxonomy Database. Bioinformatics 36, 1925–1927 (2019).
pmcid: 7703759
Seemann, T. Prokka: rapid prokaryotic genome annotation. Bioinformatics 30, 2068–2069 (2014).
pubmed: 24642063
doi: 10.1093/bioinformatics/btu153
Aramaki, T. et al. KofamKOALA: KEGG Ortholog assignment based on profile HMM and adaptive score threshold. Bioinformatics 36, 2251–2252 (2020).
pubmed: 31742321
doi: 10.1093/bioinformatics/btz859
Eddy, S. R. Accelerated profile HMM searches. PLoS Comput. Biol. 7, e1002195 (2011).
Bushnell, B.BBMap. 1. Bushnell, B. BBMap. https://sourceforge.net/projects/bbmap/ (2018).
Caro-Quintero, A. & Konstantinidis, K. T. Bacterial species may exist, metagenomics reveal. Environ. Microbiol. 14, 347–355 (2012).
pubmed: 22151572
doi: 10.1111/j.1462-2920.2011.02668.x
Sievers, F. et al. Fast, scalable generation of high-quality protein multiple sequence alignments using Clustal Omega. Mol. Syst. Biol. 7, 539 (2011).
Jaffe, A. L., Castelle, C. J., Dupont, C. L. & Banfield, J. F. Lateral gene transfer shapes the distribution of RuBisCO among candidate phyla radiation bacteria and DPANN archaea. Mol. Biol. Evol. 36, 435–446 (2019).
pubmed: 30544151
doi: 10.1093/molbev/msy234
Aylward, F. O. & Santoro, A. E. Heterotrophic Thaumarchaea with small genomes are widespread in the dark ocean. mSystems 5, e00415-20 (2020).
pubmed: 32546674
pmcid: 7300363
doi: 10.1128/mSystems.00415-20
Price, M. N., Dehal, P. S. & Arkin, A. P. FastTree: computing large minimum evolution trees with profiles instead of a distance matrix. Mol. Biol. Evol. 26, 1641–1650 (2009).
pubmed: 19377059
pmcid: 2693737
doi: 10.1093/molbev/msp077
Alves, R. J. E., Minh, B. Q., Urich, T., Von Haeseler, A. & Schleper, C. Unifying the global phylogeny and environmental distribution of ammonia-oxidising archaea based on amoA genes. Nat. Commun. 9, 1–17 (2018).
doi: 10.1038/s41467-018-03861-1
Katoh, K. & Standley, D. M. MAFFT multiple sequence alignment software version 7: improvements in performance and usability. Mol. Biol. Evol. 30, 772–780 (2013).
pubmed: 23329690
pmcid: 3603318
doi: 10.1093/molbev/mst010