A major grant from the John Templeton Foundation expands the frontier of evolutionary science.Back in 2008, a small scientific conference at the Konrad Lorenz Institute, located in the sleepy Austrian village of Altenberg, made headlines around the world. The conference was titled “The Extended Evolutionary Synthesis (EES)” and was organized by the philosopher and evolutionary biologist Massimo Pigliucci and the theoretical biologist Gerd B. Müller.Just another conference out of many that people like me are accustomed to attending. Even the title was sedate. Nothing revolutionary here—just an extension of what has been taking place since the 1940’s, which was dubbed “The Modern Synthesis” at the time. But then science journalist Suzan Mazur published a scoop several months before the event titled Altenberg! The Woodstock of Evolution?, which began this way:

It’s not Yasgur’s Farm, but what happens at the Konrad Lorenz Institute in Altenberg, Austria this July promises to be far more transforming for the world than Woodstock. What it amounts to is a gathering of 16 biologists and philosophers of rock star stature – let’s call them “the Altenberg 16″ – who recognize that the theory of evolution which most practicing biologists accept and which is taught in classrooms today, is inadequate in explaining our existence. It’s pre the discovery of DNA, lacks a theory for body form and does not accommodate “other” new phenomena. So the theory Charles Darwin gave us, which was dusted off and repackaged 70 years ago, seems about to be reborn as the “Extended Evolutionary Synthesis.

Well! All of a sudden, the Altenberg 16 (I was one of them) were in the middle of a media circus. Creationists and Intelligent Design theorists gloated that Darwin’s theory had finally been revealed as dead. We began to be described as some sort of cabal. Why were these 16 invited and not others? Conservative evolutionary biologists sniffed that the word “extended” in the title was unnecessary. The Modern Synthesis was good enough for them and should be good enough for the next generation. Some of us were dismayed by the hype while others rather enjoyed our 15 minutes of fame as rock asteroids, if not rock stars.Fast forwarding to the present, the phrase “The Extended Evolutionary Synthesis” has gained a tiny toehold in the scientific literature. According to Google Scholar, it has appeared in the title of only 12 articles and books between 2008 and 2016 (listed at the end of this report). That is about to change, however, because the John Templeton Foundation has just awarded one of the largest grants in its history—approaching 10 million dollars—to put the EES to the test, including over 20 inter-related projects involving a team of over 50 scientists headed by Kevin N. Laland at the University of St. Andrews in Scotland and Tobias Uller at Lund University in Sweden.As a science communication outlet run by scientists, TVOL is in a unique position to report on how this group of scientists, now empowered by a major private foundation, is attempting to define the frontier of evolutionary science (Full disclosure: TVOL is also partially funded by JTF). We begin with an interview with Laland and proceed with a series of articles that explore the philosophical foundation of the EES and the scientific research projects that will be funded.DSW: Welcome, Kevin, to This View of Life and congratulations on your success with the JTF grant.KNL: Thank you, David. The grant is particularly pleasing given our vision of the EES as delivering a complementary research program. It lends confidence to the view that the EES can make a constructive contribution by inspiring novel research.DSW: You were not one of the Altenberg 16, but you were the lead author of a major 2015 review article on the EES published in the Proceedings of the Royal Society of London. Tell us about your background and how you acquired your leadership role, first for that article.KNL: My interest in the EES arose in the aftermath of the Altenberg meeting. It was clear that the notion of an extended synthesis divided the evolutionary biology community, generating both enormous excitement and strong negative responses. However, I held the view that the negativity arose primarily from the absence of a clear rationale for an EES, and the mistaken perception that the EES was a rejection of neo-Darwinism. If it were possible to harness the enthusiasm and new ideas, whilst at the same time circumventing the concerns of more orthodox evolutionists, then the EES could prove a stimulant to the field.I was drawn to thinking about these issues through my research on niche construction, with John Odling-Smee (Oxford) and Marc Feldman (Stanford). John was a participant at the Altenberg meeting, and he and I, together with evolutionary ecologist Tobias Uller (then Oxford, now Lund), began discussing whether there was a conception of the EES that could do useful work. We had no sympathy with the argument that evolutionary biology was undergoing a ‘paradigm shift’ – to my mind paradigm shifts are an outdated notion (sciences change more through gradual evolution than dramatic revolution). Nonetheless, we were very conscious of how academic fields possess conceptual frameworks – ways of thinking – that influenced what questions are asked, what data is collected, and how that data is interpreted. Here, alternative perspectives can be of real value to extent that they encourage researchers to generate and test novel hypotheses, or open up new lines of enquiry. That is how we envisaged the EES could be of service – as an alternative way of thinking about evolution, which could be deployed alongside traditional perspectives to stimulate innovative research.However, any such alternative needs to be formulated in a disciplined way. We noted that certain literatures, for instance, concerning developmental bias, developmental plasticity, and expanded views of inheritance, stood out as being open to both traditional and progressive interpretations, leaving them lying squarely on the fault line. In all cases, the more progressive reading emphasized an organism-centred perspective, rejected the idea that development was controlled by a genetic program, and recognized that developmental processes played important (and not fully appreciated) evolutionary roles. It made sense to conceptualize the EES as an eco-developmental perspective, and to highlight these literatures as the intellectual territory on which an EES might focus.So we pulled together an interdisciplinary team of evolutionary and developmental biologists, as well as philosophers, chosen to provide expertise in these domains, and for two years discussed the issues. Our target was to add substance to the EES, by specifying in detail what it was, why it was required, how it differed from traditional perspectives, what new predictions it made, and so forth. We were delighted when our conclusions were published as Proceedings of the Royal Society B’s annual Darwin Review, which is the journal’s special “showcase” article of 2015.DSW: How did that lead to the JTF grant?KNL: Earlier, in 2014, we had published a brief précis of our thinking in Nature, as part of the debate entitled: Does evolutionary theory need a rethink? The Nature piece was too short to spell out our position in detail, but worked well to draw attention to the project. We were deluged with feedback (I received more correspondence over that brief article than over all of my other books and papers put together) with the overwhelming majority of the many hundreds of emails highly enthusiastic. Among the interested readers were two representatives of the John Templeton Foundation who liked the ideas that we proposed, and a dialogue ensued. Eighteen months later, after our application had been well received in review, the grant was funded. We are very excited that the program of research can now go ahead, as it essentially tests the predictions specified in our Proceedings article.DSW: I agree with you that deep philosophical issues lie behind the seemingly innocuous phrase “Extended Evolutionary Synthesis” and I’m glad that you include a number of philosophers on your team. TVOL will be featuring them directly in future articles, but could you please provide a brief introduction to them and their role in the research program?KNL: I’m a big fan of philosophers of biology. David Hull, Elliott Sober, and many others have made really valuable contributions to evolutionary biology over the years. The philosopher with whom I have worked most closely is Kim Sterelny (ANU), who was a coauthor of those EES papers, and helped pioneer thinking on niche construction. However, this research project also involves several other leading philosophers, including Tim Lewens and Marta Halina at Cambridge, as well as Jonathan Birch (LSE) and Ellen Clarke (Oxford). They have a really important job to do. For instance, the recognition of multiple inheritance systems generates challenging issues concerning how we conceptualize and measure fitness, or evolutionary change. Likewise, a central claim of the EES is that developmental processes should be recognized as evolutionary processes. That raises questions like: What is an evolutionary process, and how do the structures and assumptions of the field affect what are recognized as causes of evolution? Philosophers can help biologists to think clearly about these issues, and their involvement will help promote awareness of the roles that conceptual frameworks play in science.DSW: I really look forward to digging into that! What other work will be carried out as part of the grant?KNL: We will be investigating key questions highlighted by the EES. For instance, one critical issue is whether developmental plasticity can play a central directing role in evolution, as envisaged by Mary Jane West-Eberhard (2003). This question will be addressed in experimental studies of sticklebacks, lizards and butterflies, by teams led by Susan Foster, Tobias Uller and Paul Brakefield. We also set out to evaluate when and how extra-genetic inheritance arises, how multiple inheritance systems interact, and how their evolution differs from systems solely reliant on genetic inheritance. These issues will be explored experimentally, for instance, through Armin Moczek’s studies of horned beetles, as well as through novel theoretical work by Mike Wade, Marc Feldman, Rufus Johnstone and Michael Lachmann. Devising novel theory is actually a big part of the grant, including new approaches to modeling complex genotype-to-phenotype relations, and the emergence of biological complexity. Other central issues include investigations of the circumstances under which developmental bias and niche construction impose direction on evolution, generate convergences, or help account for taxonomic diversity and adaptation. These are all effectively 'big unanswered questions' for evolutionary biology, and we hope that by addressing them in this comprehensive manner the research program will have a major impact.DSW: Sounds very exciting! This table from your Proceedings article does a good job contrasting the core assumptions of the Modern Synthesis and the EES. In my opinion, your portrayal of the Modern Synthesis is accurate. It is the view of evolution that most people were taught and continue to teach. Against that background, the core assumptions of EES look very different indeed, even deserving the label “New Synthesis” or “New Paradigm”. Why shouldn’t the EES be described in bolder terms?

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KNL: Some people do, of course. Steve Gould spoke of the need for a new evolutionary paradigm and more recently Denis Noble has argued that neo-Darwinism needs ‘replacing’. I personally find such language counterproductive. I defend people’s right to think differently, and I strongly believe that in a healthy science no assumptions, no matter how longstanding or cherished, should be beyond questioning. However, I worry that talk of ‘revolution’ and ‘replacement’ gives the wrong impression. It implies that the radicals want to rip up the textbooks and start all over, and that is simply not correct. I know of no professional biologist that wants that. What the people who use those terms want to ‘replace’ is simply how the findings are interpreted. It is certainly true that many EES sympathizers, myself included, would like to see fundamental change in how the evolutionary process is described and understood. But what this really boils down to is recognition that, in addition to selection, drift, mutation and other established evolutionary processes, other factors, particularly developmental influences, shape the evolutionary process in important ways. However, that influence is hard to see from the traditional standpoint, so we need to encourage alternative conceptual frameworks. Nevertheless, in the EES, all processes central to contemporary evolutionary theory, and all empirical and theoretical findings, remain important. That is why I am more comfortable speaking of ‘extension’ or ‘revision’.DSW: I’m all for conceptual evolution rather than revolution. After reading your grant proposal, however, one thing seems conspicuously lacking—the application of evolutionary theory to all aspects of humanity. Cultural evolution is listed as part of inclusive inheritance but all of the research projects are on non-human species and fall squarely within the biological sciences. Isn’t evolution in relation to human affairs part of the EES?KNL: Absolutely. The response to our Nature article brought home to me how much of the enthusiasm for the EES derives from what might be regarded as the periphery of evolutionary biology – fields such as evolutionary developmental biology, physiology, ecology, botany, earth sciences and, in particular, the human sciences. These fields use evolutionary principles to organize their research, but the researchers are not always entirely satisfied with what traditional evolutionary explanations have to offer. Many evolutionary anthropologists, archaeologists and developmental psychologists have already started using key EES concepts, such as niche construction, constructive development, and expanded inheritance, in their research. It is important to recognize that there remains a great deal of dissatisfaction amongst the social sciences and humanities with adaptationist, gene-centric accounts of human behavior. The EES potentially offers a richer and more attractive evolutionary framework.We did originally plan for some human work in our grant, but then realized that this is an important topic that merited special attention. Our plan is to apply for another major grant from the John Templeton Foundation in the near future that would focus specifically on the application of EES thinking to human affairs. I encourage TVOL readers to get in touch with me, if they are doing relevant work, and we could maybe work them into our application, or involve them in our activities.DSW: I’m involved in the creation of a new society for the study of cultural evolution that can perhaps play a role. We have over 1000 founding members who collectively belong to over 450 other academic societies, illustrating the scope of integration that is taking place. I look forward to featuring the various facets of the EES on TVOL at the inception of your grant, which can provide a baseline for comparison after the results of research start to emerge. We’ll also be sure to include contrarian voices. Best of luck—you have your work cut out!KNL: Thank you, David. We appreciate your interest.For further information about the EES and project see http://synergy.st-andrews.ac.uk/ees/Books and articles on the Extended Evolutionary Synthesis:

  1. Pigliucci, M. (2007). Do we need an extended evolutionary synthesis?.Evolution, 61(12), 2743-2749.
  2. Dickins, T. E., & Rahman, Q. (2012, May). The extended evolutionary synthesis and the role of soft inheritance in evolution. In Proc. R. Soc. B (p. rspb20120273). The Royal Society.
  3. Pigliucci, M., & Müller, G. B. (2010). Elements of an extended evolutionary synthesis. Evolution: The extended synthesis, 3-17.
  4. Mesoudi, A., Blanchet, S., Charmantier, A., Danchin, É., Fogarty, L., Jablonka, E., ... & Pujol, B. (2013). Is non-genetic inheritance just a proximate mechanism? A corroboration of the extended evolutionary synthesis. Biological Theory, 7(3), 189-195.
  5. Pigliucci, M. (2009). An extended synthesis for evolutionary biology. Annals of the New York Academy of Sciences, 1168(1), 218-228.
  6. Laland, K. N., Uller, T., Feldman, M. W., Sterelny, K., Müller, G. B., Moczek, A., ... & Odling-Smee, J. (2015, August). The extended evolutionary synthesis: its structure, assumptions and predictions. In Proc. R. Soc. B (Vol. 282, No. 1813, p. 20151019). The Royal Society.
  7. De Loof, A. (2015). Organic and cultural evolution can be seamlessly integrated using the principles of communication and problem-solving: The foundations for an extended evolutionary synthesis (EES) as outlined in the Mega-Evolution concept. Functional Genomics.
  8. Pigliucci, M., & Finkelman, L. (2014). The extended (evolutionary) synthesis debate: where science meets philosophy. BioScience, 64(6), 511-516.
  9. Wozniak, A., & Konstanczak, S. (2013). Evolutionary Ethics in the Light of Extended Synthesis. Ethics and Bioethics (in Central Europe) 3 (1-2):21-30.
  10. Serrelli, E. The Extended Evolutionary Synthesis: a metascientific view of evolutionary biology, and some directions to transcend its limits.
  11. Weber, B. H. (2013). The Baldwin Effect in an Extended Evolutionary Synthesis. Beyond Mechanism: Putting Life Back Into Biology, 254.
  12. Vianello, A., & Passamonti, S. (2016). Biochemistry and physiology within the framework of the extended synthesis of evolutionary biology. Biology direct,11(1), 1.