---
OA_place: publisher
OA_type: hybrid
PlanS_conform: '1'
_id: '21985'
abstract:
- lang: eng
  text: Upon infecting a bacterial cell, temperate phages make a decision between
    lysis and lysogeny. While research has previously explored how phages sense environmental
    information to make this choice, most studies have focused on modelling known
    mechanisms that impact the decision. These mechanisms tell us what environmental
    information the phage does respond to, but not what it should respond to, as the
    signals sensed by the phage may serve as proxies for other sources of information.
    Here, using a mechanism-agnostic population dynamics model, we find that irreversible
    phage binding to lysogens protects sensitive host cells from infection. This results
    in lysogens being an additional environmental factor that the phage should sense
    while making its decision to undergo lysis or lysogeny. Using this model, we derive
    a responsive lysogeny probability for phages that respond to both cell and lysogen
    densities optimized towards invading phage-occupied systems, and show that it
    is more capable of invading and resisting invasion than phage with fixed lysogeny
    probabilities across different environmental conditions.
acknowledgement: We thank Fyodor Kondrashov and Gašper Tkačik for valuable input and
  guidance in building the model, and Stephen Abedon as well as the two anonymous
  reviewers for the comments provided on the manuscript.
article_processing_charge: Yes (via OA deal)
article_type: original
author:
- first_name: Bryan
  full_name: Wu, Bryan
  id: 3C521EBA-F248-11E8-B48F-1D18A9856A87
  last_name: Wu
- 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: Wu B, Guet CC. Responsive lysogeny under nonproductive phage binding. <i>Evolution</i>.
    2026;80(6):1365-1373. doi:<a href="https://doi.org/10.1093/evolut/qpag061">10.1093/evolut/qpag061</a>
  apa: Wu, B., &#38; Guet, C. C. (2026). Responsive lysogeny under nonproductive phage
    binding. <i>Evolution</i>. Oxford University Press. <a href="https://doi.org/10.1093/evolut/qpag061">https://doi.org/10.1093/evolut/qpag061</a>
  chicago: Wu, Bryan, and Calin C Guet. “Responsive Lysogeny under Nonproductive Phage
    Binding.” <i>Evolution</i>. Oxford University Press, 2026. <a href="https://doi.org/10.1093/evolut/qpag061">https://doi.org/10.1093/evolut/qpag061</a>.
  ieee: B. Wu and C. C. Guet, “Responsive lysogeny under nonproductive phage binding,”
    <i>Evolution</i>, vol. 80, no. 6. Oxford University Press, pp. 1365–1373, 2026.
  ista: Wu B, Guet CC. 2026. Responsive lysogeny under nonproductive phage binding.
    Evolution. 80(6), 1365–1373.
  mla: Wu, Bryan, and Calin C. Guet. “Responsive Lysogeny under Nonproductive Phage
    Binding.” <i>Evolution</i>, vol. 80, no. 6, Oxford University Press, 2026, pp.
    1365–73, doi:<a href="https://doi.org/10.1093/evolut/qpag061">10.1093/evolut/qpag061</a>.
  short: B. Wu, C.C. Guet, Evolution 80 (2026) 1365–1373.
corr_author: '1'
date_created: 2026-06-10T07:38:12Z
date_published: 2026-06-01T00:00:00Z
date_updated: 2026-06-16T12:46:02Z
day: '01'
ddc:
- '570'
department:
- _id: CaGu
doi: 10.1093/evolut/qpag061
external_id:
  pmid:
  - '41968110'
file:
- access_level: open_access
  checksum: 6d0f48566a7a36cb0c469e1968c9cb1c
  content_type: application/pdf
  creator: dernst
  date_created: 2026-06-16T12:45:09Z
  date_updated: 2026-06-16T12:45:09Z
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  file_name: 2026_Evolution_Wu.pdf
  file_size: 2077781
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  success: 1
file_date_updated: 2026-06-16T12:45:09Z
has_accepted_license: '1'
intvolume: '        80'
issue: '6'
language:
- iso: eng
month: '06'
oa: 1
oa_version: Published Version
page: 1365-1373
pmid: 1
publication: Evolution
publication_identifier:
  eissn:
  - 1558-5646
  issn:
  - 0014-3820
publication_status: published
publisher: Oxford University Press
quality_controlled: '1'
related_material:
  link:
  - relation: software
    url: https://github.com/theguetlab/responsive-lysogeny
scopus_import: '1'
status: public
title: Responsive lysogeny under nonproductive phage binding
tmp:
  image: /images/cc_by.png
  legal_code_url: https://creativecommons.org/licenses/by/4.0/legalcode
  name: Creative Commons Attribution 4.0 International Public License (CC-BY 4.0)
  short: CC BY (4.0)
type: journal_article
user_id: 2DF688A6-F248-11E8-B48F-1D18A9856A87
volume: 80
year: '2026'
...
---
OA_place: publisher
_id: '20470'
abstract:
- lang: eng
  text: "Systems design has classically relied on composable systems, in which individual
    subsystems\r\nhave defined inputs, outputs, and interactions with each other;
    however, attempts at\r\ndesigning complex systems in synthetic biology has often
    run in to issues of crosstalk and\r\ninterference, given that these systems must
    function within the context of the host. In nature,\r\nmobile genetic elements
    are systems that have evolved to travel between hosts, and thus\r\nappear to be
    a good candidate with which to evaluate composability. Selecting temperate\r\nphages
    as a model system, I used mathematical modelling to identify sources of information\r\nthat
    temperate phages should respond to. I found that essential proteins of temperate
    phages\r\ncan interfere with potential hosts, indicating limitations to composability.
    I also designed a\r\nlysogeny reporter construct and characterize its behavior
    across various laboratory and\r\nenvironmental strains, finding differences in
    phage lambda lysogens, and potential\r\ninterference from prophages that already
    exist within the environmental strains. Although\r\nthe information gathered is
    not conclusive, it suggests that composability is not a key property\r\nof temperate
    phages, implying that biological systems may not be composable, and that other\r\nsystem
    design principles should be considered when designing synthetic systems."
alternative_title:
- ISTA Thesis
article_processing_charge: No
author:
- first_name: Bryan
  full_name: Wu, Bryan
  id: 3C521EBA-F248-11E8-B48F-1D18A9856A87
  last_name: Wu
citation:
  ama: Wu B. An examination on phages as a naturally composable system. 2025. doi:<a
    href="https://doi.org/10.15479/AT-ISTA-20470">10.15479/AT-ISTA-20470</a>
  apa: Wu, B. (2025). <i>An examination on phages as a naturally composable system</i>.
    Institute of Science and Technology Austria. <a href="https://doi.org/10.15479/AT-ISTA-20470">https://doi.org/10.15479/AT-ISTA-20470</a>
  chicago: Wu, Bryan. “An Examination on Phages as a Naturally Composable System.”
    Institute of Science and Technology Austria, 2025. <a href="https://doi.org/10.15479/AT-ISTA-20470">https://doi.org/10.15479/AT-ISTA-20470</a>.
  ieee: B. Wu, “An examination on phages as a naturally composable system,” Institute
    of Science and Technology Austria, 2025.
  ista: Wu B. 2025. An examination on phages as a naturally composable system. Institute
    of Science and Technology Austria.
  mla: Wu, Bryan. <i>An Examination on Phages as a Naturally Composable System</i>.
    Institute of Science and Technology Austria, 2025, doi:<a href="https://doi.org/10.15479/AT-ISTA-20470">10.15479/AT-ISTA-20470</a>.
  short: B. Wu, An Examination on Phages as a Naturally Composable System, Institute
    of Science and Technology Austria, 2025.
corr_author: '1'
date_created: 2025-10-15T13:30:21Z
date_published: 2025-10-30T00:00:00Z
date_updated: 2026-05-06T08:01:28Z
day: '30'
ddc:
- '579'
degree_awarded: PhD
department:
- _id: GradSch
- _id: CaGu
doi: 10.15479/AT-ISTA-20470
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  date_created: 2025-10-21T17:33:27Z
  date_updated: 2026-04-30T22:30:02Z
  embargo_to: open_access
  file_id: '20516'
  file_name: 2025_Wu_Bryan_Thesis.docx
  file_size: 10603235
  relation: source_file
- access_level: open_access
  checksum: 53590046fd3244c5550b4022282449d2
  content_type: application/pdf
  creator: brwu
  date_created: 2025-10-21T17:33:26Z
  date_updated: 2026-04-30T22:30:02Z
  embargo: 2026-04-30
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  file_size: 6251936
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file_date_updated: 2026-04-30T22:30:02Z
has_accepted_license: '1'
language:
- iso: eng
month: '10'
oa: 1
oa_version: Published Version
page: '102'
publication_identifier:
  issn:
  - 2663-337X
publication_status: published
publisher: Institute of Science and Technology Austria
status: public
supervisor:
- first_name: Calin C
  full_name: Guet, Calin C
  id: 47F8433E-F248-11E8-B48F-1D18A9856A87
  last_name: Guet
  orcid: 0000-0001-6220-2052
title: An examination on phages as a naturally composable system
type: dissertation
user_id: 8b945eb4-e2f2-11eb-945a-df72226e66a9
year: '2025'
...
