Muscles and movement: a memorial meeting to celebrate the life and work of Professor Roger Woledge (1938-2015).

On a cold, damp late-April day in east London, the family, friends, colleagues and admirers of muscle physiologist Roger Woledge came together to remember his life and scientific contributions at a celebratory symposium at Queen Mary University of London (QMUL). Woledge died unexpectedly in March 2015 following a riding accident, but by the end of the day-long symposium supported by The Company of Biologists, we were left in little doubt of the great intellect and infectious enthusiasm that defined Woledge and his approach to life and research. While some of the delegates had come from just along the corridor, others had travelled from as far as Australia and the USA, to share recollections, anecdotes and their affection for this giant of modern muscle physiology. ‘Roger had a depth of knowledge that was, to me, unsurpassed’, said meeting co-organiser Tim West, who was a postdoc with Woledge and one of his key collaborators, Nancy Curtin, at Imperial College, London, in the late 1990s. Saying that Woledge’s versatility was exceptional, West adds, ‘He could talk about an enzyme in a dish, understand the kinetics of it and talk to experts about it. At the same time he could turn around and talk about muscle physiology and whole-animal kinematics with as deep an understanding.’ Introducing the symposium, Curtin detailed Woledge’s early career as an undergraduate in the University College London (UCL) Physiology Department, where he was able to satisfy ‘his itch for quantitative information’ with his love for practical work. After graduating from UCL, Woledge joined the lab of Nobel Prize winning scientist A. V. Hill for a three year fellowship. ‘The years that Roger spent with A. V. must have been a remarkably good match between mentor and student; there seems to have been just the right balance between instruction by A. V. in the lab and time when Roger worked on his own’, Curtin said. However, she remembers that Hill did not encourageWoledge to apply for a PhD, because Hill did not believe in the degree. Instead, Doug Wilkie persuaded Woledge that he should register for a PhDwhile in his lab towork on the energetics of tortoise muscle, andWoledge eventually graduated in 1966, publishing his seminal paper in the Journal of Physiology two years later (Woledge, 1968). Referring to their life-long collaboration, Curtin said, ‘He was not one of these people that got discouraged when things didn’t work; he was quite optimistic. He also knew a lot, we complemented each other as we each had our own separate skills.’ Ultimately, Woledge went on to head the department where he had been an undergraduate, in addition to directing the UCL Institute of Human Performance before his retirement from university administration in 2003, although he continued to publish in journals such as Journal of Applied Biomechanics and Journal of Experimental Biology until his death. Over the course of the symposium, friends and colleagues recalled their discussions and collaborations with Woledge. Jack Rall from Ohio State University, USA, described Woledge’s innate sense of fairness. Remembering discussions with Woledge about the challenges faced by Graham Hoyle and others in the 1980s as they attempted to gain acceptance for the role of a third ‘S’ filament in muscle contraction, Rall said that Woledge encouraged him to discuss the controversy in his recently published book, Mechanism of Muscular Contraction (Rall, 2014), ‘as he felt that the third filament’s proponents were being ignored unjustifiably’, said Rall. Throughout the remainder of his talk, Rall recalled the early suppression of the role of the S filament, which had been evident in the first papers outlining the sliding filament model of muscle contraction, and then discussed the work of Koscak Maruyama and Kuan Wang, who independently identified the key S filament protein, now known as titin. In the following talk, Earl Homsher from the University of California Los Angeles, USA, acknowledged his debt of gratitude to Woledge, talking about his insight into experimental design and modest approach when discussing experiments. ‘For me, he was a leading force in muscle energetics at a time of confusion, and showed us how to use energetics to evaluate processes that occurred in muscle.... Roger’s experiments and papers on energetics set the standard for muscle energetics research in the 1970s and, I think, to the present’, said Homsher. Outlining the structure of the book Energetic Aspects of Muscle Contraction (Woledge et al., 1985) that he wrote in collaboration with Woledge and Curtin, Homsher described the immense investment of time that resulted in the chapter dedicated to the heat of chemical change – accounting for 108 of the book’s eventual 357 pages. Homsher explains that the chapter was designed ‘to tell you everything you wanted to Roger Woledge riding his horse Luke at Shoeburyness Beach ca. 2012. Photo credit: Elizabeth Dollimore.