He developed improved experimental equipment and applied numerical methods to the study of lichen ecophysiology.
After compulsory National Service in the British military, he undertook doctoral research about plant communities at the University of Wales, Bangor, supervised by Peter Greig-Smith.
He and his students also worked on lichens, characterising the genera Stereocaulon and Umbilicaria and also undertook ecophysiological investigations, including measurements of photosynthesis, carbon dioxide exchange and nitrogen fixation.
He collaborated with a climatologist, Wayne Rouse, for over a decade studying the impact of microclimates on lichen physiology.
His other research programmes focused on lichens in woodland and continued to investigate photosynthesis and nitrogen fixation.