In this article we explore the relationship between consciousness and the unconscious as it has taken shape within contemporary cognitive science - meaning by this term the mature cognitive science, which has fully incorporated the results of the neurosciences. In this framework we first compare the neurocognitive unconscious with the Freudian one, emphasizing the similarities and above all the differences between the two constructs. We then turn our attention to the implications of the centrality of unconscious processes in cognitive science (...) for the classical conception of the self. Our analysis will bring to light a bit of claustrophobic dialectic between an eliminative variety of naturalism and an anti-naturalistic form of hermeneutics. Hence we conclude by recommending the pursuit of a mediation between such extreme stances. (shrink)

Biological research on race has often been seen as motivated by or lending credence to underlying racist attitudes; in part for this reason, recently philosophers and biologists have gone through great pains to essentially deny the existence of biological human races. We argue that human races, in the biological sense of local populations adapted to particular environments, do in fact exist; such races are best understood through the common ecological concept of ecotypes. However, human ecotypic races do not in general (...) correspond with 'folk' racial categories, largely because many similar ecotypes have multiple independent origins. Consequently, while human natural races exist, they have little or nothing in common with 'folk' races. (shrink)

The Modern Synthesis (MS) is the current paradigm in evolutionary biology. It was actually built by expanding on the conceptual foundations laid out by its predecessors, Darwinism and neo-Darwinism. For sometime now there has been talk of a new Extended Evolutionary Synthesis (EES), and this article begins to outline why we may need such an extension, and how it may come about. As philosopher Karl Popper has noticed, the current evolutionary theory is a theory of genes, and we still lack (...) a theory of forms. The field began, in fact, as a theory of forms in Darwin’s days, and the major goal that an EES will aim for is a unification of our theories of genes and of forms. This may be achieved through an organic grafting of novel concepts onto the foundational structure of the MS, particularly evolvability, phenotypic plasticity, epigenetic inheritance, complexity theory, and the theory of evolution in highly dimensional adaptive landscapes. (shrink)

The so-called “New Atheism” is a relatively well-defined, very recent, still unfold- ing cultural phenomenon with import for public understanding of both science and philosophy. Arguably, the opening salvo of the New Atheists was The End of Faith by Sam Harris, published in 2004, followed in rapid succession by a number of other titles penned by Harris himself, Richard Dawkins, Daniel Dennett, Victor Stenger, and Christopher Hitchens.

Evolutionary theory is undergoing an intense period of discussion and reevaluation. This, contrary to the misleading claims of creationists and other pseudoscientists, is no harbinger of a crisis but rather the opposite: the field is expanding dramatically in terms of both empirical discoveries and new ideas. In this essay I briefly trace the conceptual history of evolutionary theory from Darwinism to neo-Darwinism, and from the Modern Synthesis to what I refer to as the Extended Synthesis, a more inclusive conceptual framework (...) containing among others evo–devo, an expanded theory of heredity, elements of complexity theory, ideas about evolvability, and a reevaluation of levels of selection. I argue that evolutionary biology has never seen a paradigm shift, in the philosophical sense of the term, except when it moved from natural theology to empirical science in the middle of the 19th century. The Extended Synthesis, accordingly, is an expansion of the Modern Synthesis of the 1930s and 1940s, and one that—like its predecessor—will probably take decades to complete. (shrink)

In recent years, biologists have increasingly been asking whether the ability to evolve — the evolvability — of biological systems, itself evolves, and whether this phenomenon is the result of natural selection or a by-product of other evolutionary processes. The concept of evolvability, and the increasing theoretical and empirical literature that refers to it, may constitute one of several pillars on which an extended evolutionary synthesis will take shape during the next few years, although much work remains to be done (...) on how evolvability comes about. (shrink)

The concept of burden of proof is used in a wide range of discourses, from philosophy to law, science, skepticism, and even in everyday reasoning. This paper provides an analysis of the proper deployment of burden of proof, focusing in particular on skeptical discussions of pseudoscience and the paranormal, where burden of proof assignments are most poignant and relatively clear-cut. We argue that burden of proof is often misapplied or used as a mere rhetorical gambit, with little appreciation of the (...) underlying principles. The paper elaborates on an important distinction between evidential and prudential varieties of burdens of proof, which is cashed out in terms of Bayesian probabilities and error management theory. Finally, we explore the relationship between burden of proof and several (alleged) informal logical fallacies. This allows us to get a firmer grip on the concept and its applications in different domains, and also to clear up some confusions with regard to when exactly some fallacies (ad hominem, ad ignorantiam, and petitio principii) may or may not occur. (shrink)

The “demarcation problem,” the issue of how to separate science from pseu- doscience, has been around since fall 1919—at least according to Karl Pop- per’s (1957) recollection of when he first started thinking about it. In Popper’s mind, the demarcation problem was intimately linked with one of the most vexing issues in philosophy of science, David Hume’s problem of induction (Vickers 2010) and, in particular, Hume’s contention that induction cannot be logically justified by appealing to the fact that “it works,” (...) as that in itself is an inductive argument, thereby potentially plunging the philosopher straight into the abyss of a viciously circular argument. (shrink)

Discussions about the biological bases (or lack thereof) of the concept of race in the human species seem to be never ending. One of the latest rounds is represented by a paper by Neven Sesardic, which attempts to build a strong scientific case for the existence of human races, based on genetic, morphometric and behavioral characteristics, as well as on a thorough critique of opposing positions. In this paper I show that Sesardic’s critique falls far short of the goal, and (...) that his positive case is exceedingly thin. I do this through a combination of analysis of the actual scientific findings invoked by Sesardic and of some philo- sophical unpacking of his conceptual analysis, drawing on a dual professional background as an evolu- tionary biologist and a philosopher of science. (shrink)

The so-called ‘‘species problem’’ has plagued evolution- ary biology since before Darwin’s publication of the aptly titled Origin of Species. Many biologists think the problem is just a matter of semantics; others complain that it will not be solved until we have more empirical data. Yet, we don’t seem to be able to escape discussing it and teaching seminars about it. In this paper, I briefly examine the main themes of the biological and philosophical liter- atures on the species problem, (...) focusing on identifying common threads as well as relevant differences. I then argue two fundamental points. First, the species problem is not primarily an empirical one, but it is rather fraught with philosophical questions that require—but cannot be settled by—empirical evidence. Second, the (dis-)solution lies in explicitly adopting Wittgenstein’s idea of ‘‘family resemblance’’ or cluster concepts, and to consider spe- cies as an example of such concepts. This solution has several attractive features, including bringing together apparently diverging themes of discussion among bio- logists and philosophers. The current proposal is con- ceptually independent (though not incompatible) with the pluralist approach to the species problem advocated by Mishler, Donoghue, Kitcher and Dupre ́, which implies that distinct aspects of the species question need to be emphasized depending on the goals of the researcher. From the biological literature, the concept of species that most closely matches the philosophical discussion pre- sented here is Templeton’s cohesion idea. (shrink)

Genes are often described by biologists using metaphors derived from computa- tional science: they are thought of as carriers of information, as being the equivalent of ‘‘blueprints’’ for the construction of organisms. Likewise, cells are often characterized as ‘‘factories’’ and organisms themselves become analogous to machines. Accordingly, when the human genome project was initially announced, the promise was that we would soon know how a human being is made, just as we know how to make airplanes and buildings. Impor- tantly, (...) modern proponents of Intelligent Design, the latest version of creationism, have exploited biologists’ use of the language of information and blueprints to make their spurious case, based on pseudoscientific concepts such as ‘‘irreducible complexity’’ and on flawed analogies between living cells and mechanical factories. However, the living organ- ism = machine analogy was criticized already by David Hume in his Dialogues Concerning Natural Religion. In line with Hume’s criticism, over the past several years a more nuanced and accurate understanding of what genes are and how they operate has emerged, ironically in part from the work of computational scientists who take biology, and in particular developmental biology, more seriously than some biologists seem to do. In this article we connect Hume’s original criticism of the living organism = machine analogy with the modern ID movement, and illustrate how the use of misleading and outdated metaphors in science can play into the hands of pseudoscientists. Thus, we argue that dropping the blueprint and similar metaphors will improve both the science of biology and its understanding by the general public. (shrink)

Mayr’s proximate–ultimate distinction has received renewed interest in recent years. Here we discuss its role in arguments about the relevance of developmental to evolutionary biology. We show that two recent critiques of the proximate–ultimate distinction fail to explain why developmental processes in particular should be of interest to evolutionary biologists. We trace these failures to a common problem: both critiques take the proximate–ultimate distinction to neglect specific causal interactions in nature. We argue that this is implausible, and that the distinction (...) should instead be understood in the context of explanatory abstractions in complete causal models of evolutionary change. Once the debate is reframed in this way, the proximate–ultimate distinction’s role in arguments against the theoretical significance of evo-devo is seen to rely on a generally implicit premise: that the variation produced by development is abundant, small and undirected. We show that a “lean version” of the proximate–ultimate distinction can be maintained even when this isotropy assumption does not hold. Finally, we connect these considerations to biological practice. We show that the investigation of developmental constraints in evolutionary transitions has long relied on a methodology which foregrounds the explanatory role of developmental processes. It is, however, entirely compatible with the lean version of the proximate–ultimate distinction. (shrink)

Evolutionary biology is a field currently animated by much discussion concerning its conceptual foundations. On the one hand, we have supporters of a classical view of evolutionary theory, whose backbone is provided by population genetics and the so-called Modern Synthesis (MS). On the other hand, a number of researchers are calling for an Extended Synthe- sis (ES) that takes seriously both the limitations of the MS (such as its inability to incorporate developmental biology) and recent empirical and theoretical research on (...) issues such as evolvability, modularity, and self-organization. In this article, I engage in an in-depth commentary of an influential paper by population geneticist Michael Lynch, which I take to be the best defense of the MS-population genetics position published so far. I show why I think that Lynch’s arguments are wanting and propose a modification of evolutionary theory that retains but greatly expands on population genetics. (shrink)

In a now classic paper published in 1991, Alberch introduced the concept of genotype–phenotype (G!P) mapping to provide a framework for a more sophisticated discussion of the integration between genetics and developmental biology that was then available. The advent of evo-devo first and of the genomic era later would seem to have superseded talk of transitions in phenotypic space and the like, central to Alberch’s approach. On the contrary, this paper shows that recent empirical and theoretical advances have only sharpened (...) the need for a different conceptual treat- ment of how phenotypes are produced. Old-fashioned metaphors like genetic blueprint and genetic programme are not only woefully inadequate but positively misleading about the nature of G!P, and are being replaced by an algorithmic approach emerging from the study of a variety of actual G!P maps. These include RNA folding, protein function and the study of evolvable soft- ware. Some generalities are emerging from these disparate fields of analysis, and I suggest that the concept of ‘developmental encoding’ (as opposed to the classical one of genetic encoding) provides a promising computational–theoretical underpinning to coherently integrate ideas on evolvability, modularity and robustness and foster a fruitful framing of the G!P mapping problem. (shrink)

Twenty years have passed since Gould and Lewontin published their critique of ‘the adaptationist program’ – the tendency of some evolutionary biologists to assume, rather than demonstrate, the operation of natural selection. After the ‘Spandrels paper’, evolutionists were more careful about producing just-so stories based on selection, and paid more attention to a panoply of other processes. Then came reactions against the excesses of the anti-adaptationist movement, which ranged from a complete dismissal of Gould and Lewontin’s contribution to a positive (...) call to overcome the problems. We now have an excellent opportunity for finally affirming a more balanced and pluralistic approach to the study of evolutionary biology. (shrink)

In addition to considerable debate in the recent evolutionary literature about the limits of the Modern Synthesis of the 1930s and 1940s, there has also been theoretical and empirical interest in a variety of new and not so new concepts such as phenotypic plasticity, genetic assimilation and phenotypic accommodation. Here we consider examples of the arguments and counter- arguments that have shaped this discussion. We suggest that much of the controversy hinges on several misunderstandings, including unwarranted fears of a general (...) attempt at overthrowing the Modern Synthesis paradigm, and some fundamental conceptual confusion about the proper roles of phenotypic plasticity and natural selection within evolutionary theory. (shrink)

In this paper I outline a theory of legitimacy that grounds the state’s right to rule on a natural duty not to harm others. I argue that by refusing to enter the state, anarchists expose those living next to them to the dangers of the state of nature, thereby posing an unjust threat. Since we have a duty not to pose unjust threats to others, anarchists have a duty to leave the state of nature and enter the state. This duty (...) correlates to a claim-right possessed by those living next to them, who also have a right to act in self-defence to enforce this obligation. This argument, if successful, would be particularly attractive, as it provides an account of state legitimacy without importing any normative premises that libertarians would reject. (shrink)

This paper outlines a critique of the use of the genetic variance–covariance matrix (G), one of the central concepts in the modern study of natural selection and evolution. Specifically, I argue that for both conceptual and empirical reasons, studies of G cannot be used to elucidate so-called constraints on natural selection, nor can they be employed to detect or to measure past selection in natural populations – contrary to what assumed by most practicing biologists. I suggest that the search for (...) a general solution to the difficult problem of identifying causal structures given observed correlation’s has led evolutionary quantitative geneticists to substitute statistical modeling for the more difficult, but much more valuable, job of teasing apart the many possible causes underlying the action of natural selection. Hence, the entire evolutionary quantitative genetics research program may be in need of a fundamental reconsideration of its goals and how they correspond to the array of mathematical and experimental techniques normally employed by its practitioners. (shrink)

Provided that traditional jus ad bellum principles are fulfilled, military humanitarian intervention to stop large scale violations of human rights (such as genocide, crimes against humanity or war crimes) is widely regarded as morally permissible. In cases of “supreme humanitarian emergency”, not only are the victims morally permitted to rebel, but other states are also permitted to militarily intervene. Things are different if the human rights violations in question fall short of supreme humanitarian emergency. Because of the importance of respecting (...) political self-determination, in cases of “ordinary oppression”, we normally think that rebellion might be permissible, but not military humanitarian intervention. Thus, according to the received view, the conditions for the permissibility of intervention coincide with the conditions for the permissibility of revolution in cases of supreme humanitarian emergency, but not in cases of ordinary oppression. In cases of ordinary oppression there is an asymmetry between the conditions for the permissibility of revolution and intervention (call this the Asymmetry View). Should we accept the Asymmetry View? I answer this question by outlining an account of political self-determination and by illustrating the complex role that this notion should play in discussing the morality of revolution and intervention. (shrink)

The idea of phenotypic novelty appears throughout the evolutionary literature. Novelties have been defined so broadly as to make the term meaningless and so narrowly as to apply only to a limited number of spectacular structures. Here I examine some of the available definitions of phenotypic novelty and argue that the modern synthesis is ill equipped at explaining novelties. I then discuss three frameworks that may help biologists get a better insight of how novelties arise during evolution but warn that (...) these frameworks should be considered in addition to, and not as potential substitutes of, the modern synthesis. †To contact the author, please write to: Departments of Ecology and Evolution and Philosophy, Stony Brook University, Stony Brook, NY 11794; e‐mail: pigliucci@platofootnote.org. (shrink)

Science and philosophy have a very long history, dating back at least to the 16th and 17th centuries, when the first scientist-philosophers, such as Bacon, Galilei, and Newton, were beginning the process of turning natural philosophy into science. Contemporary relationships between the two fields are still to some extent marked by the distrust that maintains the divide between the so-called “two cultures.” An increasing number of philosophers, however, are making conceptual contributions to sciences ranging from quantum mechanics to evolutionary biology, (...) and a few scientists are conducting research relevant to classically philosophical fields of inquiry, such as consciousness and moral decision-making. This article will introduce readers to the borderlands between science and philosophy, beginning with a brief description of what philosophy of science is about, and including a discussion of how the two disciplines can fruitfully interact not only at the level of scholarship, but also when it comes to controversies surrounding public understanding of science. (shrink)

Ever since Socrates, philosophers have been in the business of asking ques- tions of the type “What is X?” The point has not always been to actually find out what X is, but rather to explore how we think about X, to bring up to the surface wrong ways of thinking about it, and hopefully in the process to achieve an increasingly better understanding of the matter at hand. In the early part of the twentieth century one of the most (...) ambitious philosophers of sci- ence, Karl Popper, asked that very question in the specific case in which X = science. Popper termed this the “demarcation problem,” the quest for what distinguishes science from nonscience and pseudoscience (and, presumably, also the latter two from each other). (shrink)

The study of phenotypic plasticity has progressed significantly over the past few decades. We have moved from variation for plasticity being considered as a nuisance in evolutionary studies to it being the primary target of investigations that use an array of methods, including quantitative and molecular genetics, as well as of several approaches that model the evolution of plastic responses. Here, I consider some of the major aspects of research on phenotypic plasticity, assessing where progress has been made and where (...) additional effort is required. I suggest that some areas of research, such the study of the quantitative genetic underpinning of plasticity, have been either settled in broad outline or superseded by new approaches and questions. Other issues, such as the costs of plasticity are currently at the forefront of research in this field, and are likely to be areas of major future development. (shrink)

The debate about the levels of selection has been one of the most controversial both in evolutionary biology and in philosophy of science. Okasha’s book makes the sort of contribution that simply will not be able to be ignored by anyone interested in this field for many years to come. However, my interest here is in highlighting some examples of how Okasha goes about discussing his material to suggest that his book is part of an increasingly interesting trend that sees (...) scientists and philosophers coming together to build a broadened concept of “theory” through a combination of standard mathematical treatments and conceptual analyses. Given the often contentious history of the relationship between philosophy and science, such trend cannot but be welcome. (shrink)

A counter analysis of David Chalmers' claims about the possibility of mind uploading within the context of the Singularity event.

‘‘Theoretical biology’’ is a surprisingly heter- ogeneous field, partly because it encompasses ‘‘doing the- ory’’ across disciplines as diverse as molecular biology, systematics, ecology, and evolutionary biology. Moreover, it is done in a stunning variety of different ways, using anything from formal analytical models to computer sim- ulations, from graphic representations to verbal arguments. In this essay I survey a number of aspects of what it means to do theoretical biology, and how they compare with the allegedly much more restricted (...) sense of theory in the physical sciences. I also tackle a recent trend toward the presentation of all-encompassing theories in the biological sciences, from general theories of ecology to a recent attempt to provide a conceptual framework for the entire set of biological disciplines. Finally, I discuss the roles played by philosophers of science in criticizing and shap- ing biological theorizing. (shrink)

Are science and religion compatible when it comes to understanding cosmology (the origin of the universe), biology (the origin of life and of the human species), ethics, and the human mind (minds, brains, souls, and free will)? Do science and religion occupy non-overlapping magisteria? Is Intelligent Design a scientific theory? How do the various faith traditions view the relationship between science and religion? What, if any, are the limits of scientific explanation? What are the most important open questions, problems, or (...) challenges confronting the relationship between science and religion, and what are the prospects for progress? These and other questions are explored in Science and Religion: 5 Questions--a collection of thirty-three interviews based on 5 questions presented to some of the world's most influential and prominent philosophers, scientists, theologians, apologists, and atheists. Contributions by Simon Blackburn, Susan Blackmore, Sean Carroll, William Lane Craig, William Dembski, Daniel C. Dennett, George F.R. Ellis, Owen Flanagan, Owen Gingerich, Rebecca Newberger Goldstein, John F. Haught, Muzaffar Iqbal, Lawrence Krauss, Colin McGinn, Alister McGrath, Mary Midgley, Seyyed Hossein Nasr, Timothy O'Connor, Massimo Pigliucci, John Polkinghorne, James Randi, Alex Rosenberg, Michael Ruse, Robert John Russell, John Searle, Michael Shermer, Victor J. Stenger, Robert Thurman, Michael Tooley, Charles Townes, Peter van Inwagen, Keith Ward, Rabbi David Wolpe. (shrink)

Just war theory is currently dominated by two positions. According to the orthodox view, provided that jus in bello principles are respected, combatants have an equal right to fight, regardless of the justice of the cause pursued by their state. According to “revisionists” whenever combatants lack reasons to believe that the war they are ordered to fight is just, their duty is to disobey. I argue that when members of a legitimate state acting in good faith are ordered to fight, (...) they acquire a pro-tanto obligation to obey which does not depend for its validity on the justice of the cause being pursued. However, when the war is unjust, this obligation may be overridden, under certain conditions, by the obligation not to contribute to the unjustified killing of innocents. This is because the pro-tanto force of the duty to obey the law is best understood in terms of “presumptive”, rather than “exclusionary” reasons for action. This approach captures the insights of both the orthodox and the revisionist view, while avoiding the problems that afflict each of them. (shrink)

The idea of genetic assimilation, that environmentally induced phenotypes may become genetically fixed and no longer require the original environmental stimulus, has had varied success through time in evolutionary biology research. Proposed by Waddington in the 1940s, it became an area of active empirical research mostly thanks to the efforts of its inventor and his collaborators. It was then attacked as of minor importance during the ‘‘hardening’’ of the neo-Darwinian synthesis and was relegated to a secondary role for decades. Recently, (...) several papers have appeared, mostly independently of each other, to explore the likelihood of genetic assimilation as a biological phenomenon and its potential importance to our understanding of evolution. In this article we briefly trace the history of the concept and then discuss theoretical models that have newly employed genetic assimilation in a variety of contexts. We propose a typical scenario of evolution of genetic assimilation via an intermediate stage of phenotypic plasticity and present potential examples of the same. We also discuss a conceptual map of current and future lines of research aimed at exploring the actual relevance of genetic assimilation for evolutionary biology. (shrink)

Why Jerry Fodor and Massimo Piattelli-Palmarini are wrong about Darwin and evolution.

What Jerry Fodor and Massimo Piattelli-Palmarini got wrong about Darwin and evolution.

The creation-evolution “controversy” has been with us for more than a century. Here I argue that merely teaching more science will probably not improve the situation; we need to understand the controversy as part of a broader problem with public acceptance of pseudoscience, and respond by teaching how science works as a method. Critical thinking is difficult to teach, but educators can rely on increasing evidence from neurobiology about how the brain learns, or fails to.

The relationship between ethics and science has been discussed within the framework of continuity versus discontinuity theories, each of which can take several forms. Continuity theorists claim that ethics is a science or at least that it has deep similarities with the modus operandi of science. Discontinuity theorists reject such equivalency, while at the same time many of them claim that ethics does deal with objective truths and universalizable statements, just not in the same sense as science does. I propose (...) here a third view of quasi-continuity (or, equivalently, quasi-discontinuity) that integrates ethics and science as equal partners toward the uncovering of new knowledge. In this third way, a program envisioned by William James but made practicable only by contemporary scientific advancement, science can and must inform ethics at a deep level, and ethical theory— while going beyond science—cannot do without it. In particular, I identify four areas of ethics-science collaboration: neurobiological research into the basis of moral judgment, comparative anthropol- ogy, comparative evolutionary biology of primates, and game-theo- retical modeling. I provide examples within each of these fields to show how they link to ethical theories (including prescriptive work) and questions. The essay concludes with a brief discussion of the light that a scientifically informed ethics can shed on some classical problems in moral theory, such as the relationships between rational- ity and selfishness, egoism and altruism, as well as the concept of social contract. A joint research program involving both philosophers and scientists is called for if we wish to move ethical theory into the twenty-first century. (shrink)

Sewall Wright introduced the metaphor of evolution on “adaptive landscapes” in a pair of papers published in 1931 and 1932. The metaphor has been one of the most influential in modern evolutionary biology, although recent theoretical advancements show that it is deeply flawed and may have actually created research questions that are not, in fact, fecund. In this paper I examine in detail what Wright actually said in the 1932 paper, as well as what he thought of the matter at (...) the very end of his career, in 1988. While the metaphor is flawed, some of the problems which Wright was attempting to address are still with us today, and are in the process of being reformulated as part of a forthcoming Extended Evolutionary Synthesis. (shrink)

How micro- and macroevolutionary evolutionary processes produce phenotypic change is without question one of the most intriguing and perplexing issues facing evolutionary biologists. We believe that roadblocks to progress lie A) in the underestimation of the role of the environment, and in particular, that of the interaction of genotypes with environmental factors, and B) in the continuing lack of incorporation of development into the evolutionary synthesis. We propose the integration of genetic, environmental and developmental perspectives on the evolution of the (...) phenotype in the form of the concept of the developmental reaction norm (DRN) The DRN represents the set of multivariate ontogenies that can be produced by a single genotype when it is exposed to environmental variation. It encompasses: 1) the processes that alter the phenotype throughout the ontogenetic trajectory, 2) the recognition that different aspects of the phenotype are (and must be) correlated and 3) the ability of a genotype to produce phenotypes in different environments. This perspective necessitates the explicit study of character expression during development, the evaluation of associations between pairs or groups of characters (e.g., multivariate allometries), and the exploration of reaction norms and phenotypic plasticity. We explicitly extend the concept of the DRN to encompass adjustments made in response to changes in the internal environment as well. Thus, ‘typical’ developmental sequences (e.g., cell fate determination) and plastic responses are simply manifestations of different scales of ‘environmental’ effects along a continuum. We present: (1) a brief conceptual review of three fundamental aspects of the generation and evolution of phenotypes: the changes in the trajectories describing growth and differentiation (ontogeny), the multivariate relationships among characters (allometry), and the effect of the environment (plasticity); (2) a discussion of how these components are merged in the concept of the developmental reaction norm; and (3) a reaction norm perspective of major determinants of phenotypes: epigenesis, selection and constraint. (shrink)

Recent debates between proponents of the modern evolutionary synthesis (the standard model in evolutionary biology) and those of a possible extended synthesis are a good example of the fascinating tangle among empirical, theoretical, and conceptual or philosophical matters that is the practice of evolutionary biology. In this essay, we briefly discuss two case studies from this debate, highlighting the relevance of philosophical thinking to evolutionary biologists in the hope of spurring further constructive cross-pollination between the two fields.

During the last two decades the role of quantitative genetics in evolutionary theory has expanded considerably. Quantitative genetic-based models addressing long term phenotypic evolution, evolution in multiple environments (phenotypic plasticity) and evolution of ontogenies (developmental trajectories) have been proposed. Yet, the mathematical foundations of quantitative genetics were laid with a very different set of problems in mind (mostly the prediction of short term responses to artificial selection), and at a time in which any details of the genetic machinery were virtually (...) unknown. In this paper we discuss what a model is in population biology, and what kind of model we need in order to address the complexities of phenotypic evolution. We review the assumptions of quantitative genetics and its most recent accomplishments, together with the limitations that such assumptions impose on the modelling of some aspects of phenotypic evolution. We also discuss three alternative appr oaches to the theoretical description of evolutionary trajectories (nonlinear dynamics, complexity theory and optimization theory), and their respective advantages and limitations. We conclude by calling for a new theoretical synthesis, including quantitative genetics and not necessarily limited to the other approaches here discussed. (shrink)

Research in ecology and evolutionary biology (evo-eco) often tries to emulate the “hard” sciences such as physics and chemistry, but to many of its practitioners feels more like the “soft” sciences of psychology and sociology. I argue that this schizophrenic attitude is the result of lack of appreciation of the full consequences of the peculiarity of the evo-eco sciences as lying in between a-historical disciplines such as physics and completely historical ones as like paleontology. Furthermore, evo-eco researchers have gotten stuck (...) on mathematically appealing but philosophi- cally simplistic concepts such as null hypotheses and p-values defined according to the frequentist approach in statistics, with the consequence of having been unable to fully embrace the complexity and subtlety of the problems with which ecologists and evolutionary biologists deal with. I review and discuss some literature in ecology, philosophy of science and psychology to show that a more critical methodological attitude can be liberating for the evo-eco scientist and can lead to a more fecund and enjoyable practice of ecology and evolutionary biology. With this aim, I briefly cover concepts such as the method of multiple hypotheses, Bayesian analysis, and strong inference. (shrink)

Biologists are increasingly reexamining the conceptual structure of evolutionary theory, which dates back to the so-called Modern Synthesis of the 1930s and 1940s. Calls for an Extended Evolutionary Synthesis (EES) cite a number of empir- ical and theoretical advances that need to be accounted for, including evolvability, evo- lutionary novelties, capacitors of phenotypic evolution, developmental plasticity, and phenotypic attractors. In Biological Emergences, however, Robert Reid outlines a theory of evolution in which natural selection plays no role or—worse—actually impedes evo- lution (...) by what Reid calls “natural experimentation.” For Reid, biological complexity emerges because of intrinsic mechanisms that work in opposition to natural selection, a view that would reopen old questions of orthogenesis and Lamarckism.This review outlines why we do need an EES, but also why it is unlikely to take the shape that Reid advocates. (shrink)

Ever since Darwin a great deal of the conceptual history of biology may be read as a struggle between two philosophical positions: reductionism and holism. On the one hand, we have the reductionist claim that evolution has to be understood in terms of changes at the fundamental causal level of the gene. As Richard Dawkins famously put it, organisms are just ‘lumbering robots’ in the service of their genetic masters. On the other hand, there is a long holistic tradition that (...) focuses on the complexity of developmental systems, on the non-linearity of gene– environment interactions, and on multi-level selective processes to argue that the full story of biology is a bit more complicated than that. Reductionism can marshal on its behalf the spectacular successes of genetics and molecular biology throughout the 20th and 21st centuries. Holism has built on the development of entirely new disciplines and conceptual frameworks over the past few decades, including evo-devo and phenotypic plasticity. Yet, a number of biologists are still actively looking for a way out of the reductionism–holism counterposition, often mentioning the word ‘emergence’ as a way to deal with the conundrum. This paper briefly examines the philosophical history of the concept of emergence, distinguishes between epistemic and ontological accounts of it, and comments on conceptions of emergence that can actually be useful for practising evolutionary biologists. (shrink)

The concept of reaction norms plays a crucial role in connecting molecular and evolutionary biology.

It is an unfortunate fact of academic life that there is a sharp divide between science and philosophy, with scientists often being openly dismissive of philosophy, and philosophers being equally contemptuous of the naivete ́ of scientists when it comes to the philosophical underpinnings of their own discipline. In this paper I explore the possibility of reducing the distance between the two sides by introducing science students to some interesting philosophical aspects of research in evolutionary biology, using biological theories of (...) the origin of religion as an example. I show that philosophy is both a discipline in its own right as well as one that has interesting implications for the understanding and practice of science. While the goal is certainly not to turn science students into philoso- phers, the idea is that both disciplines cannot but benefit from a mutual dialogue that starts as soon as possible, in the classroom. (shrink)

Public discussions of science are often marred by two pernicious phenomena: a widespread rejection of scientific findings (e.g., the reality of anthropogenic climate change, the conclusion that vaccines do not cause autism, or the validity of evolutionary theory), coupled with an equally common acceptance of pseudoscientific notions (e.g., homeopathy, psychic readings, telepathy, tall tales about alien abductions, and so forth). The typical reaction by scientists and science educators is to decry the sorry state of science literacy among the general public, (...) and to call for more science education as the answer to both problems. But the empirical evidence concerning the relationship between science literacy, rejection of science and acceptance of pseudoscience is mixed at best. In this chapter I argue that—while certainly important—efforts at increasing public knowledge of science (science education) need to be complemented by attention to common logical fallacies (philosophy), cognitive biases and dissonance (psychology), and the role of ideological commitments (sociology). Even this complex, multi-disciplinary approach to science education will likely only yield measurable results in the very long term. Meanwhile science remains, as Carl Sagan famously put it, a candle in the dark, delicate and in need of much nurturing. (shrink)

The scientific status of evolutionary theory seems to be more or less perennially under question. I am not referring here (just) to the silliness of young Earth creation- ism (Pigliucci 2002; Boudry and Braeckman 2010), or even of the barely more intel- lectually sophisticated so-called Intelligent Design theory (Recker 2010; Brigandt this volume), but rather to discussions among scientists and philosophers of science concerning the epistemic status of evolutionary theory (Sober 2010). As we shall see in what follows, this debate (...) has a long history, dating all the way back to Darwin, and it is in great part rooted in the fundamental dichotomy between what French biologist and Nobel laureate Jacques Monod (1971) called chance and necessity—i.e., the inevitable and inextricable interplay of deterministic and stochastic mechanisms operating during the course of evolution. (shrink)

Few metaphors in biology are more enduring than the idea of Adaptive Landscapes, originally proposed by Sewall Wright (1932) as a way to visually present to an audience of typically non- mathematically savvy biologists his ideas about the relative role of natural selection and genetic drift in the course of evolution. The metaphor, how- ever, was born troubled, not the least reason for which is the fact that Wright presented different diagrams in his original paper that simply can- not refer (...) to the same concept and are therefore hard to reconcile with each other (Pigliucci 2008). For instance, in some usages, the landscape’s non- fitness axes represent combinations of individual genotypes (which cannot sensibly be aligned on a linear axis, and accordingly were drawn by Wright as polyhedrons of increasing dimensionality). In other usages, however, the points on the diagram represent allele or genotypic frequencies, and so are actually populations, not individuals (and these can indeed be coherently represented along continuous axes). (shrink)

Metaphors play a crucial role in both science in particular and human discourse in gen- eral. Plato’s story of the cave—about people shackled to a wall and incapable of perceiv- ing the world as it really is—has stimulated thinking about epistemology and the nature of reality for more than two millennia. But metaphors can also be misleading: being too taken with Plato’s story has cost philosophers endless discussions about how to access the world “as it is,” until Kant showed us (...) that it is just not going to happen, ever. (shrink)

The scientific study of living organisms is permeated by machine and design metaphors. Genes are thought of as the ‘‘blueprint’’ of an organism, organisms are ‘‘reverse engineered’’ to discover their func- tionality, and living cells are compared to biochemical factories, complete with assembly lines, transport systems, messenger circuits, etc. Although the notion of design is indispensable to think about adapta- tions, and engineering analogies have considerable heuristic value (e.g., optimality assumptions), we argue they are limited in several important respects. In (...) particular, the analogy with human-made machines falters when we move down to the level of molecular biology and genetics. Living organisms are far more messy and less transparent than human-made machines. Notoriously, evolution is an oppor- tunistic tinkerer, blindly stumbling on ‘‘designs’’ that no sensible engineer would come up with. Despite impressive technological innovation, the prospect of artificially designing new life forms from scratch has proven more difficult than the superficial analogy with ‘‘programming’’ the right ‘‘software’’ would sug- gest. The idea of applying straightforward engineering approaches to living systems and their genomes— isolating functional components, designing new parts from scratch, recombining and assembling them into novel life forms—pushes the analogy with human artifacts beyond its limits. In the absence of a one-to-one correspondence between genotype and phenotype, there is no straightforward way to imple- ment novel biological functions and design new life forms. Both the developmental complexity of gene expression and the multifarious interactions of genes and environments are serious obstacles for ‘‘engi- neering’’ a particular phenotype. The problem of reverse-engineering a desired phenotype to its genetic ‘‘instructions’’ is probably intractable for any but the most simple phenotypes. Recent developments in the field of bio-engineering and synthetic biology reflect these limitations. Instead of genetically engi- neering a desired trait from scratch, as the machine/engineering metaphor promises, researchers are making greater strides by co-opting natural selection to ‘‘search’’ for a suitable genotype, or by borrowing and recombining genetic material from extant life forms. (shrink)

Is integrative biology a good idea, or even possible? There has been much interest lately in the unifica- tion of biology and the integration of traditionally separate disciplines such as molecular and develop- mental biology on one hand, and ecology and evolutionary biology on the other. In this paper I ask if and under what circumstances such integration of efforts actually makes sense. I develop by example an analogy with Aristotle’s famous four “causes” that one can investigate concerning any object (...) or phenomenon: material (what something is made of), formal (what distinguishes that particular object from others), efficient (how was the object made) and final (why was the object made). The example is provided by ongoing research on different aspects of flowering time in the model system Arabidop- sis, a small weed belonging to the mustard family. I show that understanding how flowering time is controlled is an epistemologically different sort of question from why and how it evolved, and that the two research agendas can be pursued largely independently of each other. Toward the end, I propose that the real goal of integrative biology is to understand the boundary layers between levels of biologi- cal analysis, something to which modern philosophy of science can contribute significantly. (shrink)