We investigate how the brain produces behaviour by looking at how movements are generated and adjusted to meet animals’ needs.


We are interested in the neuroscience of how animals move to meet their needs. We explore the mechanisms that underlie the generation of movements using a combination of multi-disciplinary approaches that include imaging, electrophysiology, and connectomics. We are also a maker lab, building custom microscopes and other setups to help us do the experiments we want to do.

 We use the zebrafish and fruit fly larva model organisms to see how the nervous systems of these diverse animals have evolved to move around in their environment.

About us

Cameron Burton, PhD student

I received my bachelor’s in neuroscience and biology in the United States at the University of Nebraska at Omaha. During my undergraduate studies, I worked in the lab of Dr. Ryan Wong investigating the effects of gene expression on variation in stress coping styles in zebrafish. After receiving my bachelor’s, I decided to come to St Andrews to join the Zwart lab as a part of a cotutelle PhD program between here and the University of Bonn. My project will investigate the descending control of behavior in Drosophila larvae.

Eleonora Gagliardi, PhD student

I graduated in General and Applied Biology at the University of Naples “Federico II”. Thanks to my bachelor thesis project, I had my first experience with the zebrafish model organism, studying prenatal stress. Then, I pursued a master’s degree in Molecular Biology at the same University. During the Master program, I experienced a one year exchange at the University of Cadiz, Spain. I based my master studies at Zoological Station of Naples, in the Locascio lab, focusing on BDNF zebrafish mutants and their locomotion behaviour. Then, I decided to expand my research field to neuroscience moving to Scotland for my PhD in the Zwart lab. I live in beautiful St Andrews, by the sea which I love.

Pierce Mullen, Leverhulme Early Career Fellow

I received my Bachelor in Pharmacology from the University of Leeds and stayed to pursue a PhD in Neuroscience in the Deuchars lab under the supervision of Dr Susan Deuchars. My PhD focused on the contribution of voltage-gated potassium channels to excitability in the spinal cord. At Leeds I further expanded my research towards the peripheral nervous system in Nikita Gamper’s lab, with a focus on decoding somatosensory information in peripheral nerves. I joined the Zwart lab to investigate how sensory information and motor drive are integrated to produce behaviour. I live with Sîan in picturesque Falkland.

Valentina Saccomanno, PhD student

After my BCs in Biological Science at the University of Sannio, I pursued a Master’s in Neuroscience between the University of Trieste and SISSA, in Italy. My interests for motor circuits brought me first in St Andrews for my master’s internship, which was carried out as exchange student in Dr Wenchang Li’s lab. I am now very excited to be back for my PhD! My project is co-supervised by Dr Maarten Zwart and Dr Wenchang Li, and will involve both zebrafish and frog tadpoles. Besides neuroscience, I like listening to rock music, reading classical books and walking by the sea.

Hesho Shaweis, PhD student

I received my Bachelor in Neuroscience at King’s College London. Later I moved to the Netherlands for my postgraduate studies at the Vrije Universiteit (VU), Amsterdam. During my masters I did an internship at the Centre for Neurogenomics and Cognitive Research (CNCR) in Matthijs Verhage’s labas well as at the Netherlands Institute for Neuroscience (NIN, KNAW) in Chris de Zeeuw’s lab. Here I worked on the role of the cerebellum in motor learning and memory processes. I’ve joined the Zwart lab for my PhD to further explore this system. When I’m not behind a microscope, I’ll be discovering the eastern shores.

Sarah Suber, PhD student

I graduated from the College of Charleston in South Carolina with a bachelor’s in molecular biology and minor in neuroscience. While studying at Charleston, I worked in Dr. Yukiko Sugi’s lab at the Medical University of South Carolina, investigating the interaction between BMP and Notch signalling during atrioventricular valvulogenesis. I then joined Dr. Mike Ruscio’s lab, where I studied FMRFamide distribution in snapping shrimp, Alpheus angulosus. After graduating, I moved across the pond to obtain a master’s in neuroscience at the University of Manchester, where I was supervised by Prof David Brough and explored the effects of squaramide-based ion transporters on apoptosis. I am very excited to have joined the community at St Andrews for my PhD. I am co-supervised by Dr. Maarten Zwart, Prof Karen Spencer, and Prof David Dritschel. My project explores whether zebrafish embryos can sense their environment and how neural development can be tuned to this environment.

Bella Xu Ying, Undergraduate Intern

I am currently undertaking my BSc (Hons) in Neuroscience right here at the University of St Andrews. During my 1st year I worked as a research assistant for PhD candidate Nick Maragkos at Dr Stefan Pulver’s lab, studying the effects of modulating environmental temperature on the pathophysiology of Drosophila ALS models. Later, I performed data analysis for Stefan and Maarten’s research looking into L-dopa-induced reconfiguration of Drosophila larval tunnelling behaviour. Soon I became an undergraduate researcher at both labs. Currently at the end of my 2nd year, I am co-supervised by Maarten and Stefan researching a new potential model organism in motor systems, Tribolium castaneum, characterising various larval motor programs and locomotor behaviours. When not in the lab, I like playing video games, drawing, goalkeeping and drumming to rock/metal music, but never all 4 at once!.

Hong Yan Zhang, Research Associate

My scientific journey began with a foundation in general medicine and human physiology at Qingdao University in China. Then I pursued a PhD in Animal Physiology at Radboud University Nijmegen, the Netherlands, focusing on voltage-gated calcium channels and electrophysiological techniques. Following my PhD, I joined Prof. Keith Sillar’s lab at the University of St Andrews, where I delved into the fascinating world of spinal locomotor control using young tadpoles as a model. This experience enabled me to further develop my career as a Chancellor’s Fellow at the University of Edinburgh. After the pandemic, I went back to St Andrews and joined Dr. Wenchang Li’s lab to investigate the role of spinal neurons that control tadpole struggling behaviour. Currently, I am thrilled to be part of the Zwart lab, exploring the intricate neural circuitry underlying sensory-motor integration in zebrafish larvae. Beyond the lab, I find joy in gardening and exploring the breathtaking landscapes of Scotland.

Maarten Zwart, PI

After my undergraduate degree in bioengineering at the University of Wageningen, I moved to the UK for my graduate studies at the University of Cambridge with Daniel St Johnston, Gurdon Institute. I started working on motor systems during my PhD in Matthias Landgraf’s lab in the department of Zoology, followed by postdocs with Albert Cardona and Misha Ahrens, both at the HHMI Janelia Research Campus. I’m currently at the School of Psychology and Neuroscience, University of St Andrews, in beautiful Scotland. I live with Sonja, Freya, and Otto in St Andrews.



Xu Ying B, Zwart MF*, Pulver SR*. Context-dependent coordination of movement in Tribolium castaneum larvae. BioRxiv doi: 10.1101/2024.06.17.598650 (*co-corresponding)

Jonaitis J, Hibbard K, McCafferty Layte K, Hiramoto A, Cardona A, Truman JW, Nose, A, Zwart MF, Pulver SR. Steering From the Rear: Coordination of Central Pattern Generators Underlying Navigation by Ascending Interneurons. BioRxiv doi: 10.1101/2024.06.17.598162


Liu Y, Hasegawa E, Nose A, Zwart MF*, Kohsaka H*. Synchronous multi-segmental activity between metachronal waves controls locomotion speed in Drosophila larvae. eLife. 2023 Aug 8; 12:e83328. doi:10.7554/eLife.83328 (*co-corresponding)

Mullen PN, Bowlby B, Armstrong HC, Zwart MF. PoseR – A deep learning toolbox for decoding animal behavior. BioRxiv doi: 10.1101/2022.09.08.507222


Yang E, Zwart MF, Rubinov M, James B, Wei Z, Narayan S,  Vladimirov N, Mensh BD, Fitzgerald JE, Ahrens MB. A brainstem integrator for self-localization and positional homeostasis. Cell. 2022 Dec 22; 185(26):5011-5027.e20.  doi: 10.1038/s41467-019-12693-6

Liu Y, Hasegawa E, Nose A, Zwart MF*, Kohsaka H*. Synchronous multi-segmental activity between metachronal waves controls locomotion speed in Drosophila larvae. BioRxiv doi: 10.1101/2022.09.08.507222 (*co-corresponding)


Yang E, Zwart MF, Rubinov M, James B, Wei Z, Narayan S,  Vladimirov N, Mensh BD, Fitzgerald JE, Ahrens MB. A brainstem integrator for self-localization and positional homeostasis. BioRxiv doi: 10.1101/2021.11.26.468907


Babski H, Jovanic T, Surel C, Yoshikawa S, Zwart MF, Valmier J, Thomas JB, Enriquez J, Carroll P, Garcès A. A GABAergic Maf-expressing interneuron subset regulates the speed of locomotion in Drosophila. Nat Commun. 2019 Oct 22;10(1):4796. doi: 10.1038/s41467-019-12693-6

Kohsaka H, Zwart MF, Fushiki A, Fetter RD, Truman JW, Cardona A, Nose A. Regulation of forward and backward locomotion through intersegmental feedback circuits in Drosophila larvae. Nat Commun. 2019 Jun 14;10(1):2654. doi:10.1038/s41467-019-10695-y.


Oswald MC, Brooks PS, Zwart MF, Mukherjee A, West RJ, Giachello CN, Morarach K, Baines RA, Sweeney ST, Landgraf M. Reactive oxygen species regulate activity-dependent neuronal plasticity in Drosophila. Elife. 2018 Dec 17;7. pii: e39393. doi:10.7554/eLife.39393.


Kawashima T, Zwart MF, Yang CT, Mensh BD, Ahrens MB. The Serotonergic System Tracks the Outcomes of Actions to Mediate Short-Term Motor Learning. Cell. 2016 Nov 3;167(4):933-946.e20. doi:10.1016/j.cell.2016.09.055.

Zwart MF*, Pulver SR, Truman JW, Fushiki A, Fetter RD, Cardona A, Landgraf M*. Selective Inhibition Mediates the Sequential Recruitment of Motor Pools. Neuron. 2016 Aug 3;91(3):615-28. doi:10.1016/j.neuron.2016.06.031 (*co-corresponding).

Schneider-Mizell CM, Gerhard S, Longair M, Kazimiers T, Li F, Zwart MF, Champion A, Midgley FM, Fetter RD, Saalfeld S, Cardona A. Quantitative neuroanatomy for connectomics in Drosophila. Elife. 2016 Mar 18;5. pii: e12059. doi:10.7554/eLife.12059.

Fushiki A, Zwart MF, Kohsaka H, Fetter RD, Cardona A, Nose A. A circuit mechanism for the propagation of waves of muscle contraction in Drosophila. Elife. 2016 Feb 15;5. pii: e13253. doi:10.7554/eLife.13253.


Heckscher ES, Zarin AA, Faumont S, Clark MQ, Manning L, Fushiki A, Schneider-Mizell CM, Fetter RD, Truman JW, Zwart MF, Landgraf M, Cardona A, Lockery SR, Doe CQ. Even-Skipped(+) Interneurons Are Core Components of a Sensorimotor Circuit that Maintains Left-Right Symmetric Muscle Contraction Amplitude. Neuron. 2015 Oct 21;88(2):314-29. doi:10.1016/j.neuron.2015.09.009.


Zwart MF, Randlett O, Evers JF, Landgraf M. Dendritic growth gated by a steroid hormone receptor underlies increases in activity in the developing Drosophila locomotor system. Proc Natl Acad Sci U S A. 2013 Oct1;110(40):E3878-87. doi:10.1073/pnas.1311711110.


Hörnberg H, Wollerton-van Horck F, Maurus D, Zwart M, Svoboda H, Harris WA, Holt CE. RNA-binding protein Hermes/RBPMS inversely affects synapse density and axon arbor formation in retinal ganglion cells in vivo. J Neurosci. 2013 Jun 19;33(25):10384-95. doi:10.1523/JNEUROSCI.5858-12.2013.

Pulver SR, Cognigni P, Denholm B, Fabre C, Gu WX, Linneweber G, Prieto-Godino L, Urbancic V, Zwart M, Miguel-Aliaga I. Why flies? Inexpensive public engagement exercises to explain the value of basic biomedical research on Drosophila melanogaster. Adv Physiol Educ. 2011 Dec;35(4):384-92. doi:10.1152/advan.00045.2011.

Barraud P, Seferiadis AA, Tyson LD, Zwart MF, Szabo-Rogers HL, Ruhrberg C, Liu KJ, Baker CV. Neural crest origin of olfactory ensheathing glia. Proc Natl Acad Sci U S A. 2010 Dec 7;107(49):21040-5. doi:10.1073/pnas.1012248107.

Doerflinger H, Benton R, Torres IL, Zwart MF, St Johnston D. Drosophila anterior-posterior polarity requires actin-dependent PAR-1 recruitment to the oocyte posterior. Curr Biol. 2006 Jun 6;16(11):1090-5. doi:10.1016/j.cub.2006.04.001.

Join the lab

We’re always looking for keen postdocs, PhD students, and technicians to join the team! Just drop Maarten a line. Anyone from a wide range of backgrounds including neuroscience, physics, engineering, and computer science who is interested in working in the lab is encouraged to get in touch. Funding for PhD students is available through:

the BBSRC Eastbio DTP

the School of Psychology and Neuroscience

the University of St Andrews


Maarten Zwart
Reader in Neuroscience
tel +44 01334 462086
email mfz[at]st-andrews.ac.uk

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