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. & Simulating Effects of Variable Stoichiometry and Temperature on Mixotrophy in the Harmful Dinoflagellate Karlodinium veneficum. Frontiers in Marine Science 5
  2. & The simultaneous assimilation of ammonium and l-arginine by the marine diatom Phaeodactylum tricornutum Bohlin. Journal of Experimental Marine Biology and Ecology 95(3), 257-269.
  3. & Effects of N deprivation and darkness on composition of free amino acid pool in and on amino acid release from diatom Phaeodactylum tricornutum Bohlin. Journal of Experimental Marine Biology and Ecology 119(2), 131-143.
  4. & The ratio of glutamine:glutamate in microalgae: a biomarker for N-status suitable for use at natural cell densities. Journal of Plankton Research 11(1), 165-170.
  5. Composition of intracellular and extracellular pools of amino acids, and amino acid utilization of microalgae of different sizes. Journal of Experimental Marine Biology and Ecology 139(3), 151-166.
  6. & Changes in intracellular and extracellular ?-amino acids in Gloeothece during N2-fixation and following addition of ammonium. Archives of Microbiology 153(6), 574-579.
  7. & Relationships between photopigments, cell carbon, cell nitrogen and growth rate for a marine nanoflagellate. Journal of Experimental Marine Biology and Ecology 153(1), 87-96.
  8. Algal carbon–nitrogen metabolism: a biochemical basis for modelling the interactions between nitrate and ammonium uptake. Journal of Plankton Research 13(2), 373-387.
  9. Non-steady state ammonium-limited growth of the marine phytoflagellate, Isochrysis galbana Parke.
  10. & Changes in fatty acids, amino acids and carbon/nitrogen biomass during nitrogen starvation of ammonium- and nitrate-grownIsochrysis galbana. Journal of Applied Phycology 4(2), 95-104.
  11. Relations between carbon and nitrogen during growth of Nannochloropsis oculata (Droop) Hibberd under continuous illumination.
  12. & Predator-prey interactions between Isochrysis galbana and Oxyrrhis marina. II. Release of non-protein amines and faeces during predation of Isochrysis. Journal of Plankton Research 15(8), 893-905.
  13. & Predator-prey interactions between Isochrysis galbana and Oxyrrhis marina. I. Changes in particulate δ 13C. Journal of Plankton Research 15(4), 455-463.
  14. & Modelling temporal decoupling between biomass and numbers during the transient nitrogen-limited growth of a marine phytoflagellate. Journal of Plankton Research 15(3), 351-359.
  15. Changes in dinoflagellate intracellular amino acids in response to diurnal changes in light and N-supply.
  16. & Carbon-nitrogen relations during batch growth ofNannochloropsis oculata (Eustigmatophyceae) under alternating light and dark. Journal of Applied Phycology 5(4), 465-475.
  17. Use of intracellular amino acid analysis as an indicator of the physiological status of natural dinoflagellate populations.
  18. Changes in toxin content, biomass and pigments of the dinoflagellate Alexandrium minutum during nitrogen refeeding and growth into nitrogen or phosphorus stress.
  19. Carbon-nitrogen relations at whole-cell and free-amino-acid levels during batch growth of Isochrysis galbana (Prymnesiophyceae) under conditions of alternating light and dark.
  20. Predator-prey interactions between Isochrysis galbana and Oxyrrhis marina. III.Mathematical modelling of predation and nutrient regeneration.
  21. Modelling interactions between phytoplankton and bacteria under nutrient-regenerating conditions.
  22. & Modelling interactions between phytoplankton and bacteria under nutrient-regenerating conditions. Journal of Plankton Research 17(6), 1395-1395.
  23. & Prey selection and rejection by a microflagellate; implications for the study and operation of microbial food webs. Journal of Experimental Marine Biology and Ecology 196(1-2), 357-372.
  24. Comparisons among species of Alexandrium (dinophyceae) grown in nitrogen- or phosphorus-limiting batch culture.
  25. Changes in toxins, intracellular and dissolved free amino acids of the toxic dinoflagellate Gymnodinium catenatum in response to changes in inorganic nutrients and salinity.
  26. & An automated HPLC method for the rapid analysis of paralytic shellfish toxins from dinoflagellates and bacteria using precolumn oxidation at low temperature. Journal of Experimental Marine Biology and Ecology 197(1), 145-157.
  27. & Modelling the interactions between ammonium and nitrate uptake in marine phytoplankton. Philosophical Transactions of the Royal Society B: Biological Sciences 352(1361), 1625-1645.
  28. Evidence for production of paralytic shellfish toxins by bacteria associated with Alexandrium spp. (Dinophyta) in culture.
  29. A short version of the ammonium-nitrate interaction model.
  30. & Release of nitrite by marine dinoflagellates: development of a mathematical simulation. Marine Biology 130(3), 455-470.
  31. Estimation of kinetic parameters for the transport of nitrate and ammonium into marine phytoplankton.
  32. Variations in the maximum transport rates for ammonium and nitrate in the prymnesiophyte Emiliania huxleyi and the raphidophyte Heterosigma carterae.
  33. & Utilization of dissolved inorganic carbon (DIC) and the response of the marine flagellateIsochrysis galbanato carbon or nitrogen stress. New Phytologist 144(3), 463-470.
  34. The loss of organic nitrogen during marine primary production may be significantly overestimated when using 15N substrates.
  35. Nitrate transport and ammonium-nitrate interactions at high nitrate concentrations and low temperature.
  36. & Interactions between nitrate and ammonium in Emiliania huxleyi. Journal of Experimental Marine Biology and Ecology 236(2), 307-319.
  37. Interactions between iron, light, ammonium, and nitrate: Insights from the construction of a dynamic model of algal physiology.
  38. An investigation of non-steady-state algal growth. I. An experimental model ecosystem.
  39. & Amino acid uptake by the toxic dinoflagellate Alexandrium fundyense. Marine Biology 133(1), 11-19.
  40. The relationship between the dissolved inorganic carbon concentration and growth rate in marine phytoplankton.
  41. & Modelling phosphate transport and assimilation in microalgae; how much complexity is warranted?. Ecological Modelling 125(2-3), 145-157.
  42. Modelling Si-N-limited growth of diatoms.
  43. & Growth dynamics and toxicity of Alexandrium fundyense (Dinophyceae): the effect of changing N∶P supply ratios on internal toxin and nutrient levels. European Journal of Phycology 35(1), 11-23.
  44. & A mechanistic model of photoinhibition. New Phytologist 145(2), 347-359.
  45. A comparison of two N-irradiance interaction models of phytoplankton growth.
  46. Toxin production in migrating dinoflagellates: a modelling study of PSP producing Alexandrium. Harmful Algae 1(2), 147-155.
  47. & The large capacity for dark nitrate-assimilation in diatoms may overcome nitrate limitation of growth. New Phytologist 155(1), 101-108.
  49. & Modelling suggests that optimization of dark nitrogen-assimilation need not be a critical selective feature in phytoplankton. New Phytologist 155(1), 109-119.
  50. Modelling changes in paralytic shellfish toxin content of dinoflagellates in response to nitrogen and phosphorus supply.

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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.


  • 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 (current)

    Student name:
    Other supervisor: Dr Aditee Mitra