My interest in biology seems to have come from a childish desire to collect, which in my case took the form of squashing half the British flora. By my mid-teens the immobility of plants began to bore me, and I switched to zoology when I went to Cambridge. In those days (ca. 1960) the sex ratio in Cambridge was so dire as to preclude normal life, but the resulting monastic existence did mean I got a decent degree. Among my teachers there I would single out Carl Pantin, the professor of Zoology. His course on the Invertebrates – a mixture of taxonomy, anatomy, physiology, ecology and behaviour – left me with a life-long enthusiasm for the seafood end of the animal kingdom. It helped that there was a good museum in the department, where I developed a kind of nineteenth century naturalist’s fascination for the more bizarre products of natural selection.

After Cambridge I went to do a PhD at University College London, where I worked on the eyes of scallops and finally enjoyed some female company. I had persuaded my supervisor, John Gray (later to become secretary of the Medical Research Council and ultimately Sir John) that scallops had a nice ganglion that all these eyes fed into, and a clear – if dull- behaviour (shutting), and that this ganglion clearly needed its input/output relationships sorting out. In fact I didn’t get past the eye. Quite early on I made a chance observation that when one looks into a scallop’s eye, one sees an inverted image of oneself. At first I thought, fine, all eyes have images, but then in eyes like ours the image, as viewed through the lens, isn’t in the eye itself, but back where the object was – somewhere near infinity. Scallops’ eyes turned out to be unique; the image was indeed not formed by a lens, but by the reflecting mirror that lined the back. This was a great stroke of luck. A nicely identifiable discovery that made me think “hey, I can do this science stuff”, and I haven’t really looked back.

After London, to Berkeley, on a Miller fellowship sponsored by Gerald Westheimer, who taught me about eye movements and magic optical arrangements such as the Maxwellian view, and Dick Eakin, who by then had looked at the ultrastructure of the eyes of just about everything. As the Chinese curse has it: “May you live in interesting times”. Berkeley in the late 60’s was certainly interesting. We had Eldridge Cleaver and the black panthers, people’s park, the Alameda County Sherriff’s deputies and finally the National Guard, with the Vietnam war in the background as the engine for much of the unrest. At the same time there was a wonderful gathering of eye people. Apart from my sponsors, Horace Barlow, Jack Pettigrew, Colin Blakemore and Bill Levick were there, as were Barry Frost, Ron Hoy and several other human physiologists. After one or two false starts I decided to work on jumping spiders, inspired by some remarkable work on their vision by Heinrich Homann in the 1920’s and set about collecting them by bashing branches of trees into an umbrella earning me a reputation as a harmless lunatic. Jumping spiders have movable principal eyes, and my most interesting discovery from that era was that when a spider sees a small moving target the eyes go into a very repeatable scanning routine involving both side to side and torsional movements of the retina. At least one function of this scanning routine was suggested already; it permitted the spider to make the distinction between objects that were edible and others that it would be more appropriate to mate. Oscar Drees had shown in the 1950’s that spiders really like legs: enough legs at the right angle and a male jumping spider will start his highland-fling-like mating dance.

I returned to England in 1971, to a temporary post in the University of Sussex, a new and attractive campus that had managed to get most  of its buildings completed before the money ran out in the mid 1970’s. My first collaborations were with Tom Collett, on the ways flies chase each other, based on the observation that flies under lampshades seemed to go in for small-scale Snoopy and the Red Baron dog-fights, and if this was the case there would have to be appropriate control systems involved (I had earlier read some of Horst Mittelstaedt’s papers and become quite interested in control theory). Switching from houseflies to hoverflies Tom and I were particularly impressed by the similarities of some hoverfly behaviours to human eye movements – the saccades, tracking, optokinetic stabilizing, etc – except of course that these were brought about by flight manoeuvres not eye muscles. The thought that much behaviour, especially in insects, is to be explained as subserving the active uptake of visual information, is an idea that has remained with me ever since. Tom and I also looked at visual memory in hoverflies, and he developed ideas that were to lead to the celebrated “snapshot model” of bee landmark memory that he and Brian Cartwright developed in 1982.

Meanwhile, I had deserted behaviour, temporarily, and returned to optics. I was lucky enough to have a friend, Peter Herring, who organised deep sea cruises on the RRS Discovery, and I went on 4 of these altogether. In 1975 I found that shrimp eyes work on a mirror mechanism, in which an array of corner reflector boxes, at right angles to the cornea, form a superposition image on the retina. Later I found out that I was only the belated co-discoverer of this system; Klaus Vogt, then in Tübingen, had found the same mechanism in crayfish the year before. Still, by then I had already had my celebratory drink! Subsequently this mechanism turned out to be useful for focussing X-rays (for telescopes) and collimating them (for sub-micron circuit etching). It was satisfying to see biology leading engineering, for a change. There were many other good things in the sea: the spectacular double eyes of hyperiid amphipods and euphausiids, the scanning eyes of heteropod molluscs, and the strange eyes of pontellid copepods – each using a different optical principle. Sadly, going to sea now involves a lengthy bureaucratic hassle – in the old days Peter simple called me on the phone.