Bee on a flower

How do changing environmental conditions impact insect energy budgets?


Funded PhD


3 years (full-time)

Application deadline:

1 March 2024

About this project

We are a multi-disciplinary team (including behavioural biologists and physiologists) working on UKRI (BBSRC) funded project that is aiming to understand and predict the effects of the thermo-nutritional developmental environment upon the life history of two key UK wild bee species, and the consequent implications for pollination services.

We are now inviting applications for a fully-funded, full time PhD studentship aimed at developing state-of-the-art predictions of how environmental variables affect the energy budgets and life histories of two key pollinators for UK crops – bumblebees and mason bees.



Bees are vital for ecosystem stability and global food security – providing pollination services worth hundreds of billions of pounds annually. The UK has ~245 species of wild bees, collectively performing more pollination than managed honeybees and bumblebees. All animals need a balance of key nutrients, such as protein, carbohydrate and fat, in their diets for growth, maintenance and reproduction. However, different environmental conditions, such as temperature, may necessitate different balances. Temperature affects animals’ metabolic rate, physiology, digestion, and nutrient assimilation. For bees, this is an unassessed and possibly serious threat both to their health and their pollination services upon which we depend for our food security. Bees' health depends on access to a mix of nutrition appropriate for the conditions - but the climate is heating up, so this ideal mix may change. Crucially, if this affects bees’ flower choices, then the pollination services bees provide today may not be the same at higher temperatures.

To address these issues and achieve the central aim of the project, we will study two wild pollinator species representing two distinct contrasting lifestyles of UK bees - social-nesting buff-tailed bumblebees (Bombus terrestris), and solitary-nesting red mason bees (Osmia bicornis). Both are commercially important pollinators, but they have important differences in life history that may result in different responses to nutrition and temperature.


The PhD Studentship

The successful candidate will either have a background in a quantitative or computational scientific discipline (e.g. Informatics, Mathematics, Physics, Engineering) with an interest in applying these skills to biology or be a biologist with an interest in quantitative analysis. They will use these skills to apply cutting-edge modelling techniques to create and parameterise computational models using a combination of existing experimental data gathered by the project team and previous literature. The models will be used to: (i) test hypotheses relating bee development, health and reproduction to achieve changes in practice (such as the sowing of wildflower strips of different nutritional composition in the context of predicted temperature shifts); and (ii) augment existing published models of bee reproduction and effectiveness as pollinators across conditions, landscapes and climates, making them nutritionally and thermally explicit.

The student will use, among others, Dynamic Energy Budget (DEB) models that permit modelling of multiple physiological parameters across bees’ life histories. Such models use reaction kinetics and physico-chemical processes to dynamically estimate the uptake, allocation and usage of nutrients into and out of different functional pools (structure, reserve, maturity, reproduction) across organisms' lifetimes [1]. The models will be implemented in a programming language such as Python, R or MATLAB.


The Project Team

The broader project team will consist of three main investigators who will act as supervisors for the PhD student. Dr James Gilbert is the principal investigator, based at the University of Hull. He is a behavioural ecologist who has pioneered rearing protocols for the economically and ecologically important solitary bee, Osmia bicornis [2,3], providing an unprecedented window onto their nutritional ecology. Prof. Jeremy Niven, a co-supervisor based at the University of Sussex, is an insect physiologist with extensive experience in computational modelling of physiological systems [4-6]. Dr Beth Nicholls, a co-supervisor also based at the University of Sussex is a behavioural biologist and ecologist whose work has primarily focussed on bumblebees [7]. Nicholls and Niven used in vitro feeding of honeybee larvae to show that dietary protein:carbohydrate ratio affected growth, survival, and the scaling of metabolic rate [8].


Research environment

The student will be based in the Department of Biology in the School of Natural Sciences at the University of Hull. The University of Hull is in the Times Higher Education's top global 100 for research impact and is one of the highest climbers in the REF 2022, ranking 55th. The research will be based at the Department of Natural Sciences, which has multiple research groups focussed on monitoring and management of environmental change at the molecular, metabolic, individual, landscape, social and commercial levels. The two co-supervisors will be based in the School of Life Sciences at the University of Sussex, which is ranked in the top 20 biological sciences departments in the UK (REF 2022) and includes >10 highly interactive research groups focussed on insect behaviour, conservation, evolution and ecology.

The successful candidate will capitalise on this opportunity to synthesize the research interests of these research groups and create collaborative links between institutions. The candidate will be integrated into both institutions and will benefit from the infrastructure and connections at both universities.

For details please contact Dr James Gilbert.



1. Sousa T, Domingos T, Poggiale J-C, Kooijman SALM. Dynamic energy budget theory restores coherence in biology. Philos Trans R Soc Lond B Biol Sci. 2010;365: 3413–3428.

2. Tainsh F, Woodmansey SR, Austin AJ, Bagnall TE, Gilbert JDJ. Sporopollenin as a dilution agent in artificial diets for solitary bees. Apidologie . 2020. doi:10.1007/s13592-020-00801-1

3. Austin AJ, Gilbert JDJ. Solitary bee larvae prioritize carbohydrate over protein in parentally provided pollen. Funct Ecol. 2021. doi:10.1111/1365-2435.13746

4. Niven JE, Anderson JC, Laughlin SB. Fly photoreceptors demonstrate energy-information trade-offs in neural coding. PLoS Biol. 2007 Apr;5(4):e116. doi: 10.1371/journal.pbio.0050116.

5. Sengupta B, Stemmler M, Laughlin SB, Niven JE. Action potential energy efficiency varies among neuron types in vertebrates and invertebrates. PLoS Comput Biol. 2010 Jul 1;6(7):e1000840. doi: 10.1371/journal.pcbi.1000840.

6. Sengupta B, Laughlin SB, Niven JE. Consequences of converting graded to action potentials upon neural information coding and energy efficiency. PLoS Comput Biol. 2014 Jan;10(1):e1003439. doi: 10.1371/journal.pcbi.1003439. Epub 2014 Jan 23.

7. Nicholls E, de Ibarra NH. Bees associate colour cues with differences in pollen rewards. J Exp Biol 217 (15): 2783–2788. doi:10.1242/jeb.106120

8. Nicholls E, Rossi M, Niven JE. Larval nutrition impacts survival to adulthood, body size and the allometric scaling of metabolic rate in adult honeybees. J Exp Biol. 2021;224. doi:10.1242/jeb.242393

How to apply

Open to UK applicants only.

You will need to supply a personal statement when applying for this scholarship position. Find out more about writing a personal statement.

Please also ensure you include the following information:

  1. What motivates you to pursue PhD study

  2. A short (half page) research proposal in at least one of the areas of interest outlined above

  3. How your skill set matches the requirements for your choice of project and/or any additional training you will need

  4. The wider significance of research in this area and potential future research directions for the project

Apply for the scholarship.

Closing date for applications

1 March 2024

Scholarship commences

27 May 2024

Postgraduate Rebecca Humphries & Georgia Sykes walking by HUBS Wharfe Building

University of Hull's Doctoral College provides support to postgraduate research students. Offering skills development opportunities and dedicated facilities, the college is here to help you achieve your potential.


  • The successful applicant will receive a Home fee waiver and a maintenance grant for three years (full-time), which covers the research period of the PhD. The fee waiver for 23/24 is £4712 (Home fee) and the maintenance grant is £18622 per annum. This rises each year in line with the UKRI’s recommended stipend allowance.

  • If you need to move into a fourth year to complete your thesis, you will not receive a tuition fee waiver or maintenance grant during this period and you will be required to pay a continuation fee.

Entry requirements

Applicants should have a minimum 2:1 degree in biological, physical, mathematical or computational sciences, ideally with a demonstrable interest in mathematical and/or modelling approaches to biological questions. A taught MSc or Masters by Research in computational modelling, applied mathematics or related discipline would be an advantage.

Submission of thesis

Submission of your final thesis is expected within three years and three months from the start of your PhD scholarship.

Research training

As a PhD candidate here at the University of Hull you will undertake the Postgraduate Training Scheme alongside your main degree, to help you develop the research skills and knowledge you’ll need in your future career. You will gain an additional Certificate or Diploma in Research Training in addition to your PhD.