Second Part

0:09:07 Collaborated with John Griffith and published at least one paper together; taking forward how the two neurones interacted electrically because if you could understand that you would begin to understand how networks and nerve cells interact; no mathematics available to study it so he had to design the mathematical tools for studying it; there were no computers then; these were electrical pulses and you couldn't analyse the time elapsing between each pulse so had to film it and then the film was analyzed by a woman who was also measuring film of pulsars at that time; took months for the measuring the intervals; became a fellow in 1962 and we spent much time in my room here, in his room, and my laboratory, and formed a great friendship; and going beyond our own work together he was a very original scientist and I think would have got a Nobel Prize

3:11:19 Working on attention where there was always two stimuli I had found some nerve cells in the visual part of the brain that responded not only to visual but also tactile stimulation; there were very few of them and the responses were quite weak but visual physiologists did not like this; there was a group in Germany let by Baumgärtner who had found similar things for auditory input, neurones in the visual cortex responding; Dave Hubel and Vernon Mountcastle really didn't like that and they were very influential figures in the US at the time; coming from it from attention and finding nerve cells in the visual part of the brain responding to a tactile stimulus whether the animal was attending or not; there was a basic level of interaction going on which was independent; Peter Venables and I did an experiment in humans with similar results; not only did you get switching between modalities, like audio and visual, but they actually interfere with each other at quite a low level of the cerebral cortex; in 1963 Charles Shute and Peter Lewis in the Department of Anatomy where I was working had shown that there were pathways that had never been discovered that were actually not visual but they went into the visual pathways from other parts of the brain; I thought this was the key; there was a lot of evidence that this extraordinary high level of brain function, attention switching, for example, and what one would call clinically, consciousness, is controlled deep down in the very primitive parts of the brain called the brain stem; I wanted to spend some time putting electrodes into the brain stem and seeing whether there were lots of nerve cells there that responded to more than one sensory modality as that is what I would predict; I had written a chapter on some neurocorrelates of perception and gave it to Horace Barlow to read; he liked it and when he could not go immediately to take up a Chair at Berkeley he suggested I go for a term teaching on the physiology of vision; when I arrived I was told I would have to work with Dick Hill as there was not enough equipment for me to work by myself on brain stem research; working on anaesthetised rabbits we put an electrode on the brain stem and came across nerve cells that responded beautifully to visual stimulus; I remember one day dropping a pin on the floor while we were monitoring the activity of the nerve cells and getting an immediate response; further research found lots of nerve cells that responded to sound and touch as well as to visual input; however, when we came to look at where the electrode had been it was not in the core of the brain stem but in the roof; the roof of the brain stem is the most ancient visual centre in vertebrates called the optic tectum; we looked through the literature and found nothing published and so the world didn't believe us; remember coming back to Cambridge and telling Dixon Boyd who asked for proof which I gave him; then a lot of people started working on this; I then also found that neurones in that part of the brain responded differently to infrequent and frequent stimuli; for the former there was always a response but it declined with frequency, showing habituation; this had been observed in behaviour and I had heard about from Bill Thorpe at the Thorpe-Zangwill club; in animals, they give a response to a novel stimulus but if not rewarded with food, for example, they don't respond; I suddenly realized that we were dealing with a nerve neural counterpart to behavioural habituation so I saw my life retrospectively opening up in two directions; saw I must follow this elementary form of learning as here was a clue to understanding what the neural basis of learning might be; the other was sensory interaction

14:19:03 In 1967-68 Dick Hill wrote to me and said he would like to come and spend a summer in Cambridge; I had read about the vestibular stimuli in the visual cortex and I'd already been to Makerere to work with Hugh Fraser-Rowell where I'd worked on locust brains; came across the work of a Dutch man, Wiersma, whom I met, who was recording from the optic nerve of crayfish and found that the nerve fibre was looking at the field of vision above the horizon; every time a visual stimulus was below the horizon there was no response; found that if he tilted the animal the nerve cell went on responding in the same area and didn't tilt; suggested to Dick that we see if something like this existed in the visual cortex of a mammal; we found this was so and we sent off a letter to 'Nature' which was published; it caused intense controversy; the following year I was joined by Gerry Steckler and we spent a year working on it once again and we published the full paper in 1972; could never understand the vitriol that was heaped on me in meetings in the visual forum and it took many years for other people to confirm; it was a difficult experiment to perform and also took courage to do it in the States; there was a group there which published a paper in 1981 saying in a sense that I was wrong in underestimating the number of these as there were more, but it still went dead and is a lacuna in the scientific system that this still isn't quite respectable

19:07:00 [PB: I will tell you why. It is because people trained in a certain kind of science do an experiment where you vary one thing and keep everything else constant, and if you get an effect that is the cause. People still think in this very linear way and if someone like you comes along and says it is not the cause but one of the causes, people hate this systems approach to biology. I have encountered many physiologists who hate this]

19:42:13 I remember I had some interaction with Colin Blakemore about this and he said 'The trouble is we physiologists don't know how to handle parallel systems and therefore it's not popular'; I don't know if that's it or the explanation for it; I read a review by someone in Stirling University about this kind of influence and she had never heard of this earlier work; I wrote to her and she said she was pleased to know about it but was facing terrible trouble getting her work published; I felt then that the quicker I get out of the field of visual physiology the happier I should be in another field

20:40:17 You and I had already started in 1965 before I went to Makerere; when I came back we began to think of ways of dealing with that problem; on the habituation front, I went on to study it in the insect brain and work out its characteristics and a year after Dick Hill and I published in 'Nature' showing these lovely habituating curves of neurones and showing that these nerve cells had so many of the properties of behaviour; this paper was 1964 and in 1966 a paper appeared from Tauk and Brunaire in France showing that you could get this kind of response decrement and showing many of the properties of behavioural habituation across a single junction between nerve cells and the next neurone; the trouble was they didn't know it was any one cell to the next nerve cell and it could have been many nerve cells interacting and they hadn't got any system that did it; I thought that I needed a system to do that; I knew there was one system where there was unequivocally the case that there was one nerve cell sending a signal to another nerve cell and that was the squid giant ganglion; thought that is the place if I can show this habituation then I know its occurring in the synapse and you have got to look within the synapse to understand it, you don't need to look much further; it happened and I remember giving a presentation of these results to a conference Robert Hinde and I organised in 1969 on short term change and neural activity; Eric Kandel was there and I showed these slides of this lovely waning of a response and it had to be on the pre-synaptic side not on the post-synaptic side and I said I thought that calcium was probably involved; Kandel showed later on that calcium was involved and it did happen at the synaptic junction but he never referred to that work, except when I met him after he got the Nobel Prize referring to my great paper on the squid stellate ganglion

24:26:23 I had been in the Department of Anatomy here in Cambridge for eighteen years and I really wasn't interested in running anything but just doing my research; I'd been pulled onto the research committee by Robert Hinde and began to see that the way the University was running was important for my research too so I began to look beyond it; I was then teaching six hours a week and had been for years here in King's; I had been appointed Reader; was not very impressed by the way Professor Harrison ran the Department; used to go on family holidays to Cornwall and Devon, driving through Bristol which I loved; when the Chair of Anatomy came up at Bristol I applied; I was extremely lucky in that Barry Cross had not been a traditional anatomist and had done an extremely good job in organising that department; he had allowed a young junior lecturer to revise the teaching of topographical anatomy so it was going extremely well when I arrived; the teaching looked good so I was able to concentrate on research; I always had a love of developmental biology and embryology; the field of molecular developmental biology was just beginning; I had a vacancy in the department and I managed to get a very bright young man, John Knowland who was working with Fred Sanger at the MRC Laboratories here in Cambridge; he came and began molecular developmental biology; sadly he only stayed about as long as I did in Bristol; felt that if I am going to populate a department of anatomy with scientists who had no medical training they ought to know a bit about anatomy; needed a clinical input and thought I could get John Knowland to get familiar with these things; too big a job and he opted out; I brought in quite a lot of neuroscientists to the department; I encouraged the development of biomechanics with research by Lance Lanyon; became quite strong in oral biology with Bernie Moxham; really a matter of appointing the right people to jobs and encouraging them; also, I as head of department did research; felt that no head of a science department should just be a manager; there was some discretionary money that I could use as head of department; one thing it brought me was the opportunity to bind the people in the department together so I used to spent some of that money on a party once or twice a term for all the staff, including technical and secretarial, and their partners, which was appreciated; two of the people I appointed, one Malcolm Brown who became Fellow of the Royal Society and remain in Bristol; Bernie Moxham went to become head of department at Cardiff; both those two departments got five stars in the Research Assessment Exercise, the only two to do so

33:52:22 Had a telephone conversation with Robert Hinde about applying for the chair in the Department of Zoology in Cambridge; thought it was absurd as I was not a zoologist and had no real interest in natural history and knowledge of biology was limited; gave all the reasons why I shouldn't apply; he countered with my work on squid, locust, all sorts of animals and even humans; I had great esteem for the department with which I had had links in the past; thought there were brilliant people there; in the end agreed that Robert could put my name forward and finally I was offered the Chair and it was far from clear that I should take it; I was well known in the department for my work with neuroscientists there; my research cut across boundaries and the one thing I saw as positive in going to zoology was the fights I continually had to mount in the medical faculty and the science faculty in Bristol about intellectual territory; to me they were senseless arguments as I didn't see the world that way and being forced to teach student that way was anathema; I then got the reputation in Bristol of wanting to take over the physiology department which was the last thing on my mind; the attraction of zoology in my discussions with you and Robert was the awareness that it didn't matter whether you worked on behaviour or cells it was all one; zoology cut across everything to do with animals which included humans; at that time the department was not in good shape so I knew it would be an uphill task while everything in Bristol was simply wonderful; remember going away with the family and we discussed it endlessly; in the end they left it to me to decide; on the basis of this challenge I suddenly thought what an opportunity to abandon all these constraints and step into the wide world of real biology