No field showcases the virtues of philosophy of biology more effectively than evolutionary science (although I recently learned from Gregory Rupik, 2024, that this observation is not necessarily a virtue). As a graduate student and would-be evolutionist in London in the late 1980s, I was introduced to the pioneering writings of David Hull on evolutionary epistemology, to Elliott Sober’s brilliant ‘The Nature of Selection’ (1984), which stood ever since as a benchmark of great philosophy of science, to Susan Oyama’s wonderfully rich and thought-provoking ‘The Ontogeny of Information’ (1985), and to Philip Kitcher’s (1985) devastating ‘Vaulting Ambition’, which deconstructed human sociobiology. To a young researcher, like me, drawn to ‘big questions’ in science, these works were like nectar from the gods.
The stuff of my philosophy of biology upbringing may not qualify as philosophy ‘in’ biology, but it left me with a deeply entrenched admiration for what philosophers could contribute to my field, and that admiration has spawned many subsequent collaborations that brought philosophers directly into the scientific arena. Even that tribute short-changes the contributions that philosophers have made to my intellectual development, as over the decades an endless stream of savants have acted like a free consultancy service, ever willing to chew the intellectual cud. My career has entailed a journey, from reading and appreciating philosophy of biology, to consulting with philosophers, and finally to collaborating with them. In fact, my thoughts have been so enriched by philosophers that I could almost write a book on the subject. Perhaps one day I will, but just now I have a different book in mind.
Come September, Princeton University Press will publish a new book of mine, written together with Tobias Uller, Nathalie Feiner, Marcus Feldman and Scott Gilbert, entitled ‘Evolution Evolving: The Developmental Origins of Adaptation and Biodiversity’. You can think of it as an Extended Evolutionary Synthesis (EES) view of how adaptive evolution occurs, and the central role that developmental processes play in that. Naturally, this means we address many conceptual issues – the origins of biological information, the utility of the genotype-phenotype distinction, the nature of biological causation, the sources of inheritance, and more – that I imagine would be of interest to readers of this site. Given the strong philosophical thread running through our tome, I am very happy to get this opportunity to acknowledge the multiple contributions that philosophers of science have made to its inception. While here I share my story, I my coauthors too have had extensive and productive interaction with philosophers, over many years: Scott Gilbert, for instance, has worked closely with Sahotra Sarkar, Marc Feldman with Lisa Lloyd, and Tobias Uller (with Kostas Kampourakis) recently edited a book on ‘Philosophy of Science for Biologists’ (2020). I have no doubt that these prior experiences of philosophy in biology have also enriched our writing.
The roots of ‘Evolution Evolving’ I trace back at least to my time as a postdoc at UC Berkeley in the early 1990s, and the subsequent decade I spent in the Zoology department at Cambridge University. During that period, my co-conspirators John Odling-Smee, Marc Feldman and I were developing the theoretical foundations of niche construction theory, and John and I made regular visits to Stanford to work with Marc. I remember those typically month-long visits as being incredibly intense, with us laboring long hours, occasionally interspersed with visits to sit at the knee of other Stanford researchers, and pick their brains.
One such luminary with whom we had productive dialogue was Peter Godfrey-Smith, then on the Stanford philosophy faculty. Godfrey-Smith (1996) had distinguished between different types of explanation for the relationships between organisms and environments, which he had labelled ‘externalist’, ‘internalist’, ‘constructive’, and so forth. ‘Externalist explanations’ accounted for the internal properties of organisms in terms of environmental properties, while ‘internalist explanations’ described one set of internal properties of a system in terms of another set of internal properties, and ‘constructive explanations’ interpreted environmental properties in terms of the properties of organisms. These distinctions helped us to conceptualise the key differences between how standard evolutionary theory explained the organism-environment relationship and how we wanted to describe that relationship in niche-construction terms. Standard theory provided externalist explanations, since the adaptations of organisms were explained relative to the properties of selective environments. In contrast, by stressing that the selective environments of organisms are part-dependent on the niche-constructing activities of organisms, we were proposing a mix of externalist and constructivist explanations, that Godfrey-Smith labelled ‘interactionist’. Eventually we were to see niche construction theory as fitting into a bigger picture – a broader vision of the causal structure of evolution that is now often labelled the EES. With that came the recognition, now articulated in Evolution Evolving, that that the roots of adaptation are perhaps even more interactionist than we envisaged in our early works on niche construction, and that developmental processes always bias the phenotypic variation exposed to selection:
“In continual interactive cycles, developmental processes bias what gets selected, but then selection modifies the developmental processes that create developmental bias. This process of reciprocal causation guides the evolution of morphology, and indeed all aspects of the phenotype. To disregard the causal role of development in evolution on the grounds that it is a product of selection, as is common, is questionable reasoning”
(Evolution Evolving, p16).
Nowadays, we tend to speak of a commitment to ‘reciprocal causation’, but our thinking through the issues was greatly facilitated by those conversations with PGS. With the benefit of hindsight, I can see now that part of our conceptual struggle concerned the requirement to untangle a complex of interrelated questions, including: How should biologists explain adaptation? What is an evolutionary cause? And Why do offspring resemble parents?. These issues took us years to resolve, but we were abetted by many philosophers of biology along the way. If PGS gets credit for helping us to get our heads around adaptation, then Kim Sterelny was likewise a catalyst for our views of evolutionary causation, and Paul Griffiths and Karola Stotz greatly aided our thinking on the nature of inheritance. Lurking beneath these issues was a tentative and barely conscious commitment on our part to what is now labelled ‘process ontology’, which we picked up from Conrad Waddington’s writings. In passing, we had flirted with concepts like ‘homeorhesis’ and ‘autopoiesis’, but – rightly or wrongly – had decided these labels had little explanatory utility to an audience of hard-nosed and often sceptical biologists that by-and-large were disappointedly uninterested in conceptual issues (at least, those conceptual issues we cared about).
In the early 2000s, we were invited to contribute to Susan Oyama, Russell Gray and Paul Griffiths’ important ‘Cycles of Contingency. Developmental Systems and Evolution’ (2001) volume, and also to give talks at a symposium on Richard Dawkins’ extended phenotype concept organised by Kim Sterelny and Mateo Mameli at ISHPSSB in Vienna in 2003. Between them, these events allowed for extensive conversations with Kim Sterelny, but also Paul Griffiths, Susan Oyama, Evelyn Fox Keller, Mateo Mameli and Eva Jablonka. In those discussions the distinction between niche construction and the extended phenotype would always come up. The two ideas are superficially similar – after all, they are both concerned with how organisms construct artefacts and modify environmental states – but intellectually they are miles apart. Dawkins’ treatment offers a traditional, gene-centric adaptationist account in which extended phenotypes are adaptations and the only selective feedback considered is to the genes that underly them. Conversely, niche construction theory offered a treatment that we can now regard as aligned with the EES: niche-constructed environments may (e.g. beaver dams) or may not (e.g. earthworm-processed soil) be adaptations, are often the product of extra-genetic inheritance (e.g. cultural niche construction), and can generate ecological feedback affecting selection at other loci, and in later generations (ecological inheritance). Philosophers of biology seemed to get very excited about this particular juxtaposition, and indeed in 2004 Sterelny invited me to debate Dawkins in the pages of Biology & Philosophy (Laland 2004; Dawkins 2004).
These exchanges with philosophers of biology – perhaps just over a decade of close interaction and dialogue – led to what was my first direct collaboration with a philosopher. Sold, as I was, on the insight and clarity of thought that philosophers could bring to my arguments, I invited Kim Sterelny to coauthor an article that rebutted some commonly mooted criticisms of niche construction theory. The article, entitled “Seven reasons (not) to neglect niche construction”, was published in the journal Evolution in 2006. Why I chose Kim, on reflection, was partly because, more-so than any other philosopher I knew, Kim was keen to be ‘in’ biology: that is, he wanted to be a participant in evolutionary debates, rather than just an observer and analyser. Over-and-above the intellectual assets that Kim brought to our partnership, the collaboration was a lot of fun. Kim, of course, is great company, and I had long had a weakness for staying up late drinking with him and discussing mighty topics. However, it was also fun because, as I said at the outset, I have always been searching for answers to fundamental questions, and my experience had been that philosophers, far more so than biologists, were often so inclined.
At that juncture in my career, I had really only been interacting with philosophers on the topic of niche construction, and within the bounds that perspective set. Not that those bounds were particularly restrictive: our work on niche construction had led us from population genetics, to ecosystem and community ecology, to various branches to the human sciences. I vividly remember a review of our book on niche construction from Paul Griffiths, that I can paraphrase as “8/10 for effort, but you really need to get into developmental biology”. Having published a monograph that to me seemed almost suicidally broad, Paul’s assessment seemed harsh. However, he was right of course, and further exchanges with Paul and Karola at conferences regarding developmental niche construction eventually led to John and I collaborating with Scott Gilbert to explore the parallels between niche construction and evo devo (Laland et al 2008). I think we were just beginning to suspect that the issues with which advocates of niche-construction theory were wrestling might be broader, and perhaps even more foundational, than we had thus far appreciated. Around 2011, those experiences primed me for a ‘revelatory experience’ that occurred while sitting with Gerd Müller under a tree at the KLI, and which was to shape the rest of my career.
It had become apparent to us that a major conceptual barrier to the acceptance of niche construction theory was Ernst Mayr’s distinction between proximate and ultimate causation. In 1961, Mayr had published “Cause and effect in biology”, a now classic article in Science that had influenced how most contemporary biologists understood causality. We had found that niche construction was frequently categorized as a proximate mechanism, and as a consequence was automatically disregarded as being evolutionarily significant. Worse, we were being told that we had muddled ‘proximate’ and ‘ultimate’ causation, when instead we were working with a different understanding of causation. Gerd and I both got very excited when I related this to him, and he explained that Mayr’s distinction was no less a barrier to recognizing a role for developmental bias in evolution. Suddenly, we each realized that we had been fighting the same battle all these years!
Further discussion with John Odling-Smee and Tobias Uller led us to the view that several prominent current debates in biology – not just over evolution and development, and niche construction, but also over cooperation, and the evolution of language – were linked by a common axis of acceptance or rejection of Mayr’s model of causation. As that year just happened to be the 50th anniversary of Mayr’s paper, we saw an opportunity, and hurriedly pulled together a team of researchers to write up our thoughts as an article for Science. We argued that Mayr’s formulation has acted to stabilize the dominant evolutionary paradigm against change but may now hamper progress in the biological sciences. Of course, for any such team to write with authority on causation we required a philosopher, so it seemed natural for us to invite Kim Sterelny on board.
That article brought us firmly into the terrain of wider debates about the explanatory utility of the dominant causal structure of evolutionary theory. Following our success in publishing “Cause and Effect in Science Revisited” (Laland et al 2011), Tobias Uller, John Odling-Smee and I got together a small interdisciplinary working group to think about and evaluate the concept of an extended evolutionary synthesis . The initiative was born partly out of frustration. There had been a KLI workshop on the topic, organized by Massimo Pigliucci and Gerd Müller, at which John had participated, and the proceedings had been published as an edited volume (Pigliucci & Müller 2010). However, a lot of questions remained: What was the EES?, Why was it needed?, How did it differ from the Modern Synthesis?, What findings motivated it?, What were its key assumptions and predictions?. We wanted answers to these questions, and since it seemed no one else was providing them, we set out to generate them ourselves, again assembling a team of experts in what we thought were the relevant fields. As our previous collaborations had been so successful, Sterelny was again our go-to philosopher – one brave (or foolhardy) enough to get ‘into’ biology and stick his neck out. After a couple of years of discussion, we eventually published our deliberations as a high-profile pair of articles: a short piece in Nature (Laland et al, 2014), and a more-protracted elaboration in Proceedings of the Royal Society B – the latter as the prestigious annual Darwin review (Laland et al 2015).
It is hard to overstate the impact of these papers: both have been cited well over 1000 times. While they were controversial at the time, I can’t help but feel that that level of citation must mean something. Virtually all the ideas that the EES championed – plasticity-led evolution, extra-genetic inheritance, developmental bias, niche construction, evolvability – are now becoming, or have become, mainstream. Without those high-profile papers we probably would not have got a multi-million-pound grant from the John Templeton Foundation entitled “Putting the EES to the test”. Tim Lewens and Massimo Pigliucci were key member of that EES research program grant, and many other philosophers were involved in more modest capacities, including Marta Halina, Jonathan Birch, Ellen Clarke, Kim Sterelny, Andrew Buskell, and Lynn Chiu. That grant, in turn, was phenomenally productive, leading to over 200 other scientific articles, many in top journals (see Chiu 2023 for a summary).
Yet for all those papers’ success, it was apparent to us from an early stage that it would not be possible to make a truly compelling case for an extended evolutionary synthesis in article form. This was a subject that required consideration of the history of biology, the philosophy of science, as well as an understanding of how development works, knowledge of evolutionary genetics, an appreciation of insights from various branches of theoretical biology, and comprehension of the avalanche of ‘new biology’ findings – particularly emanating from the burgeoning field of epigenetics. No, we would need a book length treatment. A decade later, ‘Evolution Evolving’ is the result.
As I hinted above, there are several juicy topics in our ‘Evolution Evolving’ book that I believe philosophers of science might like to sink their teeth into, and I obviously hope that readers will consider reading it for themselves and perhaps even engaging with us in further discussion. I think I can speak for all my co-authors in saying we would welcome that. However, as a taster, here I pick out three issues from the book to whet your appetites.
Chapter 9 of our book focuses on natural selection, and revisits Elliott Sober’s (1984) important distinction between the selection of objects and selection for properties. Sober famously illustrated this using “a toy that my niece once enjoyed playing with before it was confiscated to serve the higher purposes of philosophy” (an elegant quip that remains one of my favourite lines of academic writing). You will recall the toy is a “selection machine”: a transparent container, structured into levels by dividing partitions, and containing balls of differing sizes and colours. Sober explains how the smallest balls are the objects that are selected, but green balls have been selected at the same time. By highlighting the distinction between selection of green balls and selection for smallness, Sober’s selection toy provides a helpful analogy illuminating the causes and effects of selection. Here, however, having acknowledged our debt to Sober, we deliberately stretch the analogy to encourage further consideration of the role of the organism in selection:
Helpful though this analogy may be, there are important respects in which the selection toy gives a misleading impression of the action of natural selection – a characterization that portrays organisms in overly passive terms. For instance, the structural features of the toy – the balls and the perforated partitions – were factory made. The balls played no part in building the partitions. The partitions simply exist, and the balls are selected according to the partition’s properties (i.e., the size and shape of the holes). However, termites construct mounds, and regulate temperature and humidity within them, which redirects selection away from internal organs that deal with water scarcity, such as thick cuticles, towards selection for effective behavioral strategies for humidity control, such as digging down to the water table to retrieve moisture. And desert rhubarb collect water next to their primary root, generating selection for huge, heavily-ridged leaves, as opposed to selection for small leaves or spines. The actions of living organisms determine what is selected for. It is as if the balls in the selection toy themselves build the partitions and cut out the holes.
We go on to quibble that the balls in the selection toy only move when acted upon by external forces, with no ability to choose which level of the toy they will occupy, nor change their shape. Yet, in the real world, organisms can evade extreme conditions through behavioral plasticity, “as if the balls … are made of a squidgy … jelly”. We also mention other pathways to adaptive fit, for instance, how bacterial symbionts can detoxify poisons “analogous to the balls releasing an acid to eat away the partition”. Finally, we stress how the seemingly arbitrary association between ball size and colour misleads:
There is no reason why small balls need to be green – we are implicitly conscious that they could just as easily be red. This gives the impression that, in selecting for smallness, greenness has been randomly chosen. But here again the analogy is misleading. Floppy ears are not linked to the tameness of domesticated animals by chance: the components of domestication syndrome are connected by joint developmental regulation through the neural crest GRN. Nor are the fragmented bones of cavefish coupled with an improved ability to sense vibrations in the water through historical contingency. Fragmentation is a by-product of the increased production of sensory cells on these bones; cells that form the lateral line and are responsible for this enhanced mechanosensory capability. In the real world, developmental mechanisms connect the selection of traits to the selection for traits, with a powerful implication. Through investigating developmental bias, evolutionary biologists could understand and predict which traits there would be selection of, alongside the character selected for.
Our agenda here is, of course, not to give Sober a hard time over the limitations of his analogy, but rather to use it, as he did, to highlight the ‘nature of selection’ and maybe the habits of evolutionary biologists too. Just as the selection toy starts with a pre-existing partition and round balls (with biologists and philosophers taking as given smallness to be the trait selected for, and greenness as a coincident character) so evolutionary biology often begins with the identification of fitness differences, and works through the ramifications of these for character evolution, adaptation, and speciation, detecting correlated change in other characters in the process. In this respect, the selection toy still provides an excellent analogy for how natural selection is conceptualized by biologists. However:
This focus, while productive, comes at a cost, in the form of limitations on the power of evolutionary explanations. The nonchalant attribution to ‘selection’ of all evolution arising from fitness differences masks hidden determinants of the sources of fitness differences. This would not matter if fitness differences arose by chance, or if those characters selected for were packaged with the selection of other characters in a coincidental or unknowable manner, but that is not the case. In the real world, balls are green for a reason.
Moving on to my second example, in chapter 10 of ‘Evolution Evolving’ we turn to consider inheritance, where our treatment owes a debt to the prescient writings of Susan Oyama. We stress how inheritance is a time-distributed developmental process by which diverse developmental resources become available to the next generation. After reviewing extensive evidence for diverse forms of inheritance, we conclude that many extra-genetic inheritance processes are (contra Wray et al, 2014) not luxury “add ons” but rather essential tools for short-term, rapid-response adaptation:
Heredity is more than a package of genes and cellular resources handed over at conception like the baton in a relay race: it is a continuous process of developmental reconstruction that spans the life cycle. All forms of inheritance collectively guide offspring development by contributing to the production of a phenotype predicted to match the expected environment, where that ‘prediction’ is based on transmitted genes and updated by inherited extra-genetic information accrued through both detection and selection mechanisms. Parents construct offspring developmental environments by transferring resources internally and externally, choosing and building structures, and regulating conditions, in ways that enable reliable implementation of genetically inherited predispositions.
What is transmitted across generations, we suggest, is “the developmental means to construct phenotypes that are predicted to match anticipated environmental conditions”. However, we emphasize that those ‘means’ not include genes and other resources passed from parents to offspring, but also the activities that parents engage in to construct the environmental context in which their offspring develop:
If there are similarities between the traits of parents and offspring it is because, within lineages, phenotypes are reliably re-constructed across generations. That reconstruction is consistent because it is informed by processes of environmental detection and selection, operating at a range of temporal and spatial scales, including, but not restricted to, the selection of genetic variation.
My third example again delves deep into the nature of selection. Richard Lewontin famously decomposed natural selection into three subprocesses: phenotypic variation, differential fitness and heredity. However, as another philosopher from whom I have learned a great deal, Denis Walsh (2015, 2019), has emphasized, the dominant view that selection is the sole cause of adaptive evolution is tied to an additional assumption that the three subprocesses are effectively ‘quasi-autonomous’. That is, they are thought to feed into each other, but not to modify each other’s operation. Following Walsh, in our book we stress how much of the debate over the role that developmental processes play in evolution relates to instances where the three subprocesses become intertwined. By contributing to all three subprocesses simultaneously, and by modifying how subprocesses operate, developmental processes can violate the assumption that Lewontin’s components are independent.
An example is provided by the niche construction of dung beetles. We first suggest:
A traditional evolutionary explanation for the adaptive fit between dung beetles and their environments would invoke mutations that change, say, the beetles’ brood-ball-processing in a manner that enhances fitness, and are inherited by the next generation. This allows brood-ball processing to be construed as a proximate mechanism and – since the generation of variation by mutation is assumed to be random and inheritance is assumed to be unbiased – allows fitness differences to explain the brood-ball processing adaptation.
However, we go on to describe experiments carried out by Armin Moczek and the members of his laboratory, that show how the underlying causation is actually more complicated (and interesting!). This is illustrated by Figure 1, reproduced from the book.
Figure 1. The intertwining of variation, fitness, and inheritance in dung beetles (reproduced from Figure 14 in ‘Evolution Evolving’)
The experiments show that the extent to which dung beetle traits, such as body size and developmental time, contribute to fitness depends on variation in the niche-constructing behavior of mothers and larvae:
Experiments quantify how the contribution to fitness of offspring traits depends critically on the properties of the brood ball, which is constructed by the mother and modified by the larvae to act as an external rumen (i.e., depends on niche construction). Also, inheritance is not independent of phenotypic variation and fitness differences, as the brood ball is a parental effect that is ecologically inherited by the larvae. Again, niche construction brings about qualitative changes in how inheritance occurs. For instance, experiments show this inheritance depends critically on whether or not the mother incorporates into that brood ball a pedestal containing a sample of her microbiome.… Further, phenotypic variation is not independent of inheritance since beetles develop in the beetle-constructed environment of the brood ball, which experiments show fundamentally influences both the developing larvae’s traits and relationships among them.
These interactions between the components of natural selection mean that beetle niche construction is no longer just a proximate mechanism, and attributing the complementarity between beetles and their environments solely to fitness differences becomes open to question. Drawing on another distinction brought to prominence by Elliott Sober, between ‘variational’ and ‘transformational’ explanations, we suggest that analysis of the interactions between sub-processes reveals a poorly appreciated role for transformational explanations in adaptive evolution:
The ‘fittedness’ or ‘match’ of dung beetles to their immediate local environment arises partly because those beetles experiencing a poor fit have died or failed to reproduce, but also partly because individual beetles inherit a maternally modified brood ball that is well-suited to other aspects of the larval phenotype, and partly because how the larvae develop inside that brood ball is sensitive to the brood-ball’s properties. Thus, three processes operate here to bring about an organism-environment match: the standard variational explanation of the selective survival of fit individuals, the transformation of the developmental environment experienced by the larvae arising through both maternal and larval activities, and the transformation of the larvae through the development of the focal phenotype in a specialised organism-constructed medium. Moreover, these three processes interact, both in the present and in the past, and cannot be traced to just one original cause.
It should also, by now, be apparent why we have called our book ‘Evolution Evolving’. The title illustrates our core claim that how organisms develop – including their behaviour, physiology and plasticity – and what organisms do – including their niche construction – influence the rate, pattern, and direction of evolution. Development matters in evolution; and it is precisely because development matters that evolution is evolving. This is an exciting time for evolutionary biology, and we have tried to write a book that communicates some of that excitement, but is written in a non-technical style that people can understand. At the same time, we hope that there is enough of substance in our work to be of interest to both professional biologists and philosophers. There is much more that I could mention – I have coauthored articles, or have worked closely with several other philosophers, including Robert Richardson, John Dupre, Nancy Cartwright, Lynn Chiu and Thomas Pradeu – but I don’t want this article to get too long. Suffice to say that a large number of philosophers have contributed to my intellectual development and that of my coauthors, and to arguments presented in our book. I hope one day that ‘Evolution Evolving’ will come to be regarded as a prime example of how the close interaction between biologists and philosophers can be productive.
To find out more about ‘Evolution Evolving’, and access a 30% discount if you pre-order a copy, see https://www.evolutionevolving.org
References
Chiu L 2023. https://www.issuelab.org/resources/40950/40950.pdf
Dawkins R. 2004. Biology & Philosophy
Godfrey-Smith,P. 1996. Complexity and the Function of Mind in Nature. Cambridge: Cambridge University Press.
Hull, D. L. 1988. Science as a Process: An Evolutionary Account of the Social and Conceptual Development of Science Chicago: University of Chicago Press
Kampourakis K & Uller T 2020. Philosophy of Science for Biologists. Cambridge: CUP.
Kitcher P 1987. Vaulting Ambition. Sociobiology and the Quest for Human Nature. MIT Press
Lala KN, Uller T, Feiner N, Feldman MW & Gilbert S 2024. Evolution Evolving: The Developmental Origins of Adaptation and Biodiversity. Princeton University Press
Laland, K.N. 2004. Extending the extended phenotype. Biology & Philosophy 19: 313–325.
Laland, K. N., F. J. Odling-Smee, & S. F. Gilbert. 2008. Evo-devo and niche construction: Building bridges. Journal of Experimental Zoology Part B 310:549–566.
Laland, K. N., K. Sterelny, F. J. Odling-Smee, W. Hoppitt, and T. Uller. 2011. Cause and effect in biology revisited: Is Mayr’s proximate-ultimate dichotomy still useful? Science 334:1512–1516.
Laland, K.N. & Sterelny, K. 2006. Seven reasons (not) to neglect niche construction. Evolution 60: 1751–1762.
Laland, K. N., T. Uller, M. W. Feldman, K. Sterelny, G. B. Müller, A. Moczek, E. Jablonka, et al. 2014. Does evolutionary theory need a rethink? Yes. Nature 514:161–164.
Laland, K. N., T. Uller, M. W. Feldman, K. Sterelny, G. B. Müller, A. Moczek, E. Jablonka, et al. 2015. The extended evolutionary synthesis: Its structure, assumptions and predictions. Proceedings of the Royal Society B 282:20151019.
Mayr, E. 1961. Cause and effect in biology. Science 134:1501–1506.
Oyama, S. 1985. The Ontogeny of Information: Developmental Systems and Evolution, 2nd ed. Durham, NC: Duke University Press.
Oyama, S., P. E. Griffiths, and R. D. Gray. 2001. Cycles of Contingency: Developmental Systems and Evolution. Cambridge, MA: MIT Press.
Pigliucci, M., and G. B. Müller. 2010. Evolution, the Extended Synthesis. Cambridge, MA: MIT Press.
Rupik, G. 2024. Remapping biology with Goethe, Schelling, and Herder. Romanticizing Evolution. London: Routledge.
Sober, E. 1984. The Nature of Selection: Evolutionary Theory in Philosophical Focus. Cambridge, MA: MIT Press.
Walsh, D. M. 2015. Organism, Agency, and Evolution. New York: Cambridge University Press.
Walsh, D. M. 2019. The paradox of population thinking: First order causes and higher order effects. In: Evolutionary Causation: Biological and Philosophical Reflections, ed. T Uller and K. N. Laland, 227–246. Cambridge, MA: MIT Press.
Wray, G. A., D. A. Futuyma, R. E. Lenski, T.F.C. MacKay, D. Schluter, J. E. Strassman, and H. E. Hoekstra. 2014. Does evolutionary biology need a rethink? Counterpoint: no, all is well. Nature 5:161–164.
Comments