Global Doctoral Scholarship - St Andrews and Cologne - Psychology and Biophotonics

Application period opens
Wednesday 15 February 2023
Application period closes
Saturday 15 April 2023

The University of St Andrews and the University of Cologne are pleased to offer a scholarship to support an exceptional student undertaking doctoral research in the following area:

Moving softly: how the interplay of neuronal circuits and biomechanics enables complex movement in soft-bodied animals

The student may start their degree at any point in the academic year 2023-2024 prior to a final entry date of 27 May 2024 subject to agreement with the supervisory team.

About St Andrews

As a doctoral student at the University of St Andrews you will be part of a growing, vibrant, and intellectually stimulating postgraduate community. The Pulver laboratory is based within the School of Psychology and Neuroscience at St Andrews. The Pulver lab supports a wide range of neuroethology experiments including Drosophila neurogenetics, electrophysiology, live imaging of neural activity, and animal behaviour. Research and teaching within the School bridges the disciplines of psychology, animal behaviour, and neuroscience. The School benefits from being embedded in an interdisciplinary research environment at St Andrews that includes the Institute for Behavioural and Neural Sciences (IBANS) and the Centre for Biophotonics (CoB).

About Cologne

Prof. Malte Gather is the director of the newly established Humboldt Centre for Nano- and Biophotonics. The centre provides state-of-the-art optics labs, a newly built clean-room, as well as in-house cell culture, fly keeping, and nanofabrication facilities. The interdisciplinary research benefits from the high-profile biomedical research environment established at the University of Cologne, including the Cluster of Excellence - Cellular Stress Responses in Aging-Associated Diseases (CECAD), and the Centre for Molecular Medicine Cologne (CMMC).

Commitment to equality, diversity, and inclusion

The Universities of St Andrews and Cologne are committed to promoting inclusivity, diversity, and equal opportunities for students and staff. Women and individuals from underrepresented minorities are especially encouraged to apply for this scholarship. All applicants will be considered in accordance with University of St Andrew’s equality, diversity and inclusion policies, as well as the Equal Opportunities Act of North Rhine-Westphalia (Landesgleichstellungsgesetz – LGG NRW).


Soft bodied animals can radically change their body shape, squeeze through cracks, then crawl, roll, tunnel, or even jump. Controlling soft bodies is difficult because they have highly non-linear physical properties and virtually unlimited degrees of freedom. Moving at all poses a challenge for soft animals as their muscles have no rigid levers to work against. How the brains of these animals solve these problems, and how these animals seem to sometimes even defy Newton’s mechanics remains relatively poorly understood. Apart from advancing fundamental understanding, solving these mysteries can contribute to the design of future (micro)robots with enhanced capabilities.

Progress in soft animal biomechanics has been limited by lack of suitable model organisms and tools for measuring animal behaviour and forces exerted on substrates while also manipulating neural activity. Working together, the Gather lab (University of Cologne) and the Pulver lab (University of St Andrews) have recently developed methods for measuring the forces exerted by freely behaving Drosophila larvae1. In this project, we will use Drosophila optogenetics coupled with a novel microscopy modality for measuring ground reaction forces to explore how neural activity in a soft animal is transformed into complex patterns of mechanical actions.

The student will first build instrumentation that integrates mechanical force measurement with behavioural imaging and optogenetic stimulation. This will enable us to remotely control brain regions with light, while also measuring behavioural repertoires and surface biomechanical forces generated during larval movements. Next, the student will use Drosophila optogenetic tools to turn on and off specific circuits within behaving larvae while measuring substrate interactions. We will systematically search for circuits that act as ‘linch-pins’ for enabling specific substrate interactions underlying soft-bodied movement. By following this up with anatomical analyses of those circuits and associated musculature, we can uncover direct functional relationships between particular neural circuits, muscles and actual surface forces generated by larvae. This type of fully integrated neuro-biomechanical approach will allow the student to gain training and expertise in state-of-the-art experimental approaches in optical physics, biomechanics, and neuroethology.

The project will be managed jointly between the School of Psychology and Neuroscience at St Andrews and the Humboldt Centre for Nano- and Biophotonics at University of Cologne, Germany. The student will be supervised by Dr Stefan Pulver (St Andrews) and Prof. Malte C. Gather (Cologne).

Informal enquiries regarding this scholarship may be addressed to the co-supervisors ( and


  1. Booth J, Meek AT, Kronenberg NM, Pulver SR, Gather MC. Optical mapping of ground reaction force dynamics in freely behaving Drosophila melanogaster larvae. BioRxiv 2022.10.21.513016. doi:

Value of award (per year)

The funding comprises a scholarship equivalent of a full-fees award and stipend for a period of up to 3.5 years. It is expected that the student will spend approximately half of the scholarship term at the University of St Andrews and half at the University of Cologne:

  • For the period spent at the University of St Andrews, the scholarship will comprise a full fees award and a stipend paid at the current UK Research Council rate (£17,668 each year in 2022–2023)
  • For the period spent at the University of Cologne, the scholarship will comprise a stipend of €1800/month.

Duration of award

Up to 3.5 years. The student will be expected to spend approximately half of the award term at the University of St Andrews and half at the University of Cologne. The successful candidate will be expected to have completed the doctorate degree by the end of the award term. The award term excludes the continuation period and any extension periods.

Application restrictions

Study level

Available to students studying at:


Domicile for fee status

No restrictions


Available to faculty members from:

Psychology and Neuroscience

Application assessment

Academic merit

Available to

Prospective students

Mode of study

Full time

Geographical criteria

No restrictions

Additional criteria

Admission and scholarship criteria of both universities must be met.

Applicants must not already (i) hold a doctoral degree; or (ii) be matriculated for a doctoral degree at the University of St Andrews or another institution.

How to apply

Please indicate in your application that you wish to be considered for this Global Doctoral scholarship (reference Pulver-Gather).

Terms and conditions of scholarships for St Andrews can be found here: Awards general terms and conditions.

Please contact the scholarships team and the supervisors if you have any questions regarding the scholarship:, and

Next steps

Once notified, successful candidates should proceed to fulfil the application requirements for each University and must meet all normal entry requirements for admission – please see the advice on applying for research degree programmes. Applications should indicate that the applicant is the recipient of a Global Doctoral Scholarship St Andrews (reference Pulver-Gather).

Terms and conditions

Please read the University of St Andrews scholarships terms and conditions (opens in new tab).

These are applicable during the St Andrews duration of the award; please consult the partner institution for their terms and conditions relating to scholarships.

When will I know the outcome?

We will aim to notify you of the outcome of your application by 15 May 2023. Awards are subject to final signatures of contractual relationships between the parties, and are not an indication of admission to the doctoral programme. Successful scholarship applicants must apply to both institutions and meet all relevant entry requirements for admission including any immigration requirements that may be in place.


Please contact us should you have any questions regarding the scholarship:

Informal enquiries regarding this scholarship may be addressed to the co-supervisors ( and