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Unusual biology across a group comprising more than 15% of domain Bacteria
Open accessAbstract
A prominent feature of the bacterial domain is a radiation of major lineages that are defined as candidate phyla because they lack isolated representatives. Bacteria from these phyla occur in diverse environments and are thought to mediate carbon and hydrogen cycles. Genomic analyses of a few representatives suggested that metabolic limitations have prevented their cultivation. Here we reconstructed 8 complete and 789 draft genomes from bacteria representing >35 phyla and documented features that consistently distinguish these organisms from other bacteria. We infer that this group, which may comprise >15% of the bacterial domain, has shared evolutionary history, and describe it as the candidate phyla radiation (CPR). All CPR genomes are small and most lack numerous biosynthetic pathways. Owing to divergent 16S ribosomal RNA (rRNA) gene sequences, 50-100% of organisms sampled from specific phyla would evade detection in typical cultivation-independent surveys. CPR organisms often have self-splicing introns and proteins encoded within their rRNA genes, a feature rarely reported in bacteria. Furthermore, they have unusual ribosome compositions. All are missing a ribosomal protein often absent in symbionts, and specific lineages are missing ribosomal proteins and biogenesis factors considered universal in bacteria. This implies different ribosome structures and biogenesis mechanisms, and underlines unusual biology across a large part of the bacterial domain.
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“Trees visualized at https://itol.embl.de/itol.cgiusing Netwick files from Hug et al. (2016) concatenated ribosomal proteins in place of 16S rRNA genes (Fig. 3B) indicate that a large radiation of predominantly uncultured mesophilic taxa, which are comprised of potentially symbiotic organisms with small cells and genomes and are referred to as the bacterial candidate phyla radiation (CPR) and the”
From The Emergence of Life (2019) · cites this paper as Brown et al. 2015
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“Amplicon sequencing andmetagenomic eorts directed towards marine bacteria (using the 16S rRNA gene) have revealed the possibility of more bacterial species than previously recognized (Sogin et al., 2006; Brown et al., 2015).”
BackgroundInfluentialNovel Microorganisms and Their Function on Anaerobic Digesters
Kirkegaard, R. (2016) · doi:10.5278/vbn.phd.engsci.00180 · Cited by 1
“The sequences deposited in the reference databases are mostly PCR derived and it is thus likely that the conservation of primer sites is overestimated (Brown et al., 2015).”
BackgroundStream–lake network position structures freshwater microbial communities across multiple catchments
Kirtane, A. A., Deiner, K., & Hall, A. R. (2026) · bioRxiv · doi:10.64898/2026.10.29.762755
“In contrast, stream cores included nitrifying and redox-associated taxa such as Nitrospira (Daims et al., 2015), freshwater methylotrophs such as Methylotenera (Kalyuzhnaya et al., 2012), soil-associated genera including Cellvibrio (DeBoy et al., 2008), and groundwater-linked candidate phyla radiation lineages such as…”
A life-history axis of resource acquisition in bacteria: depolymerisation versus uptake
Fu, Y.-Y., Rodriguez, L. C., Ramoneda, J., Shek, K. L., Martiny, J. B., Allison, S. D., Luo, Y., & Malik, A. A. (2026) · bioRxiv · doi:10.64898/2026.10.25.761621
Reconstructing spore evolution recalibrates the timescale of land plant diversification.
Hooper, J., Worth, A. M., Wellman, C. H., Clark, J. W., & Donoghue, P. C. J. (2026) · Nature Plants · doi:10.1038/s41477-026-02432-3
Host Cell Wall Polysaccharides Modulate Nanosynbacteria Recognition and Attachment
Grossman, A. S., Souza, J. G., Lei, L., Anochshenko, K., Xia, M.-Y., Wang, Y.-X., Reyre, J.-L., Lowe, E. C., He, X.-S., Moynihan, P. J., et al. (2026) · bioRxiv · doi:10.64898/2026.10.06.756255
Hidalgo-Martinez, K., dos Santos, R. A. C., Novoletti, J. V. L., Capelini, T. D. L., Oliveira, A. L. M., Riaño-Pachón, D., Zafra, G., de Sousa, M. D., Martins, L. F., & de Oliveira, V. D. (2026) · Microorganisms · doi:10.3390/microorganisms14102197
“Interestingly, nearly 3% of the genomes were affiliated with the recently proposed Patescibacteria phylum [36], with greater prevalence in activated sludge, wastewater, and tailing ponds (Figure S3).”
Microbial ecology of shallow groundwater ecosystems.
Karwautz, C., & Griebler, C. (2026) · Nature Reviews Microbiology · doi:10.1038/s41579-026-01362-3
Ali, A., Saisho, D., Mochida, K., & Tani, A. (2026) · Microorganisms · doi:10.3390/microorganisms14102213
“Patescibacteria have repeatedly been reported at high relative abundance in subsurface and groundwater environments [14,50,51], providing an important point of reference for the patterns observed here.”
A hidden archaeal lineage with an ultra-reduced genome retaining only its replicative core
Harada, R., Nishimura, Y., Nomura, M., Yabuki, A., Shiba, K., Inaba, K., Allers, T., Inagaki, Y., & Nakayama, T. (2026) · Current Biology · doi:10.1016/j.cub.2026.09.042 · Cited by 9
Beaver, R. C., & Neufeld, J. D. (2026) · Current Biology · doi:10.1016/j.cub.2026.03.074
Liu, P.-Y., Lv, Y.-X., Zhang, Y., Wang, Y.-Z., Zhu, K.-L., Cao, Z.-Y., Dong, L., Xiao, X., Pan, Y., & Li, J.-H. (2026) · Nature Communications · doi:10.1038/s41467-026-77994-z
“For Patescibacteria , quality metrics were based on marker gene sets 65 .”
Hu, H.-F., Kitzinger, K., Nierychlo, M., Liu, L., Lopatina, A., Kristensen, J. M., Peces, M., Herbold, C., Dueholm, M. K. D., Speth, D., et al. (2026) · bioRxiv · doi:10.64898/2026.09.16.751625
Reservoir fluids versus produced water as proxies for oil reservoir microbiomes
Alibrandi, A., Plewka, J., Vuillemin, A., Bartholomäus, A., Primio, R. D., Probst, A. J., & Kallmeyer, J. (2026) · Metabarcoding and Metagenomics · doi:10.3897/mbmg.10.194096
“Length-filtered scaffolds were binned into MAGs using Abawaca (v. 1.0.0; Brown et al. 2015) and MaxBin2 (v. 2.2.7; Wu et al. 2016) with default parameters, bins were aggregated with DASTool (v. 1.1.6; Sieber et al. 2018), and the resulting selection was manually curated with uBin (v. 0.9.14;…”
InfluentialRomano, R., Morita, T., Peres, F., Lizárraga, R., Padoan, L., Nascimento, M., Rocha, L., Bendia, A., Karmann, I., & Pellizari, V. (2026) · Frontiers in Microbiology · doi:10.3389/fmicb.2026.1883366
“…is now recognized as one of the defining evolutionary features of the CPR radiation and has been consistently reported across phylogenetically diverse lineages occupying groundwater, subsurface and sedimentary environments (Brown et al., 2015; Castelle and Banfield, 2018; Tian et al., 2020).”
InfluentialKumar, A., Nahar, N., Chouhan, D., McLean, J. S., Bor, B., Dong, P.-T., & He, X.-S. (2026) · Proceedings of the National Academy of Sciences of the United States of America · doi:10.1073/pnas.2615890123
Liu, Z., Zheng, M.-Y., Ren, Y.-B., Wu, P.-C., Song, Y., Pei, Y.-L., Ma, M., Feng, Z.-H., & Jia, T. (2026) · International Biodeterioration & Biodegradation · doi:10.1016/j.ibiod.2026.106445
Liu, Y.-Y., He, Q.-Q., Wang, G., Wu, Z.-Y., Du, M., Bai, D.-Y., Dugarjaviin, M., & Zhang, X.-Z. (2026) · Frontiers in Veterinary Science · doi:10.3389/fvets.2026.1922191
“Candidatus_Saccharimonas , a representative genus of the Candidate Phyla Radiation (CPR) group Patescibacteria , is a specialized fiber-degrading taxon enriched in the EL period (66–68).”
Du, Y., Wu, M., Wu, H., Wu, Y., Fang, C., Li, L., Zhang, H.-T., & Wang, Y. (2026) · Frontiers in Physiology · doi:10.3389/fphys.2026.1883010
“…, although less studied, have recently been reported to show increased abundance in colorectal cancer and liver fi brosis, suggesting that these underexplored taxa may also contribute to host metabolic dysregulation and immune perturbation (Brown et al., 2015; Wong and Yu, 2019; Iwaki et al., 2021).”
Barra, P., Parra-Almuna, L., Pontigo, S., Paredes, C., Larama, G., Valdebenito, D., Jerez-Quezada, C., Menezes-Blackburn, D., & Mora, M. D. L. L. (2026) · Plant physiology and biochemistry : PPB · doi:10.1016/j.plaphy.2026.111618
Leclercq, L., Kientz, G., Lloret, E., Palaric, C., Taminiau, B., Daube, G., Molinié, R., Fontaine, J.-X., Mathiron, D., Drider, D., et al. (2026) · Frontiers in Microbiology · doi:10.3389/fmicb.2026.1855632
Zhang, B.-S., Wang, X., Kuang, C.-Y., Wu, J.-P., & Hong, Y.-G. (2026) · Marine Environmental Research · doi:10.1016/j.marenvres.2026.108247
Wang, Y., Ye, L., Cao, C., Che, G., Zhang, C.-Z., Wei, Q., Hong, Y., & Jiang, K. (2026) · Systematic and Applied Microbiology · doi:10.1016/j.syapm.2026.126751
Sun, Y., Tang, J., Jia, L., Sun, X., Huang, Z.-T., Li, M.-Z., & Yang, X.-H. (2026) · Water Research · doi:10.1016/j.watres.2026.126588
Tryptophan became part of the universal genetic code post-LUCA
Wehbi, S., Ly-Trong, N., Wheeler, A. L., Bui, M., Lauretta, D., & Masel, J. (2026) · bioRxiv · doi:10.64898/2026.05.28.728509
“Wehbi et al. 8 also classified 2234 modern clans, for which all Pfams emerged in a single prokaryotic supergroup, using previously classified supergroups 61,81-83 .”
Ståhle, M., Johnson, A., Turner, S., Mårtensson, P., Kalinowski, B., & Dopson, M. (2026) · Microbial Ecology · doi:10.1007/s00248-026-02812-4
“The BM004 family (Desulfobacterota) is from the order Desulfobulbales that mostly encompasses sulfate reducing bacteria [ 52] and along with UBA2206 have been identified in groundwaters [53 , 54 ].”
Museum specimens reveal evolutionary drivers and functions of oral microbiome in mammalsPreprint
Moraitou, M., Rozwalak, P., Richards, J., Saliari, K., Gillissen, E., Timmons, Z., Kitchener, A. C., Pauwels, O. S. G., Sabin, R., Kokkini, P., et al. (2026) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.64898/2026.06.19.733412 · Cited by 2
The ultrasmall ocean microbiome: a reservoir of microbial diversity and nitrogen fixation
Hollerer, C., Miravet-Verde, S., Ustick, L. J., Bork, P., Acinas, S., Pelletier, E., Sunagawa, S., & Bowler, C. (2026) · bioRxiv · doi:10.64898/2026.06.16.732551
“…Nanoarchaeota, and Nanohaloarchaeota) and the Patescibacteria (also known as Candidate Phyla Radiation, CPR), are both known to include lineages with ultrasmall cell sizes (100–400 nm) 7,8 , and are characterized by highly reduced genomes and symbiotic or host-dependent lifestyles 7–13 .”
A Novel Pilus System in Candidate Phyla Radiation BacteriaPreprint
Troman, L. A., Kim, J., Rose, J. J. A., Johnson, M. D., Banfield, J., Petrovski, S., & Ghosal, D. (2026) · bioRxiv · doi:10.64898/2026.03.03.709456
“The advancement of high-throughput sequencing and genome-resolved metagenomics led to the recognition of Candidate Phylum Radiation (CPR) bacteria and prediction of their symbiotic lifestyles (1, 2).”
Self-citationRuiz-González, C., Andrés-Barrao, C., Lanzén, A., Alonso-Sáez, L., Ducklow, H. W., Gasol, J., & Morán, X. A. G. (2026) · Molecular Ecology · doi:10.1111/mec.70434
Chiniquy, D., Diamond, S., Carlson, H. K., Kazakov, A., Coleman-Derr, D., Northen, T., Banfield, J., & Deutschbauer, A. (2026) · bioRxiv · doi:10.64898/2026.05.14.725246
“The association of CAZyme-poor taxa with Terriglobus could reflect: (i) vitamin/cofactor exchange that relieves auxotrophies [15,16,97], (ii) removal or detoxification of inhibitory intermediates generated during polymer turnover [88], and/or (iii) spatial or metabolic niche partitioning that…”
Self-citationCurrent Understanding of Taxonomy and Ecology of the Phylum Minisyncoccota
Fujii, N., Nakajima, M., Narihiro, T., Kuroda, K., & Kindaichi, T. (2026) · Microbes and Environments · doi:10.1264/jsme2.me25084
“Additional lineages, such as Kazan and SM2F11, were also reported, but without prior documentation linking them to Patescibacteria .”
InfluentialZhang, Q., Gentekaki, E., Leger, M. M., Zou, S., Zhang, G.-A.-T., Omar, A., Fu, Y., & Gong, J. (2026) · Marine Life Science & Technology · doi:10.1007/s42995-026-00382-5 · Cited by 1
“2; Supplementary Table S4)—largely uncultured microbial lineages known primarily through molecular data (Brown et al. 2015; Liu et al. 2022; Murug-kar et al. 2020; Rinke et al. 2013).”
Shi, S., Wang, S.-Y., Li, S.-Y., Lu, R.-F., Pan, S.-X., Chen, F., & He, X.-S. (2026) · Frontiers in Cellular and Infection Microbiology · doi:10.3389/fcimb.2026.1814983
“…our data revealed between the two groups a signi fi cant difference in the relative abundance of those non-conventional bacteria, such as Saccharibacteria and SR1, which have long been considered microbial “ dark matter ” with limited cultured representatives (Brown et al., 2015; Hug et al., 2016).”
da Silva, A. K., de Paula, C. H., de Matos, J. P., Ribeiro, D. F., Cordeiro, I. F., Lemes, C. G. D. C., Sanchez, A. B., Garcia, C. C. M., de Abreu, V. D., Alves, R. M., et al. (2026) · Symbiosis · doi:10.1007/s13199-026-01124-y
“This emphasizes the value of genome-resolved metagenomics for uncovering novel microbial diversity in non-model tropical systems (Brown et al. 2015; Hug et al. 2016; Parks et al. 2017).”
Deep-sea siliceous sponges harbor distinct and functionally diverse microbiomes
Lane, K. R., Meyer, K. S., Collens, A. B., Leal, C. V., Collins, A. G., Herrera, S., & Hansel, C. M. (2026) · Applied and Environmental Microbiology · doi:10.1128/aem.01747-26
“Patescibacteria are ultrasmall and have symbiotic and parasitic relationships with bacterial hosts (115–117).”
Hu, D., Wang, H., Groß, V., Burns, A. M., & Urich, T. (2026) · ISME Communications · doi:10.1093/ismeco/ycag169
“Patescibacteria [ 28, 29].”
Hu, B., An, L.-Y., Wu, M., Xu, J.-B., Nie, Y., & Wu, X.-L. (2025) · mSystems · doi:10.1128/msystems.01204-25
“Based on three previous studies (7, 8, 11), a total of 82 groundwater metagenomic data sets were downloaded from NCBI Sequence Read Archive and NCBI BioSample databases (Data S1).”
Mapping the oral microbiome opens links to periodontitis.
Sempeck, C., & Olm, M. (2025) · Cell Host and Microbe · doi:10.1016/j.chom.2025.10.009
Gabashvili, E., Küsel, K., Pratama, A. A., Wang, H., & Taubert, M. (2025) · Microbiome · doi:10.1186/s40168-025-02244-1
“Although its exact phylogenetic placement as one or multiple phyla is still debated, it represents a large fraction of bacterial diversity [3–6].”
Millán-Aldana, N., González, N. C., & Vanegas, J. S. (2025) · Revista de Biología Tropical · doi:10.15517/8pkmas90
“Amplification of the 16S rRNA gene via PCR primers shows a bias towards certain taxa, resulting in the inability to detect others that could be identified using a metagenomic approach (Brown et al., 2015; Eloe-Fadrosh et al., 2016).”
Jobe, N. E., Jones, D. S., Hernandez, J. C., Nisson, D., Skaar, C. H., Onstott, T., & Kieft, T. (2025) · bioRxiv · doi:10.1101/2025.10.27.684889
“…the virome study at the Äspö hard rock laboratory include dominance of Pseudomonadota hosts and also of viruses infecting members of the candidate phyla radiation, now termed Patescibacteria , a group of divergent microorganisms first identified in an aquifer near Rifle, Colorado in the U.S. [67].”
ChloroScan: Recovering Plastid Genome Bins From Metagenomic Data
Tong, Y., Marcelino, V., Turnbull, R., & Verbruggen, H. (2025) · Molecular Ecology Resources · doi:10.1111/1755-0998.70143
Boruah, J. L. H., Borah, B., Yadav, A., Gogoi, P., Dutta, P., Goswami, M., Dasgupta, A., Gogoi, P., Sarma, R. K., Debnath, R., et al. (2025) · International Journal of Environmental Science and Technology · doi:10.1007/s13762-025-06769-8
“Syntrophorhabdus and Candidate Division OD1 are syntrophic bacteria and engage in interspecies hydrogen and acetate transfer with methanogens, optimizing energy yield from complex substrates under thermophilic, anoxic conditions (Brown et al. 2015).”
Zhu, J., Li, Z., Han, C.-H., Ma, L., Liu, X.-J., Qin, Y., & Yang, Y.-Z. (2025) · Urban Forestry & Urban Greening · doi:10.1016/j.ufug.2025.129062
A prevalent huge phage clade in human and animal gut microbiomes
Chen, L.-X., Qin, Y., Duan, Y., Li, H., Koonin, E., Camargo, A. P., Wang, H.-Y., & Zou, Y.-Q. (2025) · Research Square · doi:10.21203/rs.3.rs-7356405/v1
“They have a genome size of 540-660 kbp, which is comparable to that of some prokaryotes 37 , and were detected mostly in gut metagenomes of humans on non-Western diets as well as various animals 35,36 .”
A prevalent huge phage clade in human and animal gut microbiomes
Chen, L.-X., Camargo, A. P., Qin, Y., Koonin, E., Wang, H.-Y., Zou, Y.-Q., Duan, Y., & Li, H. (2025) · bioRxiv · doi:10.1101/2025.08.10.669567
“They have a genome size of 540-660 kbp, which is comparable to that of some prokaryotes 37 , and were detected mostly in gut metagenomes of humans on non-Western diets as well as various animals 35,36 .”
Razzouk, R., Azour, A., Rolain, J., & Bittar, F. (2025) · Journal of Global Antimicrobial Resistance · doi:10.1016/j.jgar.2025.06.024
Plewka, J., Alibrandi, A., Bornemann, T., Esser, S., Stach, T., Sures, K., Becker, J., Moraru, C., Soares, A., Primio, R. D., et al. (2025) · microLife · doi:10.1093/femsml/uqae027
Tuomela, N., Elovaara, S., Hultman, J., Kaartokallio, H., & Thomas, D. N. (2025) · ISME Communications · doi:10.1093/ismeco/ycaf240 · Cited by 1
“While this bacterial phylum is relatively unknown, it has been detected in high abundances in, e.g. groundwater [87, 88] wastewaters [89] and thermokarst ponds [35, 40].”
Samylina, O., Gabyshev, V., Kosyakova, A. I., Kadnikov, V. V., Beletsky, A., & Pimenov, N. (2024) · Microbiology · doi:10.1134/s0026261724609813
“The presumed physiology of “ Candidatus Patescibacteria” is based solely on metagenomic data (Brown et al., 2015; Anantharaman et al., 2016; Castelle et al., 2018; Tian et al., 2020).”
First detection of fungal diversity associated with Prunus salicina Lindl malformation syndrome
Huang, D., Pu, P., He, X., Lan, M., Du, B., Zhu, L., Li, Q., Yang, Y., Xiang, R., Wang, L., et al. (2024) · All Life · doi:10.1080/26895293.2024.2417460
Metadata
- Authors
- Brown, C. T., Hug, L. A., Thomas, B. C., Sharon, I., Castelle, C. J., Singh, A., Wilkins, M. J., Wrighton, K. C., Williams, K. H., & Banfield, J. F.
- Affiliations
- University of California, Berkeley (US)
- The Ohio State University (US)
- Lawrence Berkeley National Laboratory (US)
- Corresponding author
- Banfield, Jillian F.
- ORCID
- Venue
- Nature
- Published
- 2015-06-15
- Volume
- 523
- Issue
- 7559
- Pages
- 208-211
- ISSN
- 0028-0836 (ISSN-L)
- 1476-4687
- OpenAlex
- W1528357204
- Cited by
- 1,216
- References
- 63
- Citations by year
- 2026: 56 · 2025: 100 · 2024: 95 · 2023: 125 · 2022: 108 · 2021: 120 · 2020: 144 · 2019: 121 · 2018: 116 · 2017: 104 · 2016: 101 · 2015: 26
- FWCI
- 50.17
- Type
- journal article
- Language
- en
- Open access
- green
- www.osti.gov
- Licence
- www.springer.com/tdm (tdm)
- Locations
- Nature (publishedVersion)
- PubMed (publishedVersion)
- OSTI OAI (U.S. Department of Energy Office of Scientific and Technical Information) (submittedVersion)
- RePEc: Research Papers in Economics (submittedVersion)
- eScholarship (California Digital Library) (submittedVersion)
- Funders
- U.S. Department of Energy (DESC0004918, DE AC02 05CH11231)
- Joint Genome Institute
- Office of Science
- Biological and Environmental Research
- Pacific Northwest National Laboratory
- Topics
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Genomics and Phylogenetic Studies
- Physical Sciences › Environmental Science › Ecology › Microbial Community Ecology and Physiology
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Genetics › Bacterial Genetics and Biotechnology
- Keywords
- Candidate Phyla Radiation, uncultured bacteria, bacterial evolution, ribosome composition, ribosome biogenesis, self-splicing introns, small genomes, groundwater bacteria, metagenome-assembled genomes, symbionts
- MeSH terms
- Bacteria, Environmental Microbiology, Introns, Phylogeny, Ribosomal Proteins, RNA, Ribosomal, 16S, Genome, Bacterial
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