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

   School of Psychology

  ,  Applications accepted all year round  Funded PhD Project (Students Worldwide)

About the Project

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 hosted jointly by the 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) and will have opportunities to work in both Scotland and Germany.

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

Biological Sciences (4) Engineering (12) Materials Science (24) Physics (29)

Funding Notes

This project is jointly funded by the University of St Andrews and the University of Cologne.
See here for further information on the project and the hosts:
View Website
Applicants should submit their application to the co-supervisors. Applications will be accepted on a rolling basis until June 30 2023. Please send your application to Dr Stefan Pulver (St Andrews) and Professor Malte Gather (Cologne) at the following email addresses: ;
Your application should include the following:
1) Statement why you are the right candidate for the project (max. 500 words)
2) CV
3) Names and contact information for 2 References
4) Transcripts from undergraduate or postgraduate study


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:

How good is research at University of St Andrews in Psychology, Psychiatry and Neuroscience?

Research output data provided by the Research Excellence Framework (REF)

Click here to see the results for all UK universities

Register your interest for this project