---
DOAJ_listed: '1'
OA_place: publisher
OA_type: gold
_id: '23090'
abstract:
- lang: eng
  text: 'Homologous recombination is ubiquitous across evolution, generating variation
    through genetic reshuffling. Although bacteria lack meiotic recombination, homologous
    recombination has been known to extensively shape their genomes. However, the
    mechanisms and ecological conditions sustaining frequent recombination in bacteria
    remain unclear. Using an Escherichia coli–P1 phage model system, we reveal how
    frequent recombination emerges from CRISPR-based herd immunity to a generalized
    transducing phage (DNA vector). Herd immunity maintains phage-susceptible donor
    bacteria, recipient immune bacteria, and transducing phage, which promotes chromosomal
    gene flow that exceeds spontaneous mutation by up to two orders of magnitude.
    Notably, recombination begins as bacterial cultures enter stationary phase, when
    canonical mutation supply through population growth and DNA replication is strongly
    reduced, and recombination can be reinitiated by brief fresh nutrient pulses.
    Genetic analysis showed that stationary-phase recombination requires recA, recB,
    and recG, connecting this process to a defined homologous recombination pathway,
    and suggests a plausible transcriptional regulatory link to transition to nutrient
    limitation. Bacterial herd immunity therefore has an unexpected evolutionary consequence:
    it can promote homologous recombination by turning phages into de facto pollinators.
    This mechanism can help explain how high rates of homologous recombination may
    be sustained across bacterial populations and suggests that bacterial immune systems
    can shape molecular evolution not only by blocking infection but also by regulating
    genetic exchange.'
acknowledgement: We thank Stuart Baird, Nick Barton, Max Joesch, Meriem El Karoui,
  Mato Lagator, Ivana Mašlaňová, Stepan Ovchinnikov, Roman Pantůček, Maroš Pleška,
  Jitka Polechová, and Bryan Wu for their feedback on the manuscript. We are grateful
  to the Redlinger Hut for hosting many discussions about the project away from the
  daily buzz. We acknowledge the core facility CELLIM supported by the Czech-BioImaging
  large RI project (LM2023050 funded by MEYS CR) for their support. This work was
  supported by H2020 European Research Council EVOLHGT No. 648440 to JPB, European
  Commission Programme EXCELES project LX22NPO5103 “National Institute of Virology
  and Bacteriology” to PPl, and a bilateral Czech-Austrian mobility grant 8J3AT003
  funded by MEYS CR and OeAD AT to PPa and CCG.
article_number: msag238
article_processing_charge: Yes
article_type: original
author:
- first_name: Pavel
  full_name: Payne, Pavel
  id: 35F78294-F248-11E8-B48F-1D18A9856A87
  last_name: Payne
  orcid: 0000-0002-2711-9453
- first_name: Pavel
  full_name: Plevka, Pavel
  last_name: Plevka
- first_name: Jonathan P
  full_name: Bollback, Jonathan P
  id: 2C6FA9CC-F248-11E8-B48F-1D18A9856A87
  last_name: Bollback
  orcid: 0000-0002-4624-4612
- first_name: Calin C
  full_name: Guet, Calin C
  id: 47F8433E-F248-11E8-B48F-1D18A9856A87
  last_name: Guet
  orcid: 0000-0001-6220-2052
citation:
  ama: Payne P, Plevka P, Bollback JP, Guet CC. Herd immunity underlies homologous
    recombination in stationary phase bacteria. <i>Molecular Biology and Evolution</i>.
    2026;43(10). doi:<a href="https://doi.org/10.1093/molbev/msag238">10.1093/molbev/msag238</a>
  apa: Payne, P., Plevka, P., Bollback, J. P., &#38; Guet, C. C. (2026). Herd immunity
    underlies homologous recombination in stationary phase bacteria. <i>Molecular
    Biology and Evolution</i>. Oxford University Press. <a href="https://doi.org/10.1093/molbev/msag238">https://doi.org/10.1093/molbev/msag238</a>
  chicago: Payne, Pavel, Pavel Plevka, Jonathan P Bollback, and Calin C Guet. “Herd
    Immunity Underlies Homologous Recombination in Stationary Phase Bacteria.” <i>Molecular
    Biology and Evolution</i>. Oxford University Press, 2026. <a href="https://doi.org/10.1093/molbev/msag238">https://doi.org/10.1093/molbev/msag238</a>.
  ieee: P. Payne, P. Plevka, J. P. Bollback, and C. C. Guet, “Herd immunity underlies
    homologous recombination in stationary phase bacteria,” <i>Molecular Biology and
    Evolution</i>, vol. 43, no. 10. Oxford University Press, 2026.
  ista: Payne P, Plevka P, Bollback JP, Guet CC. 2026. Herd immunity underlies homologous
    recombination in stationary phase bacteria. Molecular Biology and Evolution. 43(10),
    msag238.
  mla: Payne, Pavel, et al. “Herd Immunity Underlies Homologous Recombination in Stationary
    Phase Bacteria.” <i>Molecular Biology and Evolution</i>, vol. 43, no. 10, msag238,
    Oxford University Press, 2026, doi:<a href="https://doi.org/10.1093/molbev/msag238">10.1093/molbev/msag238</a>.
  short: P. Payne, P. Plevka, J.P. Bollback, C.C. Guet, Molecular Biology and Evolution
    43 (2026).
corr_author: '1'
das_tickbox: '1'
dataavailabilitystatement: The data underlying this article are available in the article
  and in its online material.
date_created: 2026-10-11T22:02:04Z
date_published: 2026-10-03T00:00:00Z
date_updated: 2026-10-12T10:51:54Z
day: '03'
ddc:
- '570'
department:
- _id: CaGu
doi: 10.1093/molbev/msag238
external_id:
  pmid:
  - '42827080'
fulldoi: https://doi.org/10.1093/molbev/msag238
has_accepted_license: '1'
intvolume: '        43'
issue: '10'
keyword:
- homologous recombination
- defense systems
- CRISPR-Cas
- herd immunity
- stationary phase
- bacterial evolution
language:
- iso: eng
main_file_link:
- open_access: '1'
  url: https://doi.org/10.1093/molbev/msag238
month: '10'
oa: 1
oa_version: Published Version
pmid: 1
publication: Molecular Biology and Evolution
publication_identifier:
  eissn:
  - 1537-1719
  issn:
  - 0737-4038
publication_status: epub_ahead
publisher: Oxford University Press
quality_controlled: '1'
researchdata_availability: no
scopus_import: '1'
status: public
supplementarymaterial: yes
title: Herd immunity underlies homologous recombination in stationary phase bacteria
tmp:
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  name: Creative Commons Attribution-NonCommercial 4.0 International (CC BY-NC 4.0)
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type: journal_article
user_id: 2DF688A6-F248-11E8-B48F-1D18A9856A87
volume: 43
year: '2026'
...
---
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abstract:
- lang: eng
  text: Herd immunity, a process in which resistant individuals limit the spread of
    a pathogen among susceptible hosts has been extensively studied in eukaryotes.
    Even though bacteria have evolved multiple immune systems against their phage
    pathogens, herd immunity in bacteria remains unexplored. Here we experimentally
    demonstrate that herd immunity arises during phage epidemics in structured and
    unstructured Escherichia coli populations consisting of differing frequencies
    of susceptible and resistant cells harboring CRISPR immunity. In addition, we
    develop a mathematical model that quantifies how herd immunity is affected by
    spatial population structure, bacterial growth rate, and phage replication rate.
    Using our model we infer a general epidemiological rule describing the relative
    speed of an epidemic in partially resistant spatially structured populations.
    Our experimental and theoretical findings indicate that herd immunity may be important
    in bacterial communities, allowing for stable coexistence of bacteria and their
    phages and the maintenance of polymorphism in bacterial immunity.
article_processing_charge: No
author:
- first_name: Pavel
  full_name: Payne, Pavel
  id: 35F78294-F248-11E8-B48F-1D18A9856A87
  last_name: Payne
  orcid: 0000-0002-2711-9453
- first_name: Lukas
  full_name: Geyrhofer, Lukas
  last_name: Geyrhofer
- first_name: Nicholas H
  full_name: Barton, Nicholas H
  id: 4880FE40-F248-11E8-B48F-1D18A9856A87
  last_name: Barton
  orcid: 0000-0002-8548-5240
- first_name: Jonathan P
  full_name: Bollback, Jonathan P
  id: 2C6FA9CC-F248-11E8-B48F-1D18A9856A87
  last_name: Bollback
  orcid: 0000-0002-4624-4612
citation:
  ama: 'Payne P, Geyrhofer L, Barton NH, Bollback JP. Data from: CRISPR-based herd
    immunity limits phage epidemics in bacterial populations. 2018. doi:<a href="https://doi.org/10.5061/dryad.42n44">10.5061/dryad.42n44</a>'
  apa: 'Payne, P., Geyrhofer, L., Barton, N. H., &#38; Bollback, J. P. (2018). Data
    from: CRISPR-based herd immunity limits phage epidemics in bacterial populations.
    Dryad. <a href="https://doi.org/10.5061/dryad.42n44">https://doi.org/10.5061/dryad.42n44</a>'
  chicago: 'Payne, Pavel, Lukas Geyrhofer, Nicholas H Barton, and Jonathan P Bollback.
    “Data from: CRISPR-Based Herd Immunity Limits Phage Epidemics in Bacterial Populations.”
    Dryad, 2018. <a href="https://doi.org/10.5061/dryad.42n44">https://doi.org/10.5061/dryad.42n44</a>.'
  ieee: 'P. Payne, L. Geyrhofer, N. H. Barton, and J. P. Bollback, “Data from: CRISPR-based
    herd immunity limits phage epidemics in bacterial populations.” Dryad, 2018.'
  ista: 'Payne P, Geyrhofer L, Barton NH, Bollback JP. 2018. Data from: CRISPR-based
    herd immunity limits phage epidemics in bacterial populations, Dryad, <a href="https://doi.org/10.5061/dryad.42n44">10.5061/dryad.42n44</a>.'
  mla: 'Payne, Pavel, et al. <i>Data from: CRISPR-Based Herd Immunity Limits Phage
    Epidemics in Bacterial Populations</i>. Dryad, 2018, doi:<a href="https://doi.org/10.5061/dryad.42n44">10.5061/dryad.42n44</a>.'
  short: P. Payne, L. Geyrhofer, N.H. Barton, J.P. Bollback, (2018).
date_created: 2021-08-09T13:10:02Z
date_published: 2018-03-12T00:00:00Z
date_updated: 2025-04-15T08:17:50Z
day: '12'
department:
- _id: NiBa
- _id: JoBo
doi: 10.5061/dryad.42n44
fulldoi: https://doi.org/10.5061/dryad.42n44
main_file_link:
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  url: https://doi.org/10.5061/dryad.42n44
month: '03'
oa: 1
oa_version: Published Version
publisher: Dryad
related_material:
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    relation: used_in_publication
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status: public
title: 'Data from: CRISPR-based herd immunity limits phage epidemics in bacterial
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type: research_data_reference
user_id: 6785fbc1-c503-11eb-8a32-93094b40e1cf
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_id: '423'
abstract:
- lang: eng
  text: Herd immunity, a process in which resistant individuals limit the spread of
    a pathogen among susceptible hosts has been extensively studied in eukaryotes.
    Even though bacteria have evolved multiple immune systems against their phage
    pathogens, herd immunity in bacteria remains unexplored. Here we experimentally
    demonstrate that herd immunity arises during phage epidemics in structured and
    unstructured Escherichia coli populations consisting of differing frequencies
    of susceptible and resistant cells harboring CRISPR immunity. In addition, we
    develop a mathematical model that quantifies how herd immunity is affected by
    spatial population structure, bacterial growth rate, and phage replication rate.
    Using our model we infer a general epidemiological rule describing the relative
    speed of an epidemic in partially resistant spatially structured populations.
    Our experimental and theoretical findings indicate that herd immunity may be important
    in bacterial communities, allowing for stable coexistence of bacteria and their
    phages and the maintenance of polymorphism in bacterial immunity.
acknowledgement: "We are grateful to Remy Chait for his help and assistance with establishing
  our experimental setups and to Tobias Bergmiller for valuable insights into some
  specific experimental details. We thank Luciano Marraffini for donating us the pCas9
  plasmid used in this study. We also want to express our gratitude to Seth Barribeau,
  Andrea Betancourt, Călin Guet, Mato Lagator, Tiago Paixão and Maroš Pleška for valuable
  discussions on the manuscript. Finally, we would like to thank the \r\neditors and
  reviewers for their helpful comments and suggestions."
article_number: e32035
article_processing_charge: No
author:
- first_name: Pavel
  full_name: Payne, Pavel
  id: 35F78294-F248-11E8-B48F-1D18A9856A87
  last_name: Payne
  orcid: 0000-0002-2711-9453
- first_name: Lukas
  full_name: Geyrhofer, Lukas
  last_name: Geyrhofer
- first_name: Nicholas H
  full_name: Barton, Nicholas H
  id: 4880FE40-F248-11E8-B48F-1D18A9856A87
  last_name: Barton
  orcid: 0000-0002-8548-5240
- first_name: Jonathan P
  full_name: Bollback, Jonathan P
  id: 2C6FA9CC-F248-11E8-B48F-1D18A9856A87
  last_name: Bollback
  orcid: 0000-0002-4624-4612
citation:
  ama: Payne P, Geyrhofer L, Barton NH, Bollback JP. CRISPR-based herd immunity can
    limit phage epidemics in bacterial populations. <i>eLife</i>. 2018;7. doi:<a href="https://doi.org/10.7554/eLife.32035">10.7554/eLife.32035</a>
  apa: Payne, P., Geyrhofer, L., Barton, N. H., &#38; Bollback, J. P. (2018). CRISPR-based
    herd immunity can limit phage epidemics in bacterial populations. <i>ELife</i>.
    eLife Sciences Publications. <a href="https://doi.org/10.7554/eLife.32035">https://doi.org/10.7554/eLife.32035</a>
  chicago: Payne, Pavel, Lukas Geyrhofer, Nicholas H Barton, and Jonathan P Bollback.
    “CRISPR-Based Herd Immunity Can Limit Phage Epidemics in Bacterial Populations.”
    <i>ELife</i>. eLife Sciences Publications, 2018. <a href="https://doi.org/10.7554/eLife.32035">https://doi.org/10.7554/eLife.32035</a>.
  ieee: P. Payne, L. Geyrhofer, N. H. Barton, and J. P. Bollback, “CRISPR-based herd
    immunity can limit phage epidemics in bacterial populations,” <i>eLife</i>, vol.
    7. eLife Sciences Publications, 2018.
  ista: Payne P, Geyrhofer L, Barton NH, Bollback JP. 2018. CRISPR-based herd immunity
    can limit phage epidemics in bacterial populations. eLife. 7, e32035.
  mla: Payne, Pavel, et al. “CRISPR-Based Herd Immunity Can Limit Phage Epidemics
    in Bacterial Populations.” <i>ELife</i>, vol. 7, e32035, eLife Sciences Publications,
    2018, doi:<a href="https://doi.org/10.7554/eLife.32035">10.7554/eLife.32035</a>.
  short: P. Payne, L. Geyrhofer, N.H. Barton, J.P. Bollback, ELife 7 (2018).
date_created: 2018-12-11T11:46:23Z
date_published: 2018-03-09T00:00:00Z
date_updated: 2025-03-31T16:00:24Z
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ddc:
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department:
- _id: NiBa
- _id: JoBo
doi: 10.7554/eLife.32035
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title: CRISPR-based herd immunity can limit phage epidemics in bacterial populations
tmp:
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  legal_code_url: https://creativecommons.org/licenses/by/4.0/legalcode
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type: journal_article
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...
---
OA_place: publisher
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abstract:
- lang: eng
  text: Bacteria and their pathogens – phages – are the most abundant living entities
    on Earth. Throughout their coevolution, bacteria have evolved multiple immune
    systems to overcome the ubiquitous threat from the phages. Although the molecu-
    lar details of these immune systems’ functions are relatively well understood,
    their epidemiological consequences for the phage-bacterial communities have been
    largely neglected. In this thesis we employed both experimental and theoretical
    methods to explore whether herd and social immunity may arise in bacterial popu-
    lations. Using our experimental system consisting of Escherichia coli strains
    with a CRISPR based immunity to the T7 phage we show that herd immunity arises
    in phage-bacterial communities and that it is accentuated when the populations
    are spatially structured. By fitting a mathematical model, we inferred expressions
    for the herd immunity threshold and the velocity of spread of a phage epidemic
    in partially resistant bacterial populations, which both depend on the bacterial
    growth rate, phage burst size and phage latent period. We also investigated the
    poten- tial for social immunity in Streptococcus thermophilus and its phage 2972
    using a bioinformatic analysis of potentially coding short open reading frames
    with a signalling signature, encoded within the CRISPR associated genes. Subsequently,
    we tested one identified potentially signalling peptide and found that its addition
    to a phage-challenged culture increases probability of survival of bacteria two
    fold, although the results were only marginally significant. Together, these results
    demonstrate that the ubiquitous arms races between bacteria and phages have further
    consequences at the level of the population.
alternative_title:
- ISTA Thesis
article_processing_charge: No
author:
- first_name: Pavel
  full_name: Payne, Pavel
  id: 35F78294-F248-11E8-B48F-1D18A9856A87
  last_name: Payne
  orcid: 0000-0002-2711-9453
citation:
  ama: Payne P. Bacterial herd and social immunity to phages. 2017.
  apa: Payne, P. (2017). <i>Bacterial herd and social immunity to phages</i>. Institute
    of Science and Technology Austria.
  chicago: Payne, Pavel. “Bacterial Herd and Social Immunity to Phages.” Institute
    of Science and Technology Austria, 2017.
  ieee: P. Payne, “Bacterial herd and social immunity to phages,” Institute of Science
    and Technology Austria, 2017.
  ista: Payne P. 2017. Bacterial herd and social immunity to phages. Institute of
    Science and Technology Austria.
  mla: Payne, Pavel. <i>Bacterial Herd and Social Immunity to Phages</i>. Institute
    of Science and Technology Austria, 2017.
  short: P. Payne, Bacterial Herd and Social Immunity to Phages, Institute of Science
    and Technology Austria, 2017.
corr_author: '1'
date_created: 2019-04-09T15:16:45Z
date_published: 2017-02-01T00:00:00Z
date_updated: 2026-04-08T14:16:28Z
day: '01'
ddc:
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degree_awarded: PhD
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- _id: JoBo
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language:
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month: '02'
oa: 1
oa_version: Published Version
page: '83'
publication_identifier:
  issn:
  - 2663-337X
publication_status: published
publisher: Institute of Science and Technology Austria
status: public
supervisor:
- first_name: Jonathan P
  full_name: Bollback, Jonathan P
  id: 2C6FA9CC-F248-11E8-B48F-1D18A9856A87
  last_name: Bollback
  orcid: 0000-0002-4624-4612
- first_name: Nicholas H
  full_name: Barton, Nicholas H
  id: 4880FE40-F248-11E8-B48F-1D18A9856A87
  last_name: Barton
  orcid: 0000-0002-8548-5240
title: Bacterial herd and social immunity to phages
type: dissertation
user_id: ba8df636-2132-11f1-aed0-ed93e2281fdd
year: '2017'
...
