Matthew Jarron: the life of D'Arcy Thompson
D’Arcy Thompson on a biological field trip, (Courtesy of the University of St Andrews Library, via ongrowthandform.org)
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Dr Kat Arney sat down with the curator of the University of Dundee’s D’Arcy Thompson Zoology Museum, Matthew Jarron, to find out more about the larger-than-life Scottish geneticist, D’Arcy Thompson…
Matthew: Thompson is really one of the most extraordinary characters, I think, in late 19th, early 20th century science, and indeed in many other fields as well. So he was Professor of Biology here in Dundee in its very early years. He came here in 1885. He was here for just over 32 and a half years, and then moved over to the University of St. Andrews as a Professor of Natural History, and was there for the remaining 31 years of his life. So he was a professor for a very long time, and amazingly was still teaching right up until he died at the age of 88, and during that time became very renowned in many different areas.
Matthew: He's of course best known for his work in applying mathematics to biology. But in his lifetime, he was probably much better known for his work in fisheries. He was the scientific advisor to the Scottish fishery board. And so a lot of his sort of day-to-day work research concerned doing statistical analysis of fish catches and fairly mundane things like that.
Matthew: And it was part of that process that, of course, is still very controversial today of trying to, on the one hand, balance fish stocks in the ocean and on the other, the livelihoods of fishermen. And so he was involved in a lot of international negotiations on fishery rights and so on. And so that's really the work that at the time he would have been best known for, but we've kind of forgotten that side of it because of this extraordinary legacy of this amazing book that he wrote called On Growth And Form.
Kat: What was On Growth and Form, where did it grow out of?
Matthew: Yeah, it's really interesting looking back at the origins of this book, because I don't think D'Arcy ever really intended to make this his life's work in a sense, although to some extent that's what it became. I think because University College Dundee was a very small institution and so there were a fairly small number of staff, they worked very closely together. And I think that was one of the things that really encouraged him to think in an interdisciplinary way at a time when increasingly science was getting more and more specialist. And so he would regularly be having conversations with fellow professors in physics and in mathematics and in other subjects. And I think that encouraged him really to think in those terms.
Matthew: He was also very inspired by an organisation in Austria called the Vienna Vivarium, where they had a whole team of researchers from all different disciplines working together to study animals in different ways. And that was something that really inspired him as well.
Matthew: We know that in the late 1880s, he started to get interested in the idea of applying mathematics to biology. He actually wrote a letter to one of his former students in 1889, I think it was, saying that he was looking at foraminifera and was really interested in the spiral patterns on them and thinking about the mathematics of that.
Matthew: And then he certainly started sharing that with his students, but I think he very quickly realised that this was quite a controversial area and when he wrote to other noted biologists, they were saying, "Well, you can't possibly invoke mathematics here. This is a totally different subject." And so it took him quite some time to start to actually write this up and publish papers on this subject. But he gradually did that in the 1900s.
Matthew: And then at the same time, Cambridge University Press were publishing a series of popular science volumes. And they had actually approached him in 1911 to write a book about whales. Dundee, of course, had been one of the major centres of the whaling industry.
Matthew: He said, "Well, I don't really want to write a book about whales, but I've been thinking for some time about doing something about mathematics and biology." And so they said, "Fine off you go." And then this took several years and ended up being considerably longer than anyone actually anticipated.
Kat: We've all been there.
Matthew: Well, indeed! I think it somewhat alarmed Cambridge University Press, who were obviously expecting this little, slight, popular book and they've got this massively erudite tome. But eventually the book was actually more or less finished by 1915, but because of delays during the war and paper shortages and so on it didn't actually get published until 1917.
Matthew: It is, for anyone that hasn't seen it, a fairly weighty tome, although of course the second edition then got even weightier. And effectively it really tries to look at living organisms and understand their growth and development in the same way as inorganic forms. So this was really one of the great innovations. Whereas previously, most biologists felt that the living world had to be dealt with in a completely separate way from the non-living world, D'Arcy was saying, "Actually, no, we can look at a lot of basic forms in biology and explain their growth and explain why they take on the shapes that they take on according to fundamental laws of physics and mathematics. A living thing may grow in exactly the same way as a non-living thing because they're just obeying these fundamental laws."
Matthew: And that was a very controversial idea at the time, it really was not widely accepted for quite a long time afterwards.
Kat: I find that surprising because you look at things like these foraminifera, these seashells, and you can see that there are geometric shapes, and you know, you look at humans and humans all come out in the same basic pattern and a dog is the same basic dog pattern. And you know, a plant is the same sort of plant pattern. Things do have their form.
Kat: I'm intrigued as to why people thought that the two domains must never meet . You just can't come up with physical and mathematical laws to explain squishy biology. What did that idea grow out of, that these two must never meet?
Matthew: The kind of main division within biology at the time was this kind of debate between on the one hand, so-called vitalists who believed that because God had created all living things therefore ultimately the secrets of life could not be understood by basic man-made laws, and there had to be something that we couldn't understand, that was the sort of "secret of life".
Matthew: And then on the other hand, the so-called mechanists who basically saw organisms as machines, that they were basically a bunch of parts and you put them together and you've got a working machine. And the interesting thing is that D'Arcy, although it would seem that On Growth and Form comes down more on the mechanist side that he's definitely looking at the mechanical basis through mathematical patterns and physical laws and so on, actually he's kind of in the middle of those two.
Matthew: That on the one hand, yes, he recognised that there were certain rules that structured how things are put together, but then on the other hand, he very much recognised the idea that organisms acted as systems. That effectively they were greater than the sum of their parts.
Matthew: And so again, that was a really challenging and pioneering idea at the time that was not something that other people were really thinking about.
Kat: It is incredible because systems biology really seems to have emerged actually, quite recently, as this whole discipline of, "These are self-organising systems. Everything is a system that works together" and the episode we've just done recently on Turing patterning in development and particularly limb development, this sort of synthesis now that, "Oh, we can have these mathematical pattern generation processes that do generate squishy body parts in the right place and at the right time."
Kat: And it feels like this is very much an idea ahead of its time. He's writing in 1917 that there are these kind of systems at work.
Matthew: That's right. He undoubtedly was ahead of his time. And in a way he came at just the wrong time because, well, there were two problems. One was of course that you couldn't actually test any of his theories at that time because the kind of mathematical models you would need simply just didn't exist. And, you know, it took incredibly brilliant mathematicians lecturing to at least start coming up with some of those models. And of course we know that Turing was much influenced by D'Arcy's work.
Matthew: The other problem, of course, was that at just the time this book was coming out, there were these huge advances in genetics which suddenly everyone latched on to that, and thought, "Right. Here's the thing that explains everything in evolution and we don't need all this mathematical stuff now because we've got this."
Matthew: And so had that not happened, biology may well have gone more towards this mathematical and systems approach, but it sort of suddenly took a left turn and people started focusing much more on evolutionary biology, looking at genetics as the mechanism that explains that.
Matthew: And so for most of D'Arcy's lifetime, there were really just a small number of people that were actually following his ideas and trying to take that further. But it certainly was the case that it wasn't really until, well, long after D'Arcy's death, sadly, that partly through the development of computers and the ability to have more complex mathematical modeling, and partly because of, I think, a greater interest in developmental biology and new ideas coming out in that, that people started to realise that actually it wasn't a case that it was either Darwin was right or D'Arcy was right. Actually they're both right. But actually we can bring the two together and we can do modeling that will actually help both.
Kat: We'll come back to the whole, "Was Darwin wrong?" thing because I want to drill into the genetics angle a bit. Because you're thinking, you know, 1910: Bateson, around 1914 -1915 you've got the fruit fly geneticists and people like Thomas Hunt Morgan, who are doing all these incredibly complicated crosses with one sort of fly crossing with another sort of fly and seeing what mutations are there and how they look, and then trying to actually map out genes for the very first time and it feels to me that the genetics of that time it's almost like a paper science. It's like drawing maps, these things aren't almost real. These are just quite theoretical maps that you could draw a line on paper and go "This gene's here, this gene's here, the gene's here,"
Kat: That's very separate from where we start to get to with the biochemists and the people in the wet labs, the molecular biologists, who were understanding proteins and nucleic acids. And that starts to really take off in the 1950s where people are drilling into the structures of all these molecular machines and enzymes and all these things completely separate from the genes and the mapping.
Kat: And then you've got this separate whole, sort of, structural "How do you bring all these things together to actually make physical beings, guys?" In case you forgot, we're not test tubes or maps of genes.
Matthew: That's absolutely right. And I think that one of the issues that we have with On Growth and Form is it's very easy to dismiss what D'Arcy was doing as if he just wasn't interested in what was going on in all these developments. I think he was interested, and he does briefly refer to it. But he felt, "This isn't relevant for the specific thing I'm trying to explain in this book, so I'm just not going to bother with it."
Kat: It's like it's the wrong theoretical framework. It's like, "This just doesn't explain what I'm seeing, does it?"
Matthew: Exactly. He kind of just ignores it and then unfortunately he then carries on ignoring it. And I think perhaps the problem is not so much that he ignores it in the first edition, but by the time he eventually published his second edition in 1942, of course there'll be massive developments by that point. And he just carries on ignoring them.
Matthew: A lot of people felt, "Well, that's just ridiculous because clearly you really should be thinking about this." Of course by that time he was 82, so I think we can forgive him not being quite at the cutting edge of what was going on. But, it does mean that to a lot of biologists, he was just seen as some kind of archaic dinosaur that wasn't really up with the latest trends.
Kat: That's very unfortunate.
Kat: I wanted to get into the Darwin evolution thing because it's sometimes set up as a controversy. What was his feeling then about Darwin and Darwinian evolution? Was it right? Was it wrong? Was it again, not quite the right theoretical framework for what I'm thinking about?
Matthew: It's really interesting because D'Arcy was not in any way, anti-Darwinian. He certainly wasn't saying that there was no evolution or anything like that. And in fact, it's worth noting that the very first book D'Arcy ever published, he actually got Darwin to write the foreword for it.
Kat: That's a goal!
Matthew: Yeah, it was, it was D'Arcy's first publication and it was Darwin's last publication. It was a translation of a German text about the fertilisation of flowers and Darwin agreed to write this foreword. They never actually met, but they did correspond.
Matthew: Then there's a rather lovely letter where Darwin sends his foreword to D'Arcy and says, "I hope it's okay." I think the exact words that he uses are, "I'm worried it might be a bit rubbish." Not the kind of thing you imagine great biologists will say.
Kat: No, but it's a very, very real thing where you're like, you're writing to a mate and you're like, "Here it is, I hope it's not rubbish. Let me know."
Matthew: Needless to say it wasn't rubbish and D'Arcy was only too pleased to have this.
Matthew: He never actually met Darwin, although they did correspond, but he did regularly meet Alfred Russell Wallace. And several times Wallace actually came up to Dundee to stay with him, in fact.
Matthew: So there's no doubt that he was entirely in agreement with the ideas of Darwinian evolution. The problem was that he felt that there were lots of occasions where people looked to evolution to provide an answer to a problem. And he was saying, "Well, you don't need to look at that because actually there are other things."
Matthew: One of the things that he was really fascinated by, for example, were Radiolaria; these tiny little single- celled creatures that create these amazingly geometrically precise forms. But there's a whole myriad of different shapes. Now they'll have the same evolutionary pressures upon them. So theoretically Darwin would say they should all evolve into the same shape, but of course they don't.
Matthew: So D'Arcy was saying, "Well, we don't need evolution to explain that because it's purely the physical forces that are acting on that organism during its development that will trigger particular mathematical growth patterns that will lead to those to happen."
Matthew: The problem was of course that a lot of On Growth and Form was interpreted as being anti-Darwinian. And in fact, he had actually himself set himself up for that. Way back in 1894, he'd actually given a paper at a British Association meeting called Some Difficulties with Darwinism in which he first starts to propose some of these kind of issues. Again, he's not saying it's wrong. He's just saying that these are not the things we need to use to explain certain issues.
Matthew: So I think people were already kind of a bit cautious when they started reading this book and there's obviously a lot of it where he just keeps on saying, "We just don't need that. We just don't need that. You can forget that. We don't need to evoke evolution."
Matthew: And then there was the most famous chapter, certainly in terms of the illustrations, the final chapter on the theory of transformations, where he does these extraordinary diagrams, where he bends fish and that kind of thing.
Matthew: Whereas Darwin was saying that evolution was always this very slow, gradual process where maybe an eye would gradually get bigger over many generations, D'Arcy felt that that wasn't always the case. He felt there must've been times when there was a whole body transformation. This idea of thinking of organisms as systems, and there was this whole system of forces that caused this kind of transformation. And so again, that was seen as being anti-Darwinian. Again, I don't think he necessarily intended as such.
Matthew: I think it was a bit frustrating to him that people thought in those very black and white terms. He did get labelled as an anti-Darwinian and actually that is a problem that persists to this day. Cause you still get fans of intelligent design and creationism and so on evoking his work as some kind of scientific proof of their ideas.
Kat: Yeah, that seems like a bit of a mistake.
Kat: I'm fascinated by the fish bending thing, because when I was writing my first book, Herding Hemingway's Cats, one of the stories that absolutely fascinated me was the work of David Kingsley and his team where they're looking at sticklebacks and they find a tiny piece of DNA that if you remove it in sticklebacks their entire pelvis disappears. And you find these different types of sticklebacks living in lakes - they'd sort of been trapped by the retreat of the glaciers - and it's a tiny DNA change. And these sticklebacks that are stuck in the lakes: no pelvis! Their bum's just gone compared to the fish that are still in the sea that need this big bony pelvis to protect themselves.
Kat: And it's a really powerful example of how you can have a very, very dramatic change in the form of an organism, which is driven by the selective pressures on it, the functional pressures on it, just driven by this tiny single change in DNA.
Kat: I looked a lot at the idea that you've got all these control switches that are turning genes on and off and that's really like evolution's playground. You only have to make a little change in one of these, rather than an inching, tiny change in something getting a bit bigger or a bit smaller or a bit spottier or whatever, you just change one of these switches and you make really big changes to organisms that fit with their function. And if it works, brilliant! Selection have at it!
Matthew: And actually, this is precisely where you wish that D'Arcy had been more on board with what was going on in terms of ideas of DNA and genetics and so on, that he would have then understood, "Actually, this backs up a lot of my theories and I can use it in a different way."
Matthew: But I think he was so concerned with trying to play devil's advocate and trying to prove as much as he could without evolution that he kind of in a way didn't always come up with the right answers, even if he was asking the right questions. And the theory of transformation is a great example of that because in a sense, a lot of it's rubbish, you know, clearly these things don't actually happen the way that he presents them.
Matthew: I mean, its obviously problematic that he uses adult fish and then tries to stretch the adults rather than the embryo. And that he's using two fish that were around at the same time, so it's not like one fish evolved into the other fish. So there are a lot of things that are problematic about that, but it is, as I say, it's definitely asking the right questions.
Kat: Is there any relevance of D'Arcy's book today, On Growth and Form? Is it worth reading? I assume it's pretty hefty coffee table book. What can it say to us today?
Matthew: I think there's quite a lot that On Growth and Form can say to us today. It's definitely not an easy book to read, not just because it's full of mathematical equations and fairly dense prose. He does have a habit of quoting extensively in Greek, Latin, French, German, Italian, and just assuming you know all these languages and so doesn't bother to translate them.
Matthew: But at the same time, it's full of wonderful images, which, of course, a lot of earlier textbooks weren't. It's really one of the first great scientific textbooks to be so heavily based on images. And the influence that it has had is absolutely extraordinary and quite beyond biology.
Matthew: It's had a huge influence in art and design and engineering and architecture. In cybernetics, in systems theory, in geography, in philosophy, in anthropology. There are so many different areas and even whole new areas of science like nano chemistry, for example, Geoffrey Ozin who's one of the key pioneers of that, really credits D'Arcy Thompson as being absolutely crucial to his thinking. So I think there are a lot of ways in which that book continues to grow and evolve and have a really interesting relevance.
Matthew: Specifically of biology, much of it doesn't have a direct relevance in the sense that mathematical biologists today wouldn't use the same mathematics. They're obviously using vastly more complicated things than D'Arcy was using. But, in a sense, that whole discipline is really based on the fundamental building blocks that D'Arcy set up.
Matthew: I think the fundamental idea that we should think of organisms as systems, that we should think of them as greater than the sum of their parts, that we shouldn't lose sight of the whole organism, when so many scientists spend their entire careers looking at a single gene, and that we should continue to think in interdisciplinary terms, those are all really important ideas.
Kat: I think it's wonderful. You almost wish that he could be here to see now we've got finally the synthesis of the mathematical modeling with the computers that can do it. We've got the synthesis of genetics and evolution and the molecular biology, so we can see how all these systems actually interrelate together. And finally being able to visualise, to really look at organisms and also to measure things like physical forces now in a way that we never could.
Kat: It's really struck me. I think it was a Genetic Society meeting a couple of years ago that was really going into physical forces in biology that we don't think about, the constraints when things are pulling and tucking and folding and one side's growing faster than the other side. What do you think D'Arcy would have said if he could have sat in meetings like this today?
Matthew: I'm sure he would be absolutely delighted. I think it did frustrate him that there wasn't that of immediate rush of biologists to embrace mathematics after he published his first edition of his book. He'd totally be pleased that here at the University of Dundee, for example, mathematical biology is now a major research area. And the fact that there are whole areas where work is done in some ways inspired by his original ideas that he would never even have imagined, I think is wonderful for someone who was such a great interdisciplinary thinker. I think he would just be dead chuffed.
Kat: We've talked a lot about D'Arcy as the scientist and his ideas, but what was he actually like as a person? Because I get the idea that he was not maybe your run-of-the-mill scientist for the time.
Matthew: I think that's definitely fair to say. He was, I think one of the great eccentrics of his age, really. Particularly in his old age, he was famous for wandering around St Andrews with a parrot on his shoulder. His lectures were very entertaining. He used a whole variety of different props and he would regularly pull out you know, soap bubbles, bits of string and frogs and all sorts of different things.
Matthew: I remember being told actually by one of his former students of a time when they were due to have a lecture from him and they were waiting for him to arrive and he was kind of late and they were sitting around and it was nearly getting to the end of the allotted time and he still hadn't turned up. About two minutes before the lecture was due to end he finally bustled in with this enormous coat on, bulging pockets and so on and sort of rushed up to the podium and said, "My lecture today." And then he said, "In my left pocket, I have a dead frog." He pulls out this frog and dropped it onto the lecture theatre bench. "And then in my right pocket, I have a live frog," and then he pulls out this frog and it hopped about all over the place. And then he said, "Observe, if you will, the difference between the two,” then he walked out.
Matthew: And that was the sum total of his lecture.
Kat: I mean that's science communication, isn't it? How was he communicating his ideas more broadly? Did he try and engage more widely in talking about science and life with the public?
Matthew: Very much so, and he loved speaking in public, actually. He gave a lot of public lectures. He did, for example, some of the Royal Institution Christmas Lectures for children. He did a lot of radio broadcasts, although frustratingly, none of those recordings seems to survive today. And he wrote a lot of popular articles in magazines and so on.
Matthew: So he definitely was a great science communicator. I think it frustrated him to see scientists who weren't able to write good English and communicate properly. But then at the same time, there is this other side where, as I've mentioned he expected people to be equally as intellectual as he was and his great command of languages and so on.
Matthew: So another wonderful story by one of his former students. Towards the end of his life, he was giving tutorials in his own home to save him having to go onto the campus. And one of his students went to visit him there and he was reading from a textbook, this fairly ancient book, to this student. And he was reading quite slowly and obviously he's quite elderly at this point. So she was a bit worried and there would be these long pauses. These pauses gradually got longer and longer and she thought, "Gosh, I hope he's okay."
Matthew: And then eventually after a particularly long pause, she plucked up the courage to say, "Professor Thompson, are you all right?" And he looks up rather cross, and he said, "My dear girl, I'm translating from 15th century Italian, and it does occasionally make me pause."
Kat: Yeah. That's ... I can't make out if he'd be like the most amazing lecturer or just... I think probably in retrospect, you look back and go, that was incredible.
Matthew: Absolutely. I think one of the things that's interesting is very few of his students went on to become great biologists, but they went on to become lots of other things. I think he opened up a sense of wonder. He greatly inspired people without necessarily actually teaching them an awful lot of hardcore biology.
Kat: And you're the curator of his collection. So, he was collecting things as well. What sort of things did he collect?
Matthew: When he first came to the university, he very quickly realised that what he needed in order to teach his subject was a museum of zoology. So in a relatively short space of time and with very little money, he actually built up what became one of the largest natural history museums of its kind anywhere in the UK. And so he was collecting very widely all over the world. He swapped specimens with other collectors and other museums around the world.
Matthew: We mentioned whales earlier because Dundee was the centre of the whaling industry. He was able to make friends with the whaling captains and get them to bring back specimens from the Arctic, he sent a couple of his students on whaling expeditions, which is an incredibly dangerous thing do because very often whaling ships would get trapped in the ice and never returned.
Matthew: Obviously there were no risk assessment or health and safety in those days. So you could send your students off to almost certain death and nobody really worried about it, but thankfully they did return and they brought lots of specimens with them.
Matthew: And so that way he was able to get specimens that other collectors couldn't get and so he could then swap those for other things around the world. So we built up a huge collection. Sadly, his original museum was demolished in the 1950s when they were building new buildings on the campus and so a lot of the collection was lost at that point, but we still have the kind of basis of his collection in what we now call the D'Arcy Thompson Zoology museum after him.
Matthew: We do get D'Arcy fans coming from every corner of the world to come and see his original collection. And certainly there are specimens that directly relate to things that are in On Growth and Form, and some of the specimens that appear in the transformation diagrams. For example, we have a lot of his original microscope slides with foraminifera on them and that kind of thing. So it very much does relate to his research as well as his teaching.
Thanks to Matthew Jarron, curator of the University of Dundee’s D’Arcy Thompson Zoology Museum for sharing his story with us.
