In a paper published in the Proceedings of the Royal Society B in 2015, Kevin Lala, Tobias Uller, Marcus Feldman, Kim Sterelny, Gerd Müller, Armin Moczek, Eva Jablonka and John Odling-Smee laid out their vision for an “Extended Evolutionary Synthesis” (EES), a new framework for Evolution that took on board (then) recent findings in evo-devo, developmental plasticity, inclusive inheritance and niche construction. Lala and colleagues argued that standard evolutionary theory was too narrowly focussed on genetic inheritance and that these new findings demanded a recognition of processes through which organisms grow and develop as causes of evolution. An earlier version of this paper was reviewed and rejected at Nature, but soon afterwards Lala and colleagues were invited by the Nature editor to contribute a much shorter version of the article as one part of a debate, the other part of which would be a response to their ideas by more traditionally-minded evolutionary biologists. You can read more about the backstory to the Nature exchange here. In this interview, conducted nearly ten years after the paper was published, we spoke with Kevin Lala about the backstory to the Proceedings paper, and more generally about the origins and current status of the EES.
Citation: Laland, Kevin N., Tobias Uller, Marcus W. Feldman, Kim Sterelny, Gerd B. Müller, Armin Moczek, Eva Jablonka, and John Odling-Smee. Laland, K. N., Uller, T., Feldman, M. W., Sterelny, K., Müller, G. B., Moczek, A., … & Odling-Smee, J. (2015). The extended evolutionary synthesis: its structure, assumptions and predictions. Proceedings of the Royal Society B: Biological Sciences, 282(1813), 20151019.
Date and place of interview: 7 June 2024; Konrad Lorenz Institute for Evolution and Cognition Research (KLI), Klosterneuburg, Austria.
Credits: Hari Sridhar (background research, planning and conducting interview); Joyshree Chanam (background research, planning and editing of interview)
HS: Kevin, thank you for agreeing to do this interview.
KL: No problem. I’m very happy to do it.
HS: It’s fitting that this interview is happening at the KLI, because the origins of these papers trace their way back to a meeting that happened, not in this building, but in the old KLI. Maybe we can start by talking a little bit about that meeting.
KL: Yeah, well, I wasn’t actually at that meeting. But I do have a good visual memory of the first time I heard the label Extended Evolutionary Synthesis. It was because my colleague, John Odling-Smee, had been invited to that particular meeting. He and I, of course, had been working on niche construction theory with Marc Feldman. I remember being in John’s dining room – I suppose this must have been 2008, something like that – and he told me that he’d been invited to this KLI workshop on the Extended Evolutionary Synthesis. And I remember saying to him, “what’s that?” He replied words to the effect that “He didn’t know. But, you know, there were a lot of interesting people going to the workshop, and he thought it would be fun, so he was gonna go anyway”. And I remember saying to him, when you get back, let me know how it goes and what that Extended Evolutionary Synthesis is. And so, we kind of repeated the conversation a few months later, when he got back from the meeting, I said to him, “so, what is the Extended Evolutionary Synthesis?” And he again said, “I don’t know!” It’d been a very interesting meeting, and he met some very fascinating scientists and enjoyed it, and, I think, got a lot out of it. But he still really wasn’t any the wiser as to what the label Extended Evolutionary Synthesis, or EES, stood for. And there were some really key questions like, what findings motivated the EES, and what assumptions went into it, and how it was distinctively different from a more traditional viewpoint of the Modern Synthesis. And these kinds of questions hadn’t yet been resolved. But I think the workshop was successful in bringing together a bunch of people that were doing work that was, in some sense, relevant to that. So, we looked forward to the book coming out, where we anticipated we will get answers to those questions. And then, in 2010, the book was published, and we read it, but we still felt none the wiser! There were lots of very interesting articles in it and plenty of food for thought, but it wasn’t clear how it all fit together. And I guess, we were at a stage in our intellectual development, where we were looking for commonalities between the perspective we’d developed in niche construction theory and other strands of evolutionary thought. So, around that time, I’d had an inspirational conversation with Gerd Müller, which had brought home to me that there were certain parallels between how we were thinking about niche construction and how he was thinking about evo-devo. And in particular, the idea that what we might think of as ‘devo-evo’, that developmental processes might play a role in evolution. And while both evo-devo and niche construction theory were thriving, we nonetheless resonated over the fact that we both sometimes encountered a certain resistance or hostility, or a failure to appreciate the arguments that we’re making, which we could trace back to some conceptions of biological causation that were widespread among scientists. I had traced it all the way back to an article written by Ernst Mayr in 1961 in the pages of Science, “Cause and effect in biology”. I told Gerd that I think a lot of the problem is because people think in this way about causation, using that distinction between proximate and ultimate causation that Ernst Mayr had championed. And from that perspective niche construction can, of course, then be dismissed as being of evolutionary importance because it’s viewed as a proximate mechanism, therefore not of evolutionary significance. Gerd literally did a double take and sort of said, “but it’s the same for evo-devo!!” And we got very, very excited!
HS: When and where did this meeting happen?
KL: That happened at the old KLI, in Altenberg. I’m guessing around 2010, something like that. It was around that time, maybe 2010-2011. I don’t remember exactly. But I do remember that it prompted me to really start thinking – particularly with John Odling-Smee, and Tobias Uller who we’d started working with – about the idea that we were pushing for a different model of biological causation. And that this was relevant to some other debates that were taking place in biology concerning the evolution of language, concerning evolution of cooperation, plasticity-led evolution, and, as I mentioned, evo-devo… where in all cases you had two sides of the debate. One side that sort of embraced this Mayr formulation of a distinction between proximate and ultimate causation – a more traditional camp – and then another more radical camp that we’re pushing against that, and were arguing that developmental processes played important evolutionary roles. So that just, sort of, hinted to us that we might be, with our thinking about niche construction, part of a broader picture. And we’d also written an article with Scott Gilbert, which was published in 2008 exploring some parallels between niche construction and evo-devo, since there is a ‘constructivist’ tradition within developmental biology — where you can kind of think of what’s going on inside an organism when it develops as self-construction. John, Marc and I had obviously focused on those constructive activities that occur outside the organism, in the environment, such as when spiders spin webs or beavers build dams, but there’s really no need to make a big thing about the physical boundary of the organism. Similar processes are going on inside the organism as well. And some of that was being emphasized within the field of evo-devo. That was again, a sort of a prompt, that there might be commonalities between our ‘struggle’, and the struggles that other researchers were experiencing in different subfields of biology, who were pushing against the grain of the more traditional way of thinking.
HS: When you met Gerd at the KLI were you there for a meeting or conference of some sort?
KL: I would have been there for a workshop. We were sitting under a tree in the sunshine with a glass of wine, after a day’s discussions. It might have been a workshop on cultural niche construction, or something like that. And yeah, Gerd had just said, “So how are things going with niche construction?” I was asking him, “How are things going with evo-devo?” And we suddenly realized that we’re fighting the same fight. To sort of bring it back to the Extended Evolutionary Synthesis papers, I think our motivation to explore these issues was primed by those experiences. But it was also partly born out of frustration, that we hadn’t got the answers that we wanted to those questions: What is the Extended Evolutionary Synthesis? What biological findings motivated it? What was making people think we might need one? What assumptions did it make? How are those different from the more traditional assumptions? What predictions did it make? Can you actually juxtapose those different viewpoints and then generate hypotheses that will put them to the test? That was the kind of motivation that underpinned it. John and I, coming from this niche construction tradition, had always been hugely influenced by Richard Lewontin. And one of his books is, of course, The Dialectical Biologist, which he wrote with Richard Levins, where he endorses dialectics. And, I guess I’d, slightly unthinkingly – because Lewontin was such a hero to me – embraced this dialectical perspective. So, from my perspective, it’s always a good thing for scientists to propose an alternative way of thinking and to lay it out and say this is the sort of logical expectations of that way of thinking, and juxtapose it against another. And then, you know, that encourages scientists to think about the assumptions they’re making and try and put them to the test with empirical theoretical work. It protects the field against dogmatic views. So, I think, when writing that pair of papers about the Extended Evolutionary Synthesis, John and I, at least, envisaged that we were providing a service to the community. I don’t think we had any idea about the storm of controversy we were about to get into!
HS: We’ll come back to the papers in a minute. I first want to step back a little bit in your own trajectory and ask you about your interest in niche construction. From looking at your publication record, it looks like a paper in 1996 might have been around the time when you first started thinking about it with that frame. Could you trace how you arrived at this? Your PhD and a lot of your early work was on social transmission, especially on rats. How did you get interested in niche construction?
KL: My interest in niche construction actually traces back a lot before that. I actually wrote to John Odling-Smee, I suppose, in 1987. I was interested in the kind of topics that he was exploring, and I was looking to do a PhD. He was one of the people that I wrote to and said, I’d be interested in doing a PhD under your supervision. In the end, I ended up doing a PhD with Henry Plotkin on animal social learning. But Henry and John had worked together. And so, one of the attractions of going and working with Henry was that I would also be allowed to work with John on the side – which we did. We carried out experiments on niche construction in earthworms. But to give you more background, I’d written to John, as I say, around ’87, and he sent me a manuscript that he was working on, which eventually came out in a book that Henry edited in 1988, which was almost the first paper on niche construction theory, and certainly the first paper that uses the phrase ‘niche construction’ – although Dick Lewontin had talked about ‘constructing niches’. I was living in Southampton at the time. I recall, I was moving house, and a friend of mine had an open-back truck. He was kind enough to take all my furniture and belongings from one side of Southampton to another in the back of his truck on a sunny summer’s day. I put everything in the truck, it was about to leave, and I thought, “Oh well, I’ll just check whether any mail has arrived”. And this big fat envelope had arrived with this manuscript from John Odling-Smee! I often think, my life could have been very different had I not gone back to the mail that day! But yeah, I started reading it in the back of this truck, driving across town, one Saturday morning. The traffic was quite heavy, so the journey took a long time, and by the time I got there, I was hooked. I was sold. I’ve always had a weakness for what you might think of as big theories, theories that might be really fundamental and important and interesting and potentially impactful across a number of different domains or academic fields. And so, niche construction, being such a fundamental idea, attracted me immediately. But on the other hand, John didn’t really have the resources to support me to do a PhD, so I ended up doing this work with Henry Plotkin on social learning, and working with John on earthworms on the side.
I remember at an early stage in my PhD — I don’t know to what extent you’re interested in all these discursions — I remember getting some financial advice. In fact, a lot of the students, were given financial advice. And it was standard financial advice, sort of, if you have money, hedge your bets, make sure you invest wisely and don’t put all your money in a high-risk share, and all this kind of thing. Of course I didn’t have any money. I was a graduate student: I didn’t have two pennies to rub together. But what struck me was, what I did value was time. And as a graduate student I could adopt the same policy that this financial advisor was advising with respect to time, in the sense that I would have a balanced portfolio with my time. I wouldn’t invest all of my time one high-risk idea, like niche construction – that wouldn’t be prudent. But I could allocate some portion to that. And I could also allocate some portion to something which was a lot safer as a bet, which was more likely to generate a steady stream of papers, like animal social learning, which was also interesting. And actually, I had a third sort of horse that I was backing, which was an interest in gene-culture coevolution. So, I thought of the social learning as kind of like a 2:1 bet – it might not win me much money, but it will come in fairly reliably. And then there were 10:1 odds against gene-culture coevolution. And niche construction was like the crazy 1000:1 outsider, but it was genuinely fun to think about such a fundamental idea.
HS: I have to go back one more step to ask you: did you write to John Odling-Smee at the time when you’d have just completed your masters?
KL: Undergraduate.
HS: At that point, how did you even know about John Odling-Smee, and how were you interested in all of this already?
KL: Yeah. So, I had a sort of random walk to my early career. I, for not very good reasons, went to university and studied engineering, realized I didn’t want to be an engineer, switched to psychology, again, for fairly random reasons, and realized I didn’t want to be a psychologist. But towards the end of my undergraduate degree in psychology, I took a course called ‘Psychology, as a Post-Darwinian Science’. It was run by a brilliant guy called Alan Costall. He essentially introduced me to evolution. Because I’d never taken biology at school, or in any of my classes, I’d not really come across the idea of evolution, and not really got an opportunity to get excited about it. And I became very, very interested, in particular, in human cognitive evolution — how could we understand the evolution of the human mind, the human condition? So that sort of instigated my interest in evolution, in particular, the kind of ideas about evolution that could account for humanity. Some of the works that Costall introduced me to were papers by Henry Plotkin and John Odling-Smee. So, I’d known about some of their early work. It’s on evolutionary epistemology – the idea that there is a hierarchy of Darwinian processes that are operating, and you can think of selection as occurring on cultural variation, on development variation, and so on, and so forth. I could see I could see some virtue in that perspective. So that’s why I wrote to John. Also, why I wrote to Henry.
HS: Coming back to the EES: in the backstory to the Nature exchange, you talk about this working group that you constituted with Tobias Uller and John Odling-Smee. How did that happen?
KL: I think I mentioned, didn’t I, that John, Tobias and I had written this paper in 2011 on reciprocal causation – ‘Cause and Effect in in Biology Revisited’ is the title – it effectively reevaluates Mayr’s distinction between proximate-ultimate causation, 50 years after the original paper was published. That had brought together some members of our team. Tobias, John and I were part of the group that wrote the causation paper. Kim Sterelny was also a member of that team. I’d worked with Kim previously. He was a philosopher who is very interested in niche construction. I valued the clarity of thought that he brought to our discussions. He was very smart but also he was really interesting. Slightly crazy guy with a huge beard and hair all over the place. He was an eccentric character. He’d stay up late drinking and discussing stuff. He was a lot of fun to be with it. For someone like me who just likes talking about big challenging ideas he was great to be around. We knew that causation was a big issue. And philosophers have got a track record of thousands of years of studying causation so, we thought, if we’re going to actually talk about these issues, we really should have at least one philosopher involved, to kind of wise us up about that tradition. And Kim was the philosopher that we knew. But also, I think it’s more than that. It’s that a lot of the philosophers I’d encountered — I’d actually had quite a lot of interaction with philosophers by that point — most of them were content to be observers of biology. They would analyze what biologists were doing and saying, and how they were thinking and reflect on that. But they wouldn’t want to be participants. Kim actually wanted to be a participant. He was willing to join our team, and stick his neck out, which was a major factor as well. So, yeah, I guess publishing that paper in Science triggered us to think more broadly about this idea of an Extended Evolutionary Synthesis. And since nobody else—I just said this idea was born out of frustration— since nobody else seemed to be really trying hard to work out what it meant – What was the EES, what wasn’t EES, and so on – and since we had this intuition that our work fit into a broader picture, but we hadn’t quite clearly worked out what it was, we just thought it’d be an interesting exercise to pull some people together and try and work it out. And if we could get some of the world’s leading people in each of these domains to discuss with us, what a luxury that would be! What a privilege to sit around the table with those people and try and thrash these ideas out. And again, I sort of come back to the thinking that if we could work this out, this would, in all honesty, be a service to the community. So, we just reached out people, like Mary Jane West-Eberhard, who I knew only vaguely at that point – but I knew she was doing really interesting work on developmental plasticity and plasticity-led evolution and we thought that was important. We reached out to Marc Kirschner. We were very interested in Kirchner and Gerhart’s work on facilitated variation. We liked their emphasis on exploratory processes and thought that was important. We knew that evo-devo was a big part of our story. Armin Moczek had written some really interesting articles that seemed to be sympathetic to the view of evolution that we were developing. So, we reached out to him. I’d not met him previously, but he responded positively. In fact, everybody responded positively, initially. Doug Erwin was another one. He’s someone that we had worked with by that point. He’s very interested in niche construction, but from a macroevolutionary perspective. He is a palaeontologist, and we thought it would be good to get a macroevolutionary perspective on our working group. Who else is there?
HS: Gerd
KL: It seemed natural to involve Gerd because we’ve had those conversations where, clearly, we were thinking along the same lines. And also, he was, at the time, President of the European Evo-Devo Society, and so he was a man of authority and knowledge of evo-devo, which was clearly central to our interests. But there was another factor, which is that we wanted to write something on the EES, but we didn’t want to be perceived to be sort of splitters, who were distancing themselves from what Gerd and Massimo had put together in their book on the EES. It was important that we had at least one of them involved, just so that there was some sort of continuity with the original EES writings and thinking. Otherwise, there would be too many factions. I don’t think that’s healthy. That’s another reason why Gerd was involved although he might easily have been involved anyway. And Eva Jablonka, of course, who we’d known for a long time. Clearly, we needed somebody who had expertise on extra-genetic inheritance, and there was nobody we knew of who had more expertise on that topic than Eva. In particular, we were all being stunned by the findings that were coming out on epigenetics and epigenetic inheritance. And it was – it still is, I guess – sort of mind boggling. You’re constantly being challenged by new findings, and new mechanisms. And, you know, the molecular biology is, sometimes, quite complex and Eva had her head around all that, and we didn’t. So, Eva was, again, a very natural choice. We’d known Eva for a long time and, I suppose, could even think of her as a good friend. So yeah, I suppose, we just reached out to the people that we thought were necessary to cover certain bases that we had an intuition were likely to be important. We just chose the best person we knew, reached out to them and said, we’d like to put together a working group and think about this and see whether a case can be made for an Extended Evolutionary Synthesis. I think, at that stage, we weren’t committed to it at all. It was like: Can a case be made for it? What would it look like?
HS: One other name is Marc Feldman.
KL: Well, yeah. Marc was, again, a very natural choice for us because he was a co-author on our niche construction work. But also, Marc, again, covered some important bases. He knows a huge amount about the history of population genetics and the field of evolutionary biology. And also, he’s a very strong theoretical biologist and one of the cofounders of the field of cultural evolution. So again, he covered those bases. He was our kind of go-to theoretician and historian of evolutionary biology as well.
HS: These people that you brought together, they all come from different disciplines. How were you aware of their work, in the sense that, what was the connection to what you were doing?
KL: To answer your second question first – what was the connection to what we’re doing – that’s, in a way, what we wanted to find out. We were pulling them together because we had an intuition that what they were doing was central to what we were doing. That the work on niche construction was part of a bigger picture which we now think of as the Extended Evolutionary Synthesis. But we hadn’t got it worked out at that point. And we were open to the idea that our intuitions might be wrong, that maybe we were all tugging in different directions. It wasn’t that they were chosen because they were all like-minded. They were chosen because we thought the subject in which they were an expert might be relevant. How did we know about their work? Well, I suppose we had been to a lot of meetings, or workshops about a broader understanding of evolution. Pat Bateson, at Cambridge, who I’d worked with very closely, would often organize these workshops. He was somebody who was pushing for a more organism-centred view of evolution than the traditional view of evolution provided. And he would invite people like Eva and Mary Jane to participate in those workshops, so we’d come across them in those ways. We would read their works, and we found those works interesting. And I think we did see some points of contact with all of them. There were enough points of contact, to make them really interesting and for us to make an effort to read what they were writing. But at that stage, it wasn’t clear that we really were completely intellectually aligned.
HS: Can you talk a little bit about how this working group actually worked? Firstly, was it something formal, in the sense that, did you write a proposal to raise funds to set up this group? Or was it just an informal collection of people?
KL: It was just informal. There was a kind of inner core of myself, Tobias and John Odling-Smee. We would meet regularly, because we were all based in in the UK. I was at Cambridge, Tobias and John were at Oxford. I would usually go to Oxford, and meet them there, at John Odling-Smee’s house typically. We’d have intense discussions for a few days. And then, come up with ideas and produce some written account of those ideas and circulate them to the other people in our team. Some of them like Eva Jablonka and Kim Sterelny would regularly come to the UK. When they did, we’d corner them, we’d have further intense discussions with them. Marc Feldman, we were regularly seeing because of the niche construction work. So, when we went to Stanford, we would also talk about this work. Some, like Armin Moczek, we never met at all in person until after the papers were published. Everything was done over email.
HS: Were there no meetings when the entire group came together?
KL: There’s no meeting where the entire group was present in the same room. But that was okay. I think it worked fine. It was actually a really good collaboration. There were certainly times when I felt like it was a little bit like herding cats. Because, you know, people were tugging in different intellectual directions and did have different views. I mean, for instance, Gerd and Eva were probably much more radical in sort of pushing for talking about an Extended Evolutionary Synthesis as a kind of intellectual scientific revolution. They were comfortable with that language, and Eva was comfortable in talking about Lamarckian ideas being a core component of the EES. Other people thought that would be counterproductive. So, it was certainly was not the case that we all agreed on everything from the start. But I think there was a lot of good spirit in these discussions, and a convergence over time, in what we did have in common and what we could write. I think we, perhaps over a period of a year —the discussions probably took two or three years— but within the first year, I think we got to the point where we all agreed that there was a notion of the Extended Evolutionary Synthesis that could have heuristic value, which would be distinctively different from the Modern Synthesis, which would have different assumptions, which would have a different focus, different themes, make different predictions, and so on, and so forth. And that all started gradually to crystallize. And I suppose we were working on, at that time, on what would eventually become the Proceedings article. We never at that point had any notion of the Nature-style very short piece.
HS: Was the decision to submit to Nature unanimous? Did you have discussions on where you should submit this piece?
KL: Yeah, I mean, we did have discussions, because there was always going to be a tension between needing the space in order to spell out these complex ideas, and valuing the impact that publication in Nature or Science could bring. And I guess one way in which scientists resolve those tensions is to submit to Nature or Science but to have a huge electronic supplementary material where the bulk of the argument is placed. That’s what we tried to do. And whether that was ever going to work, I don’t know. I suspect probably not. But that’s a common strategy, and that’s the strategy we adopted. I think we just saw Nature as our first choice, the obvious place which could have an impact. And we did suspect there’s potentially a lot of interest in these ideas. So, we settled on that as an idea because we thought we had something to say. It wasn’t in our minds a huge stretch to go for Nature. We genuinely believed we had something really interesting and important to say. So yeah, submitting to Nature, I think, is what many scientists in our position would have done.
HS: What are your memories of actually putting this article together? Do you remember what each person brought to the article, who was taking the lead in writing it etc.?
KL: As I said, there was this inner core of myself, John and Tobias. I suppose I’d gotten into the habit of often finding myself the leader of these kinds of things. Scientists bring different talents to a collaboration, and different skills. There are many scientists who know more stuff than me, or are better mathematicians than me, or who’re more fluent speakers than me. But what I can do, I think relatively well, is write. I can write pretty clearly and in a way which people seem to appreciate and understand. And I can do it relatively quickly and easily. So, when it came to writing stuff, I was usually the person who took the lead. And so, given that I was taking the lead with the writing, it just naturally fell into place that I would be first author of the paper, and a leader. I think, in practice, probably we should think of myself, Tobias and John as the three leaders of the team.
HS: If you don’t mind my asking: some of the people who are part of the working group didn’t end up as authors on the papers. What happened?
KL: Yeah. Well, I think that’s quite interesting. Doug Erwin was one. Doug put a lot of time and effort into our manuscript, participated in discussions, gave us a lot of really good feedback. But in the end, he decided not to be a co-author. And I think, I mean – I haven’t actually asked him this – but I suspect he’s just a pretty astute guy. I don’t think all of us – certainly I didn’t – have this intuition that this could be very controversial. I mean, we had a sense it might have been controversial, because niche construction was a little bit controversial, but no intuition of the scale of controversy that would arise from that submission to Nature. I think Doug might have had that intuition. He might have just got a sense that some prominent evolutionary biologists would be quite critical or hostile of the EES, and maybe it was not a good idea for him to be seen to be a co-author. I suspect that might be the case for him. He is pretty savvy. And he was well-connected and had his ear to the ground. He was politically astute. So, I kind of think that might be why he wasn’t involved – but that is just a guess and I might be completely wrong. Marc Kirschner, we had more limited interactions with. He did give us feedback on at least one manuscript. But I think, bottom line, he didn’t like the idea of an Extended Evolutionary Synthesis. I think he wrote to us saying he wasn’t completely comfortable with the idea of synthesis at all, whether it was appropriate to characterize the Modern Synthesis as a synthesis and whether, to the extent that there was a synthesis, whether that was a virtue. And he was saying, I’m not sure that you should be pushing for an Extended Evolutionary Synthesis, and that maybe just encouraging more pluralism in how we think about evolution would be a more constructive way to think. To come up with some other monolithic solution that was replacing a pre-existing monolithic solution was, in his eyes, problematic, was not the answer, not the way to go. So, I think, even though he was intellectually aligned with our views, he didn’t like the idea of that kind of big theory. I might be putting ideas into his mouth, but that was what I vaguely remember was going on. Mary Jane is the most interesting one of the ones who declined in the end, because she went a very long way with us. She gave very detailed feedback on several rounds of manuscripts and took part in a lot of discussions. I think she really could have been brought on board. But there were certain sticking points, which we could never get over. One I remember was our definition of evolution. As I see it, from an EES perspective, if we’re recognizing extra-genetic inheritance as being important in evolution, and we recognize that things like epigenetic inheritance and cultural inheritance can affect phenotypes and can be subject to natural selection, that selection can operate on those additional forms of inheritance and bring about phenotypic change in that way, then we have to think about evolution as going beyond changes in gene frequencies. And I think everybody else in the group kind of reached the same conclusion. That was a logical endpoint of where we were with our discussions. But if I remember correctly, Mary Jane was committed to the idea that evolution was change in gene frequencies. I think she was perhaps more traditional, more conservative, in her view as to what evolution is, or should be, what was likely to be useful for the community, than some of us were. And, yeah, so there were two or three little things like that. Even though with 95% of what we’re saying she was on board, there were just some sticking points that we could never quite get over. And yeah, I suppose we might have, if we had a choice, we might have made compromises in the content to keep her on board. But that’s not quite how it worked out.
HS: Within your group of authors, was there agreement that this was something that needed a new name, i.e. Extended Evolutionary Synthesis? In the Nature exchange, one of the points that the other group brings up is: “We, too, want an extended evolutionary synthesis, but for us, these words are lowercase’”. What do you recollect about the discussions on this within your group? Were there any disagreements?
KL: A lot of discussion of precisely that. I mean, partly, our hands were tied, or at least I felt at the time my hands were tied, because we had sort of inherited the term. And, as I said, I thought it was appropriate to align ourselves with that initial push for an Extended Evolutionary Synthesis that Gerd and Massimo put together, and it wasn’t healthy to just constantly create new terms and new groups. We actually wanted to do the opposite. We wanted to create some sense of community. What we were hoping – I think we did achieve this – was in writing these papers and publishing them in a high profile journal that all these people who had hitherto been slightly alienated from a more traditional view of evolution would see that there are like-minded others around and this would sort of be a rallying call and that some sense of an EES community might emerge from all these different fields, all these different perspectives, in different countries and so on. And I think we did achieve that with the EES papers. So, we were reluctant to not use the term EES because we didn’t want to just constantly be creating divisions. We wanted to create a community and that was just the label that happened to be coined. It didn’t seem problematic to me. I was very naive, I think, in that regard. Clearly what we wanted to do was in some sense to ‘extend’ our understanding of how evolution works. Clearly, we’re interested in ‘evolution’. And clearly, we wanted some kind of ‘synthesis’ – a synthesis of developmental perspectives with evolutionary perspectives. So Extended Evolutionary Synthesis just seemed like an appropriate label for what we were trying to do. It never struck me that it was going to be that problematic. I can look back with the benefit of hindsight. Some people did seem to get very angry about the coining of a particular label, and a lot of significance was given to the fact that there were capital letters as opposed to small letters (EES vs ees). For me, that hardly is a big thing. But I think there’s also a sense in which those claims —that we don’t need to make a thing of it, we don’t need to give it a label—are also naive. Let’s say, we had call it ‘a developmental perspective’. It wouldn’t have the same impact. We couldn’t have got a paper in Nature. It had to have that sense of being something that could be appropriately juxtaposed with the Modern Synthesis. We went on to get a multi-million-pound grant from the John Templeton Foundation. We wouldn’t have got that with “an eco-developmental perspective on evolution”. It had to be a movement. It had to come across as a major departure from how people were thinking. So, there were pragmatic virtues in the label Extended Evolutionary Synthesis as well.
HS: Is this view – that the label was helpful in raising money, etc. – only in retrospect, or even as you were developing this, were you thinking about how this would make a difference in attracting research, in raising funds, in creating the space for research within this framework?
KL: I don’t think when we were writing papers, we were thinking about getting the grants and all that. We never imagined at that point that we might be able to get a multi-million pound grant from this abstract theoretical project. We were of course, very conscious of the fact that the label we used could impact whether we could get a publication in a high-profile journal, and whether or not people from different backgrounds would rally around that. So yes, we had a lot of discussion. And there were certainly some people who were pushing for not using the term Extended Evolutionary Synthesis. But if you don’t use that, you have to use something, and there was no clearly better alternative that we could agree on. And as I say, it seemed like an appropriate label. It wasn’t obvious to us that it was going to be hugely controversial.
HS: There are a few more names in the Acknowledgments section of the Proceedings paper that I would like to go over.
KL: Wallace Arthur: he’d done some nice work on developmental bias and he was a good person to consult. Some of these people we simply consulted, who weren’t part of the working group. Wallace Arthur fell into that category. The more people involved, the harder it is to get anything written, or to get anything done. We already had, I think, 10 or 11 people. We didn’t want to get too many more than that. And there was a sense in which someone like Wallace would have overlapping expertise with Gerd Müller and with Armin Moczek. So, we didn’t feel we needed him to be a central part of our discussions. But it was also helpful to get outsiders to kind of comment and critique what we had written. So, Wallace fell into that into that category. Patrick Bateson, again, falls into that category. Gillian Brown, again, falls into that category. Doug Futuyma: we invited to comment on this Proceedings manuscript. That was at a relatively late stage, after the Nature piece had been published.
HS: Had the others commented on the original manuscript?
KL: Well, prior to publication of the Nature piece, there was only one manuscript. We hadn’t anticipated that it will be split into two. And, in its structure and organization, it was more like the Proceedings article. And so, yeah, most of those people commented on that early manuscript. But Doug Futuyma we invited after the Nature piece had been published, to provide a commentary from the opposition, if you like. A couple of us knew him, Gerd knew him quite well. And we thought, clearly, he won’t agree with what we’re saying, but he’s a decent man, and he has integrity, and he will give us useful feedback, and make sure that we don’t say anything that misrepresents the standard view. That was our thinking in inviting him to comment. He was kind enough to do so and that was great. Scott Gilbert, again, someone we’d worked with previously. People like Pat Bateson, Scott Gilbert and Wallace Arthur were sort of friendly critics who could provide constructive feedback, but weren’t part of our discussions. Tom Morgan was a graduate student of mine, who was just interested in the issues, super keen and very bright, and provided some feedback.
HS: You have already written a backstory about the Nature exchange, so we won’t go into that so much. But if you don’t mind sharing one detail: who was the editor who commissioned this Nature piece?
KL: Patrick Goymer. He’s now the editor of Nature Ecology & Evolution. At the time, he was one of the editors at Nature. We’d written to him and submitted our manuscript. And, as I said, it was a relatively short manuscript with a huge ESM [Electronic Supplementary Material]. That was always slightly problematic, because so much of it was in the ESM. It’s kind of hard to make a case for an Extended Evolutionary Synthesis in just two and a half thousand words! We got these four reviews back. Two of the reviews were the most hostile reviews I’ve ever seen, of the 300 papers I ever submitted. And two of them were amongst the most glowing. So, it was an incredibly polarized response. But of course, with that kind of feedback, the editor is always going to reject the paper. But then, curiously, a few days after, he reached out to me and said, I’d been thinking about your paper, and how there were such polarized responses, would you be interested in doing a debate? And we ummed and aawed about it, because you know, there was something really paradoxical. The bottom line was he’d been not willing to publish our paper because so much of it was in the ESM, and we needed more space, but then he was inviting us to take part in a debate, where we would have even less space to present our views! But it’s hard to turn down Nature. And we knew there’d be a big impact associated with it being published in Nature. And I think, at the time, we didn’t necessarily think the debate would be polarizing. We didn’t know how it would pan out. We had no idea that we wouldn’t actually know who the other side would be, nor that we would not see what they’ve written, nor get any opportunity to comment on what they’ve written. It was not clear what the rules of the debate were. And so, we said “Yes”.
HS: Till that point, in conversations you were having with people, did you have any inkling that this would be so controversial, so polarizing, even?
KL: No. I look back and think, how could I have been so naive? But I honestly don’t think I did. I think we still had this mindset, that, you know, we were providing a service to the community – and I still believe that. We, of course, knew that there would be a bunch of people who wouldn’t agree with our position. But we were of the mindset that, even those people that don’t agree will sort of see virtue in us laying out what the position is, so there is clarity as to what it was, and then it can be put to the test, you can do experiments, you can do theory, and you can sort of sort between these positions, and people would see that’s a useful exercise, even if they didn’t agree with the views. And so, in that respect, I think I was quite naive.
HS: Were the reviews you received of the initial submission to Nature, the first time you got a sense of how…
KL: Yeah, the sheer emotion that this could generate surprised me. Took me aback. It didn’t make any sense. It was odd – you know, why are people so angry? Why is this such an emotive issue? I just couldn’t make head nor tail of it. It’s something I’ve obviously given a lot of reflection to since, but at the time, it was confusing. I just didn’t understand why people got so emotive about it, as if we were behaving in an unethical way.
HS: What’s your view now?
KL: Since that time, I have talked a little bit more to philosophers and historians and sociologists of science. I’m just a bit wiser about how this is actually not that uncommon. People have ways of thinking in science, and, oftentimes, they’re not so good at seeing the virtue in there being alternative perspectives. Not everybody has that dialectical biologist’s viewpoint that I have. It is much more common for scientists to think that they’re right, and that people who have different views than them are actually damaging to the field because they’re propagating misinformation, and that this is, in some sense, kind of ‘corrupting the minds’ of other people, particularly the next generation. And so, people with different views are often perceived to be kind of dangerous. And they wonder why would they be doing that when they’re clearly wrong, and jump to the conclusion that they must have some vested interests, some agenda, some political motive, some religious motive – as to why they espouse these clearly wrong views. And so, I guess I was naive in my understanding of how most scientists see the world.
HS: After this was rejected at Nature did you have a discussion about where to submit instead? How did you decide to submit to Proceedings?
KL: I guess we had these intense few months of working on the short exchange for Nature, but once the Nature piece had come out, we then turned to publishing the longer article. And I think, by that time, we’d seen the impact of the Nature debates, and that it clearly was very polarizing. I’d been avalanched with emails from supporters – over 1000 emails, and I got sent 50-60 manuscripts from people who sort of said “my views support yours”. It was really striking how almost all of those people that contacted me were on what you might think of as on the periphery of evolutionary biology. They weren’t evolutionary geneticists; they weren’t mainstream in that sense. They were developmental biologists, they were evo-devo researchers, they were palaeontologists, or botanists, or philosophers of biology, or human scientists. All kinds of people who were using evolutionary theory and you might think of professional evolutionists in some sense, but they were not mainstream evolutionary geneticists or quantitative geneticists. And so, to the extent that we had support, it seemed to us that it came from that fringe of evolutionary biology rather than from the center, which was very interesting. But that strong support also made us double down in our thinking that EES could be a useful concept, that we did have a constituency, that there were people out there who liked what we said, and who could unite under that banner. There was some misinformation that had been put out there in the Nature debate because the opposition had sort of characterized us as ‘revolutionaries’, as believing that the field had “ossified”, and certain things like that. We were really keen to correct that misinformation. A lot of that view had been propagated on social media. No one in our team was on social media. We didn’t have a means at that time to counter those views, so we were keen to publish the second paper, the more substantive paper. The Nature piece is very brief, it’s just a few hundred words, and it might give the impression that this was a bit of a back-of-an-envelope idea that had been thrown out by a few people who’d one weekend come up with this EES stuff. We were determined that we get across all the hard work that had gone into it – in fact, over three years of this working group – to sort of flesh out what the EES was. We needed to get that big paper out and we really pushed hard, once the Nature piece was done, to do so as soon as possible. I think we initially submitted it to Biological Reviews, because we thought, well, okay, we need a large amount of space in order to publish this work. And again, we got an incredibly hostile response from the editor who didn’t even send it out to review. He just didn’t like what we were doing, thought it was inappropriate and it wasn’t going to be published in his journal! So, our thoughts turned to Proceedings of the Royal Society. We wanted to publish in a general journal. Proceedings of the Royal Society B was a very good one, very prestigious one. The Reviews Editor, Per Lundberg, who was a colleague of Tobias’s at Lund, an evolutionary ecologist, I think, was without any particular vested interest in this debate and seemed fairly open minded on these issues. On one hand, he didn’t really agree what we were saying, but on the other hand, he thought there was virtue in spelling it all out. I think Tobias sounded him out. And yeah, he was kind of open to the idea of us publishing a review in Proceedings of the Royal Society. And, in fact, he pointed out that there was this category of Darwin Review. They have one review every year, which is called the Darwin Review, which is like their most prestigious article. And this would be helpful to us, because it would allow us a little bit more space to present our ideas, as well as having that prestigious label. So, we submitted our article. We had to go through an extra kind of level of checks and balances to just ensure it was suitable as the Darwin Review, but we passed that particular hurdle. We again had four reviews. Two were solicited from people who were not sympathetic to EES and two were solicited from people who were likely to be EES sympathetic. The reviews were useful and constructive, and it was eventually published. And then, we had another round of people being furious. Some very traditional minded fellows of the Royal Society thought it was outrageous that these ‘charlatans’ had been allowed to publish in the Proceedings of the Royal Society; they shouldn’t be allowed! Some of them actually contacted the Managing Editor of the journal complaining about the Reviews Editor in allowing these idiots to have the Darwin review, and that it was a terrible mistake! I heard that some of these eminent geneticists wrote to Per Lundberg and complained about his behavior. And he said, well, how about I let you write the next Darwin review, next year. And you can present your views and it’d be an interesting pair of papers in juxtaposition. And so, they agreed and it got published. This was the Charlesworth, Charlesworth and Barton paper. And I heard that the reviews editor took exactly the same balanced approach with their paper that he had taken with ours. He invited a couple of people on the EES side to review their paper (I was one), and he invited two other reviewers who were likely to be sympathetic to their viewpoint. So, the same sort of even-handed, balanced perspective. I was shocked by what was in the original draft, and later learned that the other EES-sympathetic reviewer had been too. I remember, in the original draft of their manuscript, they had compared a belief in animal social learning and animal culture as equivalent to Lysenkoism! I was stunned by such an extraordinary claim. There’s a part of me that sort of regrets that I complained about it and that eventually it ended up being removed from their paper in the published form; had it been published, it would have illustrated the extremity of their reactions. What was kind of interesting about this was that, when they received our feedback on their draft, they were not happy that the Reviews Editor was expecting them to respond to our reviews! And they complained to the Managing Editor, who was a mate of theirs, who then decided to take over responsibility for that paper and told them that they didn’t really need to consider the reviews from us, because we were clearly wrong. Subsequently, I heard that the Reviews Editor had resigned over this issue. He felt undermined by the Managing Editor jumping in and telling the authors that they don’t need to consider the reviewers’ views. So, there was another interesting dynamic going on in the backstory of that particular paper too!
HS: You said you received a lot of responses in support of your position in the Nature exchange. Did you also receive negative reactions?
KL: No. Like I said, I got I think over 1000 emails, but only one negative email. I don’t mean to imply for a second that there weren’t people out there who did hold negative views. But I think those people wrote to the other side and said “we support you” rather than write to us saying “we disagree with you”.
HS: And another part of the story is the meeting you organized subsequently…
KL: The Royal Society meeting. Yeah. Denis Noble was the lead organizer of that. Pat Bateson was also an organizer, as well as Nancy Cartwright, John Dupré and me. There were five of us. And yeah, so again, Dennis Noble thought it was useful to have a discussion of these issues, and we got together a list of potential speakers representing a breadth of views. And it went through the usual review process at the Royal Society. In fact, because we had proposed a joint meeting of the Royal Society and British Academy, it actually went through a far more stringent review process, because we had to satisfy both academies with our plans. However, we got glowing reviews. Everybody thought the meeting was a good idea, that it was timely, and so on. And, yeah, well, obviously, some suggestions were made as to how we could improve it and tweak the program to accommodate this or that, but, you know, by and large, the reviewers were positive. We had high-caliber feedback from people, and our application went completely through the review process in the appropriate ways. The meeting was announced, and again, this same group of traditionally minded evolution geneticists got very upset that a Royal Society discussion meeting had been given to these ‘obviously wrong’ people! And they wrote to the President of the Royal Society and said this meeting should not be allowed to happen. And I’ve seen the letter that they wrote, because it was shared with Denis and with the other organizers, and it described the meeting that we were organising as equivalent to the Royal Society organizing a meeting on homeopathy! That again illustrates the extremity of their position. But of course, eventually, the Royal Society said: look, this has been through all the procedures, it’s been appropriately reviewed, there’s no reason for it not to get ahead. We ended up having a couple of meetings with one or two of the hostile people. Some of them were so angry that they refused to meet with us. And we ended up inviting people like Russ Lande – actually Russ Lande was always invited. But we invited Doug Futuyma to come along and speak just so that those more traditional views were more represented at the meeting. But yeah, that’s another illustration of the strength of feeling that the EES generated.
HS: It’s been 10 years since these papers were published. Do you think that the strength of feeling has sort of dampened down now or do people still feel strongly about it?
KL: Some people still feel strongly. I think in the wider community, a large fraction at least, must now see some virtue in the EES papers, and particularly the Proceedings paper. I think there is a subsection of the community who did respond the way we kind of anticipated, who see virtue in laying out – these are two alternative views, these are the expectations or predictions, these are assumptions, let’s put them to the test. And the fact that these papers have both been cited over 1000 times – the Proceedings one maybe 1500 times – must mean something. Almost all the citations are positive. It must show that there are a lot of people out there who see some virtue in those papers and in laying out what the EES is, or might be. And I think that the views expressed in those papers, in large part, either have become mainstream, or they’re on the way to becoming mainstream. The various ideas that we champion in those papers – the importance of plasticity-led evolution, of developmental bias, of extra-genetic inheritance, and niche construction, and evolvability – all of those ideas, I would say, are either mainstream now, or on the way to becoming mainstream, or at least mainstream in some parts of evolutionary biology. So even if the label ‘Extended Evolutionary Synthesis’ is not in itself fully embraced by the community, and even if there is some residual hostility to that label, the ideas in the papers are not evoking anywhere near the same level of hostility. It’s also true that there’s still some way to go for the community to embrace the developmental perspective. If you take plasticity-led evolution, for instance, there’s a huge amount of work on this topic in the field, but how much of it really embraces that Mary Jane West-Eberhard way of thinking? So much of it is re-characterized as the natural selection of genetically specified reaction norms, which allow for plastic responses to various different aspects of the environment. And so then, if you’re thinking of plasticity-led evolution as just selection on genes, that is not really bringing development into the story. So, there’s a little bit of that that’s going on – some of these ideas have been taken on board, but they’ve also been reorganized to fit with the traditional gene-centric viewpoint a lot of the time.
HS: In the concluding section of the paper, you hope that the EES will have value in bringing together researchers from diverse fields who share an “ecological developmental perspective”. How successful do you think the EES has been in this regard in the last 10 years?
KL: I think it has been successful in that regard. In those communities, the EES is not contentious. John, Marc, and I, because of our slightly eclectic backgrounds, have always had a particular interest in humans. Most evolutionary biologists have not got particular interest in humans, but we do. And that’s where niche construction came from. Even Dick Lewontin had a particular interest in humans. He wasn’t, I think, always comfortable about using evolutionary arguments to interpret aspects of human behavior and cognition, but he was very conscious of those potential implications and that was part of his sensitivity. Conrad Waddington, again, another sort of hero of development plasticity and niche construction theory, was interested in understanding humans and the human condition and had very prescient writings on that. So, I think it’s always been a part of that niche construction perspective, that we were very interested in understanding human evolution. Because of that, we’ve engaged with human scientists from a number of disciplines —anthropology, archaeology, evolutionary psychology, primatology, and one or two other fields, — and we came across a lot of dissatisfaction with the kind of evolutionary explanations that were being provided by a more traditional viewpoint on evolution. Human scientists often saw those traditional accounts as too gene-centric, too gene-deterministic. And conversely, the kinds of things that we were offering, with our emphasis on niche construction and developmental plasticity, was much more appealing to them. We were at pains to point out we weren’t arguing for genetic determinism in explaining human niche construction. We talked in terms of cultural niche construction, how it could modify select environments and generate feedback to influence genetic evolution. That was a kind of story that I think a lot of human scientists felt comfortable with. I think that experience of interacting with humans scientists in a number of fields prior to engaging in this work on Extended Evolutionary Synthesis, has given us this perspective: that there was a need in the eyes of human scientists for a richer vision of how evolution works, one that recognizes that development was important to the production of phenotypes and the patterns of selection that would operate on populations, but also the capacity to modify environments and the feedbacks from those self-constructed features, and the importance of extra-genetic inheritance – I mean, clearly, you can’t study humans without recognizing the importance of extra-genetic inheritance because ‘culture’ is right there in your face. But also, humans are a case where you can’t ignore niche construction. Again, it’s right there in your face: humans are nowadays often characterized as champion niche constructors. And clearly for humans any crude genetic determinism, where you are explaining that capacity of a niche construction in terms of naturally selected genes, and that alone, doesn’t work. So, the virtues of the developmental perspective, are immediately apparent, I think, to many researchers who study humans, in a way that they might not be to somebody who studies mice, Drosophila, or yeast.
HS: In hindsight, do you think it was a good thing that this got published as two papers – the exchange in Nature and the Proceedings paper, instead of just one paper in Nature?
KL: Yeah, I do. I think we ended up getting the best of both worlds. I mean, there were negatives to it, for sure. I think the principal virtue of the Nature piece is that we reached a huge audience with that article. And, in a sense, it allowed us to put the EES on the map and to get people to take it seriously and rally around it. It was probably necessary, in a way, to write in relatively simple terms, non-technical terms, in order to achieve that. The negative, of course, was that it polarized the community. I have some misgivings about that, but that was something that perhaps couldn’t have been avoided. But returning to the benefits of having two papers. The Proceedings paper allowed us to really lay out these ideas in depth. It was a more technical piece, so we could reach that evolutionary biology community, and show that there are real well-thought-out ideas behind this concept of an Extended Evolutionary Synthesis. It wasn’t just some fluffy notion. And actually, that paper has been even more successful. So, I do think we ended up benefiting by having a pair of papers. It was kind of like a boxer going wham-bam, with two punches, you know. There was more impact from two punches that followed quickly after each other than a single punch. Neither of those things would work on their own. I think the Nature piece on its own, would have polarized the community, but then the whole thing would have died down. And there would have been no impact because there was no clear substance to the ideas. The fact that that Nature piece was followed swiftly by the Proceedings piece, where the substance was put there right in front of peoples’ noses, meant that it had to be taken seriously.
HS: You mentioned that there are some negatives.
KL: Sorry, I didn’t mean to say that there are negatives associated from having two pieces instead of one, but rather the negatives associated with the Nature piece sort of polarizing the community. I suppose it didn’t have to be that way. It was partly the way that the Nature editors chose to do this. I mean the fact that we didn’t know what the opposition was saying, so we didn’t have an opportunity to correct some of the claims they were making about our position. So, they were able to present us as extreme outsiders who wanted to throw everything out and start again, who were calling for a revolution, who thought that evolutionary biology was stagnant. These were not our views. These were their false characterization of our views. But I suppose if this kind of debate was ever going to take place, it had to be done that way. You couldn’t allow us to respond to them, and then they attempt to respond to us. It would have led to tit-for-tat, re-editing, and rewriting of each piece that would have gone on forever. And I can say that with some experience because I have written collaborative pieces with people who disagreed with me. It’s not easy. In that case, we went through something like 20 rounds of revision, where you sort of adjust your position, then they adjust theirs in response, and then you read just yours, because you kind of want to win the debate! And you have to get past that and say, actually, it’s not about who wins, it’s just about presenting these two positions and let other people judge. I think Nature were never going to let us respond to each other’s pieces. So, we never saw the opposition’s argument until it was published. But clearly, they too, can’t have seen our final version because the final version of ours was finished after theirs had been written. We were told that by the editor. So, they never saw what we wrote in the end. So inevitably, that meant there was a certain amount of talking past each other. And they were reacting to what they thought we were saying, rather than what we were actually saying.
HS: Do you have a sense of whether these two pieces have attracted different audiences? Have you looked at which paper cite them, or do papers tend to cite both of them?
KL: I don’t know. I suppose some historian of science could do an interesting project, analyzing the citations of these papers. I do think there are biases still in the claims of people who are citing both the papers. It’s probably still the case that hardcore evolutionary geneticists are not citing either of those papers, or at least not so much, and probably not in a flattering way. I do think that the Proceedings piece is often cited by practitioners in the field – evolutionary biologists who are doing tests of plasticity-led evolution, developmental bias, whatever. So, certainly a significant fraction of those citations of the Proceedings piece would be people we would think of as professional evolutionary biologists.
HS: I wanted to ask you about a pair of citations in a sentence in a section about niche construction in the Proceedings paper: “these findings have led to the claim that niche construction should be recognized as an evolutionary process through its guiding influence on selection [citation], a position that is contested by some evolutionary biologists [citation]”. I was curious because you’re an author on both the paper! Is there an interesting story here?
KL: Oh yeah. I have written an article with critics of niche construction theory. That ended up being published in Evolution, led by Thom Scott-Phillips. There are five authors on that paper, and the other four are critics of niche construction theory. Yeah, so the story, if you want to know the story of that one, is that the other authors originally wrote a manuscript critiquing niche construction theory, submitted it for publication, and sent it to me, after they sent it to be reviewed – partly because, I think, one of the authors Tom Dickens, I had worked with very closely. He was an evolutionary psychologist, and I’d worked with him in setting up a new academic society, the European Human Behaviour and Evolution Association. We were friends, we got along well, I liked him. There was a kind of mutual respect, even if we disagreed on a lot of stuff. So out of courtesy – I imagine that that’s what it was – out of courtesy, he felt he should send me the paper. So, he sent me the paper, or arranged for it to be sent to me. And I felt like this was not a fair reflection on what niche construction theory was or claimed. They were knocking down a straw man. I wrote back to him saying, look, I really think you are being unfair; and pointed out my concerns with that article. I said, look, why don’t we just try and do something constructive, and work together to write a paper saying what niche construction is, and what your concerns about it are, and how I would respond to those concerns. Why don’t we just write that and then that’s useful for people, rather than you writing something which I think most people who are sympathetic to niche construction theory would think is just a straw man critique, or people talking past each other. I think there’d been a similar big debate in the pages of Nature or Science over group selection and kin selection, where there were two sides that just went hammer and tongs with each other and they talked past each other. That wasn’t very helpful to people to try and understand what was going on. So, I tried to do something that was more useful. And to their credit, they responded positively to my invitation, to collaborate. They withdrew their original paper. And we ended up having protracted discussions and the paper in Evolution was the product of that. I think I’m as proud of that paper as pretty much anything I’ve done in my career, because it was so hard. It really was very, very hard. And I think, it was like 21 rounds of revision!
HS: Interesting story. I’m glad I asked you about it.
KL: But it shows it can be done. And I wish more people would try and do it.
HS: Kevin, I just have a couple more questions. One is, after the publication of these two papers, if you look back, how much has EES influenced the trajectory of your own research, and also what impact has it had on your career? Has it helped? In what ways has it helped?
KL: To answer the first question, I think it has definitely shaped the direction of my research and my thinking, and I still I don’t feel like I’ve finished with – you know, you see this as a pair of papers – but going right back to 2014, it was really clear to us that you couldn’t make a case for the Extended Evolutionary Synthesis in a convincing way with 800 words or even the six or eight thousand words that we had in the Proceedings piece. You needed much more space, because these debates are about more than the science. You needed to get into the philosophy of science. You needed to know the history of evolutionary biology to make sense of it. You needed to know about evolutionary genetics, but you also needed to know about all these new ideas and new findings that are pouring out of the research laboratories on epigenetics, and holobionts, and inherited symbionts, and horizontally transmitted genetic information. There was so much that went into whether you would be sympathetic to an EES perspective, that clearly a book length treatment would be required. And so, since those papers were published, I’ve been working on that book, and it’ll come out this year. That book – ‘Evolution Evolving: the Developmental Origins of Adaptation and Biodiversity’ – is really the third part of this EES story, at least, with respect to my involvement in it.
HS: Has it been 10 years in the making?
KL: It’s been 10 years in the making. It took a lot of work to pull together. Even though I think that the causal structure of evolution theory that we outline in the Proceedings paper is not really very different from how I think now – it’s pretty much identical – it took a long time to really think through how we can make that case. Of course, this has coincided with other things going on, with Covid, with health issues I’ve had, with various complications which have delayed it. But yeah, the third part of my EES journey will eventually come to fruition. It has kind of continued to dominate my thought, my research, since that time.
It’s kind of hard to judge its impact on my career. Certainly, it would have had mixed impacts on my career because of that polarized response. There were some very eminent biologists who sadly seem to ostracize me and other members of the EES community, who reacted in that negative way, and that might have had impacts on our chances of getting key publications and grants and winning prizes, and all those kinds of things. You never know. It seems likely that there would have been some negative impacts. But also, the success of those papers created the opportunity to apply for a mega million pound grant from the John Templeton Foundation, and several subsequent grants as well, which have supported the writing of the book. We got this big grant from the John Templeton Foundation that eventually ended up in over 200 papers being published. This huge consortium of something like 60 scientists, eight institutions, doing cutting edge research, related to the Extended Evolutionary Synthesis. The grant is called “Putting the Extended Evolutionary Synthesis to the test”. Loads of great papers came out of that, including in some of the best high-profile journals. It was incredibly successful. Plus, through the grant we organized successful conferences and workshops.
I mean, there’s a sense in which those people who eventually united under the EES banner were already there, but they didn’t know that there were other people who were like-minded, and with the publication of these papers, a sense of community, I think, emerged. Certainly, a lot of people identified with that developmental perspective on evolution. I think that’s been a very positive outcome spanning across a whole bunch of academic fields. It still remains the case that sympathy for the EES is dominated by people on the periphery of evolutionary biology. But I think even in evolutionary genetics – I’ve had some dialogue with some of those people – you do get the sense that there is a little bit of a coming together of the two communities and less polarization. There was an edited volume that Tom Dickens, the guy I was talking about earlier, and others put together, which was kind of critical of the EES and defending the Modern Synthesis, the traditional viewpoint. That was published last year I think, and my take on that is actually there’s been a shift in the critics’ position towards the EES. It’ll be interesting to see how, when our book is published this year, whether they similarly see some shift towards their position; or perceived shift, at least. So, yeah, I do hope that there is an opportunity down the line for reconciliation. I can actually foresee two kinds of futures, and both are quite positive: one would be in which the Extended Evolutionary Synthesis viewpoint, the kind of views that we champion in those papers, become mainstream. And that people take on board the points we were making about the importance of development in evolution. Another is sort of watered down version of that. Actually, it’s kind of already clear that – plasticity-led evolution, extra-genetic inheritance, developmental bias – these things are here to stay within evolutionary biology. That’s a watered down version, in the sense that development is still not quite really on the table; people aren’t thinking in a developmental way. And if that’s what we achieve, well, that’s still something. Of course, we don’t know how much we contributed to that journey, but given the sheer impact of the papers, the amount of citation, suggests that we have contributed to those ideas that we championed becoming more part of the mainstream. Either way, it seems like positive end to the story. It’s just a question of how positive.
HS: Final question. For someone who has published over 300 papers, this is probably a difficult one: do you have a favourite paper?
KL: Oh, gosh. I’d have to think. Maybe that article I mentioned earlier “Cause and Effect in Biology Revisited” which we published in Science. I’m proud of that one because it helps makes sense of so many debates spanning the biological and social sciences. It also set me on a path to thinking about the developmental bases of adaptation and biodiversity.
HS: Not one of these EES papers?
KL: No. It will probably not be one of those. But it’s true that I’m very proud of those two papers too. I can look back and think about the huge amount of work went into them. And that – certainly in the Nature piece – that might not come across, but a huge amount of work went into, not just the writing of them, but also the thinking through of the ideas that preceded that, and pulling those different themes together, extracting those themes. I do feel proud that we made those contributions.
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