---
OA_place: publisher
_id: '20811'
abstract:
- lang: eng
  text: "\tThis thesis is organized into two parts, each comprising two chapters:
    Chapter 1 and 2 offer models for the evolution of vaccine resistance in response
    to diverse vaccination strategies. Chapter 3 and 4 review the statistics of records,
    their connection to models of innovation and an application to the cultural evolution
    of sports.\r\n\tIn chapter 1 we present a modelling study from 2021 on the evolution
    of SARS-CoV-2. At that time the vaccine-resistant Omicron variant had not yet
    evolved. In our model we consider a population that is becoming vaccinated over
    time, while a pathogen is spreading in the population and eventually becoming
    resistant to the vaccine. We explore effective pharmaceutical and non-pharmaceutical
    interventions to prevent the emergence of vaccine resistance. \r\n\tIn chapter
    2 we model a particular set of complex vaccination strategies, mosaic and pyramid
    vaccination, where an immunologically diverse portfolio of vaccines is considered.
    We find that a bet-hatching strategy, in which vaccine types are distributed in
    the population, is effective at hindering the evolution of vaccine resistance
    if mutation rates are high. \r\n\tIn chapter 3 we switch gears and present a review
    on the statistics of records. We highlight similarities and analogies to other
    models in the fields of statistical physics, evolution and innovation. This offers
    interesting complimentary perspectives on well-known models. \r\n\tIn chapter
    4 we apply models of record statistics and innovation to study cultural evolution
    in sport. We propose a model of sport evolution that combines deterministic improvements
    in performance and stochastic bursts of improvements due to innovation. "
alternative_title:
- ISTA Thesis
article_processing_charge: No
author:
- first_name: Simon
  full_name: Rella, Simon
  id: B4765ACA-AA38-11E9-AC9A-0930E6697425
  last_name: Rella
citation:
  ama: 'Rella S. Adaptive processes in biology and culture: Models of evolving vaccine
    resistance and the record statistics of innovation. 2025. doi:<a href="https://doi.org/10.15479/AT-ISTA-20811">10.15479/AT-ISTA-20811</a>'
  apa: 'Rella, S. (2025). <i>Adaptive processes in biology and culture: Models of
    evolving vaccine resistance and the record statistics of innovation</i>. Institute
    of Science and Technology Austria. <a href="https://doi.org/10.15479/AT-ISTA-20811">https://doi.org/10.15479/AT-ISTA-20811</a>'
  chicago: 'Rella, Simon. “Adaptive Processes in Biology and Culture: Models of Evolving
    Vaccine Resistance and the Record Statistics of Innovation.” Institute of Science
    and Technology Austria, 2025. <a href="https://doi.org/10.15479/AT-ISTA-20811">https://doi.org/10.15479/AT-ISTA-20811</a>.'
  ieee: 'S. Rella, “Adaptive processes in biology and culture: Models of evolving
    vaccine resistance and the record statistics of innovation,” Institute of Science
    and Technology Austria, 2025.'
  ista: 'Rella S. 2025. Adaptive processes in biology and culture: Models of evolving
    vaccine resistance and the record statistics of innovation. Institute of Science
    and Technology Austria.'
  mla: 'Rella, Simon. <i>Adaptive Processes in Biology and Culture: Models of Evolving
    Vaccine Resistance and the Record Statistics of Innovation</i>. Institute of Science
    and Technology Austria, 2025, doi:<a href="https://doi.org/10.15479/AT-ISTA-20811">10.15479/AT-ISTA-20811</a>.'
  short: 'S. Rella, Adaptive Processes in Biology and Culture: Models of Evolving
    Vaccine Resistance and the Record Statistics of Innovation, Institute of Science
    and Technology Austria, 2025.'
corr_author: '1'
date_created: 2025-12-12T14:39:56Z
date_published: 2025-12-15T00:00:00Z
date_updated: 2026-10-02T10:12:19Z
day: '15'
ddc:
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- '576'
- '614'
- '519'
degree_awarded: PhD
department:
- _id: GradSch
- _id: GaTk
- _id: FyKo
doi: 10.15479/AT-ISTA-20811
doi_confirm: '1'
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fulldoi: https://doi.org/10.15479/AT-ISTA-20811
has_accepted_license: '1'
language:
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license: https://creativecommons.org/licenses/by-nc-nd/4.0/
month: '12'
oa: 1
oa_version: Published Version
page: '95'
publication_identifier:
  issn:
  - 2663-337X
publication_status: published
publisher: Institute of Science and Technology Austria
related_material:
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status: public
supervisor:
- first_name: Gašper
  full_name: Tkačik, Gašper
  id: 3D494DCA-F248-11E8-B48F-1D18A9856A87
  last_name: Tkačik
  orcid: 0000-0002-6699-1455
- first_name: Fyodor
  full_name: Kondrashov, Fyodor
  id: 44FDEF62-F248-11E8-B48F-1D18A9856A87
  last_name: Kondrashov
  orcid: 0000-0001-8243-4694
title: 'Adaptive processes in biology and culture: Models of evolving vaccine resistance
  and the record statistics of innovation'
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  short: CC BY-NC-ND (4.0)
type: dissertation
user_id: 8b945eb4-e2f2-11eb-945a-df72226e66a9
year: '2025'
...
---
OA_place: publisher
OA_type: hybrid
_id: '18307'
abstract:
- lang: eng
  text: Vaccination is the most effective tool to control infectious diseases. However,
    the evolution of vaccine resistance, exemplified by vaccine resistance in SARS-CoV-2,
    remains a concern. Here, we model complex vaccination strategies against a pathogen
    with multiple epitopes—molecules targeted by the vaccine. We found that a vaccine
    targeting one epitope was ineffective in preventing vaccine escape. Vaccine resistance
    in highly infectious pathogens was prevented by the full-epitope vaccine, that
    is, one targeting all available epitopes, but only when the rate of pathogen evolution
    was low. Strikingly, a bet-hedging strategy of random administration of vaccines
    targeting different epitopes was the most effective in preventing vaccine resistance
    in pathogens with the low rate of infection and high rate of evolution. Thus,
    complex vaccination strategies, when biologically feasible, may be preferable
    to the currently used single-vaccine approaches for long-term control of disease
    outbreaks, especially when applied to livestock with near 100% vaccination rates.
acknowledgement: We thank Raimundo Julian Saona Urmeneta, Maike Morrison, Sergey Kryazhimskiy,
  Hiroki Ishikawa, Simone Pigolotti, and Shingo Miyauchi for fruitful discussions.
  We also thank the participants of the FRISBI seminar at ISTA for useful comments.
article_processing_charge: Yes (via OA deal)
article_type: original
author:
- first_name: Simon
  full_name: Rella, Simon
  id: B4765ACA-AA38-11E9-AC9A-0930E6697425
  last_name: Rella
- first_name: Yuliya A.
  full_name: Kulikova, Yuliya A.
  last_name: Kulikova
- first_name: Aygul
  full_name: Minnegalieva, Aygul
  id: 87DF77F0-1D9A-11EA-B6AE-CE443DDC885E
  last_name: Minnegalieva
- first_name: Fyodor
  full_name: Kondrashov, Fyodor
  id: 44FDEF62-F248-11E8-B48F-1D18A9856A87
  last_name: Kondrashov
  orcid: 0000-0001-8243-4694
citation:
  ama: 'Rella S, Kulikova YA, Minnegalieva A, Kondrashov F. Complex vaccination strategies
    prevent the emergence of vaccine resistance. <i>Evolution: International journal
    of organic evolution</i>. 2024;78(10):1722-1738. doi:<a href="https://doi.org/10.1093/evolut/qpae106">10.1093/evolut/qpae106</a>'
  apa: 'Rella, S., Kulikova, Y. A., Minnegalieva, A., &#38; Kondrashov, F. (2024).
    Complex vaccination strategies prevent the emergence of vaccine resistance. <i>Evolution:
    International Journal of Organic Evolution</i>. Oxford University Press. <a href="https://doi.org/10.1093/evolut/qpae106">https://doi.org/10.1093/evolut/qpae106</a>'
  chicago: 'Rella, Simon, Yuliya A. Kulikova, Aygul Minnegalieva, and Fyodor Kondrashov.
    “Complex Vaccination Strategies Prevent the Emergence of Vaccine Resistance.”
    <i>Evolution: International Journal of Organic Evolution</i>. Oxford University
    Press, 2024. <a href="https://doi.org/10.1093/evolut/qpae106">https://doi.org/10.1093/evolut/qpae106</a>.'
  ieee: 'S. Rella, Y. A. Kulikova, A. Minnegalieva, and F. Kondrashov, “Complex vaccination
    strategies prevent the emergence of vaccine resistance,” <i>Evolution: International
    journal of organic evolution</i>, vol. 78, no. 10. Oxford University Press, pp.
    1722–1738, 2024.'
  ista: 'Rella S, Kulikova YA, Minnegalieva A, Kondrashov F. 2024. Complex vaccination
    strategies prevent the emergence of vaccine resistance. Evolution: International
    journal of organic evolution. 78(10), 1722–1738.'
  mla: 'Rella, Simon, et al. “Complex Vaccination Strategies Prevent the Emergence
    of Vaccine Resistance.” <i>Evolution: International Journal of Organic Evolution</i>,
    vol. 78, no. 10, Oxford University Press, 2024, pp. 1722–38, doi:<a href="https://doi.org/10.1093/evolut/qpae106">10.1093/evolut/qpae106</a>.'
  short: 'S. Rella, Y.A. Kulikova, A. Minnegalieva, F. Kondrashov, Evolution: International
    Journal of Organic Evolution 78 (2024) 1722–1738.'
corr_author: '1'
das_tickbox: '1'
dataavailabilitystatement: Results and figures derive from the model, described in
  detail in the main text. The code is available at github.com/Simon-Re/complex-vaccination.
date_created: 2024-10-13T22:01:50Z
date_published: 2024-10-01T00:00:00Z
date_updated: 2026-10-02T10:12:20Z
day: '01'
ddc:
- '570'
department:
- _id: GaTk
doi: 10.1093/evolut/qpae106
external_id:
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  - '38990788'
file:
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intvolume: '        78'
isi: 1
issue: '10'
language:
- iso: eng
month: '10'
oa: 1
oa_version: Published Version
page: 1722-1738
pmid: 1
publication: 'Evolution: International journal of organic evolution'
publication_identifier:
  eissn:
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publication_status: published
publisher: Oxford University Press
quality_controlled: '1'
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title: Complex vaccination strategies prevent the emergence of vaccine resistance
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volume: 78
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...
---
_id: '14901'
abstract:
- lang: eng
  text: Global services like navigation, communication, and Earth observation have
    increased dramatically in the 21st century due to advances in outer space industries.
    But as orbits become increasingly crowded with both satellites and inevitable
    space debris pollution, continued operations become endangered by the heightened
    risks of debris collisions in orbit. Kessler Syndrome is the term for when a critical
    threshold of orbiting debris triggers a runaway positive feedback loop of debris
    collisions, creating debris congestion that can render orbits unusable. As this
    potential tipping point becomes more widely recognized, there have been renewed
    calls for debris mitigation and removal. Here, we combine complex systems and
    social-ecological systems approaches to study how these efforts may affect space
    debris accumulation and the likelihood of reaching Kessler Syndrome. Specifically,
    we model how debris levels are affected by future launch rates, cleanup activities,
    and collisions between extant debris. We contextualize and interpret our dynamic
    model within a discussion of existing space debris governance and other social,
    economic, and geopolitical factors that may influence effective collective management
    of the orbital commons. In line with previous studies, our model finds that debris
    congestion may be reached in less than 200 years, though a holistic management
    strategy combining removal and mitigation actions can avoid such outcomes while
    continuing space activities. Moreover, although active debris removal may be particularly
    effective, the current lack of market and governance support may impede its implementation.
    Research into these critical dynamics and the multi-faceted variables that influence
    debris outcomes can support policymakers in curating impactful governance strategies
    and realistic transition pathways to sustaining debris-free orbits. Overall, our
    study is useful for communicating about space debris sustainability in policy
    and education settings by providing an exploration of policy portfolio options
    supported by a simple and clear social-ecological modeling approach.
acknowledgement: The authors would like to thank the special issue co-editors, Marco
  Janssen and Xiao-Shan Yap, and the anonymous reviewers for their comments that helped
  improve the manuscript. The paper also benefited from suggestions by other author
  participants in this special issue. We would also like to thank the 2022 Santa Fe
  Institute Complex Systems Summer School for providing space to initiate this study.
article_processing_charge: Yes
article_type: original
author:
- first_name: Keiko
  full_name: Nomura, Keiko
  last_name: Nomura
- first_name: Simon
  full_name: Rella, Simon
  id: B4765ACA-AA38-11E9-AC9A-0930E6697425
  last_name: Rella
- first_name: Haily
  full_name: Merritt, Haily
  last_name: Merritt
- first_name: Mathieu
  full_name: Baltussen, Mathieu
  last_name: Baltussen
- first_name: Darcy
  full_name: Bird, Darcy
  last_name: Bird
- first_name: Annika
  full_name: Tjuka, Annika
  last_name: Tjuka
- first_name: Dan
  full_name: Falk, Dan
  last_name: Falk
citation:
  ama: Nomura K, Rella S, Merritt H, et al. Tipping points of space debris in low
    earth orbit. <i>International Journal of the Commons</i>. 2024;18(1). doi:<a href="https://doi.org/10.5334/ijc.1275">10.5334/ijc.1275</a>
  apa: Nomura, K., Rella, S., Merritt, H., Baltussen, M., Bird, D., Tjuka, A., &#38;
    Falk, D. (2024). Tipping points of space debris in low earth orbit. <i>International
    Journal of the Commons</i>. Ubiquity Press. <a href="https://doi.org/10.5334/ijc.1275">https://doi.org/10.5334/ijc.1275</a>
  chicago: Nomura, Keiko, Simon Rella, Haily Merritt, Mathieu Baltussen, Darcy Bird,
    Annika Tjuka, and Dan Falk. “Tipping Points of Space Debris in Low Earth Orbit.”
    <i>International Journal of the Commons</i>. Ubiquity Press, 2024. <a href="https://doi.org/10.5334/ijc.1275">https://doi.org/10.5334/ijc.1275</a>.
  ieee: K. Nomura <i>et al.</i>, “Tipping points of space debris in low earth orbit,”
    <i>International Journal of the Commons</i>, vol. 18, no. 1. Ubiquity Press, 2024.
  ista: Nomura K, Rella S, Merritt H, Baltussen M, Bird D, Tjuka A, Falk D. 2024.
    Tipping points of space debris in low earth orbit. International Journal of the
    Commons. 18(1).
  mla: Nomura, Keiko, et al. “Tipping Points of Space Debris in Low Earth Orbit.”
    <i>International Journal of the Commons</i>, vol. 18, no. 1, Ubiquity Press, 2024,
    doi:<a href="https://doi.org/10.5334/ijc.1275">10.5334/ijc.1275</a>.
  short: K. Nomura, S. Rella, H. Merritt, M. Baltussen, D. Bird, A. Tjuka, D. Falk,
    International Journal of the Commons 18 (2024).
das_tickbox: '0'
date_created: 2024-01-30T11:58:02Z
date_published: 2024-01-11T00:00:00Z
date_updated: 2026-10-02T11:22:03Z
day: '11'
ddc:
- '550'
department:
- _id: GradSch
- _id: GaTk
doi: 10.5334/ijc.1275
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fulldoi: https://doi.org/10.5334/ijc.1275
has_accepted_license: '1'
intvolume: '        18'
issue: '1'
keyword:
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language:
- iso: eng
license: https://creativecommons.org/licenses/by/4.0/
month: '01'
oa: 1
oa_version: Published Version
publication: International Journal of the Commons
publication_identifier:
  issn:
  - 1875-0281
publication_status: published
publisher: Ubiquity Press
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supplementarymaterial: yes
title: Tipping points of space debris in low earth orbit
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volume: 18
year: '2024'
...
---
_id: '14862'
article_number: ckad160.597
article_processing_charge: No
author:
- first_name: Simon
  full_name: Rella, Simon
  id: B4765ACA-AA38-11E9-AC9A-0930E6697425
  last_name: Rella
- first_name: Y
  full_name: Kulikova, Y
  last_name: Kulikova
- first_name: Aygul
  full_name: Minnegalieva, Aygul
  id: 87DF77F0-1D9A-11EA-B6AE-CE443DDC885E
  last_name: Minnegalieva
- first_name: Fyodor
  full_name: Kondrashov, Fyodor
  id: 44FDEF62-F248-11E8-B48F-1D18A9856A87
  last_name: Kondrashov
  orcid: 0000-0001-8243-4694
citation:
  ama: 'Rella S, Kulikova Y, Minnegalieva A, Kondrashov F. Complex vaccination strategies
    prevent the emergence of vaccine resistance. In: <i>European Journal of Public
    Health</i>. Vol 33. Oxford University Press; 2023. doi:<a href="https://doi.org/10.1093/eurpub/ckad160.597">10.1093/eurpub/ckad160.597</a>'
  apa: Rella, S., Kulikova, Y., Minnegalieva, A., &#38; Kondrashov, F. (2023). Complex
    vaccination strategies prevent the emergence of vaccine resistance. In <i>European
    Journal of Public Health</i> (Vol. 33). Oxford University Press. <a href="https://doi.org/10.1093/eurpub/ckad160.597">https://doi.org/10.1093/eurpub/ckad160.597</a>
  chicago: Rella, Simon, Y Kulikova, Aygul Minnegalieva, and Fyodor Kondrashov. “Complex
    Vaccination Strategies Prevent the Emergence of Vaccine Resistance.” In <i>European
    Journal of Public Health</i>, Vol. 33. Oxford University Press, 2023. <a href="https://doi.org/10.1093/eurpub/ckad160.597">https://doi.org/10.1093/eurpub/ckad160.597</a>.
  ieee: S. Rella, Y. Kulikova, A. Minnegalieva, and F. Kondrashov, “Complex vaccination
    strategies prevent the emergence of vaccine resistance,” in <i>European Journal
    of Public Health</i>, 2023, vol. 33, no. Supplement_2.
  ista: Rella S, Kulikova Y, Minnegalieva A, Kondrashov F. 2023. Complex vaccination
    strategies prevent the emergence of vaccine resistance. European Journal of Public
    Health. vol. 33, ckad160.597.
  mla: Rella, Simon, et al. “Complex Vaccination Strategies Prevent the Emergence
    of Vaccine Resistance.” <i>European Journal of Public Health</i>, vol. 33, no.
    Supplement_2, ckad160.597, Oxford University Press, 2023, doi:<a href="https://doi.org/10.1093/eurpub/ckad160.597">10.1093/eurpub/ckad160.597</a>.
  short: S. Rella, Y. Kulikova, A. Minnegalieva, F. Kondrashov, in:, European Journal
    of Public Health, Oxford University Press, 2023.
corr_author: '1'
date_created: 2024-01-22T12:02:28Z
date_published: 2023-10-01T00:00:00Z
date_updated: 2026-10-02T10:12:19Z
day: '01'
ddc:
- '570'
department:
- _id: GaTk
doi: 10.1093/eurpub/ckad160.597
file:
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file_date_updated: 2024-01-24T11:12:33Z
fulldoi: https://doi.org/10.1093/eurpub/ckad160.597
has_accepted_license: '1'
intvolume: '        33'
issue: Supplement_2
keyword:
- Public Health
- Environmental and Occupational Health
language:
- iso: eng
license: https://creativecommons.org/licenses/by-nc/4.0/
month: '10'
oa: 1
oa_version: Published Version
publication: European Journal of Public Health
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publication_status: published
publisher: Oxford University Press
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title: Complex vaccination strategies prevent the emergence of vaccine resistance
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type: conference_abstract
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volume: 33
year: '2023'
...
---
_id: '9905'
abstract:
- lang: eng
  text: Vaccines are thought to be the best available solution for controlling the
    ongoing SARS-CoV-2 pandemic. However, the emergence of vaccine-resistant strains
    may come too rapidly for current vaccine developments to alleviate the health,
    economic and social consequences of the pandemic. To quantify and characterize
    the risk of such a scenario, we created a SIR-derived model with initial stochastic
    dynamics of the vaccine-resistant strain to study the probability of its emergence
    and establishment. Using parameters realistically resembling SARS-CoV-2 transmission,
    we model a wave-like pattern of the pandemic and consider the impact of the rate
    of vaccination and the strength of non-pharmaceutical intervention measures on
    the probability of emergence of a resistant strain. As expected, we found that
    a fast rate of vaccination decreases the probability of emergence of a resistant
    strain. Counterintuitively, when a relaxation of non-pharmaceutical interventions
    happened at a time when most individuals of the population have already been vaccinated
    the probability of emergence of a resistant strain was greatly increased. Consequently,
    we show that a period of transmission reduction close to the end of the vaccination
    campaign can substantially reduce the probability of resistant strain establishment.
    Our results suggest that policymakers and individuals should consider maintaining
    non-pharmaceutical interventions and transmission-reducing behaviours throughout
    the entire vaccination period.
acknowledgement: We thank Alexey Kondrashov, Nick Machnik, Raimundo Julian Saona Urmeneta,
  Gasper Tkacik and Nick Barton for fruitful discussions. We also thank participants
  of EvoLunch seminar at IST Austria and the internal seminar at the Banco de España
  for useful comments. The opinions expressed in this document are exclusively of
  the authors and, therefore, do not necessarily coincide with those of the Banco
  de España or the Eurosystem. ETD is supported by the Swiss National Science and
  Louis Jeantet Foundation. The work of FAK was in part supported by the ERC Consolidator
  Grant (771209-CharFL).
article_number: '15729'
article_processing_charge: Yes
article_type: original
author:
- first_name: Simon
  full_name: Rella, Simon
  id: B4765ACA-AA38-11E9-AC9A-0930E6697425
  last_name: Rella
- first_name: Yuliya A.
  full_name: Kulikova, Yuliya A.
  last_name: Kulikova
- first_name: Emmanouil T.
  full_name: Dermitzakis, Emmanouil T.
  last_name: Dermitzakis
- first_name: Fyodor
  full_name: Kondrashov, Fyodor
  id: 44FDEF62-F248-11E8-B48F-1D18A9856A87
  last_name: Kondrashov
  orcid: 0000-0001-8243-4694
citation:
  ama: Rella S, Kulikova YA, Dermitzakis ET, Kondrashov F. Rates of SARS-CoV-2 transmission
    and vaccination impact the fate of vaccine-resistant strains. <i>Scientific Reports</i>.
    2021;11(1). doi:<a href="https://doi.org/10.1038/s41598-021-95025-3">10.1038/s41598-021-95025-3</a>
  apa: Rella, S., Kulikova, Y. A., Dermitzakis, E. T., &#38; Kondrashov, F. (2021).
    Rates of SARS-CoV-2 transmission and vaccination impact the fate of vaccine-resistant
    strains. <i>Scientific Reports</i>. Springer Nature. <a href="https://doi.org/10.1038/s41598-021-95025-3">https://doi.org/10.1038/s41598-021-95025-3</a>
  chicago: Rella, Simon, Yuliya A. Kulikova, Emmanouil T. Dermitzakis, and Fyodor
    Kondrashov. “Rates of SARS-CoV-2 Transmission and Vaccination Impact the Fate
    of Vaccine-Resistant Strains.” <i>Scientific Reports</i>. Springer Nature, 2021.
    <a href="https://doi.org/10.1038/s41598-021-95025-3">https://doi.org/10.1038/s41598-021-95025-3</a>.
  ieee: S. Rella, Y. A. Kulikova, E. T. Dermitzakis, and F. Kondrashov, “Rates of
    SARS-CoV-2 transmission and vaccination impact the fate of vaccine-resistant strains,”
    <i>Scientific Reports</i>, vol. 11, no. 1. Springer Nature, 2021.
  ista: Rella S, Kulikova YA, Dermitzakis ET, Kondrashov F. 2021. Rates of SARS-CoV-2
    transmission and vaccination impact the fate of vaccine-resistant strains. Scientific
    Reports. 11(1), 15729.
  mla: Rella, Simon, et al. “Rates of SARS-CoV-2 Transmission and Vaccination Impact
    the Fate of Vaccine-Resistant Strains.” <i>Scientific Reports</i>, vol. 11, no.
    1, 15729, Springer Nature, 2021, doi:<a href="https://doi.org/10.1038/s41598-021-95025-3">10.1038/s41598-021-95025-3</a>.
  short: S. Rella, Y.A. Kulikova, E.T. Dermitzakis, F. Kondrashov, Scientific Reports
    11 (2021).
date_created: 2021-08-15T22:01:26Z
date_published: 2021-07-30T00:00:00Z
date_updated: 2026-07-29T12:57:49Z
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doi: 10.1038/s41598-021-95025-3
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title: Rates of SARS-CoV-2 transmission and vaccination impact the fate of vaccine-resistant
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