[1] His PhD was supervised by Michael Waterfield at the Imperial Cancer Research Fund where he investigated Epidermal growth factor receptor, establishing in 1984 the close similarity between this cellular growth regulatory protein and the avian retroviral oncogene, v-erbB.

His work on the Ras GTPase has made seminal contributions to our understanding of how cellular signal transduction pathways are subverted in oncogenic transformation.

He showed that transformation by Ras requires interaction with multiple effectors, which contribute differentially to cell cycle progression, cytoskeletal regulation and apoptosis.

[13] He is a member of the European Molecular Biology Organisation (1995) and an honorary Fellow of the Royal College of Physicians (2012).

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