1987 PhD. Department of Physiology University of Edinburgh 1987 - 1989 Postdoctoral Fellow, Department of Physiology University of Edinburgh 1989 - 1992 Rockefeller Postdoctoral fellow in Reproductive Biology, The Jackson Laboratory , Bar Harbor, Me, USA 1992 - 2000 Lecturer Institute of Ecology and Resource Management, Division of Biological Sciences, University of Edinburgh 2000 - Senior Lecturer, Institute of Cell and Molecular Biology, University of Edinburgh

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