The Navigational Capabilities of Migratory Hoverflies - Biosciences - NERC GW4+ DTP PhD Studentship Ref: 2790

About the award

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 six Research Organisation partners:  British Antarctic Survey, British Geological Survey, Centre for Ecology and Hydrology, the Met Office, 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

The studentships will provide funding for a stipend which is currently £14,553 per annum for 2017-2018, research costs and UK/EU tuition fees at Research Council UK rates for 42 months (3.5 years) for full-time students, pro rata for part-time students.


Lead supervisor: Dr Karl Wotton, Biosciences, University of Exeter
Co-Supervisor: Professor Nicholas Roberts, School of Biological Sciences, University of Bristol

Location: Penryn Campus, Cornwall

Project description:

The sky contains multiple visual cues that can be used as compass information by migratory insects. Migratory butterflies have been shown to use the position of the sun as a navigational cue and to offset its movement using an internal circadian clock to maintain a constant heading; this is the ‘time compensated sun compass’. But how do migrants orientate when the sun is not visible? One visual cue is the pattern of polarized light in the sky created by atmospheric scattering of sunlight. Photoreceptors that detect the polarization of light are commonly found in the dorsal rim area (DRA) of the compound eye in a wide range of insect orders, suggesting polarized light detection is a general adaptation in insect eyes.  Yet the existence of a ‘time compensated polarized light compass’ has yet to be convincingly demonstrated in any insect.


Image 1: The long hoverfly, Sphaerophoria scripta. The dorsal rim area (DRA) of the compound eye is the expected site for polarized light vision. Image 2: A portable flight simulator recording flight headings of migratory hoverflies at the Col de Coux, a migration hotspot in the Swiss Alps.

Project Aims and Methods

The overall aim of this project is to provide the first unambiguous evidence for a time-compensated polarized light compass for migratory navigation. It will exploit an emerging model in the field of migration biology, the long hoverfly, Sphaerophoria scripta. First you will investigate the photoreceptors of the DRA in the eyes of the long hoverfly. You will use both electron microscopy (Labhart & Meyer 1999) and molecular markers (Wernet et al. 2014) to the detail structure of the DRA. Navigational characterisation will take place predominantly during the migration season (Aug-Oct) at migration hotspots in the Alps and Pyrenees. Building on previous work in the lab you will investigate orientation responses to the polarization of light. Portable flight simulators (modified from Weir & Dickinson 2012), coupled with controllable LCD generated skylight polarization patterns (Wang at al. 2016) will be used to record flight headings. You will manipulate polarized light, or the ability of the long hoverfly to sense it, to investigate if they use this cue for directional information during migration. Finally, you will study the time compensated response to polarized light by time-shifting the internal clock of the long hoverfly and recording flight headings.


This project would suit a candidate interested in taking a multidisciplinary approach to understanding migration with particular interests in sensory and behavioural ecology, anatomy, and genetics. An interest in physics and programming would also be beneficial but not strictly necessary. The candidate should also be comfortable in the field and with prolonged periods in isolated mountainous locations.


In the lab of KW, the student will be provided with training in insect handling, flight simulator construction and coding, behavioural experimental design and molecular biology, including in situ hybridization. NR will provide training in electron microscopy, microspectrophotometry, polarization stimulus creation using LCDs and coding. Both KW and NR will provide training for the analysis and interpretation of results. We will encourage the student to attend courses in R, Linux and Python. We will also encourage the student to engage in NERC's media training, as the project is likely to generate results of direct interest to the public. Training will also include the 2-week postgraduate International Sensory Ecology Course, regarded as the world’s best sensory ecology training course and attracting internationally leading authorities in sensory ecology.


1. Labhart & Meyer (1999). Detectors for polarized skylight in insects: a survey of ommatidial specializations in the dorsal rim area of the compound eye. Microscopy Research and Technique, 47(6), 368–79.
2. Reppert at al. (2004). Polarized light helps monarch butterflies navigate. Current Biology : CB, 14(2), 155–8.
3. Stalleicken, at al. (2005). Do monarch butterflies use polarized skylight for migratory orientation? The Journal of Experimental Biology, 208(Pt 12), 2399–408.
4. Wang, X., Gao, J., Fan, Z., Roberts, N.W. (2016) An analytical model for the celestial distribution of polarized light, accounting for polarization singularities, wavelength and atmospheric turbidity. Journal of Optics 18(6):065601.
5. Weir & Dickinson (2012). Flying drosophila orient to sky polarization. Current Biology, 22(1), 21–27.
6. Wernet & Desplan (2014). Homothorax and Extradenticle alter the transcription factor network in Drosophila ommatidia at the dorsal rim of the retina. Development, 141(4), 918–28.

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. 

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

Applicants who are classed as International for tuition fee purposes are not eligible for funding.


Application deadline:7th January 2018
Value:£14,553 per annum for 2017-18
Duration of award:per year
Contact: PGR Recruitment

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. 

You will be asked to name 2 referees as part of the application process however we will not contact these people until the shortlisting stage. Your referees should not be from the prospective supervisory team.

The closing date for applications is midnight on 7 January 2018.  Interviews will be held at the University of Exeter between 5 - 16 February 2018.

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

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.