Dr Carrie Willcox

Institute of Immunology and Immunotherapy
Senior Research Fellow

Contact details

Institute of Immunology and Immunotherapy
Robert Aiken Institute for Clinical Research
University of Birmingham
B15 2TT

Dr Carrie Willcox is a molecular immunologist with a strong interest in antigen recognition by unconventional lymphocytes, and wide experience in molecular and cellular techniques.


  • PhD in Immunology 2002, University of London
  • MSc in Immunology 1998, Yale University
  • BA in Biology 1997, Northwestern University


Carrie has focussed on understanding unconventional T cell recognition for most of her career, and completed her PhD training in Professor Adrian Hayday’s laboratory at King’s College London.

Although she has worked on Natural Killer and Natural Killer T cells, much of her research has been focussed on understanding gamma delta T cell recognition, an area she works on with Prof Ben Willcox. A particular emphasis is on understanding the core immunobiology underlying human gamma delta T cell function, and characterising ligands recognised by the gamma delta T cell receptor.


  • MBChB - Tutor for year 2 Cancer: Causes to Cure
  • MRes - Cancer Immunology and Immunotherapy Module

Postgraduate supervision

In recent years, Carrie has supervised several MSc and PhD students in the field of antigen recognition requirements of gamma delta T cells.

If you are interesting in studying any of these subject areas please contact Dr Carrie Willcox directly, or for any general doctoral research enquiries, please email mds-gradschool@contacts.bham.ac.uk.

For a full list of available Doctoral Research opportunities, please visit our Doctoral Research programme listings.


Carrie is currently funded by Prof Ben Willcox’s Wellcome Trust Award focussing on human gamma delta (γδ) T cells. Through analysis of human γδ T cell receptor (TCR) repertoires, the group has examined two classes of adaptive-like γδ T cells in humans: Vd2neg T cells and Vg9negVd2+ T cells. These subsets express diverse TCRs, and ongoing studies focus on identifying adaptive ligands for this subset. Previously, in collaboration with Dr. Julie Dechanet-Merville (Bordeaux), Carrie identified endothelial protein C receptor (EPCR) as a ligand for a γδ T cell clone that expanded in a patient with acute CMV infection. EPCR remains a key example of a ligand for an adaptive-like γδ T cell clone.

In parallel, it has recently become clear that some human γδ T cell populations are more innate-like, akin to iNKT or MAIT cells in the αβ T cell compartment. Butyrophilin (BTN) family molecules seem to be required for regulation of these innate-like γδ T cells, including BTN3A1 for the human Vg9Vd2+ T cell subset, and BTNL3 for the intestinal Vg4+ compartment. We have recently shown that BTNL3 binds directly to human Vg4+ TCRs in a superantigen-like manner, distinct from clonally-restricted antigen. Future studies are aimed at understanding how BTN-like molecules influence other subsets of human and mouse γδ T cells.

In addition, Carrie is interested in NKG2D ligand recognition, collaborating with Professor Moss’ group on this work in this area. She combines cellular assays with BIAcore binding analyses to understand how the strength of NKG2D-ligand interactions influences anti-tumour immunity.

Other activities

  • Member of the British Society for Immunology
  • Member of the Cancer Immunotherapy and Immunology Centre Advisory Board
  • Reviewer for several Journals


Recent publications


Karunakaran, MM, Subramanian, H, Jin, Y, Mohammed, F, Kimmel, B, Juraske, C, Starick, L, Nöhren, A, Länder, N, Willcox, CR, Singh, R, Schamel, WW, Nikolaev, VO, Kunzmann, V, Wiemer, AJ, Willcox, BE & Herrmann, T 2023, 'A distinct topology of BTN3A IgV and B30.2 domains controlled by juxtamembrane regions favors optimal human γδ T cell phosphoantigen sensing', Nature Communications, vol. 14, no. 1, 7617 . https://doi.org/10.1038/s41467-023-41938-8

Willcox, CR, Salim, M, Begley, CR, Karunakaran, MM, Easton, EJ, von Klopotek, C, Berwick, KA, Herrmann, T, Mohammed, F, Jeeves, M & Willcox, BE 2023, 'Phosphoantigen sensing combines TCR-dependent recognition of the BTN3A IgV domain and germline interaction with BTN2A1', Cell Reports, vol. 42, no. 4, 112321. https://doi.org/10.1016/j.celrep.2023.112321

McMurray, J, von Borstel, A, Taher, T, Syrimi, E, Taylor, G, Sharif, M, Rossjohn, J, Remmerswaal, E, Bemelman, F, Vieira Braga, FA, Chen, X, Teichmann, SA, Mohammed, F, Berry, A, Lyke, K, Williamson, K, Stubbington, M, Davey, M, Willcox, C & Willcox, B 2022, 'Transcriptional profiling of human Vδ1 T cells reveals a pathogen-driven adaptive differentiation program', Cell Reports, vol. 39, no. 8, 110858. https://doi.org/10.1016/j.celrep.2022.110858

Faustini, S, Jossi, S, Perez-Toledo, M, Shields, A, Allen, JD, Watanabe, Y, Newby, ML, Cook, A, Willcox, C, Salim, M, Goodall, M, Heaney, J, Marcial Juarez, E, Morley, G, Torlinska, B, Wraith, D, Veenith, T, Harding, S, Jolles, S, Mark, PJ, Plant, T, Huissoon, A, O’Shea, MK, Willcox, B, Drayson, M, Crispin, M, Cunningham, A & Richter, A 2021, 'Development of a high-sensitivity ELISA detecting IgG, IgA and IgM antibodies to the SARS-CoV-2 spike glycoprotein in serum and saliva', Immunology, vol. 164, no. 1, pp. 135-147. https://doi.org/10.1111/imm.13349

Karunakaran, MM, Willcox, CR, Salim, M, Paletta, D, Fichtner, AS, Noll, A, Starick, L, Nöhren, A, Begley, CR, Berwick, KA, Chaleil, RAG, Pitard, V, Déchanet-merville, J, Bates, PA, Kimmel, B, Knowles, TJ, Kunzmann, V, Walter, L, Jeeves, M, Mohammed, F, Willcox, BE & Herrmann, T 2020, 'Butyrophilin-2A1 directly binds germline-encoded regions of the Vγ9Vδ2 TCR and is essential for phosphoantigen sensing', Immunity, vol. 52, no. 3, pp. 487-498.e6. https://doi.org/10.1016/j.immuni.2020.02.014

Faustini, SE, Jossi, SE, Perez-Toledo, M, Shields, A, Allen, JD, Watanabe, Y, Newby, ML, Cook, A, Willcox, CR, Salim, M, Goodall, M, Heaney, JL, Marcial-Juarez, E, Morley, GL, Torlinska, B, Wraith, DC, Veenith, T, Harding, S, Jolles, S, Mark, PJ, Plant, T, Huissoon, A, O'Shea, MK, Willcox, BE, Drayson, MT, Crispin, M, Cunningham, AF & Richter, AG 2020, 'Detection of antibodies to the SARS-CoV-2 spike glycoprotein in both serum and saliva enhances detection of infection', medRxiv. https://doi.org/10.1101/2020.06.16.20133025

Willcox, CR, Mohammed, F & Willcox, BE 2020, 'The distinct MHC‐unrestricted immunobiology of innate‐like and adaptive‐like human γδ T cell subsets—Nature's CAR‐T cells', Immunological Reviews, vol. 298, no. 1, pp. 25-46. https://doi.org/10.1111/imr.12928

Willcox, BE, Mohammed, F & Willcox, CR 2020, 'γδ TCR Recognition of MR1: Adapting to Life on the Flip Side', Trends in Biochemical Sciences, vol. 45, no. 7, pp. 551-553. https://doi.org/10.1016/j.tibs.2020.03.012

Willcox, C, Vantourout, P, Salim, M, Zlatareva, I, Melandri, D, Zanardo, L, George, R, Kjaer, S, Jeeves, M, Mohammed, F, Hayday, AC & Willcox, B 2019, 'Butyrophilin-like 3 directly binds a human Vγ4+ t cell receptor using a modality distinct from clonally-restricted antigen', Immunity, vol. 51, no. 5, pp. 813-825.e4. https://doi.org/10.1016/j.immuni.2019.09.006

Hunter, S, Willcox, C, Davey, M, Kasatskaya, SA, Jeffery, H, Chudakov, D, Oo, Y & Willcox, B 2018, 'Human liver infiltrating γδ T cells are composed of clonally expanded circulating and tissue-resident populations', Journal of Hepatology, vol. 69, no. 3, pp. 654-665. https://doi.org/10.1016/j.jhep.2018.05.007

Davey, M, Willcox, C, Hunter, S, Kasatskaya, SA, Remmerswaal, EBM, Salim, M, Mohammed, F, Bemelman, FJ, Chudakov, DM, Oo, YH & Willcox, B 2018, 'The human Vδ2+ T-cell compartment comprises distinct innate-like Vγ9+ and adaptive Vγ9- subsets', Nature Communications, vol. 9, no. 1, 1760. https://doi.org/10.1038/s41467-018-04076-0

Davey, MS, Willcox, CR, Hunter, S, Oo, YH & Willcox, BE 2018, 'Vδ2 T Cells-Two Subsets for the Price of One', Frontiers in immunology, vol. 9, 2106. https://doi.org/10.3389/fimmu.2018.02106


Perez-Toledo, M, Faustini, SE, Jossi, SE, Shields, AM, Marcial‐Juarez, E, Kanthimathinathan, HK, Allen, JD, Watanabe, Y, Goodall, M, Willcox, BE, Willcox, CR, Salim, M, Wraith, DC, Veenith, TV, Syrimi, E, Drayson, MT, Jyothish, D, Al‐Abadi, E, Chikermane, A, Welch, SB, Masilamani, K, Hackett, S, Crispin, M, Scholefield, BR, Cunningham, AF & Richter, AG 2021, 'SARS‐CoV‐2‐specific IgG1/IgG3 but not IgM in children with Pediatric Inflammatory Multi‐System Syndrome', Pediatric allergy and immunology, vol. 32, no. 5, pp. 1125-1129. https://doi.org/10.1111/pai.13504

Review article

Willcox, C, Davey, M & Willcox, B 2018, 'Development and selection of the human Vγ9Vδ2+ T- cell repertoire', Frontiers in immunology, vol. 9, 1501. https://doi.org/10.3389/fimmu.2018.01501

Davey, M, Willcox, C, Baker, A, Hunter, S & Willcox, B 2018, 'Recasting Human Vδ1 Lymphocytes in an Adaptive Role', Trends in Immunology, vol. 39, no. 6, pp. 446-459. https://doi.org/10.1016/j.it.2018.03.003

View all publications in research portal