Revisiting Love 2003

In the early days of evo-devo, the split between evolution and development was seen primarily as one between genetics and embryology respectively. Evo-devo, therefore, was characterised as an attempt to bring together evolutionary theory and embryology using developmental genetics. In a paper published in Biology and Philosophy in 2003, Alan Love provided an alternate way of thinking about this synthesis of evolution and development that focussed on functional morphology and comparative anatomy, which, Love argued, were important for understanding evolutionary innovation, a key concept in evo-devo. Twenty-two years after the paper was published, I spoke with Alan Love about the origins of his interest in evo-devo, his memories of the making of this paper, and his reflections on the growth and evolution of evo-devo over the last two decades.

Citation: Love, A. C. (2003). Evolutionary morphology, innovation, and the synthesis of evolutionary and developmental biology. Biology and Philosophy, 18(2), 309-345.

Interview conducted online on 3 March 2025; Alan Love was in Minneapolis, USA and Hari Sridhar in Klosterneuburg, Austria.

 

Hari Sridhar: I would like to start by talking a little bit about your motivation to doing the work presented in this paper, and also how it sits in relation to your PhD more broadly.

Alan Love: It is fun to think back to when I was working on this paper. The paper was, in many ways, reflective of some key themes in my dissertation, which at that point was not finished yet. What you’re seeing in the paper is some of those core themes starting to crystallize for me, in terms of what I wanted to emphasize, both historically and philosophically, about evo-devo. In the context of my dissertation, what I went on to do was offer more historical backdrop to some of these questions, and then a philosophical gloss on how to better understand ideas about synthesizing evolutionary and developmental biology in terms of problem structures rather than overarching theories. In a sense, we might boil down the core components to the following. One is that the history had been, at least at that time, unduly narrowed to a history of developmental genetics, in part due to the discovery of HOX genes and the amazing technological progress that was made possible through experimental developmental biology, through the late 80s and early 90s. I was trying, in particular, to put pressure on that point, through the lens of morphology, because morphology had been a major instigator of bringing evolution and development back together prior to the discovery of the HOX genes, especially through the mid-1970s. Philosophically, I wanted to leverage this insight to show that what one needed for a synthesis was going to be more than just putting together population genetics and developmental genetics. A number of people at the time had tended to frame the juncture between evolution and development as primarily constituted by those two entities. Morphology suggests that we need higher levels of organization included in the conversation. We also need to make sure that we’re taking into view function, and recognizing that there’s a structure-function relation that must be contemplated. Both of those things were arguably not as salient to a number of people who were thinking about the synthesis of evolution and development at the time. In some ways, looking back on the paper, it’s an approach where I’m quoting a lot of different individuals because I’m trying to make the case for this missing history and this reconceptualization of the philosophical basis of the synthesis. At the same time, it’s something that arguably remained a strong current, not only in my dissertation but even beyond that, in the work that I’ve done over the last two decades in thinking about evo-devo.

 

HS: Stepping back a bit, how did you get interested in this evo-devo?

AL: When I was a graduate student in the history and philosophy of science department at the University of Pittsburgh, I was casting about for interesting topics very early on, and I started reading a lot in evolution and development, but my reading was skewed towards earlier work. As I was following citations and reading more material, I was pulled not in the direction of the latest developmental genetic discoveries, but back towards some of the work from these evolutionary morphologists—people like David Wake—and seeing their ability to weave together different disciplinary approaches into a stronger overall package of explanation. That was very interesting to me, because I first of all knew that disciplines didn’t necessarily work together that well. It isn’t obvious that it should happen. Additionally, the evolutionary morphologists in these traditions seemed to recognize that the problems they were working on demanded this kind of multi-dimensional approach. As I was reading in that literature, evo-devo was exploding onto the public scene in a more visible way. You had the formation of the subdivision of evolutionary developmental biology within the Society for Integrative and Comparative Biology, and new journals like Evolution & Development that had been recently inaugurated. It was very interesting to say: ‘Wait a second, something is seemingly missing between what people are saying and what happened.’ That was the impetus for my dissertation project. And that reading came through what you might call old-fashioned reference chasing: ‘Oh, that’s interesting, they refer back to this paper, what does that paper say?’ I just followed it back and found that I was in the midst of literature that was less prominent in some ways, but very interesting conceptually and therefore attractive to me to investigate further.

 

HS: At the time when you joined graduate school, did you already know that your PhD was going to be in evo-devo or was it much more open than that?

AL: No, it was much more open than that. I was trained in yeast genetics as an undergraduate. My undergraduate major was molecular biology with a minor in philosophy. I did not have a lot of organismal biology, evolutionary biology, ecology, or any of these things. It was very much a molecular orientation. And so, you might argue that, in some ways, I was following a natural pattern: what is this thing that I’m not that familiar with, that seems to be so important for the conceptual basis of this project? A lot of my own learning process was happening in that period, as I got interested. I went on to Indiana University, Bloomington, to do a Master’s degree with Rudy Raff, working on sea urchin evo-devo, to immerse myself in some of the actual scientific work, to become more familiar with it.

 

HS: There are a couple of papers that you wrote with Rudy Raff around that time (1, 2). Was that work from your Master’s?

AL: Yeah, primarily. I took a leave of absence from the University of Pittsburgh in 2002 and spent two years at Indiana University. Rudy and I did a combination of papers there. Some of those papers were more empirical, some more conceptual. That was a very fruitful period for me, because I was both able to dive into the science, but I was also able to work with somebody who had a living memory of a lot of that history from the 70s and 80s. That was a spur to my own work. And the dissertation wasn’t finished until 2005, after I returned to Pittsburgh from Indiana University, and so I was able to fully incorporate that experience into my dissertation work. And I think that made for a good, healthy combination of history, philosophy, and science within a single dissertation, and all oriented primarily around the concept of innovation and novelty.

 

HS: You attended a seminar and workshop, both at Woods Hole, which seem to have had some role in the background to this paper. Was this paper published in a special publication that came out of these meetings?

AL: Let me say a little bit about the workshops, and then I’ll come back to the publications. The 2001 workshop, ‘From Embryology to Evo-Devo’, was a part of the ongoing series that happens at the Marine Biological Laboratory (MBL) in Woods Hole each year with different themes. I attended this one as a graduate student. It was a week-long event, so lots of sustained time for conversations and interactions. Lots of people were there, Ron Amundson, Jim Griesemer, as well as scientists like Rudy Raff and Scott Gilbert. In fact, that’s where I met Rudy and we started talking about the possibility of me going to his lab in Indiana. I didn’t present the paper there. Rather, I was an attendee, mainly listening to other people’s presentations. But I had been developing this material for my dissertation project, and a follow up workshop was held a year or so later. That second one was not at Woods Hole. It was at the Dibner Institute at MIT, and it had the same title, ‘From Embryology to Evo-Devo’. Eventually, the presentations there, in revised form, came out in the book ‘From Embryology to Evo-Devo’, which was published in 2007. That was where I was able to share some of the historical work that I had done in the dissertation, which was not in the 2003 Biology & Philosophy paper. What the workshops meant for me, in terms of developing the material—especially the 2001 MBL Woods Hole meeting—was that they helped me triangulate on the importance of morphology, the importance of paying attention to different levels of organization, and the way in which the historical story was important for recognizing how people thought about the present. In many ways, it was a great push to me as a graduate student that, yes, this sort of approach can be fruitful for thinking about contemporary science, that is, by stepping back into history and understanding how we got to where we are, and what was forgotten in some of the presentations of the field by contemporary practitioners. Those two meetings definitely were very catalytic, and came at a good time for me as a graduate student.

 

HS: And what about this paper? Did it have anything to do with the meetings?

AL: The publication in Biology & Philosophy did not have anything to do with those meetings. There was a lot of conversation going on among people about evo-devo at the time. There was a special issue planned for Biology & Philosophy, and I was invited to submit my paper, based on presentations I’d made on it at a couple of other events. In many ways, it was very much affected by all those meetings that were going on, but it was not a direct consequence of them.

 

HS: You describe the approach you took in this paper as a phylogenetic approach. At that time, was such an approach novel? More broadly, could you talk about how you chose the approach to the questions you were asking?

AL: I don’t think the phylogenetic approach was by any means unique or new. I was following how my advisor, Jim Lennox, had framed it in his own work. But it was also very compelling to me, because the phylogenetic approach suggests that there are patterns in the history that one needs to analytically recover. That is, the patterns are not obvious, they’re patterns that you have to dig out through certain kinds of methods, but once you do, those really inform the present. Just like when we think, ‘Oh, why does an organism have the structure it does?’ If we understand the history, if we understand how it got there, we really are in a much better position to analyse and understand it. So too with the history of science. It’s not a causal study; sometimes, the various causes for those past events are not necessarily reflected in the phylogenetic pattern, and in fact, can be subject to debate. That was something I was especially interested in because it seemed to me that the kinds of causal factors that were relevant could be quite variable. It could be somebody being persuaded by an argument. It could be somebody doing a key experiment. It could be somebody changing funding mechanisms at a funding agency. It could be that a particular paper is published and becomes popular and leads to a trend of research. From my vantage point, all those things were possible causes, but what was important was to understand how the pattern manifested in the present, which gives us insight into why people thought what they were doing was configured in the way it was. I don’t think that any of that is unique. There are lots of philosophers who adopt that kind of approach. I think that I was very self-conscious about it, at the time, because I was working on my dissertation and thinking very clearly about what methodology I was using to approach my subject matter.

 

HS: You cite close to 200 different articles in this paper, covering a wide range of sources: journals in different fields, conference proceedings, books, etc. How challenging was it at that time to access all of this literature?

AL: The first thing to recognize is that it was a pre-PDF period! One still went to the library and photocopied journal articles. I still have physical copies of a number of those cited articles that I read and marked up. I think that what you’re seeing in this paper is really very cumulative. It had to do with multiple years of this material really stewing in my own thinking, and obviously also in preparing to write a dissertation about it. So, in some ways, there was just standard labour invested to collect those different sources. It didn’t go fast, but it didn’t go slow; it just took time. And I think that, in some ways, you might worry that there are too many references in the paper. However, at that moment, I had felt it was important to put forward this diversity of voices and this range of material all in one place. It definitely was not written quickly; it was the slow simmering stew of scholarship. This kind of work did simply take longer than it currently does, because you couldn’t access that material online like you can today.

 

HS: What else do you remember about the writing? Did you work on it only after you’d received the invitation from the journal, or did you already have some kind of a manuscript ready when you were invited?

AL: I had some bits of a manuscript, but definitely, once I had the invitation, I was more focused on drafting it. There was an interesting moment where the special issue editors asked me to shorten the first draft I submitted because it had been too long. Then, the irony was, when I shortened it, they came back and said, “you shortened it too much, and you need to put back in some of the stuff you had.” In a funny sort of way, the special issue nature of the publication was probably helpful because they were looking to put forward papers that were really illuminating the situation with evo-devo for a larger audience, and my paper was doing that in a unique way. If you go back and look at that special issue, you will see that I’m the only one focused on morphology, so I’ve got a distinct angle that I’m offering in the space. But I don’t actually have a lot of distinct memories about the peer review process. It was peer reviewed, and there were comments, but I don’t remember anybody saying, ‘this is completely wrong,’ or something like that. It was, I think, a very standard request for, ‘more detail here,’ or ‘more clarity there,’ or, ‘shave this part down.’ The revised paper benefited from the peer review process, but it didn’t have anything that made one keep it as a memory. Some peer review processes can actually be quite salient in people’s memories!

 

HS: I’d like to go over the names in the Acknowledgements to get a sense of who these people were and how they contributed to this study. The first name is Ron Amundson.

AL: Ron was, arguably, the first philosopher to really get interested in evo-devo, and had been writing on it for quite a while. Woods Hole was the first time I met him. We subsequently became correspondents and interacted a lot over the years.

 

HS: The next name is Ingo Brigandt.

AL: Ingo was a fellow graduate student at the University of Pittsburgh who was also working on evo-devo topics. His dissertation focused on homology. He and I went on to be co-authors on a number of subsequent papers.

 

HS: And then, Graham Budd.

AL: Graham Budd is a palaeontologist and morphologist. I got into contact with him during this period precisely because he seemed to be somebody who was a practitioner of this approach that blended together different disciplinary approaches. He read an early draft of the paper.

 

HS: Greg Davis.

AL: Greg Davis is a biologist who works on aphid development and evolution. He had been at the University of Pittsburgh for a Master’s degree in history and philosophy of science before he went on to take his PhD and go on in science. He had the same advisor as me at Pitt, so we had this connection. And then, because I was starting to circulate in evo-devo settings, and he was in them himself, he and I engaged in a lot of conversations and discussions that were very generative for this paper.

 

HS: Paul Griffiths.

AL: Philosopher of biology who was at the University of Pittsburgh up until 2002, or maybe a little later, and then moved down under to Sydney. He was somebody who I’d taken graduate seminars with. He was working on questions about developmental systems theory, which was another approach at the time that was thinking about the relevance of development to evolution. He was very much a key influence on me at that time.

 

HS: Brian Hall.

AL: Evo-devo biologist and historian. Definitely somebody who I got to know in correspondence and at some meetings at the time. He was also at the ‘From Embryology to Evo-Devo’ MBL Woods Hole workshop. He’s somebody I learned a lot from, who had much of this recent history in view, and also himself represented this synthesis of different disciplinary approaches. He worked on topics like bone and cartilage, and so he was sensitive to the movement back and forth between different levels of organization in analysing evolution and development.

 

HS: Jon Hodge.

AL: Jon Hodge is a historian. I got many helpful pointers about how to think about historical work from him. He had helped me on a previous historical paper on Darwin, and was very good in guiding me to think methodologically about my historical work. I think of myself primarily as a philosopher, but wanted to do good history as a part of this phylogenetic approach.

 

HS: How did you know him? Did you just reach out to him?

AL: He was a connection from Robert Olby, the historian of genetics and molecular biology, who was on my dissertation committee, who I’d also taken seminars with. I was put in touch with Jon by Bob.

 

HS: Jim Lennox

AL: My advisor. I don’t know if I need to say much more than that. As a PhD advisor, he was very central to me taking up the topic and thinking about these things, both from a methodological standpoint, but also in terms of helping me keep in view the philosophical import of the historical work.

 

HS: Lennart Olsson

AL: Lennart Olsson is a biologist, originally from Sweden, who now works in Germany. He was another biologist who I interacted with a lot in that early period, who also clearly recognized that there were interesting historical precedents for current work. He also himself displayed this interest in morphology and how to incorporate that into contemporary approaches and not be completely skewed towards genetics.

 

HS: The next name is Jason Robert.

AL: Jason Robert was one of the special issue editors. He was collaborating with Brian Hall and some others on a number of evo-devo and developmental systems theory type projects. He was really instrumental in helping my work get attention, because he was at some of the conferences that I was at, including the MBL Woods Hole conference. Subsequently, he moved into more bioethical topics in his own research. But he was very much in the middle of everything right at that moment, in thinking about evo-devo and where it was going.

 

HS: Sahotra Sarkar

AL: Sahotra Sarkar was another one of the special issue editors, and somebody who I had been interacting with. He was a more senior philosopher, and somebody who had thought about the history, as well as the contemporary philosophical issues. He had collaborated with Scott Gilbert and some others. I had seen him at a number of conferences and received good feedback from him.

 

HS: Günter Wagner

AL: Günter Wagner is now retired, but was a central figure in evo-devo at the time (and still is). I was just getting to know him, and he would actually also end up being on my dissertation committee. He really helped clue me into the differences between the North American tradition and the more continental traditions. Even though he was trained in mathematical population genetics, he had an earlier background in comparative morphology through Rupert Riedl. And so, he was aware of exactly the kinds of issues that I was thinking about for this paper. He subsequently went on to be a collaborator, so I’ve had a really fruitful relationship with Günter Wagner over the years. At that point in time, I had just recently met him. I think I had known him for no more than 18 to 20 months, but was already interacting with him substantively around this material.

 

HS: The last name is David Wake

AL: David Wake’s scholarship plays a big role in the paper, because his group was one of those that really exemplified a multi-dimensional approach—the combination of development, morphology, systematics, ecology, all in a package around salamanders. David was very inspiring, but also was very generous in listening to philosophical  and historical work that was being done. And he himself was interested in that. One of his own papers, from 1982, in Perspectives in Biology and Medicine, was a key one for me in thinking about some of this material. So, I was glad to have his input along the way.

 

HS: I find it striking that you reached out to so many people for comments. Is this common in philosophy, to reach out to people with a manuscript and get their advice? Or was it something that you were doing on your own?

AL: That’s a good question. I think it’s probably not as typical. What you see in this Acknowledgements list is reflective of the fact that there was a lot of crisscrossing at meetings and other places with these people. Not everybody in this list read a full draft of the paper. Some of them were giving me feedback at a conference where I presented the paper, and I was having conversations with them about it. They were clearly giving me helpful input, but they were not reading a full manuscript. I think it was non-standard in part because in the background were all of these meetings—the Dibner Institute, the MBL Woods Hole meeting, plus another meeting at the University of Texas, Austin. I was seeing some of these people a couple times a year at different conferences, and so lots of conversations were even fluid across those moments. I was talking to them, and then I would email, and then see them again. It was very dynamic, and probably, in some ways, non-representative of how this kind of work is done. I’d have to go back and check, but I’m pretty sure it was also quite different from work that I was doing in parallel at the time on a couple of other papers. Those didn’t have as many eyes on them.

 

 

HS: You also thank participants at two other meetings. One of them is the one you just mentioned, which is the philosophy of biology graduate student conference in Austin in 2002. And the other is a Pittsburgh London colloquium in 2001. Can you talk a little bit about these meetings?

AL: Sahotra Sarkar is at UT Austin. He was the host of that one. That was a conference where graduate student work was foregrounded, but also evo-devo was very much in the air, and one of the central topics. That was really formative for me. At that point, I was presenting some of this work that had been well developed. The 2001 conference will be forever memorable because we were in that meeting when 9/11 happened. We were literally in a seminar room in London, and somebody said, you gotta come see what’s on TV. That meeting will never be forgotten. That was one of the earlier stages of me presenting this material. I also felt like the input was really helpful because those were the times when I was just working out the core of the of the argument, and getting some really key feedback that would help me shape it along the way.

 

HS: I wanted to also ask you a little bit about the financial support. You thank the Mustard Seed Foundation and the Harvey Fellows Program.

AL: The Mustard Seed Foundation is a broader foundation with many programs. The Harvey Fellows Program is specifically for graduate students, and it is intended to support work in fields that Christians are less likely to go into. Lots of Christians go into, say, medical fields or dental fields, but fewer Christians go into linguistics or something like that. I had won that award in 1999. It was a three year award, so it supported me and gave me some freedom from teaching as a graduate student, during the period when I was preparing a lot of this material, doing a lot of the initial research, and reading and writing. It was definitely key. At least for my graduate student experience, I came in on teaching funding—my funding was dependent on me teaching along the way. So, this award definitely freed me up to make faster progress at that particular moment in graduate school.

 

HS: You already said that you don’t remember anything striking about the peer review. I was wondering if there were responses or reactions to the paper after it was published that you remember. Did it attract any attention?

AL: In a funny way, a lot of people knew the argument, because they had heard it already. I had presented it at conferences, and a number of people had given feedback on it. So, there was a sense in which it was not out of the blue.  But I think that it definitely helped me put a kind of flag in the landscape as a scholar, in terms of what kinds of concerns I thought were really important and interesting and worth calling attention to in evo-devo. I think that people probably took this paper as an indicator that Alan’s focus in his dissertation and his scholarly beginnings are all about these questions in the context of evo-devo.

I think what’s interesting to me, as you noted in our original correspondence about this, is how much the paper continues to get attention. There are ways in which the paper was written at a particular moment, and that particular moment has passed, yet it continues to be a well-cited and ongoing element in a lot of people’s reading and thinking, which I find exciting and fun. But that was not definitely my intent at the time. I thought of it more as an intervention right then and there, and the fact that it continues to have echoes and impact 20+ years later is a bonus.

 

HS: Do you have a sense of what kind of attention it’s getting these days?

AL: I haven’t tracked it as specifically, in terms of, for example, within the last two or three years what papers were citing it. I think that a lot of people cite it because of the title, which is unfortunately how citations sometimes happen. Some of the philosophical themes about synthesis and integration and concepts and disciplinary connections are probably not as much widely cited as these broader themes about bring evo-devo together and the importance of morphology and other disciplines. It would be an interesting bibliometric analysis to go see what exactly is the citation pattern for the paper.

 

HS: At the point when you published this paper, did you already know that this was going to be like a lifelong pursuit, this question of the lineage of evo-devo? Did you know at least that you were going to continue to be in evo-devo research for the long term?

AL: I had a sense, at that time, that I was in a pretty fruitful area. There was good reason to think that working on these topics would pay returns. If there’s anything that’s uniquely interesting about my trajectory, it is that some of the themes in this paper ended up being a little submerged in my own research after the late 2000s, especially with respect to function and functional morphology. I’ve recently returned to them indirectly, and over the last couple of years have had the opportunity to begin to present again on the importance of evolutionary morphology for evolutionary theory, more broadly. I’ve also been able to dig in a little bit more deeply into functional morphology as its own kind of area, in thinking about questions of measurement and how practitioners operate in those contexts. So, in many ways, it continues to pay dividends, but not necessarily because I worked on all of it continuously the whole time. I continued with a few themes about synthesis and integration, and set aside some of the stuff about morphology, and now I’m returning more to that stuff and also the topic of function. I think the themes I stayed with throughout this period are things like levels of organization and the ways in which biological research often focuses on one or more of those, and what does it mean for there to be traffic across levels of organization in different disciplinary approaches. Those themes definitely continue to be crucial, and I think are not in any way going to be exhausted. There’s just so much there, within the space that’s broadly called evo-devo, that I don’t need to switch topics!

 

HS: This paper was published in what you might call the early days of evo-devo, about 20 years into its formal origins. In it you make an argument for the directions in which you think the field must go, or what evo-devo should be. I want to just read out some lines from the paper. You say, “Functional and evolutionary morphology are critical for understanding the development of a concept central to evolutionary developmental biology, evolutionary innovation. Highlighting the discipline of morphology and the concepts of innovation and novelty provides an alternative way of conceptualizing the ‘evo’ and the ‘devo’ to be synthesized”. And in another place you say, “since EDB [Evolutionary Developmental Biology] is a synthesis in the process of construction rather than a completed product, how the history is understood is even more critical – nothing less than the future shape of EDB is at stake”. Today, 22 years after the paper was published, I would like you to reflect on where the field stands in relation to these proposals you made in this paper.

AL: I think the message that I had there was not fully embraced, in part because the momentum behind developmental genetic approaches became so strong. And, in fact, you can see that even in my recent Cambridge Elements book, where I discuss how you can still find people who tend to frame things along that single trajectory out of developmental genetics and ignore morphology and the things that I was drawing attention to back in 2003. I do think this orientation has shaped the field in such a way that it has standardized a lot more than it was at the time. One of the things that was true of the intersection between evolution and development, especially through the 70s and the 80s, was that you had so many different voices in in the mix that those different voices generated a lot of interesting material because there was not a lot of agreement. There was a lot of diversity. There was not a single big picture that everybody got behind. Everybody was in agreement that they wanted to challenge, say, the broad picture from the modern synthesis, but there was complete disagreement about what the challenge was, where it needed to happen, and what the replacement would be. In the 2000s and 2010s, you get an aligning that eventually leads to the Pan-American Society for Evolutionary Developmental Biology, and that has a homogenization effect which is very typical when disciplines are forming. Disciplines are disciplined, right? That’s the origin of the word. There’s a way in which to become a discipline you have to narrow, because that’s part of the process.

And so, I think that some of the things that I highlighted were not taken up. Which is fine—there are lots of reasons for that. Now we’re starting to see a renewed appreciation of those needs. I was just at the Society for Integrative and Comparative Biology annual meeting, and there was a special session on evo-devo, reflecting on its history and its future. There was an awareness that we need to revisit these questions, and that something is missing. We’re starting to see new configurations of approaches that bring in a little bit more of that friction, whereby the diversity of ideas kicks up really interesting things. Maybe the nice thing is to be around long enough to watch some of that shift happen. When I was working on the 2015 book on conceptual change, even though I had a number of individuals participating in that project who were themselves morphologists or palaeontologists, it was quite clear that the predominant perspective on evo-devo was one of that developmental genetic vintage, which, back in 2003, I had said was too narrow! But at the same time, a lot of the people who I was talking about, and who even were involved in the 2015 book, were all people at the zenith of their careers, even in retirement, and many of them have now passed away. And so, arguably, the interesting thing is: what does the next generation do? What happens when that original, agitating generation that got the ball rolling is no longer there? Is everybody comfortable in their spot? Or do they continue to agitate? And what are they agitating for? I think there was a moment of standardized ‘business as usual’ for evo-devo, which might be in the process of reconfiguring even as we speak. For me, it’s a privilege to be involved in that at some level, as a historian and philosopher of science. To be able to talk with the scientists, to be able to get their input and their feedback. I really appreciate that, and enjoy those opportunities. I also realize that people are responding to a lot of different kinds of pressures. Certainly, one of the things that we see in the last 20 years is an increasing emphasis on grant funding and the need to secure that. I think that puts certain kinds of pressures on research. Even though earlier research required grant funding, the structure was not necessarily as intense as it is today, in certain ways, from institutions. We have to keep that in mind and say, okay, yes, there is a way in which the problems at the intersection of evolution and development are continuous. They’re recognizable as something that was worked on 30, 40, 50, 100 years ago. But, of course, the structure of science is changing; it’s not static. And so, we have to keep in mind what kinds of effects those changes have on scientific communities. I think that’s true also for technology and what kinds of technologies are adopted. The last five to seven years have seen a predominance of work with single-cell sequencing technology, because that has been the cutting edge that everybody was trying to apply. Some of those sorts of trends also pull people in how they organize their research and what things they focus on.

 

HS: Are there differences in these trend between the North American and the European scientific communities? Does it even make sense to think about these as different communities when it comes to evo-devo?

AL: That’s a good question. I would say that there are some differences. They’re reflected, to some degree, in the European evo-devo versus Pan-American evo-devo societies. But I’m also not sure that the differences mean you don’t have sufficient continuity between them. You have different flavours and themes, but you still have a relatively continuous community that interacts with one another. I do think there’s probably more, and always has been, in some ways, more interest from continental Europe in morphology. In that respect, the pitch for the importance of different levels of organization and analyses that take this into account seems like a harder sell for many North American research programmes. But, at the same time, it’s very clear that some of the cleanest examples of that work—like, say, David Wake or some of his students—is in North America. So, in that respect, it’s not a sharp line of some kind between North America and Europe, I think.

 

HS: What do you see as the place of this paper in the literature on evo-devo today?  What might be a reason for a student or a young researcher to pick up this paper and read it?

AL: A couple of things come to mind. One is that it captures well a kind of transition moment in the evo-devo community, from that period of time where there was intense interest in how we organize our community to tackle the problems that are of interest to us. I’m able to show that a number of these questions revolve around how practitioners understand their own history, and if they understand the history in one way they’re going to think about some of the contemporary issues in one way. Even though some of those have shifted over the last 20 years, the very fact that my paper captures that happening is helpful for the next generation of scholars to think about. I think also it continues to be salient for puncturing the relatively narrow view of evo-devo that is sometimes reflected in textbook presentations or more popular presentations; it’s just abundantly obvious that the people interested in the intersection of evolution and development were coming from lots of different disciplinary backgrounds, and specifically, many of them from these evolutionary morphology orientations. Lastly, I think that it could be seen as a key moment in thinking about how philosophers understand the organization of the sciences. Through a lot of the 20th century, philosophers had tended to think about relations between sciences in terms of reductionism and unification. Questions like, ‘Can biology be reduced to physics,’ were prominent. My paper—and, of course, other papers as well—from around this period and maybe a few years earlier, were putting pressure on these monolithic ways of relating the sciences to one another. My paper flags that by saying, look, thinking about the relationship between evolution and development is dynamic. It’s got multiple ways to be cashed out—in thinking about what is a synthesis or what is integration. Subsequently, some literatures have begun to grow a little bit on those topics, suggesting that philosophers should be paying more attention to how the sciences are dynamically organized, in ways that are more reflective of their problems and how their problems are structured, than some ‘for-all time’ relationship between the sciences, like you got in some of those earlier discourses in philosophy. I don’t know to what degree the analysis I offer in the paper is the best example of that, necessarily, but I feel like it is a very concrete illustration of that in a particular area of science. And this fits with a lot of the literature that has emerged where people have gone in and looked more closely at scientific practices and seen that some of these more monolithic approaches to relating sciences don’t map very well onto what scientists do, and that we need a richer toolkit of concepts to figure that out. And that has become, I think, a sort of cottage industry in philosophical work, to try and work out those details. That’s a couple things I think could drive somebody back to read this today.

 

HS: What does this paper mean to you personally? Would you count this as one of your favourite pieces of work?

AL: Yes, it will always hold a very special place in my work, because it is such a reminder of that early stage when I was sinking my teeth into this particular area for the first time and getting excited about the possibilities of studying evo-devo historically and philosophically. In rereading it for our interview, there are a number of things that I would change. For example, I utilized epigram-style quotations from different scientists at the beginning of each section, but I did not always unpack their meaning and significance in sufficient detail. In some cases, I relied on them (too much) for rhetorical effect. This is maybe not surprising, but at the same time, I’m quite happy with how it holds up, in the broad structure of the analysis. I think that it probably best represents where my research was going to go. Out of the things that I published as a graduate student or thereabouts, this paper really was my flag on the landscape of what my career trajectory as a scholar was going to look like. And so, in that respect, it does have a special place. And it definitely is one that I’m fond to think of, both in terms of the longevity of the analysis, but also in the personal memory of how it crystallized so much of what I was getting excited about and thinking was important, and especially the people I was interacting with at the time, some of whom are no longer with us. In that respect, it is a good reminder of just how unique of a moment it was.