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PhD projects available within the Mechanisms of Behaviour group at the University of St Andrews.

Below are a few PhD projects that we are looking to fill for the 2017 entry.  The type of funding and information about competitions is listed under each project.

Anyone interested should first contact me about application procedures: karen.spencer@st-andrews.ac.uk. Potential applicants can also email me if they are interested in these general areas too. I am happy to chat about other possible projects.



1. Gulls and garbage: the ecology, behaviour and physiology of a human-wildlife conflict

Supervisors: Dr Karen Spencer, Dr Jeroen Minderman (St Andrews) and Dr Liz Humphreys (BTO Scotland).

Deadline for applications:  13th January 2017.

How to apply: In the first instance please contact Dr Karen Spencer (kas21@st-andrews.ac.uk), to discuss your application. A formal application form will then be filled out. Initial interviews are likely to be held in January 2017, with a final interview held in February 2017.

Funding: The PhD is funded by a NERC CASE grant, hence it is open to all qualified UK residents. A relevant degree (BSc) is required and further postgraduate study is also favourable. EU residents are also eligible to apply for the full funding level (stipend of £14,296 p.a. plus fee payment) if they have been resident in the UK for three years or more at studentship commencement. EU nationals may also be eligible for part funding of this project. For more details please contact the named supervisor.

Background: In response to an increasingly urbanised world, many animals have rapidly adapted to living in built-up environments. This includes two species of gull in the UK, herring- and lesser black-backed gulls, which increasingly nest and forage in close proximity to humans. This has led to growing numbers of reports of disturbance and nuisance, including direct attacks, hazards to aircraft and pathogen transmission, and subsequent control efforts. This is problematic, as both species are currently undergoing population declines. The proposed projects takes a multidisciplinary approach to understand the behavioural and physiological mechanisms underlying the adaptation of herring and lesser black-backed gulls to anthropogenic environments, focussing on the role of early life experience. Early life experiences can have significant life-long impacts on many traits. Although this may be particularly relevant for gulls provisioning their chicks with diets increasingly sourced from urban areas, this has not been studied to date. For example, both diets of lower nutritional quality as well as variation in stress hormone levels associated with urban foraging may lead to changes in physiological and behavioural development of chicks. Studies in other species have suggested that such patterns may lead to “developmental programming” of offspring, so that they cope better in particular environments in later life. For gulls making increasing use of urban areas this may have strong effects on population dynamics, but this has not been studied.

The interdisciplinary project aims to: 1. Determine the extent to which birds breeding in coastal and urban environments utilise anthropogenic sites for feeding and how resultant dietary differences affect both egg hormone levels and physiological stress responses in chicks. 2. Determine the nature and consequences of behavioural and physiological differences between individuals hatching in different environments, specifically the ability of nestlings to cope with unfamiliar environments, and effects of this on chick development and foraging behaviour of juveniles. 3. Develop a model to predict how variation in early life experience (e.g. diet and subsequent physiological and behavioural differences) in different environments may affect site usage in later life and resulting population dynamics.

Training: The successful candidate will learn a wide range of techniques, including field survey and sampling techniques, the use of tracking technologies, modelling techniques, physiological assays and design and implementation of behavioural experiments. This project will therefore provide the student with an outstanding training experience.

2. Early life experiences and social behaviour in a gregarious species: a role for mesotcin?

Supervisor: Karen Spencer (Psychology & Neuroscience, St Andrews)

Deadline for applications: 10th January 2017

Funding: School of Psychology & Neuroscience scholarship. Student will be interviewed centrally for a funded place.

More information at: http://www.st-andrews.ac.uk/psychology/prospective/pg/apply/

Eligibility: UK and EU residents.

Mesotocin (MT) is a nonapeptide hormone and is considered the avian homologue of the mammalian hormone oxytocin. Such hormones are known to modulate many social behaviours including pair bonding, maternal bonding, social communication and aggression.  Work on highly gregarious avian species, such as the zebra finch (Taeniopyggia guttata) has shown that MT facilitates affiliative behaviours and can promote flocking. More recently a direct link has been shown between social pairing in zebra finches and the expression of both MT and vasotocin (VT) mRNA (another nonapeptide) in specific nuclei within the hypothalamus. However there are still many unresolved questions, and in particular the role of mesotocin in mediating social networks or social information use has not been tested. Recent work in the Mechanisms of Behaviour group here at St Andrews has shown that early life exposure to stress can alter an individual’s position within a social network, producing phenotypes that create more, but looser connections within a network of conspecifics. We currently do not know the mechanisms underlying this phenomenon. The aim of this PhD will be to investigate the effects of early life experiences on a range of social behaviours in later life and the potential role of MT (and other nonapeptides) as a mechanism. The student will gain training in behavioural assay, neurological techniques, such as immunohistochemistry, physiological assay of hormones and molecular techniques (qPCR).

  • Boogert, N.J., Farine, D. & Spencer, K.A. (2014) Developmental stress predicts social network position. Biology Letters,10, 20140561.
  • Lowrey, E.M., Tomaszycki, M.L. (2014) The formation and maintenance of social relationships increases nonapeptide mRNA in zebra finches of both sexes. Behavioral Neuroscience, 128(1), 61-70.
  • Goodson, J.L. & Thompson, R.R. (2010) Nonapeptide mechanismsm of social cognition, behaviour and species specific social systems. Current Opinion in Neurobiology, 20(6) 784-794.

3. Developmental stress and biological rhythms

Supervisors: Karen Spencer, Tyler Stevenson

Funding: EastBio BBSRC DTP – http://www.eastscotbiodtp.ac.uk/developmental-stress-and-biological-rhythms

Deadline December 1st 2016

Background: Seasonal changes in the environment are evident across the globe. Animals of all species use the presence of salient environmental cues to generate physiological responses, which in turn regulate seasonal rhythms in hormonal, immune and behavioural traits (1). Within a population there is significant variation in the ability or speed of individuals to respond to cues such as photoperiod, food availability or temperature and this translates into variation in the ability to match the timing of important life cycle events to appropriate environmental conditions. There is a wealth of evidence to show that such mismatches can significantly reduce health, well-being and reproductive output, however the mechanism that underpins this variation in the ability to respond to cues, and the impact across the lifespan is little understood.

Exposure to environmental stressors during development can have significant implications for many phenotypic traits in later life. So-called developmental programming has been studied in several contexts, including the role of stress in mediating long-term effects on health and wellbeing (2).  Activation of the neuroendocrine axis that underlies the response to stress (the HPA, hypothalamic-pituitary-adrenal axis) during sensitive developmental periods can alter an individual’s ability to cope with stress in later life. These events modify the HPA axis and may lead to disruptions in the ability to regulate stress hormones (3). However the impact of developmental stress on the neuroendocrine axis underlying the timing of seasonal biology is not well described. Using a comparative approach, this studentship will examine the role of developmental stress on long-term timing of neural, physiological and behavioural processes. The outcomes of the project will lead to a significant gain in our understanding of the evolutionarily conserved neural mechanisms that underlie the ability to adapt to environmental change.

Study specifics: This study aims to integrate information across these different levels: brain, physiology and behaviour to determine the influence of early life on an individual’s ability to respond to environmental cues, such as photoperiod, food availability and temperature, and maintain synchrony with internal seasonal processes. The studentship will test the hypothesis that environmental cues impart lasting molecular (i.e. DNA methylation) and hormonal effects on key regions within the hypothalamo-pituitary-adrenal/gonad axis. The project will determine the effects of early life on the neural and hormonal systems regulating the timing of behaviours. Additionally, the student may incorporate novel in vivo imaging techniques (i.e. light sheet microscopy) to identify the role of downstream hypothalamic neurosteroid and gonadal steroids on neuro-glial plasticity. Finally the project will investigate the potential epigenetic mechanisms that underlie the physiological and behavioural responses observed. The project will utilise multiple model species that have been studied extensively in terms of biological rhythms, including the Siberian hamster, Japanese quail and the potential for field based research on European starling.

Training: The successful student will have the opportunity to learn a wide range of techniques including genomic and molecular analyses, and whole animal physiology and behaviour. Specific training includes: experimental design, behavioural observation and analysis, complex statistical analyses, molecular (e.g. quantitative real time PCR, sodium bisulfite DNA/RNA methylation analysis; DNA sequencing) cellular (e.g. brain immunocytochemistry), hormone analyses (i.e. radioimmunoassay) and several novel, real-time imaging techniques. The work will mainly take place at the University of St Andrews and the student will be based within the active Mechanisms of Behaviour research group led by Dr Karen Spencer.  In addition the student will also have the opportunity to work in the Stevenson laboratory in the Institute of Biological and Environmental Sciences at the University of Aberdeen.


1.         Stevenson TJ, Visser ME, Arnold W, et al. Disrupted seasonal biology impacts health, food security and ecosystems. Proceedings of the Royal Society B: Biological Sciences. 2015;282:20151453.

2.         Bateson P, Barker D, Clutton-Brock T, et al. Developmental plasticity and human health. Nature. 2004;430(6998):419-21.

3.         Zimmer C, Boogert NJ, Spencer KA. Developmental programming: Cumulative effects of increased pre-hatching corticosterone levels and post-hatching unpredictable food availability on physiology and behaviour in adulthood. Hormones and Behavior. 2013;64(3):494-500.

Keywords: Developmental programming, hormones, stress, biological clock, seasonal biology, behavioural endocrinology, animal behaviour.

4. Dog-human relationships: attachment and communication in pet and working dogs

Supervisors: Catherine Hobaiter, Karen Spencer

Funding: EASTBio BBSRC DTP – http://www.eastscotbiodtp.ac.uk/dog-human-relationships-attachment-and-communication-pet-and-working-dogs

Introduction: Genetic evidence suggests that the domestication of dogs occurred over 30,000 years ago; today dogs play a significant role in our lives, with recent evidence showing that their presence can benefit both our mental and physical wellbeing (Stafford, 2006). To date, little is understood about how the social bonds between dogs and humans develop, or what mechanisms promote the wellbeing they provide. Previous studies have suggested these bonds resemble those of infant-attachment, with evidence for dependency and separation anxiety (Topal et al., 1998) and the involvement of oxytocin, but both the behavioural and biological mechanisms remain unclear. We will investigate dog-human bonding across a range of different relationships: pet, working (assistance-, police-, and gun-dogs), and pack dogs.

We will further explore how variation in the strength and type of dog-human social bonds impacts communication. Several aspects of communication, such as pointing or gaze-following, are argued to be uniquely human, with evidence mixed in even our closest relatives: the other great apes. In contrast, some of the strongest non-human evidence for these capacities comes from dogs, with a range of evidence suggesting that domesticated dogs outperform closely related non-domesticated canines, such as wolves. However, to date these studies were limited to referential signals, which represents a fraction of the rich range of communication humans’ employ. We plan to extend these studies to investigate whether there has been further modification of dog communication style towards that of humans, as compared to wolves.

Approach: The student will employ a range of experimental approaches from the study of human-infant interaction, now adapted for the study of human-dog interaction. For example: we recently successfully piloted use of the Still Face Paradigm (SFP) in humans and pet dogs. SFP was developed to test the hypothesis that infants are active contributors to social interaction (Tronick et al., 1987). Typically used to investigate mother-child attachment and communication it provides a reliable measure of attachment in infancy and adulthood. SFP consists of a short (<10min) face-to-face interaction with three phases: (1) baseline normal social interaction, (2) Still Face, in which the adult becomes unresponsive, and (3) reunion, in which normal interaction resumes. Together with behavioural observations from human-dog interactions, the student will use infrared thermography, and non-invasive (saliva) measures of oxytocin and cortisol to measure variation in both dog and human bonding and stress response. Infrared thermography and salvia cortisol have been established as a non-invasive measure of psychogenic stress-responses across species, including dogs. While current evidence across species, including dogs and humans, suggests that oxytocin mediates social bond formation, less is understood about its function in bond maintenance. The student will use these data to investigate (i) how type and duration of human-dog social interactions, and the caretaker’s human relationship attachment style affect the strength and type of human-dog attachment, and (ii) how bond style and strength, as measure by oxytocin, impacts working performance in both dogs and humans.

Training: This project will expose the student to wide-ranging fields of research including: animal behaviour, animal physiology, and neuroendocrinology. The student will be trained in behavioural observations, video coding, infrared thermography, hormone measurement, data collection, interpretation and writing papers for publication. The student will be expected to engage with other scientists in the institute and to attend and present their findings at local laboratory meetings as well as UK and international conferences.

As well as increasing our understanding of the formation of dog-human social bonds and communication, we hope to use these data to develop a simple, effective test of both the strength and type of human-dog attachment. In doing so we aim to provide tools to assess the bond between dog-human working pairs, and offer training tools for those working to match, train, or rehome pet and working dogs.

References: Stafford, K. (2006) The welfare of Dogs. Springer; Topal, J., et al., (1998) Attachment behaviour in dogs (Canis Familiaris): A new application of Ainsworth’s (1969) Strange Situation Test. J. Comp. Psychol. 112; Tronick, E., et al., (1978) The infant’s response to entrapment between contradictory messages in face-to-face interaction. J. Am. Acad Child Adolesc Psychiatry. 17

Keywords: attachment, communication, canine, oxytocin, cortisol, thermography, social bond

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