Telephone: (01792) 295726
Room: Academic Office - 103
First Floor
Wallace Building
Singleton Campus

I am a marine biologist with interests spanning all aspects of plankton (virus, bacteria, algae, mixotrophs, zooplankton), from experimental physiology, ecology through to (and now especially) modelling.

I have worked on projects ranging from ocean acidification and ocean-scale production, harmful algal blooms, to commercial exploitation of algal growth and aquaculture.

My main interests now are in marine plankton mixotrophy (notably via the MixITiN project; ) and in optimising commercial microalgal production.

A core component of my work is modelling, for which a series of interconnecting acclimative, multi-nutrient, variable stoichiometry models have been developed. The primary applications of these are in studies of trophic dynamics for biogeochemical and ecological analyses. A secondary application is in the guise of systems-biology applications to commercial production of microalgal biomass, cost-benefit analyses and optimization.

A free e-book to guide biologists in using systems dynamic modelling in dynamic ecology is available: please email me, or check

I work with colleagues both within the UK (notably PML, NOCS) and internationally (notably EU, USA and Australia).

Find me at ResearchGate -


MixITiN [MSCA H2020]; 2017-2021 €2.88 (PI A Mitra, Swansea)

EnhanceMicroALgae [INTERREG]; 2017-2020 (Lead ANFACO, Spain)

Pilots4U [H2020]; 2017-2019 (Lead BBEU, Belgium)



SEACAMS [ERDF]; 2010-15; £2.2M (Swansea University component)

EUROBASIN [FP7 Integrated Science]; 2011-14; £100k (Swansea University component)

Improved understanding of the potential population, community and ecosystem impacts of OA for commercially important species [NERC, Defra, DECC]; 2011-14; £700k

Placing marine mixotrophs in context: modelling mixotrophy in a changing world [Leverhulme International Network]; 2011-14; £100k

Energetic Algae (EnAlgae) [ERDF]; 2011-15; £5M

H+ fluxes in phytoplankton - a mechanistic and modelling study of their physiological roles and impact upon community responses to ocean acidification [NERC]; 2012-15; £45k to KJF

MacroBioCrude [EPSRC]; 2013-18; £1.5M

Areas of Expertise

  • plankton modelling
  • plankton physiology
  • commercial algal production
  • ocean acidification


  1. & Future HAB science: Directions and challenges in a changing climate. Harmful Algae, 101632
  2. & Dynamic changes in carbonate chemistry in the microenvironment around single marine phytoplankton cells. Nature Communications 9(1)
  3. & Simulating Effects of Variable Stoichiometry and Temperature on Mixotrophy in the Harmful Dinoflagellate Karlodinium veneficum. Frontiers in Marine Science 5
  4. Dynamic Ecology - an introduction to the art of simulating trophic dynamics.. Swansea University: Swansea University.
  5. & Effects of growth rate, cell size, motion, and elemental stoichiometry on nutrient transport kinetics. PLOS Computational Biology 14(4), e1006118
  6. & Toward a mechanistic understanding of trophic structure: inferences from simulating stable isotope ratios. Marine Biology 165(9)
  7. & Introducing mixotrophy into a biogeochemical model describing an eutrophied coastal ecosystem: The Southern North Sea. Progress in Oceanography 157, 1-11.
  8. & Nutrients from anaerobic digestion effluents for cultivation of the microalga Nannochloropsis sp. — Impact on growth, biochemical composition and the potential for cost and environmental impact savings. Algal Research 26, 275-286.
  9. & Oceanic protists with different forms of acquired phototrophy display contrasting biogeographies and abundance. Proceedings of the Royal Society B: Biological Sciences 284(1860), 20170664
  10. & Minimising losses to predation during microalgae cultivation. Journal of Applied Phycology 29(4), 1829-1840.
  11. & Physiology limits commercially viable photoautotrophic production of microalgal biofuels. Journal of Applied Phycology
  12. & Why Plankton Modelers Should Reconsider Using Rectangular Hyperbolic (Michaelis-Menten, Monod) Descriptions of Predator-Prey Interactions. Frontiers in Marine Science 3
  13. & The role of coccolithophore calcification in bioengineering their environment. Proceedings of the Royal Society B: Biological Sciences 283(1833), 20161099
  14. & Ocean Acidification Affects the Phyto-Zoo Plankton Trophic Transfer Efficiency. PLOS ONE 11(4), e0151739
  15. & What is the limit for photoautotrophic plankton growth rates?. Journal of Plankton Research
  16. & Estimating the ecological, economic and social impacts of ocean acidification and warming on UK fisheries. Fish and Fisheries
  17. & Coupling a simple irradiance description to a mechanistic growth model to predict algal production in industrial-scale solar-powered photobioreactors. Journal of Applied Phycology
  18. et. al. Defining Planktonic Protist Functional Groups on Mechanisms for Energy and Nutrient Acquisition: Incorporation of Diverse Mixotrophic Strategies. Protist 167(2), 106-120.
  19. & In silico optimization for production of biomass and biofuel feedstocks from microalgae. Journal of Applied Phycology 27(1), 33-48.
  20. & Modeling DOM Biogeochemistry. In Biogeochemistry of Marine Dissolved Organic Matter. (pp. 635-667).
  21. & Ocean acidification with (de)eutrophication will alter future phytoplankton growth and succession. Proceedings of the Royal Society B: Biological Sciences 282(1804), 20142604-20142604.
  22. & Acclimation, adaptation, traits and trade-offs in plankton functional type models: reconciling terminology for biology and modelling. Journal of Plankton Research 37(4), 683-691.
  23. & Modelling alkaline phosphatase activity in microalgae under orthophosphate limitation: the case ofPhaeocystis globosa. Journal of Plankton Research 37(5), 869-885.
  24. & Have we been underestimating the effects of ocean acidification in zooplankton?. Global Change Biology 20(11), 3377-3385.
  25. et. al. European sea bass, Dicentrarchus labrax, in a changing ocean. Biogeosciences 11(9), 2519-2530.
  26. et. al. Bridging the gap between marine biogeochemical and fisheries sciences; configuring the zooplankton link. Progress in Oceanography
  27. & Parental exposure to elevated pCO2 influences the reproductive success of copepods. Journal of Plankton Research 36(5), 1165-1174.
  28. & The role of mixotrophic protists in the biological carbon pump. Biogeosciences 11(4), 995-1005.
  29. & Influence of the N:P supply ratio on biomass productivity and time-resolved changes in elemental and bulk biochemical composition of Nannochloropsis sp.. Bioresource Technology 169, 588-595.
  30. & Variation in elemental stoichiometry of the marine diatomThalassiosira weissflogii(Bacillariophyceae) in response to combined nutrient stress and changes in carbonate chemistry. Journal of Phycology 50(4), 640-651.
  31. & Introduction to the BASIN Special Issue: State of art, past present a view to the future. Progress in Oceanography 129, 171-175.
  32. & Acquired phototrophy in Mesodinium and Dinophysis – A review of cellular organization, prey selectivity, nutrient uptake and bioenergetics. Harmful Algae 28, 126-139.
  33. & Rapid determination of bulk microalgal biochemical composition by Fourier-Transform Infrared spectroscopy. Bioresource Technology 148, 215-220.
  34. & Cutting the Canopy to Defeat the “Selfish Gene”; Conflicting Selection Pressures for the Integration of Phototrophy in Mixotrophic Protists. Protist 164(6), 811-823.
  35. & Monster potential meets potential monster: pros and cons of deploying genetically modified microalgae for biofuels production. Interface Focus 3(1), 20120037-20120037.
  36. & Erratum: Changes in pH at the exterior surface of plankton with ocean acidification. Nature Climate Change 2(10), 760-760.
  37. et. al. Review of climate change impacts on marine aquaculture in the UK and Ireland. Aquatic Conservation: Marine and Freshwater Ecosystems 22(3), 389-421.
  38. et. al. End-To-End Models for the Analysis of Marine Ecosystems: Challenges, Issues, and Next Steps. Marine and Coastal Fisheries 2(1), 115-130.
  39. & Placing microalgae on the biofuels priority list: a review of the technological challenges. Journal of The Royal Society Interface 7(46), 703-726.
  40. & Modeling of HABs and eutrophication: Status, advances, challenges. Journal of Marine Systems 83(3-4), 262-275.
  41. Ecological modelling in a sea of variable stoichiometry: Dysfunctionality and the legacy of Redfield and Monod. Progress In Oceanography 84(1-2), 52-65.
  42. & Phytoplankton in a changing world: cell size and elemental stoichiometry. Journal of Plankton Research 32(1), 119-137.
  43. & Modelling mixotrophy in harmful algal blooms: More or less the sum of the parts?. Journal of Marine Systems 83(3-4), 158-169.
  44. & Is the Growth Rate Hypothesis Applicable to Microalgae?. Journal of Phycology 46(1), 1-12.
  45. Do external resource ratios matter?: Implications for modelling eutrophication events and controlling harmful algal blooms. Journal of Marine Systems 83(3-4), 170-180.
  46. & Aldehyde-induced insidious effects cannot be considered as a diatom defence mechanism against copepods. Marine Ecology Progress Series 377, 79-89.
  47. Food-density-dependent inefficiency in animals with a gut as a stabilizing mechanism in trophic dynamics. Proceedings of the Royal Society B: Biological Sciences 276(1659), 1147-1152.
  48. & Building the "perfect beast": modelling mixotrophic plankton. Journal of Plankton Research 31(9), 965-992.
  49. & Phagotrophy in the origins of photosynthesis in eukaryotes and as a complementary mode of nutrition in phototrophs: relation to Darwin's insectivorous plants. Journal of Experimental Botany 60(14), 3975-3987.
  50. & Accounting for grazing dynamics in nitrogen-phytoplankton-zooplankton (NPZ) models. Limnology and Oceanography 52(2), 649-661.

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  • BI-E00 Bioscience Industrial Placement Year

    The industrial placement year (often referred to as the Year in Industry) takes place before the final year. Only students on schemes which explicitly includes a year in industry are eligible for industrial placements. Students may enrol on programmes with an industrial placement year at the beginning of their studies, subject to appropriate enhanced entry qualifications, or may transfer to such a programme (subject to placement availability) up to the end of Level 5. Students complete a minimum of 40 weeks in a placement in companies in the UK (or potentially outside the UK).

  • BIO227 Marine Plankton And Oceanography

    This module introduces students to the fundamental concept of plankton ecology at level 2. Students will receive 18-20 lectures and four practicals (1 wet lab, 3 IT lab, 1 boat work). The lectures will cover three key themes: oceanography pertaining to planktonic production, plankton ecophysiology, and the functioning of the planktonic foodwebs. Three reports (1 IT-based, 1 from a lab-practical, 1 from boat work) will be assessed, together with one examination consisting of 30 multiple choice questions + one essay question + one analytical question.


  • Macroalgae as Biomass (current)

    Student name:
    Other supervisor: Dr Darren Oatley-Radcliffe
  • Microbial methane production in oxic water (current)

    Student name:
    Other supervisor: Prof Kam Tang
  • Accounting for mixotrophy in marine microbial food webs (current)

    Student name:
    Other supervisor: Dr Aditee Mitra