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. & 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.
  2. & Minimising losses to predation during microalgae cultivation. Journal of Applied Phycology 29(4), 1829-1840.
  3. & Oceanic protists with different forms of acquired phototrophy display contrasting biogeographies and abundance. Proceedings of the Royal Society B: Biological Sciences 284(1860), 20170664
  4. & Physiology limits commercially viable photoautotrophic production of microalgal biofuels. Journal of Applied Phycology
  5. & Introducing mixotrophy into a biogeochemical model describing an eutrophied coastal ecosystem: The Southern North Sea. Progress in Oceanography 157, 1-11.
  6. & Modelling alkaline phosphatase activity in microalgae under orthophosphate limitation: the case ofPhaeocystis globosa. Journal of Plankton Research 37(5), 869-885.
  7. & Ocean Acidification Affects the Phyto-Zoo Plankton Trophic Transfer Efficiency. PLOS ONE 11(4), e0151739
  8. A mechanistic model for describing dynamic multi-nutrient, light, temperature interactions in phytoplankton. Journal of Plankton Research 23(9), 977-997.
  9. & Coupling a simple irradiance description to a mechanistic growth model to predict algal production in industrial-scale solar-powered photobioreactors. Journal of Applied Phycology
  10. & Why Plankton Modelers Should Reconsider Using Rectangular Hyperbolic (Michaelis-Menten, Monod) Descriptions of Predator-Prey Interactions. Frontiers in Marine Science 3
  11. & What is the limit for photoautotrophic plankton growth rates?. Journal of Plankton Research
  12. & Estimating the ecological, economic and social impacts of ocean acidification and warming on UK fisheries. Fish and Fisheries
  13. & The role of coccolithophore calcification in bioengineering their environment. Proceedings of the Royal Society B: Biological Sciences 283(1833), 20161099
  14. et. al. Defining Planktonic Protist Functional Groups on Mechanisms for Energy and Nutrient Acquisition: Incorporation of Diverse Mixotrophic Strategies. Protist 167(2), 106-120.
  15. & 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.
  16. & Ocean acidification with (de)eutrophication will alter future phytoplankton growth and succession. Proceedings of the Royal Society B: Biological Sciences 282(1804), 20142604-20142604.
  17. & Modeling DOM Biogeochemistry. In Biogeochemistry of Marine Dissolved Organic Matter. -667).
  18. et. al. Bridging the gap between marine biogeochemical and fisheries sciences; configuring the zooplankton link. Progress in Oceanography
  19. et. al. European sea bass, Dicentrarchus labrax, in a changing ocean. Biogeosciences 11(9), 2519-2530.
  20. et. al. The role of mixotrophic protists in the biological carbon pump. Biogeosciences 11(4), 995-1005.
  21. & Have we been underestimating the effects of ocean acidification in zooplankton?. Global Change Biology 20(11), 3377-3385.
  22. & Parental exposure to elevated pCO2 influences the reproductive success of copepods. Journal of Plankton Research 36(5), 1165-1174.
  23. & In silico optimization for production of biomass and biofuel feedstocks from microalgae. Journal of Applied Phycology 27(1), 33-48.
  24. & 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.
  25. & 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.
  26. & The role of mixotrophic protists in the biological carbon pump. Biogeosciences 11(4), 995-1005.
  27. & Cutting the Canopy to Defeat the “Selfish Gene”; Conflicting Selection Pressures for the Integration of Phototrophy in Mixotrophic Protists. Protist 164(6), 811-823.
  28. & Rapid determination of bulk microalgal biochemical composition by Fourier-Transform Infrared spectroscopy. Bioresource Technology 148, 215-220.
  29. & Monster potential meets potential monster: pros and cons of deploying genetically modified microalgae for biofuels production. Interface Focus 3(1), 20120037-20120037.
  30. & Modelling mixotrophy in harmful algal blooms: More or less the sum of the parts?. Journal of Marine Systems 83(3-4), 158-169.
  31. & Is the Growth Rate Hypothesis Applicable to Microalgae?. Journal of Phycology 46(1), 1-12.
  32. & Aldehyde-induced insidious effects cannot be considered as a diatom defence mechanism against copepods. Marine Ecology Progress Series 377, 79-89.
  33. 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.
  34. Modelling multi-nutrient interactions in phytoplankton; balancing simplicity and realism. , 249-279.
  35. Nutritional status and diet composition affect the value of diatoms as copepod prey. , 1457-1459.
  36. & Chlorophyll content and fluorescence responses cannot be used to gauge reliably phytoplankton biomass, nutrient status or growth rate. , 525-536.
  37. & Importance of interactions between food quality, quantity, and gut transit time on consumer feeding, growth, and trophic dynamics. , 632-646.
  38. Do external resource ratios matter?: Implications for modelling eutrophication events and controlling harmful algal blooms. Journal of Marine Systems 83(3-4), 170-180.
  39. & Modelling mixotrophy in harmful algal blooms: More or less the sum of the parts?. Journal of Marine Systems 83(3-4), 158-169.
  40. 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.
  41. & Placing microalgae on the biofuels priority list: a review of the technological challenges. Journal of The Royal Society Interface 7(46), 703-726.
  42. & Modeling of HABs and eutrophication: Status, advances, challenges. Journal of Marine Systems 83(3-4), 262-275.
  43. & IS THE GROWTH RATE HYPOTHESIS APPLICABLE TO MICROALGAE?1. Journal of Phycology 46(1), 1-12.
  44. Ecological modelling in a sea of variable stoichiometry: Dysfunctionality and the legacy of Redfield and Monod. Progress In Oceanography 84(1-2), 52-65.
  45. & Phytoplankton in a changing world: cell size and elemental stoichiometry. Journal of Plankton Research 32(1), 119-137.
  46. & Building the "perfect beast": modelling mixotrophic plankton. Journal of Plankton Research 31(9), 965-992.
  47. & 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.
  48. Going for the slow burn: why should possession of a low maximum growth rate be advantageous for microalgae?. Plant Ecology & Diversity 2(2), 179-189.
  49. & Allometry and stoichiometry of unicellular, colonial and multicellular phytoplankton. New Phytologist 181(2), 295-309.
  50. & Selection for fitness at the individual or population levels: Modelling effects of genetic modifications in microalgae on productivity and environmental safety. Journal of Theoretical Biology 263(3), 269-280.

See more...


  • 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.


  • Disposal of washed up seaweeds in an ecologically and commercially optimum way (current)

    Student name:
    Other supervisor: Dr Darren Oatley-Radcliffe
  • N/A (current)

    Student name:
    Other supervisor: Prof Kam Tang
  • Accounting for mixotrophy in marine microbial food webs: investigating the new paradigm for marine ecology. (a review is being considered) (current)

    Student name:
    Other supervisor: Dr Aditee Mitra