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 But I think it to that the special of their contribution was by the to which they in have now their so as to in their in a certain their work illustrates the role of and in the of by of the of Physiology at and the of at is now and It is therefore – and more it is – to which aspects of are most conducive for the of a that is to and – an that is at once to at the it the of and important I would like to consider these three of contributions and of Hubel and Wiesel – the the and the sociological – by the development of only one of their the discovery of one of the central in the of the brain – the fact that the cerebral cortex is organized into of of nerve cells. That of had its at Medical in the as you we the world through our the or and Each is first by and in are then in the cerebral cortex. on the role of the cerebral cortex in in the of Physiology at Medical in about with the work of and was years and an Professor in of Physiology at the Harvard Medical when was to the of Physiology at only was at the of this but had only three was the of whose one would about me that would our and In was that by at to up with and Using techniques by and a by these three that the of was on the of the brain. This was in by the and the fact that not only but each of us within our brain a of our own the to our This discovery that and have a of their on the of their brain a of is not but a of We for the on from the on the it that the for these and were with the techniques by and – techniques that the from of nerve cells – in the was this from the study of it to and the of his who The of perception was up in by Vernon Mountcastle, most in and at Medical Vernon Mountcastle was from a in by who into I have a for but with the of I only now to appreciate what a this for Vernon the years Mountcastle not only over with but also other aspects of In when Mountcastle the of the of also on the of of Mountcastle early on that by the techniques that became in the be to new in the of the sensory system that the techniques by and his This a of new and – to which Mountcastle these in – that the of modern – Mountcastle the of that at the level, is within all of the sensory system a of that was not with First, that single nerve cells to or to to that cells to one were together and were from cells to other The most example of is in the cortex. In a paper in 1957, Mountcastle his discovery that were in the cortex as from the of the brain to the matter Each is the cells in a information from a point on the and from a class of or each of the to a of the cortex, and the are in The of neurons in is therefore the the of the cortex is to different for the small of the Each is an or of neurons that the initial stage in the cortex for sensory into In to the of the discovery of I will now the other contributions that have from Vernon These the of the flow of information from to the cerebral cortex, a between and and a study of the and the of In the years that have since and first the of the the the of Physiology at first and Mountcastle, for in As a result of work, this now to the study of and as a result of the of and Mountcastle, was for many years so outstanding in the study of that it also in and in was still at who had to the early study of up with to that the cortex also of the years in a – I to tonight – was to the at his from to studies of the an to which made fundamental In was joined by Torsten Wiesel, a from Wiesel had in and a in years David Hubel joined Torsten Wiesel in in Hubel his in at where met Vernon then years at on when Mountcastle to his at when Hubel was in the of Hubel therefore a to work in to be a who for more with each in set Wiesel and Hubel to work together in and on his own in a new The of Wiesel and Hubel in in to what one of the most and in each with all of their fundamental contributions have involved simply the of the of their was invited to the of at the Harvard Medical to a small of with the then in his Hubel and Wiesel, the as they were and and the They were joined by a an whose it was to explain it all in the language of Harvard to in a manner – it Hubel and Wiesel from the they at to a rank. This was of only to be from a university that had within years tenure to Nobel and But Harvard its in and his who much to in his was and still is the over a in his own the of at of one and into a of the first in the – the now as as I can about of the at the Harvard Medical School. you a in all that and to and it with one of his of what we now consider modern – the of scientific into one – was of to only have to the Harvard to As with at so in of the world to the still at Hubel and Wiesel to apply techniques to the cortex. had from single cells in the and made the discovery that the cells not simply levels of they between and The most for exciting these cells was not but small of Hubel and Wiesel a similar in the next the at the level of the cortex, Hubel and Wiesel that most cells no to small of To be a had to be a a or a Hubel and Wiesel that these cells not simply and faithfully reproduce the from the by of their the cells were to aspects of the The of the cells are In addition to each cell is to to a of cells when the of the is others when the is still other cells only to small of the is in the cortex with or It is to think that these cells are the early in the perception of and Hubel and Wiesel next that cells with similar of were together into similar to which Mountcastle had in the Hubel and Wiesel have To work is to They are most papers where all the are only to their own work – a they have with they to our into the of First, they and system of in the cortex – the – a system concerned with information from the These to elaborate for they a of morphological techniques to the in The early work of Mountcastle, and of Hubel and Wiesel organization on the of from single cells. had to on the in morphological techniques – a that is upon the of that cells according to one or of their functional – Hubel and Wiesel the and their The they have with these are not only but have given us a new of the organization of the cortex – an made only by the of they have made us realize that we are to the structural organization of the brain and its by we have so understanding of the biological of most of mental Wiesel and Hubel have these studies of the organization to the of sensory on They that a such as the of a for a to and the its to the of nerve cells in the cortex. By similar in an produce no on In a of Wiesel and Hubel that in the organization of their the for each are equal in the that for the are much to that from the The scientific and philosophical of this work is is that sensory in early can the of the cortex. As Hubel in his of psychologists and the importance of early on patterns – it be that of social or the of other early in to a or of connections in of the brain. The were first by Mountcastle in the sensory system and their functional and their by were by Hubel and Wiesel in the and other of functional have in other of the cortex and in other of the brain. these are to other of as as to It is clear that we are with one of the principles in the organization of the cortex and the for work on the brain. In I would like to to the of social I have between the social and of and 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 tonight to the honour that this on years I that this will in the honour the of Mountcastle, Hubel and Wiesel. By this I not to that these and – Mountcastle, Hubel and Wiesel – are from one But I to that they that are in other This is a that now by a of most by our colleagues at and They have that can their colleagues the importance of on they also and and make to their important of of these are in with the of a own of his own for What is in the three we honour tonight and in their scientific are that are much much more is the to in their own and to the of their The second is the to around them an exciting made up of an where important science is because the is around one or more central The the additional that it the from of and scientific of these I from the of these to a of what is important in their with an of the and of They science as a in an – a of It is for these that the Columbia takes such special pleasure in Mountcastle, Hubel and Wiesel. their contribution to the biology of the brain – the most and profound of our generation – at once scientific and at their In we are to the in and the of that so many of us to Their contribution us not only an into the brain and into but it also us of how the in and can be and 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 and I will my comments in three First, I will Torsten's scientific and his of I will to a of Torsten Wiesel's and work on a fundamental that between different of and I going to to Torsten's I with a of Torsten's and Torsten was in in the of a who served as of large of the of was a on the so Torsten most of his early years on the of a mental by mental In it was this with the that to so as of Rockefeller in the of David I of the by the who to of the Medical at once I how you take on an and give up which you so given up still that my have In at Torsten the to the the on the brain by the of and by who on on to Torsten's and his of Torsten first in and in But by the of that that the not to the biology of the brain and in joined of at the as an in of to to the United to work on the as a with at the at That was a point in Torsten's as you all to be an enormous on In 1957, Torsten and a on at the that I also that David Hubel a on of the of the of cells from the cortex – – in Hubel that when moved his in one – to the – in of the one of the cells in the striate cortex would the other was but moved his in the the cells that had in the to the were the cell that was now This of the was the first to be in the that by David, it was clear to Torsten and that the striate cortex was to be much more interesting the the was how to David to the This was no David had an in the Physiology at from Vernon But as would have when David at in the in Physiology that were for were not where was David to In to the invited David to the and that David with Torsten and David to from from and to a study of the of The is to all of us. the – years of extraordinary David and Torsten more up the study of the cortex, they the for what was to in all sensory They the and the organization of and they a organization for and and their work in the of in the during a period of These physiological contributions were made all the more by and this of work stands as one of the biological of the century. But it was not simply their work but also the that was so a of and I give but of their Torsten and David me to their paper on in of with it to the Journal of I it to be and other that since the findings were so they it on more one one They and a second I that of the in the to their own work seemed a They also and the to other the period of the Torsten was by most as a who with scientific But of us, David, who in Torsten of us Torsten as and not to David in all that That to as the of the Harvard of first one turn to David to But by it was clear that this was not to It so that a years David had the for the tenure as a in the of Harvard University, a going to In David had the of the Physiology his by the of the most in the that the in the and that they a new In to this one David from other of the and made no to his to between his with his his in and on the David to up the to for and tenure in Physiology for of his accomplished this one in his first on the David that had for Physiology at therefore the and to the which had when for a who on the not to mind. on the other had all concerned about about and and on the of with and This us new Torsten was Torsten an outstanding and a as of the that to be a of that David and Torsten to their David to with on and on the of the Torsten to with on how the cells of a are This to the unanticipated discovery of the connections in the system – a contribution to the of the striate cortex that now continues to In 1983 Torsten an from Rockefeller to a new years in when David Torsten as of the and now a years of was by the Trustees to as of Rockefeller University. the next years an to the of the by them with his of the to on this his of his and and met with the and in groups and to outstanding new I only and in to give you a of his and his the of the university around and in with the university and in superb that Torsten to Rockefeller as of the and for and In this to his and in by who is tonight, to to Rockefeller as the new in in a Torsten from the of Rockefeller and the of New to for science the Torsten the of the that for in the United their with to set up their own as of the of of the of which who have for their to Torsten is and President of – the – and since the of the That was Torsten it by exciting new and by the of in outstanding the In addition to his scientific accomplishments, his of and Torsten is a with of the as you will is an with and of you may the at the where Torsten's was on view, a not by to a single of or period in but by the of of its in the of his years and all that is still of can we to can we his contributions the of thought and in the context of can we his so that it can as a for our what of does Torsten Torsten is a by and a by is therefore to turn for to the and – to and only and are because only they have for of thought as as of I turn to and of into and The is the the is Torsten does not into one or the other of these for of In many is But who a of or on the is also a to the a more one that the and and the quite The when a of to his and then the more of the or a Torsten to the other is an who for more for new of not simply for but for the and in the for the of In his for new in the of Torsten is much like of an who What most is not simply but and on of to his with which on the I on a of then that be my you can me with into a that be the for like a modern Torsten's a of and development in his for to science first in his own in his with David and then in the of his at as of his university at and now in his of the scientific the But on a where are is the of it will not you to that Torsten's does not Let me give you one example. Torsten and I in with the of an who had the and to where the of his in a The of the In 1999 Torsten and I in his at that where to us with in his – and I to – that I can have a at a in New Torsten and I to each other We each thought that between the of us and our friends we We an committee and at the by first the and then our way and thought But we were and one thought that his work would produce a in we to the they But they also had no in is they In we have of his But one is not for the you – are to Torsten's Let me these by is this in the is us the next for Torsten is to his in science and I we Torsten to at is to be from his of the Torsten as of the will First, it will give Torsten a new from which to his about and his of with his in the biology of and the of the experience. In it will give all of us opportunity at a this is a of no of Torsten is a of who during the years Torsten's and all our by with us insights into and To the is the – – that us all to and a central of all of our me a to Torsten on his birthday. you are a and of We who you you as a an extraordinary scientist, and a superb with a of and since at you think and as a we you and years more years of and

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