Personal profile


Click here for current PhD opportunities in CHE. But feel free to email me to discuss projects outside these areas and alternative sources of funding including the following BBSRC-DTP funded post.


Group website:
ResearcherID profile
Google Scholar; citation index

Fraser obtained a BSc in Chemistry from the University of Sheffield in 1987 and a Dr. rer. nat. from the Department of Chemistry at the Free University of Berlin (Germany) in 1993. He pursued post-doctoral research at the Technical University Berlin (Germany) (1993-1995), the Commissariat de l’Energie Atomique, Saclay, France (1995-1998) and the Johann Wolfgang Goethe-University Frankfurt (Germany) (1998-2006).

Fraser is currently a Reader in the School of Chemistry at the University of East Anglia, Norwich where he joined the faculty in 2007. To coincide with his appointment Fraser has won a Royal Society Wolfson Research Merit Award. He leads biological EPR research in the Henry Wellcome Laboratory.

His present research aims to elucidate structure / function / dynamics relationships in biomacromolecules and macromolecular complexes. Particular emphasis being on the application, and expansion, of electron paramagnetic resonance methods to study membrane proteins.

Selected publications

Jason C. Crack, Basema K. Balasiny, Sophie P. Bennett, Matthew D. Rolfe, Afonso Froes, Fraser MacMillan, Jeffrey Green, Jeffrey A. Cole, and Nick E. Le Brun
The Di-Iron Protein YtfE Is a Nitric Oxide-Generating Nitrite Reductase Involved in the Management of Nitrosative Stress
J Amer Chem Soc 144, 7129-7145 (2022)

Shiran Barber-ZuckerJenny HallAfonso FroesSofiya KolushevaFraser MacMillan* and Raz Zarivach*
The cation diffusion facilitator protein MamM’s cytoplasmic domain exhibits metal-type dependent binding modes and discriminates against Mn2+
J. Biol. Chem. 295, 16614-16629 (2020)  
DOI: 10.1074/jbc.RA120.014145

Jenny L Hall, Azmat Sohail, Eurico J Cabrita, Colin Macdonald, Thomas Stockner, Harald H. Sitte, Jesus Angulo, & Fraser MacMillan
Saturation transfer difference NMR on the integral trimeric membrane transport protein GltPh determines cooperative substrate binding
Scientific Reports, 10, 16483 (2020)
DOI 10.1038/s41598-020-73443-z 

Matthew O. Ross, Fraser MacMillan, Jingzhou Wang, Alex Nisthal, Thomas J. Lawton, Barry D. Olafson, Stephen L. Mayo, Amy C. Rosenzweig, Brian M. Hoffman
Particulate methane monooxygenase contains only mononuclear copper centers
364, 566-570 (2019)
DOI: 10.1126/science.aav2572

Elliot L. Bennett, Elliot J. Lawrence, Robin J. Blagg, Anna S. Mullen, Fraser MacMillan, Andreas W. Ehlers, Daniel J. Scott,Joshua S. Sapsford, Andrew E. Ashley, Gregory G. Wildgoose, J. Chris Slootweg,
A New Mode of Chemical Reactivity for Metal-Free Hydrogen Activation by Lewis Acidic Boranes
Angewandte Chemie International Edition 58 (25), 8362-8366 (2019)
DOI: 10.1002/anie.201900861

Hannah B. L. Jones, Rory M. Crean, Anna Mullen, Emanuele G. Kendrick, Steven D. Bull , Stephen A. Wells, David R. Carbery, Fraser MacMillan , Marc W. van der Kamp, and Christopher R. Pudney.
Exposing the Interplay Between Enzyme Turnover, Protein Dynamics, and the Membrane Environment in Monoamine Oxidase B
Biochemistry, 58, 2362-2372 (2019)
DOI: 10.1021/acs.biochem.9b00213

Shiran Barber‐Zucker, Jenny Hall, Sivasubramanyan, Venkata Mangapuram, Itamar Kass, Sofiya Kolusheva, Fraser MacMillan, Raz Zarivach, Arnon Henn,
Metal binding to the dynamic cytoplasmic domain of the cation diffusion facilitator (CDF) protein MamM induces a 'locked-in' configuration
FEBS Journal 286(11) 2193-2215 (2019)
DOI: 10.1111/febs.14795 

Despo Polyviou, Moritz M. Machelett, Andrew Hitchcock, Alison J. Baylay, Fraser MacMillan, C. Mark Moore, Thomas S. Bibby and Ivo Tews,
Structural and functional characterisation of IdiA/FutA (Tery_3377) from the ocean diazotroph Trichodesmium erythraeum
J. Biol. Chem. 293, 18099-18109 (2018)
DOI: 10.1074/jbc.RA118.001929

John Anderson, Daniel Watkins, Jonathan Jenkins, Katie Grayson, Nicola Wood, Jack Steventon, Kristian Le Vay, Matt Goodwin, Anna Mullen, Henry Bailey, Matthew Crump, Fraser MacMillan, Adrian Mulholland, Gus Cameron, Richard Sessions, and Stephen Mann
Construction and in vivo assembly of a catalytically proficient and hyperthermostable de novo enzyme
Nature Communications 8, 358 (2017)
DOI: 10.1038/s41467-017-00541-4



  • 2007 –  present: Reader in Chemistry in the School of Chemistry and Director of the Henry Wellcome Unit for Biological EPR, UEA
  • 2007 – 2012: Wolfson Research Merit Award of The Royal Society
  • 1998 – 2006: Independent/Junior Group Leader Fachbereich Chemie, Biochemie & Pharmazie, J. W. Goethe Universität, Frankfurt, Germany
  • 1995 – 1998: Marie-Curie Stipendiat (of the EU)  at the CEA Saclay, Séction de Bioénergétique, CEA Saclay, Gif-sur-Yvette, France 1993 – 1995    Post-Doc in the Max-Volmer Institut of the Technische Universität Berlin, Germany
  • 1988 – 1993: Dr. rer. nat. in the Institut für Organische Chemie of the Freie Universität Berlin, Germany
  • 1984 – 1987: B.Sc. Sp. Hons. in Pure and Applied Chemistry at the University of Sheffield, UK


Research Group or Lab Membership


  • Dr. Christian Teutloff (Royal Society, Non Stipendiary) (2007 - )
  • Dr. Jessica van Wonderen (UEA/Wellcome Trust) (2009 - )
    Dr. Megen O’Mara (Non Stipendiary) (2012 - 2014)
  • Dr. Gaye White UEA, (BBSRC) 2007 – 2008
  • Dr. Justin Bradley UEA (Royal Society), 2011 – 2012

PhD Students (pre UEA)

  • Christian Teutloff, FU Berlin, Germany, (DFG) 1998 – 2003
  • Stéphane Grimaldi, Goethe Universität Frankfurt, Germany (DFG), 1998 – 2002
  • Axel Weber, Goethe Universität Frankfurt, Germany (DFG) 1999 – 2003
  • Thorsten Maly, Goethe Universität Frankfurt, Germany (DFG) 1999 – 2004
  • Sevdalina Lyubenova, Goethe Universität Frankfurt, Germany (DFG) 2000 – 2006
  • Jörg Fritscher, Goethe Universität Frankfurt, Germany (DFG) 2001 – 2006
  • Marloes Penning de Vries, Goethe Universität Frankfurt, Germany (DFG) 2003 – 2006

PhD Students (at UEA)

  • Matthew Bawn, (BBSRC) 2007 – 2012
  • Morgan Bye (EPSRC), 2009 – 2014
  • Alexander Morley (EU), 2013 – 2014, completed in Vienna
  • Suzy Jones, (UEA), 2012 – 2013, completed in Leicester
  • Anna Mullen, (MED/SCI) 2015 – 2019
  • Jenny Hall, (BBSRC NRP-DTP) 2016 – 2021
  • Janika Diegel, (BBSRC NRP-DTP iCASE), 2017 – 2018 moved to Freiburg
  • Afonso Froes (self-funded), 2019 – 2023
  • Shona Hepworth, (BBSRC NRP-DTP iCASE), 2019 – 2023
  • Oliver Dillon, (BBSRC NRP-DTP iCASE), 2020 – 2024, secondary supervisor
  • Petros Tsalagradas (EU-MSCN, ETN), 2020 – 2024
  • Despoina Kapiki (EU-MSCN, ETN), LMU Munich, Germany, 2021 – 2024, secondary supervisor.
  • Zaid Ramdhan Anshari (EU-MSCN, ETN), RUG University Groningen, The Netherlands, 2020 – 2024, secondary supervisor.
  • Alberto Daminato (EU-MSCN, ETN), Nova University Lisbon, Portugal, 2021 – 2024, secondary supervisor.

Masters Research Students (pre UEA)

  • Louise van der Weerd (Wageningen/Saclay, Netherlands/France) 1996

  • Christian Teutloff (TU Berlin, Germany) 1997
  • Thorsten Maly (Goethe Universität Frankfurt, Germany) 1999
  • Janika Diegel (Freiburg, Germany) 2015 


Masters Research Students (at UEA)

  • Deborah Cave (BSc) 2007
  • Zoe Karthauser (MChem) 2008
  • Christopher Stevens (BSc) 2010
  • Alicia Stratford (MChem) 2010
  • Glenn Tobias (MChem) 2010
  • Ben Anderson (MChem) 2011
  • Alexander Morley (MChem) 2012, Prize for best research project 2012
  • Shivani Bhanaut (BSc) 2012
  • Thokozile Usuwana (BSc) 2013          
  • Chathurika Daysena (BSc) 2013
  • Anna Mullen (MChem) 2014
  • Alexander Pak (BSc) 2014
  • Isa Hassan (MChem) 2015     
  • Adam Fey (MChem) 2015
  • Jenny Hall (MChem) 2015     
  • Hannah Blades (BSc) 2016
  • Sophie Orme (MChem) 2016
  • Mikey Whitcutt (MChem) 2016
  • Patrick Marchington (MChem) 2017, Prize for best research project 2017
  • Shannon White (MChem) 2017
  • Afonso Froes (MChem) 2018
  • Teri Branson (MChem) 2018
  • Shona Hepworth (MChem) 2018       
  • Zara Krumins (MNatSci) 2019
  • Jack Sawyer (MChem) 2020
  • Charlie Thompson (MChem) 2020
  • NeiL Cadang (MChem) 2021
  • Rory Haughan (MChem) 2021
  • Armin Nikpour Khoshgrudi (MSci) 2021
  • Callum Eke (BSc Biochem) 2022 

Externally Funded Summer Internships at UEA

  • Alex Morley, The Biochemical Society, 2011
  • Helen Newson , EU, 2012
  • Anna Mullen, The Biochemical Society, 2014
  • Alexander Pak, Royal Society of Chemistry, 2014
  • Isa Hassan, The Wellcome Trust, 2014
  • Jenny Hall, The Wellcome Trust, 2015
  • Adam Fey, UEA funded summer studentship, 2015
  • Sophie Orme, UEA funded summer studentship, 2016
  • Patrick Marchington, EU, 2018
  • Afonso Froes, BBSRC NRP-DTP, 2018
  • Shona Hepworth, The Biochemical Society, 2018
  • Hannah Luedke, The Wellcome Trust, 2018
  • Armin Nikpour Khoshgrudi, The Wellcome Trust 2021
  • Stephanie Wise, UEA funded summer studentship 2021
  • Callum Eke, The Wellcome Trust 2022


Guest Research Visitors

  • Roberto Poddighe, University of Sassari, Italy, (Erasmus), 2012 – 2013
  • Prof. Megan O'Mara, ANC Canbera, Australia, 2013-2014
  • Janika Diegel, University of Freiburg, Germany, 2014 – 2015
  • Beatriz G R Pinheiro, University of Sau Paulo, Brazil, 2014 – 2015
  • Dr. Shiran Barber-Zucker, Ben Gurion University, Israel, 2014 & 2016
  • Prof. Yonca Yuzugullu, Kocaeli University, Turkey, 2014 – 2015
  • Dr. Chritian Teutloff, Freie Universität Berlin, Germany, 2017
  • Prof. Eurico Cabrita, Novo University Lisbon, Portugal, 2018
  • Prof. Csaba Bagyinka, BRC, Szeged, Hungary, 2018

Key Research Interests

Current Research/Technology Expertise and Interests:

  • All Aspects of Paramagnetic Resonance Spectroscopy (EPR, ESR) including Multi-Frequency (S-, X-, Q-, W-bands) EPR and Pulsed EPR (ENDOR, PELDOR, ESEEM, HYSCORE) Spectroscopy
  • Membrane and metalloproteins including photosynthetic and respiratory proteins
  • Structure/function/dynamics relationships in biomacromolecules
  • Electron transfer and (ABC-)transporter proteins

Current Methodologies/Techniques Used

  • All modern advanced EPR spectroscopies
  • MO-calculations (DFT)
  • Freeze-quenching
  • Spin-labelling

Areas for Potential Collaboration:
Anywhere where structural or structure/functional (including dynamic) information of a biomacromolecule is of importance. While endogenous spin probes (transition metal ion cofactors, organic cofactors or protein based radicals) are the probes of choice for the methods we apply, the use of exogenous spin probes (nitroxide spin labels and transition metal ions) also allows us to address inherently diamagnetic proteins and protein complexes, permitting access to similar information in these cases.

Current Research Projects:
My interests lie in the application and development of modern paramagnetic resonance spectroscopic techniques to characterise and correlate the structure and function of biomacromolecules. We apply advanced EPR techniques to study both local and global interactions of endogenous as well as exogneous spin probes providing very detailed structural information, which can be considered complementary to crystallographic and other structural techniques. In principle we are able to work with isolated proteins, protein ensembles, enriched membrane fragments, proteins reconstituted in proteoliposomes as well as in whole cells. Selected current active collaborations outside UEA include work on transporter proteins (p-Glycoprotein (Oxford), ATP-transporters (MRC Cambridge), respiratory proteins (complexes I, II, III and IV (Philadelphia, San Francisco, Freiburg, Newcastle and Frankfurt)), Colicins (York, Nottingham) and photosynthetic proteins (bioenergy systems (Tempe, Columbia)).

For Full details please see

Research Funding

EU MSCA ITN Network: 
NeuroTrans (NEUROtransmitter TRANSporters: from single molecules to human pathologies)
MacMillan F, Co-I (ca. .€4,000,000)
February 2020 – January 2023

The Wellcome Trust: Award (to Prof. Changjiang Dong/Dr. MacMillan)
“From Structural and functional studies of essential and novel lipopolysaccharide transport and assembly membrane proteins to novel drug discoveries combating emerging multi-drug resistant pathogenic bacteria (£2,000,000) 2016 - 2021

The Wellcome Trust: SEED Award
"Life’s ups and downs: probing the conformational dynamics of DASS transporters, a family of transporters associated with age-related diseases and lifespan determination." Funding to Dr. C. Mulligan with F. MacMillan for a joint project 2018-2020.

EU COST Action CM1306: 
Understanding Movement and Mechanism in Molecular Machines
MacMillan F (Chair and Grant Holder, UEA, UK),
June 2014 – December 2019

EU COST Action CM0902:
Molecular Machineries for Ion Translocation Across Biomembranes, Cellular Transport for Biotechnology
MacMillan F (Chair and Grant Holder, UEA, UK),
01 January 2010 – 17 November 2013

The Wellcome Trust
The molecular process underlying multidrug efflux by P-glycoprotein
MacMillan F, Callaghan R (University of Canberra, Australia).
January 2012 - January 2014

The John & Pamela Salter Charitable Trust
Analysis of ToIC Mutants by Electron Paramagnetic Resonance (EPR) Spectroscopy of Pulsed Electron Double Resonance (PELDOR)
Fanutti, C, MacMillan F
01 November 2010 – 30 June 2013

Synthesis of selectively spin labelled cardiolipin
MacMillan F, Hunte C (University of Freiburg, Germany)
05 February 2009 – 31 July 2013

The Royal Society; Wolfson Research Merit Award
Time domain and variable frequency EPR of membrane proteins
MacMillan F
01 April 2007 – 6 November 2012

The Biochemical Society
2011 Summer Studentship
MacMillan F
July 2011

The Royal Society; Travel Grant
52rd Annual Rocky Mountain Conference on Analytical Chemistry, Aspen, USA
MacMillan F
August 2010

The Biochemical Society Travel Grant
16th European Bioenergetics Conference – Warsaw, Poland
MacMillan F
July 2010

COST Action P15:
Advanced paramagnetic resonance methods in molecular biophysics
EU-COST Training School at the 42nd International Meeting of the RSC EPR group, Norwich
MacMillan, F
April 2009

The Wellcome Trust
The molecular mechanism of multidrug efflux by P-glycoprotein: an electron paramagnetic resonance spin label study
MacMillan F, Callaghan R (University of Oxford).
01 January 2009 - 31 July 2010

The Royal Society; Research Grant
A Q-band ENDOR investigation of ubiquinol-cytochrome c reductase (QCR)
MacMillan F
07 March 2008 – 06 July 2010

Deutsche Forschungsgemeinschaft
Multidimensional EPR-spectroscopy on protein-bound, redox-active, paramagnetic centres
MacMillan F & Prisner TP
01 January 2000 – 31 December 2005



Areas of Expertise

Magnetic resonance spectroscopy; structure and function of biomolecules, esp. proteins; protein-ligand interactions; electron transfer and biological transport (multi drug efflux) processes.

Administrative Posts

2022 –                 Deputy Chair of School of Chemistry Exam Board.
2020 – 2022        Member of the UEA External Expertise Project Group.
2019 –  2022      Chair of School of Chemistry Exam Board.
2016 –                 Course Director for the MSci Biochemistry with a Year abroad degree programme
2016 –                 Deputy-director of the BSc and MSci Biochemistry degree programmes 
2016 –                 Member of the Chemistry Learning and Teaching Committee.
2016 – 2019        Head of Year 3 in the School of Chemistry.
2016 – 2019        Member of the Faculty of Science Web Steering Group.
2015 – 2021        Creator/Lead Educator (with R Bowater) of MOOC “Biochemistry:The Molecules of Life:
2012 – 2018        Course Director for all the Biological and Medicinal Chemistry programmes
2012 – 2018        Co-director of Biochemistry degree programmes of the School of Biological Sciences 
2009 – 2013        Member of the School of Chemistry Executive Committee
2009 – 2012        Director of Admissions for the School of Chemistry at UEA

Expertise related to UN Sustainable Development Goals

In 2015, UN member states agreed to 17 global Sustainable Development Goals (SDGs) to end poverty, protect the planet and ensure prosperity for all. This person’s work contributes towards the following SDG(s):

  • SDG 3 - Good Health and Well-being

Education/Academic qualification

Doctor rerum naturalium, Free University of Berlin

1 Jan 19886 Oct 1993

Award Date: 6 Oct 1993


  • Chemistry
  • Membrane transport
  • Membrane proteins
  • Magnetic resonance
  • EPR/ESR spectroscopy
  • Bioenergetics
  • Structural biology
  • Site-directed spin labelling
  • Metalloproteins
  • Radicals
  • Electron transfer

Collaborations and top research areas from the last five years

Recent external collaboration on country/territory level. Dive into details by clicking on the dots or