|2017 Award of Merit Winner|
2017 Award of Merit Winner: Alan Palmer
The 2017 Award of Merit winner Alan Palmer has always defined himself as, ‘just a simple biologist’. But in that apparently unassuming guise, he has, over the past forty and more years, made an impressive contribution to our understanding of several fundamental questions in auditory neuroscience.
Indeed, what characterizes Alan’s work is the sheer diversity of brain regions he has studied over the course of his career. Starting at the auditory nerve for his PhD studies, he worked progressively upwards to cochlear nucleus, inferior colliculus, superior colliculus, thalamus and cortex - developing techniques as the scientific questions required. A second major facet of Alan’s career is his huge influence as a mentor to the many students and postdocs who have worked with him.
Alan was the first from a large working class family to continue in education beyond the age of sixteen. He studied Biological Sciences at the University of Birmingham graduating with first class honours in 1972. As an undergraduate he had developed an interest in vision and his first intention was to gain a PhD in that field. With that goal, he attended an interview at the Department of Communication and Neuroscience at Keele University during which he was persuaded to switch systems and work on the physiology of hearing with Ted Evans. The early 1970s were exciting times in auditory science with an explosion of pioneering measurements on basilar membrane mechanics, hair cell physiology and otoacoustic emissions. Alan’s PhD involved cochlear nucleus and auditory nerve recordings where he discovered that in the presence of noise backgrounds the dynamic range of cochlear nerve and cochlear nucleus neurons shifts to keep them with in operating range. This work has since been explored and revisited at several levels of the auditory pathway levels.
Following a postdoctoral fellowship with Ted Evans, Alan obtained a staff position at the National Institute for Medical Research in Mill Hill, London – an appointment that gave him carte blanche and to build his own laboratory from scratch. Inspired by the work of Knudsen and Konishi in the owl, and at the encouragement of his Head of Division, Mike Keating, he set up a free-field auditory stimulation system in anechoic space to explore auditory responses in the deep layers of the superior colliculus. He was joined in this work by Andrew King, then a PhD student, and they discovered and reported in Nature the now well-established map of auditory space in the mammalian superior colliculus. While at Mill Hill he also got involved in measuring the mechanics of hearing in the frog, a task that included developing with Andrew Pinder a white light method to study vibration in the frog ear, and the first publication of otoacoustic emissions in the frog.
After four hugely productive years at the Mill Hill, Alan was persuaded by Ian Russell and Chris Darwin to apply for one of the Royal Society’s new University Research Fellowships to move to the University of Sussex. Only 15 fellowships were available across all areas of science, but Alan was successful with an application to work on the encoding of speech signals in the peripheral auditory system. On arrival at Sussex, Alan was presented with Ian Winter as a PhD student in a fait accompli, since Ian had been told he could do a PhD only if Alan obtained the fellowship; fortunately the two of them hit it off! Alan’s stay at Sussex was relatively short: a tenured position became available at the MRC Institute of Hearing Research in Nottingham, and such an opportunity could not be dismissed, despite being only two years into his prestigious fellowship. Nevertheless it was a productive period that included a collaboration with Ian Russell demonstrating that the filtering of hair cell membranes is the major limitation to phase-locking in auditory nerve fibres.
Since arriving at the MRC Institute of Hearing Research (IHR) in 1986, Alan has spent the last 30 years as a Program Leader at the Institute, including periods as Deputy and Interim Director. His research has ranged widely, and he often reinvented himself as successful post docs and students continued elsewhere work they had started or developed at Nottingham, amongst others: Adrian Rees on amplitude modulation, David McAlpine on binaural coding, Jasmin Grimsley and David Green on communication sounds. Alan’s great generosity as a mentor and colleague means there are many hearing labs across the UK and the rest of the world with a first or second generation connection to Alan.
A few of the areas Alan’s laboratory has contributed to over the years are worthy of special mention. From the careful characterization of cells in the ventral cochlear nucleus, he and Ian Winter identified a possible source of the wideband inhibition necessary to explain the processing of spectral notches in the dorsal cochlear nucleus. Using juxtacellular labelling, Alan went on to provide direct evidence that these giant multipolar cells provided profuse projections to all other parts of the cochlear nucleus, including commissural connections to the opposite cochlear nucleus. His work on binaural responses in the inferior colliculus with David McAlpine, Dan Jiang and Trevor Shackleton spurred a major reassessment of the processing of interaural delays, and the neural representation of the binaural masking level difference. More recently his work with Mark Wallace has encompassed the mechanisms of tinnitus and the encoding of animal vocalizations, for which the guinea pig has a rich and varied repertoire.
Most of the data Alan has collected in his career has been hard won from the tip of a microelectrode, but when Mark Haggard, the then Director of IHR, decided that the time was right for the Institute to exploit Nottingham University’s expertise in fMRI, Alan played key role in the new imaging group, which amongst other discoveries, developed sparse imaging to overcome the contaminating effect of scanner noise.
Apart from his intellectual contribution, Alan has always been the most practical of scientists, devising often simple, but ingenious, means of solving technical problems. He was thus a natural choice as the manager of Institute’s very talented mechanical and electronic engineers. Under Alan’s guidance, his team developed a noise cancellation system for fMRI, now taken up commercially, an automated version of the Toy Test, now widely deployed in paediatric audiology clinics, and an otoacoustic emission measurement system using rapid reconstruction of responses to maximum length clicks sequences.
As with his mentoring of colleagues, Alan has always believed that the scientific endeavor depends on a sense of community. He has selflessly given of his time and energy through service on editorial boards, grant awarding bodies, advisory panels and as an examiner. At scientific meetings Alan happily spends hours quizzing PhD students in his incisive manner about their posters, but always leaving them with a deeper understanding of their work and a sense of achievement! Such activities often go unseen, but rank in importance with the extensive list of papers, reviews and other publications that bear his name.
Alan has always been dedicated to his scientific family, throughout his career, but his own family was always his top priority. He and his wife Chris have been together since undergraduate days, and he is immensely proud of his two daughters, his ‘girls’: Evelyne, a clinician, and Alice, a neuroscience PhD.
In 1997 Alan was appointed to an honorary chair in Auditory Neuroscience at the University of Nottingham, and in 2016 his contribution to hearing was recognized by the award of the William and Christine Hartmann Medal for Auditory Neuroscience from the Acoustical Society of America. He is now emeritus professor at the University of Nottingham. It is a great pleasure to see the many facets of Alan’s work celebrated by the 2017 ARO Award of Merit.