Skip to main content

Thank you for visiting nature.com. You are using a browser version with limited support for CSS. To obtain the best experience, we recommend you use a more up to date browser (or turn off compatibility mode in Internet Explorer). In the meantime, to ensure continued support, we are displaying the site without styles and JavaScript.

Advertisement

Scientific Reports
  • View all journals
  • Search
  • My Account Login
  • Content Explore content
  • About the journal
  • Publish with us
  • Sign up for alerts
  • RSS feed
  1. nature
  2. scientific reports
  3. articles
  4. article
Kerogen-rich rocks influence growth and composition of an anaerobic microbial community
Download PDF
Download PDF
  • Article
  • Open access
  • Published: 08 March 2026

Kerogen-rich rocks influence growth and composition of an anaerobic microbial community

  • Annemiek C. Waajen1,2,
  • Wessel de Wit1,3,
  • Mónica Sánchez-Román2,4,
  • John O. Edgar5,
  • Jon Telling5 &
  • …
  • Charles S. Cockell1 

Scientific Reports , Article number:  (2026) Cite this article

  • 551 Accesses

  • Metrics details

We are providing an unedited version of this manuscript to give early access to its findings. Before final publication, the manuscript will undergo further editing. Please note there may be errors present which affect the content, and all legal disclaimers apply.

Subjects

  • Astrobiology
  • Biogeochemistry
  • Environmental microbiology
  • Environmental sciences
  • Solid Earth sciences

Abstract

Kerogen, insoluble macromolecular organic matter in sedimentary rocks, is the most abundant form of organic carbon on Earth and plays a role in deep biosphere processes. It is classified into four types (I–IV) based on origin and chemical composition, yet its influence on microbial communities and carbon cycling remains poorly understood. In this study, we examined how kerogen-rich shales and coals, each containing a distinct kerogen type, shape anaerobic microbial community development and activity. CFU counts showed that kerogen types I and II did not significantly alter overall microbial abundance, while type III-rich rocks inhibited growth, and type IV-rich rocks enhanced it. 16S rRNA gene sequencing revealed that type II-rich rocks selectively enriched for Burkholderiaceae, whereas type IV-rich rocks promoted the proliferation of Cellulomonadaceae and Pleomorphomonadaceae. Gas chromatography showed that CO2 production occurred only in the presence of type II-rich rocks, likely driven by Burkholderiaceae activity. These findings suggest that kerogen structure and geochemical properties drive microbial community assembly and organic matter mobilization in the deep subsurface. Beyond Earth, kerogen type IV-like material is widespread in extraterrestrial environments. Our results indicate the enhancement of the habitability of these environments, offering new insights into the potential for life.

Data availability

The genomic data for this study have been deposited in the Sequence Read Archive (SRA) at the National Center for Biotechnology Information (NCBI) under accession number PRJNA1425804 (https://www.ncbi.nlm.nih.gov/sra/PRJNA1425804).

References

  1. Durand, B. et al. Kerogen: Insoluble Organic Matter from Sedimentary Rocks (éditions technip, 1980).

    Google Scholar 

  2. Vandenbroucke, M. & Largeau, C. Kerogen origin, evolution and structure. Org. Geochem. 38, 719–833 (2007).

    Google Scholar 

  3. Bar-On, Y. M., Phillips, R. & Milo, R. The biomass distribution on Earth. Proc. Natl. Acad. Sci. USA 115, 6506–6511 (2018).

    Google Scholar 

  4. Kallmeyer, J., Pockalny, R., Adhikari, R. R., Smith, D. C. & D’Hondt, S. Global distribution of microbial abundance and biomass in subseafloor sediment. Proc. Natl. Acad. Sci. USA 109, 16213–16216 (2012).

    Google Scholar 

  5. Inagari, F. et al. Exploring deep microbial life in coal-bearing sediment down to ∼2.5 km below the ocean floor. Science 349, 420–424 (2015).

    Google Scholar 

  6. Trembath-Reichert, E. et al. Methyl-compound use and slow growth characterize microbial life in 2-km-deep subseafloor coal and shale beds. Proc. Natl. Acad. Sci. USA 14, E9206–E9215 (2017).

    Google Scholar 

  7. Arndt, S., Brumsack, H.-J. & Wirtz, K. W. Cretaceous black shales as active bioreactors: A biogeochemical model for the deep biosphere encountered during ODP Leg 207 (Demerara Rise). Geochim. Cosmochim. Acta 70, 408–425 (2006).

    Google Scholar 

  8. Samuels, T. Microbial Weathering of Shale Rock in Natural and Historic Industrial Environments (The University of Edinburgh, 2018).

    Google Scholar 

  9. Kerridge, J. F., Chang, S. & Shipp, R. Isotopic characterisation of kerogen-like material in the Murchison carbonaceous chondrite. Geochim. Cosmochim. Acta 51, 2527–2540 (1987).

    Google Scholar 

  10. McCarthy, K. et al. Basic petroleum geochemistry for source rock evaluation. Oilfield Rev. 23, 32–43 (2011).

    Google Scholar 

  11. Royle, S. H., Salter, T. L., Watson, J. S. & Sephton, M. A. Mineral matrix effects on pyrolysis products of kerogens infer difficulties in determining biological provenance of macromolecular organic matter at Mars. Astrobiology 22, 520–540 (2022).

    Google Scholar 

  12. Matthewman, R., Martins, Z. & Sephton, M. A. Type IV kerogens as analogues for organic macromolecular materials in aqueously altered carbonaceous chondrites. Astrobiology 13, 324–333 (2013).

    Google Scholar 

  13. Eigenbrode, J. L. et al. Organic matter preserved in 3-billion-year-old mudstones at Gale crater, Mars. Science 360, 1096–1101 (2018).

    Google Scholar 

  14. Head, I. M., Jones, D. M. & Larter, S. R. Biological activity in the deep subsurface and the origin of heavy oil. Nature 426, 344–352 (2003).

    Google Scholar 

  15. Gray, N. D., Sherry, A., Hubert, C., Dolfing, J. & Head, I. M. Methanogenic degradation of petroleum hydrocarbons in subsurface environments: Remediation, heavy oil formation, and energy recovery. In Advances in Applied Microbiology Vol. 72 (eds Laskin, A. I. et al.) 137–161 (Academic Press, 2010).

    Google Scholar 

  16. Haeseler, F., Behar, F., Garnier, D. & Chenet, P.-Y. First stoichiometric model of oil biodegradation in natural petroleum systems: Part I—The BioClass 0D approach. Org. Geochem. 41, 1156–1170 (2010).

    Google Scholar 

  17. Petsch, S. T., Eglinton, T. I. & Edwards, K. J. 14C-dead living biomass: Evidence for microbial assimilation of ancient organic carbon during shale weathering. Science (1979) 292, 1127–1131 (2001).

    Google Scholar 

  18. Petsch, S. T., Edwards, K. J. & Eglinton, T. I. Microbial transformations of organic matter in black shales and implications for global biogeochemical cycles. Palaeogeogr. Palaeoclimatol. Palaeoecol. 219, 157–170 (2005).

    Google Scholar 

  19. Colosimo, F. et al. Biogenic methane in shale gas and coal bed methane: A review of current knowledge and gaps. Int. J. Coal Geol. 165, 106–120. https://doi.org/10.1016/j.coal.2016.08.011 (2016).

    Google Scholar 

  20. Li, Y., Liu, B., Chen, J. & Yue, X. Carbon-nitrogen-sulfur-related microbial taxa and genes maintained the stability of microbial communities in coals. ACS Omega 7, 22671–22681 (2022).

    Google Scholar 

  21. Li, Y. et al. Subsurface microbial invasion affects the microbial community of coal seams. Energy Fuels 35, 8023–8032 (2021).

    Google Scholar 

  22. Matlakowska, R. & Sklodowska, A. The culturable bacteria isolated from organic-rich black shale potentially useful in biometallurgical procedures. J. Appl. Microbiol. 107, 858–866 (2009).

    Google Scholar 

  23. Iram, A., Akhtar, K. & Ghauri, M. A. Coal methanogenesis: a review of the need of complex microbial consortia and culture conditions for the effective bioconversion of coal into methane. Ann. Microbiol. 67, 275–286. https://doi.org/10.1007/s13213-017-1255-5 (2017).

    Google Scholar 

  24. Magnabosco, C. et al. A metagenomic window into carbon metabolism at 3 km depth in Precambrian continental crust. ISME J. 10, 730–741 (2016).

    Google Scholar 

  25. Meslé, M., Dromart, G., Haeseler, F. & Oger, P. M. Classes of organic molecules targeted by a methanogenic microbial consortium grown on sedimentary rocks of various maturities. Front Microbiol. 6, 589 (2015).

    Google Scholar 

  26. Vick, S. H. W. et al. Who eats what? Unravelling microbial conversion of coal to methane. FEMS Microbiol. Ecol. 95, fiz093 (2019).

    Google Scholar 

  27. Waajen, A. C., Prescott, R. & Cockell, C. S. Meteorites as food source on early Earth: Growth, selection, and inhibition of a microbial community on a carbonaceous chondrite. Astrobiology 22, 495–508 (2022).

    Google Scholar 

  28. Abd-Gami, A. et al. Phenol and phenolic compounds toxicity. J. Environ. Microbiol. Toxicol. 2, 11–23 (2014).

    Google Scholar 

  29. Waajen, A. C., Lima, C., Goodacre, R. & Cockell, C. S. Life on Earth can grow on extraterrestrial organic carbon. Sci. Rep. 14, 3691 (2024).

    Google Scholar 

  30. Rappé, M. S. & Giovannoni, S. J. The uncultured microbial majority. Annu. Rev. Microbiol. 57, 369–394 (2003).

    Google Scholar 

  31. Coenye, T. The family Burkholderiaceae. In The Prokaryotes 759–776 (Springer, 2014). https://doi.org/10.1007/978-3-642-30197-1_239.

  32. Hetz, S. A. & Horn, M. A. Burkholderiaceae are key acetate assimilators during complete denitrification in acidic cryoturbated peat circles of the arctic tundra. Front Microbiol. 12, 628269 (2021).

    Google Scholar 

  33. Stackebrandt, E. & Schumann, P. The family Cellulomonadaceae. The Prokaryotes: Actinobacteria 163–184 (2014) https://doi.org/10.1007/978-3-642-30138-4_223.

  34. Hördt, A. et al. Analysis of 1,000+ type-strain genomes substantially improves taxonomic classification of Alphaproteobacteria. Front. Microbiol. 11, 493139 (2020).

    Google Scholar 

  35. Garrity, G. M., Bell, J. A. & Lilburn, T. Pseudomonadales Orla-Jensen 1921, 270AL. Bergey’s Manual® of Systematic Bacteriology 323–442 (2005). https://doi.org/10.1007/0-387-28022-7_9.

  36. Fulaz, S., Vitale, S., Quinn, L. & Casey, E. Nanoparticle-biofilm interactions: The role of the EPS matrix. Trends Microbiol. 27, 915–926 (2019).

    Google Scholar 

  37. Heinz, J. et al. Bacterial growth in chloride and perchlorate brines: Halotolerances and salt stress responses of Planococcus halocryophilus. Astrobiology 19, 1377–1387 (2019).

    Google Scholar 

  38. White, C. M. et al. Sequestration of carbon dioxide in coal with enhanced coalbed methane recovery—A review. Energy Fuels 19, 659–724 (2005).

    Google Scholar 

  39. Bergey’s Manual of Systematic Bacteriology, Volume Three: The Firmicutes. vol. 3 (Springer, 2009).

  40. Hayes, J. M. Organic constituents of meteorites-a review. Geochim. Cosmochim. Acta 31, 1395–1440 (1967).

    Google Scholar 

  41. Mautner, M. N., Conner, A. J., Killham, K. & Deamer, D. W. Biological potential of extraterrestrial materials: 2. Microbial and plant responses to nutrients in the Murchison carbonaceous meteorite. Icarus 129, 245–253 (1997).

    Google Scholar 

  42. Sephton, M. A. Organic compounds in carbonaceous meteorites. Nat. Prod. Rep. 19, 292–311 (2002).

    Google Scholar 

  43. Beaver, R. C. & Neufeld, J. D. Microbial ecology of the deep terrestrial subsurface. ISME J. 18, 91 (2024).

    Google Scholar 

  44. Montgomery, W., Bromiley, G. D. & Sephton, M. A. The nature of organic records in impact excavated rocks on Mars. Sci. Rep. 6, 30947 (2016).

    Google Scholar 

  45. Matthewman, R., Cotton, L. J., Martins, Z. & Sephton, M. A. Organic geochemistry of late Jurassic paleosols (Dirt Beds) of Dorset, UK. Mar. Pet. Geol. 37, 41–52 (2012).

    Google Scholar 

  46. Luong, D., Sephton, M. A. & Watson, J. S. Subcritical water extraction of organic matter from sedimentary rocks. Anal. Chim. Acta 879, 48–57 (2015).

    Google Scholar 

  47. Wright, M. C., Court, R. W., Kafantaris, F. C. A., Spathopoulos, F. & Sephton, M. A. A new rapid method for shale oil and shale gas assessment. Fuel 153, 231–239 (2015).

    Google Scholar 

  48. Ebukanson, E. J. & Kinghorn, R. R. F. Kerogen facies in the major Jurrasic mudrock formations of Southern England and the implication on the depositional environments of their precursors. J. Pet. Geol. 8, 435–462 (1985).

    Google Scholar 

  49. Hermitage of Braid and Blackford Hill Local Nature Reserve Management Plan 2011 - 2021. www.edinburgh.gov.uk/countrysiderangers.

  50. Pringle, C. R. & Beale, G. H. Antigenic polymorphism in a wild population of Paramecium aurelia. Genet. Res. 1, 62–68 (1960).

    Google Scholar 

  51. Boyd, C. E. Bottom Soils, Sediment, and Pond Aquaculture (Springer, 1995).

    Google Scholar 

  52. Thomas, P., Sekhar, A. C., Upreti, R., Mujawar, M. M. & Pasha, S. S. Optimization of single plate-serial dilution spotting (SP-SDS) with sample anchoring as an assured method for bacterial and yeast cfu enumeration and single colony isolation from diverse samples. Biotechnol. Rep. 8, 45–55 (2015).

    Google Scholar 

  53. Herlemann, D. P. R. et al. Transitions in bacterial communities along the 2000 km salinity gradient of the Baltic Sea. ISME J. 5, 1571–1579 (2011).

    Google Scholar 

  54. Klindworth, A. et al. Evaluation of general 16S ribosomal RNA gene PCR primers for classical and next-generation sequencing-based diversity studies. Nucleic Acids Res. 41, e1–e1 (2013).

    Google Scholar 

  55. Li, P.-E. et al. Enabling the democratization of the genomics revolution with a fully integrated web-based bioinformatics platform. Nucleic Acids Res. 45, 67–80 (2017).

    Google Scholar 

  56. Bolyen, E. et al. Reproducible, interactive, scalable and extensible microbiome data science using QIIME 2. Nat. Biotechnol. 37, 852–857 (2019).

    Google Scholar 

  57. Rognes, T., Flouri, T., Nichols, B., Quince, C. & Mahé, F. VSEARCH: A versatile open source tool for metagenomics. PeerJ 4, e2584 (2016).

    Google Scholar 

  58. Bokulich, N. A. et al. Optimizing taxonomic classification of marker-gene amplicon sequences with QIIME 2’s q2-feature-classifier plugin. Microbiome 6 (2018).

  59. Katoh, K., Misawa, K., Kuma, K. & Miyata, T. MAFFT: A novel method for rapid multiple sequence alignment based on fast Fourier transform. Nucleic Acids Res. 30, 3059–3066 (2002).

    Google Scholar 

  60. 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).

    Google Scholar 

Download references

Acknowledgements

We thank Mark Sephton at Imperial College London for providing the kerogen samples, Toby Samuels at the Heriot-Watt University and Sophie Nixon at the University of Manchester for the information and advice on the kerogen samples and Jennifer Saito at the Advanced Studies in Genomics, Proteomics and Bioinformatics (ASGPB) at the University of Hawaii at Manoa for the analysis of the DNA samples.

Funding

This research was funded by the NERC Doctoral Training Partnership grant 363 (NE/L002558/1), Principal’s Career Development Scholarship (PCDS) and NWO GoSpace (R/010809). CSC was supported through the Science and Technologies Facilities Council (STFC) Grant Number ST/Y001788/1. MSR also acknowledges financial support from the Beatriz Galindo Senior Grant (No. BG23-00132) funded by the Spanish Ministry of Science, Innovation and Universities (MICIU).

Author information

Authors and Affiliations

  1. UK Centre for Astrobiology, School of Physics and Astronomy, University of Edinburgh, Edinburgh, EH9 3FD, UK

    Annemiek C. Waajen, Wessel de Wit & Charles S. Cockell

  2. Department of Earth Sciences, Vrije Universiteit Amsterdam, 1081 HZ, Amsterdam, The Netherlands

    Annemiek C. Waajen & Mónica Sánchez-Román

  3. Faculty of Science, University of Amsterdam, 1098 XH, Amsterdam, The Netherlands

    Wessel de Wit

  4. Mineralogy and Petrology Department, Sciences Faculty, University of Granada, Avenida de Fuentenueva S/N, 18071, Granada, Spain

    Mónica Sánchez-Román

  5. School of Natural and Environmental Sciences, Newcastle University, Newcastle, NE1 7RU, UK

    John O. Edgar & Jon Telling

Authors
  1. Annemiek C. Waajen
    View author publications

    Search author on:PubMed Google Scholar

  2. Wessel de Wit
    View author publications

    Search author on:PubMed Google Scholar

  3. Mónica Sánchez-Román
    View author publications

    Search author on:PubMed Google Scholar

  4. John O. Edgar
    View author publications

    Search author on:PubMed Google Scholar

  5. Jon Telling
    View author publications

    Search author on:PubMed Google Scholar

  6. Charles S. Cockell
    View author publications

    Search author on:PubMed Google Scholar

Contributions

ACW, WW, JOE, JT and CC contributed to the conception and design of the study. ACW, WW and JOE carried out the experimental work. ACW performed data analysis and wrote the first draft of the manuscript. ACW, WW, MSR and CC contributed to the data analysis and interpretation. All authors contributed to the manuscript revision and read and approved the submitted version.

Corresponding author

Correspondence to Annemiek C. Waajen.

Ethics declarations

Competing interests

The authors declare no competing interests.

Additional information

Publisher’s note

Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

Supplementary Information

Below is the link to the electronic supplementary material.

Supplementary Material 1 (download DOCX )

Rights and permissions

Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/.

Reprints and permissions

About this article

Check for updates. Verify currency and authenticity via CrossMark

Cite this article

Waajen, A.C., de Wit, W., Sánchez-Román, M. et al. Kerogen-rich rocks influence growth and composition of an anaerobic microbial community. Sci Rep (2026). https://doi.org/10.1038/s41598-026-42062-5

Download citation

  • Received: 24 April 2025

  • Accepted: 24 February 2026

  • Published: 08 March 2026

  • DOI: https://doi.org/10.1038/s41598-026-42062-5

Share this article

Anyone you share the following link with will be able to read this content:

Sorry, a shareable link is not currently available for this article.

Provided by the Springer Nature SharedIt content-sharing initiative

Download PDF

Associated content

Collection

Next generation of geobiology

Advertisement

Explore content

  • Research articles
  • News & Comment
  • Collections
  • Subjects
  • Follow us on Facebook
  • Follow us on X
  • Sign up for alerts
  • RSS feed

About the journal

  • About Scientific Reports
  • Contact
  • Journal policies
  • Guide to referees
  • Calls for Papers
  • Editor's Choice
  • Journal highlights
  • Open Access Fees and Funding

Publish with us

  • For authors
  • Language editing services
  • Open access funding
  • Submit manuscript

Search

Advanced search

Quick links

  • Explore articles by subject
  • Find a job
  • Guide to authors
  • Editorial policies

Scientific Reports (Sci Rep)

ISSN 2045-2322 (online)

nature.com footer links

About Nature Portfolio

  • About us
  • Press releases
  • Press office
  • Contact us

Discover content

  • Journals A-Z
  • Articles by subject
  • protocols.io
  • Nature Index

Publishing policies

  • Nature portfolio policies
  • Open access

Author & Researcher services

  • Reprints & permissions
  • Research data
  • Language editing
  • Scientific editing
  • Nature Masterclasses
  • Research Solutions

Libraries & institutions

  • Librarian service & tools
  • Librarian portal
  • Open research
  • Recommend to library

Advertising & partnerships

  • Advertising
  • Partnerships & Services
  • Media kits
  • Branded content

Professional development

  • Nature Awards
  • Nature Careers
  • Nature Conferences

Regional websites

  • Nature Africa
  • Nature China
  • Nature India
  • Nature Japan
  • Nature Middle East
  • Privacy Policy
  • Use of cookies
  • Legal notice
  • Accessibility statement
  • Terms & Conditions
  • Your US state privacy rights
Springer Nature

© 2026 Springer Nature Limited

Nature Briefing Microbiology

Sign up for the Nature Briefing: Microbiology newsletter — what matters in microbiology research, free to your inbox weekly.

Get the most important science stories of the day, free in your inbox. Sign up for Nature Briefing: Microbiology