An introduction to the work of David Hubel and Torsten Wiesel

It is with enormous pleasure that I add my voice to that of others of my generation in celebrating the semicentenary of the 1959 publication of Hubel and Wiesel's first paper in The Journal of Physiology entitled: 'Receptive fields of single neurons in the cat's striate cortex' (Hubel & Wiesel, 1959). This paper set the stage for the continuous flow of outstanding papers that emerged over the next twenty-odd years from the Hubel and Wiesel collaboration. Their work and that of Vernon Mountcastle opened up the modern study of the cerebral cortex. As a result of their extraordinary accomplishments, Hubel and Wiesel received the Gross Horwitz Prize together with Vernon Mountcastle in 1975, and the Nobel Prize in Physiology or Medicine in 1981 together with Roger Sperry. It was on the occasion of the Gross Horwitz Prize, on whose committee I served, that I was invited to introduce Mountcastle, Hubel and Wiesel. My initial comments in that introduction were in fact directed toward an excellent scientist, a member of our Prize committee, who commented during our deliberations that Mountcastle, Hubel and Wiesel seemed to represent superb science, but their work had limited biological generality. To which I replied: 'You are right, it does not apply to the kidney or the spleen. It is much more restricted. It only helps to explain the workings of the mind.' Hubel and Wiesel's names are enshrined together in the Pantheon of Creative Collaborations in Biological Sciences, much like Hodgkin and Huxley, Watson and Crick, and Brown and Goldstein. In each case, equal partners joined forces bringing unique skills to their collaboration to produce a new level of science and a new family of insights. I first met Torsten and David in 1957, and we became friends in the period 1960–1965 when I overlapped with them at the Harvard Medical School. That friendship continues to this day, as does my admiration for the work, scientific and administrative, that they have accomplished since going their own directions in the early 1980s. In 1983 Torsten moved to New York, first as Professor and then in 1999 as President of Rockefeller University. I served on the Board of Trustees of the Rockefeller during much of Torsten's tenure and this provided me with the additional opportunity to toast his 80th birthday. What follows in the ensuing set of papers in this issue is an outpouring of affection, respect, and gratitude for Torsten and David, for who they are, for what they have given us, and for setting the tone of our science for my generation in the United States. President McGill, Vice President Marks, colleagues of Columbia University, honoured guests. Even the most important contributions, such as the work by Vernon Mountcastle, David Hubel and Torsten Wiesel that we honour tonight, have in addition to their obvious strengths, certain clear weaknesses. I will discuss the strengths in a moment, but I think it important at the outset to consider its weaknesses. Many contributions in biology inform us about general principles – principles that are important for understanding all the cells of the body. The contribution of Mountcastle, Hubel and Wiesel is concerned with only one class of cells – the cells of the brain. Their findings are therefore somewhat parochial. They are only important for understanding the mind. Despite that limitation, the contributions of Mountcastle, Hubel and Wiesel are nonetheless rich in meaning and have significance for many levels of thought. Their work will, I would suggest, be discussed by historians of science from three quite different vantage points. First, it will be discussed from a purely scientific point of view, as a central contribution to neurobiology; second, from a broader philosophical point of view, as an enhancement of our understanding of mental processes; and third, from a sociological point of view, as an example of the importance of scientific lineage and of small group interactions at large universities. First and foremost, as a scientific contribution to neurobiology, the work of Mountcastle, Hubel and Wiesel stands as the most fundamental advance in our understanding of the organization of the brain since the work of Ramón Y Cajal at the turn of the century. By applying morphological techniques to the cerebral cortex – the highest and most elaborate part of the brain – Cajal revealed a hitherto unanticipated precision of the interconnections between populations of individual nerve cells. Using modern cell physiological techniques, Mountcastle, Hubel and Wiesel have revealed aspects of the functional significance for perception of these patterns of interconnections between nerve cells. They have shown us that the connections filter and transform sensory information on the way to and within the cortex, that the cortex is organized into functional compartments or modules, and that this organization can be altered by experience. By any scientific criteria, these contributions are of the highest rank. But on a second level, the work of Mountcastle, Hubel and Wiesel takes on greater significance because it contributes to our understanding of mental processes, a contribution with profound physiological implications. We appreciate important science because it tells us something new and exciting about the world around us. What is at once so special and so parochial about the work that we honour tonight is that it tells us something new and exciting about the world within us, about ourselves. Let me give you an example. We have the feeling that when we interact with each other – when I speak to you and you listen to me – that we are directly experiencing one another. Hubel, Wiesel and Mountcastle have made us realize that this is an illusion, a perceptual illusion. The brain does not simply take the raw data that it receives through the senses and reproduce it faithfully in the brain. Rather, each sensory system first analyses and decomposes, and then restructures the incoming raw sensory information according to its own built-in connections and rules. These insights are not only remarkable; they are also timely. Hints that similar processes may be involved in the development of language and thought are now emerging from the studies of structural psychologists such as Chomsky and Piaget. On still a third level, the work of Mountcastle, Hubel and Wiesel is interesting because it illustrates in a unique manner the role of social context upon discovery and how small groups in a university can shape social contexts so as to make them conducive to creativity. Mountcastle, Hubel and Wiesel are exceptionally creative, bright and energetic. Each would have made an important mark on science no matter where he worked. But I think it fair to say that the special nature of their contribution was fostered by the particular collegial environment to which they belonged. They, in turn, have now restructured their environment anew so as to foster creative activity in their younger colleagues. Thus, in a certain sense, their work illustrates the role of intellectual continuity and intellectual renewal in the achievement of excellence by segments of two American universities, the Department of Physiology at Johns Hopkins and the Department of Neurobiology at Harvard. Academic life is now often challenged, beleaguered, and fragmented. It is therefore inspiring – and more important, it is instructive – to learn which aspects of academic life are most conducive for the establishment of a powerful intellectual environment that is resistant both to external and administrative pressures – an environment that is at once sensitive to historical perspective while at the same time it encourages the emergence of novel and important ideas. I would like to consider these three implications of Mountcastle's contributions and those of Hubel and Wiesel – the scientific, the philosophical, and the sociological – by tracing the development of only one aspect of their contribution: the discovery of one of the central ideas in the functioning of the brain – the fact that the cerebral cortex is organized into computational modules consisting of vertical columns of nerve cells. That particular strand of research had its origins at Johns Hopkins Medical School in the mid-1930s. Now, as you know, we experience the outside world through our five senses: touch-pressure (and the related skin or somatic sensation), sight, hearing, taste, and smell. Each sensation is first analysed by appropriate receptors and coded in lower relay stages. Most sensations are then elaborated in the cerebral cortex. Modern research on the role of the cerebral cortex in somatic sensation began in the Department of Physiology at Johns Hopkins Medical School in about 1936, with the work of Philip Bard and Clinton Woolsey. Philip Bard was 34 years old and an Assistant Professor in Walter B. Cannon's Department of Physiology at the Harvard Medical School when he was called to chair the Department of Physiology at Hopkins. Not only was he extraordinarily young at the time of this appointment but he had published only three original papers. He was the sort of person whose future one would worry about nowadays. Dean Tapley assures me that he would never pass our Appointments and Promotions Committee. In fact, Dean Tapley was surprised that he slipped by even at Hopkins. Soon after coming to Hopkins, Bard teamed up with two even younger colleagues, Clinton Woolsey and Wade Marshall. Using gross electrophysiological recording techniques developed by Marshall and a strategy developed by Woolsey, these three young men discovered that the body surface of monkeys was systematically represented on the surface of the brain. This was soon confirmed in humans by the Canadian neurosurgeon Wilder Penfield and established the fact that not only monkeys but each of us has within our brain a naked representation of our own body, the closest thing to our true self-image. This remarkable discovery that animals and man have a representation of their body on the surface of their brain raised a number of conceptual problems. Somatic sensation is not unitary but a composite of several distinct sensations called submodalities. We can, for example, readily distinguish the pressure on deep tissue from the light touch on the skin. However, it appeared that the maps for these deep and superficial submodalities were completely congruent. Clearly with the relatively gross techniques used by Marshall, Woolsey and Bard – techniques that averaged the responses from thousands of nerve cells – some critical dimension in the map was overlooked. After making this major contribution, Bard withdrew from the study of somatic sensation, leaving it to Woolsey and the younger members of his department who came along later. The particular question of submodality perception was picked up in 1948 by Vernon Mountcastle, Bard's most gifted student. Born in Virginia and educated at Johns Hopkins Medical School, Vernon Mountcastle was dissuaded from a career in Neurosurgery by Bard, who enticed him into Physiology. I have always considered Mountcastle's decision a great gain for Physiology, but having recently become familiar with the economics of neurosurgery, I am only now beginning to appreciate what a loss this decision has meant for Vernon Mountcastle. Over the years Mountcastle not only took over Bard's fascination with skin sensation but also other aspects of Bard's mantle. In 1946 when Bard retired Mountcastle assumed the directorship of the Department of Physiology. He also took on the editorship of Bard's distinguished Textbook of Physiology. Mountcastle realized early on that by using the cellular techniques that became available in the late 1940s he might be able to detect new dimensions in the map of the somatic sensory system that eluded the gross recording techniques used by Bard and his colleagues. This task required a number of major technical innovations including new microelectrodes and precise quantifiable natural stimuli – innovations to which Mountcastle contributed importantly. With these tools in hand – tools that formed the basis of modern cortical physiology – Mountcastle addressed the question of submodality specificity. He found that at the cellular level, there is within all areas of the somatic sensory system a segregation of submodalities that was not resolved with gross recordings. First, he found that single nerve cells respond specifically either to superficial touch stimuli or to deep pressure stimuli, almost never to both. Second, he found that cells responding to one submodality were located together and were segregated from cells responding to other submodalities. The most fascinating example of segregation is found in the cortex. In a classical paper published in 1957, Mountcastle described his remarkable discovery that submodalities were distributed in the cortex as vertical columns running from the surface of the brain to the white matter below it. Each column is submodality-specific. All the cells in a column receive information from a particular point on the skin and from a particular class of receptors, either superficial or deep. Thus, each region of the skin projects to a particular area of the cortex, and the separate receptor classes are distributed in adjacent columns. The distribution of neurons in columns is therefore the mechanism whereby the depth of the cortex is used to handle different functions for the same small region of the bodily map. Each column is an integrating unit, or logical module, comprising thousands of neurons that form the initial stage in the cortex for elaborating sensory experience into consciousness. In order to follow the history of the discovery of columnar organization, I will now describe the numerous other contributions that have subsequently come from Vernon Mountcastle. These include the analysis of the flow of information from skin to the cerebral cortex, a correlation between cellular responses and perception, and recently a study of the mechanisms underlying attention and the control of purposeful movements. In the 40 years that have passed since Bard, Woolsey and Marshall first mapped the representation of the body surface onto the brain, the Department of Physiology at Johns Hopkins, first under Bard and subsequently under Mountcastle, has been preeminent for research training in skin senses. As a result of Mountcastle's recent work, this preeminence has now been extended to the study of attention and behaviour. Indeed as a result of the leadership of Bard, Woolsey and Mountcastle, Hopkins was for many years so outstanding in the study of sensation that it also dominated research in hearing and in vision. For example, while he was still at Hopkins, Wade Marshall, who had already contributed importantly to the early study of skin sensation, teamed up with Samuel Talbot to demonstrate that the cortex also contains detailed maps of the retina. About 10 years later, in 1948, a young man, Stephen Kuffler – whom I am delighted to see here tonight – was recruited to the Wilmer Eye Institute at Hopkins. Upon arriving, Kuffler turned his attention from synaptic transmission to cellular studies of the retina, an area to which he immediately made fundamental contributions. In 1955 Kuffler was joined by Torsten Wiesel, a young postdoctoral fellow from Sweden. Wiesel had experience in child psychiatry and a particular interest in vision. Three years later, David Hubel joined Torsten Wiesel in Kuffler's laboratory. Born in Canada, Hubel took his residency in Neurology at Hopkins where he met Vernon Mountcastle. He then spent two years at Walter Reed working on vision when Mountcastle recruited him back to his laboratory at Hopkins. However, when Hubel arrived Mountcastle's laboratory was in the process of being renovated. Hubel therefore accepted a temporary invitation to work in Kuffler's lab. He turned out to be a guest who stayed for more than dinner. Confronted with two gifted young investigators each interested in vision, Kuffler set Wiesel and Hubel to work together in vision and went off on his own in a new direction. The largely accidental meeting of Wiesel and Hubel in Kuffler's laboratory in 1958 gave rise to what has been one of the most remarkable, sustained and productive collaborations in contemporary science. Although each has occasionally worked with another collaborator, almost all of their fundamental contributions have involved simply the two of them. Soon after the beginning of their collaboration, Kuffler was invited to join the Department of Pharmacology at the Harvard Medical School to head a small laboratory of Neurophysiology. A true pater familias, Kuffler took with him the four young faculty people then working independently in his laboratory. Hubel and Wiesel, the two 'brain boys' as they were called, and Furshpan and Potter, the two 'membrane boys.' They were soon joined by a fifth colleague, an enzymologist, Ed Kravitz, whose function it presumably was to explain it all in the universal language of biochemistry. Harvard responded to Kuffler's brilliant recruitment effort in a typical manner – it immediately demoted Hubel and Wiesel from the Assistant Professorships they held at Johns Hopkins to a non-professional rank. This was of course only to be expected from a university that had within recent years successfully denied tenure to two Nobel laureates, Georg Von Bekesey and Fritz Lipmann. But Harvard found its match in Kuffler and his boys. Unlike Bard who devoted much time to in his Kuffler was and still is the over a 10 in his own Kuffler turned the laboratory of at consisting of one and five postdoctoral into a Department of the first in the – A intellectual the department now as as I can about of the available at the Harvard Medical School. you a in all that some and young coming out to and it with one of his of what we now consider modern – the of scientific into one – was out of to Harvard. only have to the Harvard department to see As with Bard's department at Hopkins, so in Department of the world to the still at Hopkins, Hubel and Wiesel began to apply cellular techniques to the cortex. Kuffler had from single cells in the and made the discovery that the cells not simply levels of they between light and The most for exciting these cells was not light but small of Hubel and Wiesel found a similar in the next relay the However, at the level of the cortex, Hubel and Wiesel found that most cells no responded to small of To be a had to be a a or a Hubel and Wiesel found that these cortical cells not simply and faithfully reproduce the from the by of their the cortical cells were able to aspects of the The of the cortical cells are In addition to each cell is coded to respond to a of some cells respond when the of the is running others when the is still other cells respond only to small of the is represented in the cortex with or It is to think that these cells are the early in the perception of form and Hubel and Wiesel next found that cells with similar of were together into columns similar to those which Mountcastle had found in the Hubel and Wiesel have original To work is to them. They are most papers where all the are only to their own work – a they have sustained with Upon they to our into the nature of columnar First, they and found another completely system of columns in the cortex – the columns – a system concerned with information from the two These columns to elaborate vision for depth Second, they a of morphological techniques to the columns in The early work of Mountcastle, and of Hubel and Wiesel described columnar organization on the basis of from single cells. had to columnar However, on the in morphological techniques – a that is upon the of that cells according to one or another aspect of their functional activity – Hubel and Wiesel independently both the columns and their columns. The they have with these are not only but have given us a completely new of the organization of the cortex – an made only by the of Thus, they have made us realize that we are beginning to the structural organization of the brain and its by we have so understanding of the biological basis of most of mental Wiesel and Hubel have used these studies of the columnar organization to the of sensory on They found that a such as the of a for a to and the its to control the of nerve cells in the cortex. By similar in an produce no on vision. In a brilliant of Wiesel and Hubel found that in monkeys the organization of their columns. the columns for each are equal in After the columns that receive for the are much to those that receive from the The scientific and philosophical of this work is is that sensory in early life can the of the cortex. As Hubel out in his of psychologists and both the importance of early experience on patterns – it be that of social or the of other early in life to a or of connections in some of the brain. The columns were first discovered by Mountcastle in the somatic sensory system and their functional and their by experience were analysed by Hubel and Wiesel in the and other of functional modules have been in other areas of the cortex and in other of the brain. these modules are related to other of sensation as as to It is clear that we are here with one of the principles in the organization of the cortex and the for future work on the brain. In I would like to to the question of social I have already between the social and of Bard and Kuffler and the of scientific in Mountcastle, Hubel and Wiesel, who in turn have now where others can be and much as Mountcastle, Wiesel and Hubel here tonight to receive the same honour that this on Kuffler years I am that this will in the future honour the intellectual of Mountcastle, Hubel and Wiesel. By this I not to that these five and people – Bard, Mountcastle, Hubel and Wiesel – are from one But I to that they two that are often in gifted often other This is a question that has now been by a number of most by our colleagues at and They have out that can their colleagues the importance of working on they also and and make available to their important of All of these are in with the of a own of his own for What is in the three men whom we honour tonight and in their scientific are two that are much much more is the to in their own and to the of their gifted young colleagues. The second is the to around them an exciting environment made up of gifted an environment where important science is because the environment is around one or more central ideas. The has the additional that it the environment from of academic and scientific of these I from the of these to a remarkable of what is important in their with an almost of the and intellectual activity of science. They science as a in an intellectual – a of ideas. It is for these that the Columbia takes such special pleasure in Mountcastle, Hubel and Wiesel. For their contribution to the biology of the brain – the most remarkable and profound of our generation – at once scientific and at their In we are attention to the in and the of that so many of us to academic Their contribution us not only an into the brain and into but it also us of how the in and academic can be and sustained over to Torsten Wiesel's 80th Rockefeller University, I have the pleasure to as the for Torsten's 80th Let me introduce with an of Torsten's life and I will my comments in three First, I will describe Torsten's scientific and his of Second, I will to a of Torsten Wiesel's life and work on a fundamental that between different classes of and I am going to some to Torsten's I with a of Torsten's life and Torsten was in in the of a who served as of two large of the of being was a on the so Torsten spent most of his early years on the of a mental by mental In it was this experience with the that him to function so as of Rockefeller in the of David I am here of the by the great American who to become Dean of the Medical School at A once I how you take on an administrative and give up which you always so given up still see that my have In at Torsten the Institute to the he the on the brain by the of and by who worked on went on to become Torsten's great and his of Torsten worked first in and in child But by the of that he that the available not he turned to the biology of the brain and in joined department of at the Institute as an later, in of him he might to to the United to work on the as a postdoctoral fellow with

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