The physical properties of a marine microbe, its virus, and their environment on the co-evolutionary dynamics, NERC GW4+ DTP, PhD in Physics and Astronomy Ref: 3327

About the award


Lead Supervisor

Dr Wolfram Möbius, Living Systems Institute, University of Exeter

Additional Supervisors

Dr Ben Temperton, Department of Biosciences, College of Life and Environmental Sciences, University of Exeter

Dr Stefano Pagliara, Living Systems Institute, University of Exeter

Prof Mike Allen, Plymouth Marine Laboratory and University of Exeter

Location: University of Exeter, Streatham Campus, Exeter EX4 4QJ

Main Information

This project is one of a number that are in competition for funding from the NERC Great Western Four+ Doctoral Training Partnership (GW4+ DTP).  The GW4+ DTP consists of the Great Western Four alliance of the University of Bath, University of Bristol, Cardiff University and the University of Exeter plus five Research Organisation partners:  British Antarctic Survey, British Geological Survey, Centre for Ecology and Hydrology, the Natural History Museum and Plymouth Marine Laboratory.  The partnership aims to provide a broad training in earth and environmental sciences, designed to train tomorrow’s leaders in earth and environmental science. For further details about the programme please see

For eligible successful applicants, the studentships comprises:

  • An index-linked stipend for 3.5 years (currently £14,777 p.a. for 2018/19);
  • Payment of university tuition fees;
  • A research budget of £11,000 for an international conference, lab, field and research expenses;
  • A training budget of £4,000 for specialist training courses and expenses.

Up to 30 fully-funded studentships will be available across the partnership.

Students from EU countries who do not meet the residency requirements may still be eligible for a fees-only award but no stipend.  Applicants who are classed as International for tuition fee purposes are not eligible for funding.

Project details

With approximately 10 viral particles for every marine microbe in surface water, viral infection of bacterial cells plays a significant role in driving population structure and, in turn, is a fundamental component of global carbon biogeochemistry. The vast majority of marine bacteria and their viruses are extremely small and dilute. The co-evolutionary consequences thereof are poorly understood. For example, it was thought until recently that the size of ultra-small bacteria such as the ubiquitous Pelagibacter spp. enabled avoidance of viral infection through ‘cryptic escape’. Yet, in 2013, this idea was proved false by the discovery of viruses infecting Pelagibacter, which dominate global oceans.

This project will investigate the coevolution of host and virus and how they are shaped by the biological and physical consequences of their sizes as well as by life in a marine environment.

Project Aims and Methods

The aim of this project is to investigate the co-evolutionary consequences of host / virus size and complex encounter patterns due to fluid flow in the ocean. To achieve this goal, the student will compare co-evolution of Pelagibacter and its virus with co-evolution of a range of host and virus pairs with varying size. Co-evolutionary adaptations will be investigated using single-cell genomic analysis and atomic force microscopy to visualise genotypic and phenotypic changes (focusing on cell-membrane adaptations), respectively, within the environmental single cell genomics facility at Plymouth Marine Laboratory.

In a second step, co-evolution experiments will be performed in a range of different fluidic environments. In particular, different scenarios of encounters of bacteria and virus in the ocean will be mimicked in the laboratory through the exquisite control of compartmentalised microfluidic environments. The theoretical considerations underlying this approach can be used to study the consequences of fluid flow on co-evolution theoretically if the student is interested.

While the focus of the work will be on experimental co-evolution, the project can contain a varying amount of technology development and theory work. Following a period of initial training, the student will thus be encouraged to develop the project design together with the supervisors.


This project brings together fundamental biological questions with a physics-driven perspective through highly quantitative evolution experiments. The supervisors bring in complementary expertise from microbial ecology and evolution, single cell biology, microfluidics, microscopy, bioinformatics, and modelling. The student will thus obtain highly interdisciplinary training and be exposed to a range of techniques and approaches to science. Through the range of techniques, the student will learn to communicate across discipline boundaries, training that will be supported through visits to international conferences. The project will require creative approaches, allowing the student to develop their own project based on their own ideas.

References / Background reading list

Zhao et al., Nature 494, 357-360 (2013)

see also:

Koskella and Brockhurst, FEMS Microbiol Rev 38, 916-931 (2014)


Entry requirements

Applicants should have obtained, or be about to obtain, a First or Upper Second Class UK Honours degree, or the equivalent qualifications gained outside the UK.   Applicants with a Lower Second Class degree will be considered if they also have Master’s degree.  Applicants with a minimum of Upper Second Class degree and significant relevant non-academic experience are encouraged to apply.

Candidate Requirements
The candidate will have achieved at least a 2:1 Honours degree (or equivalent) in a physical or biological science. Preference will be given to candidates with experience in working in a biological laboratory and/or with experience in fluid dynamics.

All applicants would need to meet our English language requirements by the start of the  project


How to apply

In the application process you will be asked to upload several documents.  Please note our preferred format is PDF, each file named with your surname and the name of the document, eg. “Smith – CV.pdf”, “Smith – Cover Letter.pdf”, “Smith – Transcript.pdf”.

  • CV
  • Letter of application outlining your academic interests, prior research experience and reasons for wishing to undertake the project.
  • Transcript(s) giving full details of subjects studied and grades/marks obtained.  This should be an interim transcript if you are still studying.
  • If you are not a national of a majority English-speaking country you will need to submit evidence of your current proficiency in English.
  • Two References (applicants are recommended to have a third academic referee, if the two academic referees are within the same department/school).

Reference information
You will be asked to name two referees as part of the application process.  It is your responsibility to ensure that your two referees email their references to, as we will not make requests for references directly; you must arrange for them to be submitted by 7 January 2019

References should be submitted to us directly in the form of a letter. Referees must email their references to us from their institutional email accounts. We cannot accept references from personal/private email accounts, unless it is a scanned document on institutional headed paper and signed by the referee.

All application documents must be submitted in English. Certified translated copies of academic qualifications must also be provided.

The closing date for applications is midnight on 7 January 2019.  Interviews will be held between 4 and 15 February 2019.

If you have any general enquiries about the application process please email  Project-specific queries should be directed to the supervisor.

Data Sharing
During the application process, the University may need to make certain disclosures of your personal data to third parties to be able to administer your application, carry out interviews and select candidates.  These are not limited to, but may include disclosures to:

  • the selection panel and/or management board or equivalent of the relevant programme, which is likely to include staff from one or more other HEIs;
  • administrative staff at one or more other HEIs participating in the relevant programme.

Such disclosures will always be kept to the minimum amount of personal data required for the specific purpose. Your sensitive personal data (relating to disability and race/ethnicity) will not be disclosed without your explicit consent.


Application deadline:7th January 2019
Value:£14,777 per annum for 2018-19
Duration of award:per year
Contact: PGR Enquiries