Neuroscience, Physiology and Pharmacology

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4-year PhD in Neuroscience

funded by the Wellcome Trust


Why study Neuroscience at University College London?

Outstanding research opportunities

University College London (UCL) offers unrivalled opportunities for PhD research in all aspects of neuroscience. Specimen PhD projects are given below: the subjects studied range from the molecular biology of neuronal proteins, through cellular neuroscience, to the behaviour of sensory and motor systems and brain imaging. Neuroscience research is carried out in all of the College's biomedical departments, by researchers who are among the leaders of their fields, using the most modern techniques to address important problems of basic and clinical neuroscience.  Research labs are well-funded, so that PhD students have the best chance of getting off to a productive start in their research.  UCL produces the highest quality neuroscience research of any university in the country. For the field neuroscience it generates 30% of England’s contribution to the world’s most highly cited publications (documented HERE), more than twice as much as either Cambridge or Oxford, and in the sub-field of neuroimaging UCL and its hospitals produce 65% of England’s contribution to the world’s highly cited papers which is more than 4-fold larger than either of the runners up (Oxford and Kings College London). Accordingly it is an outstanding place to train the next generation of neuroscience researchers.

Cultural and social opportunities provided by London

UCL is located in Bloomsbury, close to the entertainment areas of the West End and South Bank which offer an enormous range of music, art, theatre and film, and a vast number of restaurants and bars. London is extremely socially diverse: most PhD students rapidly establish a thriving social life.

Why choose to study Neuroscience in the 4-year Programme?

Because the 4-year Programme provides a better training, equipping students with the experimental and theoretical techniques needed to do outstanding research in their future career (see tab below on Student Outcomes).

Deficiencies of 3-year PhDs

Conventional 3-year PhDs involve a student working with a supervisor who they often have little knowledge of before they start, on a project which they have little prior understanding of. The resulting training can be rather narrow (limited to learning the techniques offered by that one lab), and students sometimes select supervisors or projects which are not best suited to them.

Structure of the 4-year programme

Value of the first year

The four year programme provides a broader and deeper research training in neuroscience, and allows students to make a more informed choice of supervisor and project. This is achieved by having an initial training year in which the students attend some specialized courses, and do three brief (3 month) research projects in different labs. Out of the three broad subject areas of Molecular Neuroscience, Cellular Neuroscience and Systems Neuroscience & Imaging, students will choose laboratories from at least two areas, in order to maintain a broad expertise across neuroscience in the first year. By working in different labs, the students will have the opportunity to acquire a broader range of experimental and theoretical techniques, and to try out supervisors with whom they may wish to do research for the PhD. For the structure of the 1st year please refer to the tabbed panel 'Year 1 Structure' section below.

The 3 PhD years

After their first year, students will work in one lab doing research for the PhD (this might be one of the labs they worked in during the first year, or a different lab). During the PhD students will be encouraged to attend advanced training courses in the USA and Europe. For supervisors available for PhDs please refer to the tabbed panel 'Supervisors' section below.

Pastoral care

Throughout the 4 years, the student's progress will be monitored and assessed by a committee responsible for the training provided. Students will be integrated into the community of neuroscience researchers at UCL by participation in journal clubs and social events. Career advice will be given in the last year to prepare the student for their post doctoral career.

The 4-year PhD Committee

The committee currently comprises David Attwell, Sarah-Jayne Blakemore, Patricia Salinas and Alasdair Gibb, and students can approach any of the committee members for advice and guidance when needed.

Student experiences and outcomes

Want to read about students' experiences on the 4 year programme? - they are described in Trends in Neurosciences (July 2000) Vol 23, pages 280-283. An assessment of how well the students do scientifically on the programme is given in the tabbed panel 'Student Outcomes' section below.

If you have any questions...

Please read all of this website, especially the Detailed Instructions section and only if the question is not answered there, you can email Joanna Fryer, j.fryer@ucl.ac.uk, for an answer.

Molecular Neuroscience

Gill Bates

Molecular Basis of Huntington’s disease

Annette Dolphin
Voltage-dependent calcium channels

Vilaiwan Fernandes

Glial roles during brain development

Elizabeth Fisher
Motor neuron degeneration and genes

Matthew Gold
Molecular basis of neuronal second messaging

John Hardy

Genomic and cellular investigation of neurodegeneration

Parmjit Jat

Cellular functions of the prion protein

James Jepson
Drosophilia Neurogenetics

Nicoletta Kessaris
Forebrain neurogenesis

Josef Kittler
Cell biology of the synapse

Alison C Lloyd
Peripheral nerve regeneration and cancer

Neil Millar
Neuronal nicotinic acetylcholine receptors

Stephen Price
Neuronal development

Antonella Riccio
Transcriptional and epigenetic mechanisms in developing neurons

Bill Richardson
Neuroglial stem/progenitor cells

Jason Rihel

Patricia Salinas
Cell signalling in synaptic plasticity and synapse degeneration

Stephanie Schorge
Ion channels and disease

Lucia Sivilotti
Ion channels as single molecules

Trevor G Smart
Molecular pharmacology of GABA and glycine receptor-ion channels

Claudio Stern

CNS Development

Martin Stocker
Molecular pharmacology and physiology of potassium channels

Jernej Ule
Structure and function of neuronal protein-RNA complexes

David Whitmore
Circadian clocks in zebrafish

Stephen Wilson
Zebrafish CNS development

John Wood
Molecular genetics of sensory neurons

Cellular Neuroscience

David Attwell
Neuron-glial interactions and brain energy supply

Arantza Barrios

Genes and circuits for innate and learned behaviours in C. elegans

Marco Beato
Glycinergic inhibition in the ventral spinal cord

Issac Bianco
Zebrafish circuits and behaivour

Genes and circuits for innate and learned behaviours in C. elegans

Tiago Branco

Computation of instinctive decisions

Liam Browne

Neural circuits for nociception and pain

Mike Cheetham
Cell Biology of Neurodegeneration

Michael Duchen
Neuronal activity, mitochondrial function and cell death

Mark Farrant
Ionotropic GABA and glutamate receptor signalling

Jonathan Gale
Cell development and regeneration

Alasdair Gibb
Ion channel receptors and synaptic transmission

Linda Greensmith
The Graham Watts Laboratories for Research into Motor Neuron Disease

Michael Häusser
Dendritic Processing

Johannes Kohl

State-dependent neural processing

Martin Koltzenburg
Chronic Pain

Dimitri Kullmann

CNS synaptic transmission, epilepsy, and inherited mutations of ion channels in neurological disease

State-dependent neural processing

Andrew MacAskill

Synaptic and circuit basis of emotional behaviour

Shin-ichi Ohnuma
Cell cycle and neural development

Sandip Patel
Neuronal calcium signalling by NAADP

Rachael A Pearson
Stem Cell Therapy and Retinal Degeneration

Paola Pedarzani
Ion channels regulating neuronal excitability and firing properties

Dmitri Rusakov
Principles of synaptic signal formation in the brain

Giampietro Schiavo
Axonal transport in health and disease

Mala Shah
Voltage-gated ion channels

Angus Silver
Synaptic transmission and neural computation

Kenneth Smith
Pathophysiology of neuroinflammation

Sarah Tabrizi
Cellular mechanisms of neurodegeneration

Kirill Volynski
Mechanisms of neurotransmitter release in health and in neurological disease

Matthew Walker
Mechanisms and treatment of epilepsy

Systems/Cognitive Neuroscience

Gareth Barnes

Wearable brain scanners

Caswell Barry
Memory, space, and hippocampal networks

Tim Behrens

Storing and updating models of the world for controlling behaviour

Daniel Bendor
Neural Coding of Perception and Memory

Sven Bestmann
Decision implementation and action preparation

Jennifer Bizley
Auditory and auditory-visual perception

Sarah-Jayne Blakemore
The social brain and its development

Robert Brownstone

Neural circuits for movement

Neil Burgess
Neural networks and hippocampus

Francesca Cacucci
Hippocampal neural circuits

Matteo Carandini
Sensory representation by neuronal populations

Ray Dolan
Emotion and decision making

Elena Dreosti

A developing social brain circuit

Maria Fitzgerald
The Developmental Biology of Spinal Cord and Cortical Pain Processing

John Greenwood

Visual perception & psychophysics

Patrick Haggard
Control of human action

Kenneth Harris
Computation in cortical circuits

Soyon Hong

Microglia-synapse biology in brain function and dementia

Giandomenico Iannetti

Human sensory neuroscience

Kate Jeffery
The neural representation of space and context

Tara Keck
Cortical synaptic plasticity

Steve Kennerley
Neuronal mechanisms of learning and decision making

James Kilner
Action observation: perceptual learning and inference.

Alexander Kraskov
Cortical and subcortical mechanisms of movement generation and inhibition

Anna Kuppuswamy

Predictive sensorimotor control in central fatigue

Peter Latham
Network dynamics and neural coding
Nilli Lavie
Attention and Cognitive control

Nicholas Lesica
Population coding in sensory systems

Jennifer Linden

Brain mechanisms for perception of complex sounds

Eleanor Maguire
The neural basis of spatial and episodic memory in humans

Tamar Makin

Human brain plasticity

Benedetto De Martino

Value Computation, Uncertainty and Choice

Geraint Rees
Cognitive neuroscience of attention & awareness

Jon Roiser
Neural basis of motivation and emotion

Maneesh Sahani
Theoretical Neuroscience and Machine Learning

Aman Saleem

Neural circuits that transform visual information to spatial memories

Tali Sharot

Information Processing and Belief Formation

Vince Walsh
Visual Cognition 

Nick Ward
Plasticity and recovery in health and disease

Jason Warren
Clinical neuroscience of complex sound

How to apply and what studentships are available?

The next intake of students will be in September 2019. Students will spend the first year learning a wide range of neuroscience techniques by doing 3 month projects in different laboratories, before choosing a full research project and supervisor for the subsequent 3 years. Projects available cover the whole range of neuroscience, from molecular biology through cellular mechanisms to systems neuroscience and imaging.

Up to five PhD studentships, with a stipend starting at £22,278, will be available. These also pay fees at the EU rate, research costs to the laboratory and provide funds for travel to conferences or courses. Applicants should have, or expect to get, at least an upper 2nd class degree in any area of Biological or Physical Sciences (the course allows conversion to neuroscience from a physical sciences background). Non-UK applicants may apply, and receive the normal stipend etc., except that fees will only be paid at the EU rate (see Detailed instructions for more details)

To apply we need to receive a CV, a statement of why you want to do the PhD and at least two academic references, all e-mailed to NeuroPhD@ucl.ac.uk. Before you apply you must read these Detailed instructions on how to apply. The sum of the files you submit must be smaller than 1MB in size - files larger than that will be rejected (we do not need high resolution scans of transcripts, just type the grades into your CV).

Any questions not answered on that page should be sent to David Attwell at d.attwell@ucl.ac.uk

The deadline for receiving applications and references is 20th December 2018 to start September 2019. 

If you are qualified as a doctor or vet, then you should not apply to this programme, but instead to one of the Wellcome's programmes for clinicians: https://wellcome.ac.uk/funding/phd-programmes-clinicians

If you have done the pre-clinical part of medical/vet training and intend not to do the clinical training to become a medical or veterinary practioner, then you may apply to this programme.

Please do not send your application to NeuroPhD@ucl.ac.uk until October 1st. Applicants will hear if they are shortlisted for interview around January 14th.

If shortlisted, the interviews will take place late January, 2019.

Please note that you must activate your referees to send their references; we don't do that for you.

Information submitted as part of your application will be shared with the Wellcome Trust for data-monitoring purposes and with other UCL selection committees who may offer you a PhD place if you fail to gain a place on this programme.  Our full GDPR privacy statement is available HERE.

Structure of the First Year of the 4 Year PhD

The first year has 3 main components, compulsory courses, optional courses and (occupying most of the time) three 3 month laboratory placements spent doing research and learning techniques.

Compulsory Courses

These consist of:

An Induction course introducing you to the College
A course on Current Techniques in Neuroscience
A Topics in Neuroscience course, structured like a journal club in which you present research papers
A Statistics course
A course on Library and Database Usage
An Electronics course
A course on the Ethics of Animal Experimentation
Two Neuroscience courses chosen from the Optional list below
Attending Journal Clubs associated with the lab placements (see below)
A Science Communication course (may also be taken in the 2nd year)

Optional Courses

These consist of:

Computing Courses on E-mail, Word Processing, Internet, Spreadsheets, Powerpoint, Visual Basic, and more advanced programming
A Mechanical Workshop course
A Further Statistics course
A Radiation Safety course
Orientation for Foreigners
English for Foreigners

The following Neuroscience Courses:

Advanced Cytology 
Neural Basis of Learning + Motivation
Advanced Neuroanatomy 
Neurobiology of Neurodegenerative Disease 
Cellular + Developmental Neurobiology
Control of Movement
Neural Computation
Peripheral Nervous System
Animal Cell Biology 
Develpmantal Biology: Cell + Molecular Aspects 
Molecular + Cellular Pharmacology 
Synaptic Pharmacology
Neurobiology of Behaviour
Cell Physiology 
Visual Perception
Neurobiology of Vision
Clinical Neuroscience
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2 weeks  

Journal clubs in the rotation labs

The Laboratory Placements

Three of these are done, chosen from labs working in the broad areas of Molecular Neuroscience, Cellular Neuroscience, and Systems and Imaging Neuroscience. Students must choose 3 placements covering at least 2 of these broad areas (in order to avoid over-specialization in the first year). For example, students might do a placement in one lab which they think they might want to do their PhD research in, one in a similar lab for comparison, and one in a lab studying something quite different to gain experience in another area. Students doing these placements often publish papers on their work, or present it at scientific meetings.

For information on supervisors and projects refer to the supervisors tab


Student progress during the first year is assessed by:

(i) exams on the Topics in Neuroscience course, the Ethics of Animal Experimentation course and on the Statistics course
(ii) a write-up and 10 minute oral presentation on each lab placement
(iii) their placement supervisor’s assessment of their work
(iv) their contribution to journal clubs they attend
(v) the writing of a research plan outlining their proposed PhD project for the subsequent 3 years.


We currently receive about 400 applications per year from all over the world. From these, we select 30 applicants for interviews which assess knowledge, creativity and data interpretation. From these 30 candidates, we choose 5 for the places available on the PhD Programme. Our selection ratio of women:men closely reflects the ratio of women to men applying (over 22 years, out of 142 students selected 59.9% were women and 40.1% men). BME students, defined conventionally to include Asians and mixed race students, comprise 8.5% of accepted students.

How well do the students in the programme do scientifically?

We analysed this after 10 years of the programme being in operation.

Scientific publications

As of April 2007, the students going through the programme since 1996 have published a total of 251 journal papers since entering the programme, with 28 (11%) high profile papers in Nature family journals, Science, Cell or Neuron.

Five years after starting the programme, the 15 students in the first three years’ entry had published 80% more papers/student than the average of all 123 three year PhD students recruited to the same departments at the same time. One might hypothesise, however, that this superiority of the 4 year students could reflect the 3 year PhD students being less selected or working with weaker supervisors. To assess whether the 4 year programme confers particular benefits, we therefore carried out tougher comparisons, that were not confounded by the inclusion of weaker supervisors who we do not allow onto the programme, or by the selection bias inherent in our admission process. We did this as follows.

First we removed supervisor bias, by comparing the output of students in our programme with the output of matched 3 year PhD students who did a PhD in the same lab at approximately the same time (all except 5 of the 66 1996-2004 students could be matched). The results are plotted in Figure 1 as absolute number of papers, and in Figure 2 as the relative productivity of 4 and 3 year students (1 is added to the number of papers before taking the ratio to avoid divisions by zero). From 7 years after starting, the productivity of 4 year students increases significantly (Fig. 2) above that of 3 year PhD students with the same supervisors. Students in the first 3 entry cohorts have now received 2621 citations of their papers, compared with 1421 citations of the papers of matched 3 year students in the same labs (84% more).

Figure 1


Figure 2


Secondly, to eliminate effects of the higher selection to which our students were subjected, we compared the output of our 4 year students, with the output of students who we made an offer to but who chose to take up a place on a 3 or 4 year PhD elsewhere. This ensures that the comparator students were viewed by our committee as being at least as good as the accepted students. The results (Figure 3) show a significantly higher output, from 7 years after starting the course, by our 4 year students than by students who rejected our offer.

Figure 3


We conclude that going through the first year of the 4 year PhD adds significant long-lasting value to the students’ scientific training.

Career Path after the PhD

Most of the students follow a career in research. As of 2007, of the 37 students who have obtained their PhD, 29 (78%) went on to post-doc positions, 6 went to science-related jobs (drug companies, scientific administration, patent agency or management consultancy) and 2 left science, so overall 95% (35/37) took up positions using their science. The earlier cohorts are now starting to obtain permanent academic positions or Career Fellowships: one is a lecturer at the Royal Vet College, 3 are Royal Society Dorothy Hodgkin Fellows at UCL, one has a 5 year Faculty position at EBRI (Rome), one is an MRC CDF in Leicester, one is a Wellcome RCDF at UCL, two have permanent positions in Edinburgh.

The images at the top of this page

The pictures at the top of this page show different levels of function of the nervous system.

The left hand picture shows cellular interactions between neurons: the axon of an inhibitory interneuron (green) makes synapses onto a cerebellar Purkinje cell (red) in the brain's motor system. Image by Beverley Clark and Michael Häusser.

The middle picture shows information superhighways in the brain: the gold colour shows antibody to myelin, which speeds the conduction of information along neuronal axons in the brain's white matter. Image by Ragnhildur Káradóttir and David Attwell.

The right hand picture shows function at the whole brain level: the red and yellow colour shows areas where neurons are detected to be active using fMRI (functional magnetic resonance imaging) during a particular task, superimposed on a structural image of the brain. Image courtesy of Sarah-Jayne Blakemore and the Wellcome Trust Centre for Neuroimaging, UCL.