The study of organisms within the range of their existence from fertilization to birth is referred to as embryology. The process of progressive change during that period is called development. That development does not stop at birth but continues on throughout the entire life-span of the organism as the process of growth and decay — catabolism, anabolism, and metabolism. The study of this entire range of life has recently become known as developmental biology. The belief that the development (...) from an initial stage of a fertilized egg or zygote to a fully formed adult represents, in compressed time, the whole process of evolution that occurred over millions of years, has recently been named evo-devo, or evolutionary development. The more science advances, the more it studies Nature in its intimate details, the more it comes to realize the existence of a pervasive reason, an inherent natural intelligence that is working in even the most insignificant portions of the universe. Francis Bacon (1561–1626) said, “A little philosophy (science) inclineth man's mind to atheism, but depth in philosophy (science) bringeth men's minds about to religion.” This point is especially true today. It is not from ignorance that men come to have faith in God, but from a maturity of reason and experience. Vedanta philosophy teaches that there is a conscious intelligence that underlies all experienced existence. Being self-evident, this should hardly have to be argued. Yet modern science has failed to integrate this truth into its materialist/naturalistic paradigm. Correcting this deficiency will be the challenge of 21st century science, and the highest reward for humanity. (shrink)

Evolutionary developmental biology (Evo-Devo) is a new and rapidly developing field of biology which focuses on questions in the intersection of evolution and development and has been seen by many as a potential synthesis of these two fields. This synthesis is the topic of the books reviewed here. Integrating Evolution and Development (edited by Roger Sansom and Robert Brandon), is a collection of papers on conceptual issues in Evo-Devo, while From Embryology to Evo-Devo (edited by Manfred Laubichler (...) and Jane Maienschein) is a history of the problem of the relations between ontogeny and phylogeny. (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)

An exploration of the nexus between ecology, evolutionary biology and molecular biology, via the concept of phenotypic plasticity.

Although contemporary metaphysics has recently undergone a neo-Aristotelian revival wherein dispositions, or capacities are now commonplace in empirically grounded ontologies, being routinely utilised in theories of causality and modality, a central Aristotelian concept has yet to be given serious attention – the doctrine of hylomorphism. The reason for this is clear: while the Aristotelian ontological distinction between actuality and potentiality has proven to be a fruitful conceptual framework with which to model the operation of the natural world, the distinction between (...) form and matter has yet to similarly earn its keep. In this chapter, I offer a first step toward showing that the hylomorphic framework is up to that task. To do so, I return to the birthplace of that doctrine - the biological realm. Utilising recent advances in developmental biology, I argue that the hylomorphic framework is an empirically adequate and conceptually rich explanatory schema with which to model the nature of organisms. (shrink)

Griffiths and Russell D. Gray (1994, 1997, 2001) have argued that the fundamental unit of analysis in developmental systems theory should be a process – the life cycle – and not a set of developmental resources and interactions between those resources. The key concepts of developmental systems theory, epigenesis and developmental dynamics, both also suggest a process view of the units of development. This chapter explores in more depth the features of developmental systems theory that (...) favour treating processes as fundamental in biology and examines the continuity between developmental systems theory and ideas about process in the work of several major figures in early 20th century biology, most notable C.H Waddington. (shrink)

I analyze the importance of parts in the style of biological theorizing that I call compositional biology. I do this by investigating various aspects, including partitioning frames and explanatory accounts, of the theoretical perspectives that fall under and are guided by compositional biology. I ground this general examination in a comparative analysis of three different disciplines with their associated compositional theoretical perspectives: comparative morphology, functional morphology, and developmental biology. I glean data for this analysis from canonical (...) textbooks and defend the use of such texts for the philosophy of science. I end with a discussion of the importance of recognizing formal and compositional biology as two genuinely different ways of doing biology – the differences arising more from their distinct methodologies than from scientific discipline included or natural domain studied. Ultimately, developing a translation manual between the two styles would be desirable as they currently are, at times, in conflict. (shrink)

A strong motivation for the human genome project was to relate biological features to the structure and function of small sets of genes, and ideally to individual genes. However, it is now increasingly realized that many problems require a "systems" approach emphasizing the interplay of large numbers of genes, and the involvement of complex networks of gene regulation. This implies a new emphasis on integrative, systems theoretical approaches. It may be called 'holistic' if the term is used without irrational overtones, (...) in the general sense of directing attention to integrated features of organs and organisms. In the history of biology, seemingly conflicting reductionist and holistic notions have alternated, with bottom-up as well as top-down approaches eventually contributing to the solutions of basic problems. By now, there is no doubt that biological features and phenomena are rooted in physico-chemical processes of the molecules involved; and yet, integrated systems aspects are becoming more and more relevant in developmental biology, brain and behavioural science, and socio-biology. -/- . (shrink)

Developmental systems theory (DST) is a wholeheartedly epigenetic approach to development, inheritance and evolution. The developmental system of an organism is the entire matrix of resources that are needed to reproduce the life cycle. The range of developmental resources that are properly described as being inherited, and which are subject to natural selection, is far wider than has traditionally been allowed. Evolution acts on this extended set of developmental resources. From a developmental systems perspective, development (...) does not proceed according to a preformed plan; what is inherited is much more than DNA; and evolution is change not only in gene frequencies, but in entire developmental systems. (shrink)

Genes and the Agents of Life undertakes to rethink the place of the individual in the biological sciences, drawing parallels with the cognitive and social sciences. Genes, organisms, and species are all agents of life but how are each of these conceptualized within genetics, developmental biology, evolutionary biology, and systematics? The 2005 book includes highly accessible discussions of genetic encoding, species and natural kinds, and pluralism above the levels of selection, drawing on work from across the biological (...) sciences. The book is a companion to the author's Boundaries of the Mind (2004), also available from Cambridge, where the focus is the cognitive sciences. The book will appeal to a broad range of professionals and students in philosophy, biology, and the history of science. You can download the table of contents and the first chapter here. (shrink)

The cis-regulatory hypothesis is one of the most important claims of evolutionary developmental biology. In this paper I examine the theoretical argument for cis-regulatory evolution and its role within evolutionary theorizing. I show that, although the argument has some weaknesses, it acts as a useful example for the importance of current scientific debates for science education.

Developmental plasticity as the nexus between genetics and ecology.

There are two fundamentally distinct kinds of biological theorizing. "Formal biology" focuses on the relations, captured in formal laws, among mathematically abstracted properties of abstract objects. Population genetics and theoretical mathematical ecology, which are cases of formal biology, thus share methods and goals with theoretical physics. "Compositional biology," on the other hand, is concerned with articulating the concrete structure, mechanisms, and function, through developmental and evolutionary time, of material parts and wholes. Molecular genetics, biochemistry, developmental (...) biology, and physiology, which are examples of compositional biology, are in serious need of philosophical attention. For example, the very concept of a "part" is understudied in both philosophy of biology and philosophy of science. ;My dissertation is an attempt to clarify the distinction between formal biology and compositional biology and, in so doing, provide a clear philosophical analysis, with case studies, of compositional biology. Given the social, economic, and medical importance of compositional biology, understanding it is urgent. For my investigation, I draw on the philosophical fields of metaphysics and epistemology, as well as philosophy of biology and philosophy of science. I suggest new ways of thinking about some classic philosophy of science issues, such as modeling, laws of nature, abstraction, explanation, and confirmation. I hint at the relevance of my study of two kinds of biological theorizing to debates concerning the disunity of science. (shrink)

This chapter introduces the main research schools and paradigms along which the field of evolutionary biology has been developing. Evolutionary thinking was originally founded upon the Neo-Darwinian paradigm that combines the teachings of traditional Darwinism with those of the Modern Synthesis. The Neo-Darwinian paradigm has since further diversified into the Micro-, Meso-, and Macroevolutionary schools, and it has also started to integrate the school of Ecology. Together, these schools establish the paradigm called Ecological Evolutionary Developmental Biology (Eco-Evo-Devo). (...) A final school studies Reticulate Evolution as it occurs by means of symbiosis, symbiogenesis, lateral gene transfer, infective heredity, and hybridization. This chapter reviews the major tenets and points of differentiation that exist between these distinct evolution schools. The chapter ends by looking into how evolution can be defined and how distinct units, levels, and mechanisms underlie theorizing on evolutionary hierarchies and evolutionary causation. The following chapter examines how the different evolution schools are implemented into the symbolic sciences. (shrink)

A common view of how-possibly explanations in biology treats them as explanatorily incomplete. In addition to this interpretation of how-possibly explanation, I argue that there is another interpretation, one which features what I term “explanatory strategies.” This strategy-centered interpretation of how-possibly explanation centers on there being a different explanatory context within which how-possibly explanations are offered. I contend that, in conditions where this strategy context is recognized, how-possibly explanations can be understood as complete explanations. I defend this alternative interpretation (...) by analyzing the explanatory value of simple physical models the nineteenth century developmental biologist Wilhelm His constructed for animal development. (shrink)

Biological modalities, i.e., biologically possible, impossible, or necessary states of affairs have not received much attention from philosophers. Yet, it is widely agreed that there are biological constraints on physically possible states of affairs, such that not everything that is physically possible is also biologically possible, even if everything that is biologically possible is also physically possible. Furthermore, biologists use concepts that appear to be modal in nature, such as the concept of evolvability in evolutionary developmental biology, or (...) “evo-devo.” The present chapter investigates what kind of modality underlies the concept of evolvability. This concept tries to capture the capacity of an organism or a lineage to sustain genetic changes that enable it to evolve or to evolve adaptively. The basic idea of the proposed approach is to construe evolvability as a kind of accessibility in a modal space. The difficult part is to specify this modal space and the relevant accessibility relation. While there may not be a general way of defining such a relation, there exist model systems for which it is possible, e.g., evolving small RNAs. The modal space in such cases turns out to be quite distinct from those constructed by philosophers, e.g., David Lewis’s similarity metric for possible worlds. Even though the biological case examined here is quite special, attending to the way in which biological possibilities are modeled in this case harbors some general lessons about biological modalities, in particular their dependence on the explanatory goals of the models modeling modality. (shrink)

A growing number of linguistic attempts to explain how languages change use cultural-evolutionary models involving selection or drift. Developmental constraints and biases, which take center stage in evolutionary developmental biology or evo-devo, seem to be absent within this framework, even though linguistics is home to numerous notions of constraint. In this paper, we show how these evo-devo concepts could be applied to linguistic change and why they should. This requires some conceptual groundwork, due to important differences between (...) linguistic and biotic evolution. In biological evolution, development generates the organism's variable traits on which selection and drift act. In linguistic evolution by analogy, we say development generates the linguistic variants on which selection and drift can act. “Linguistic development” then picks out how individual speakers produce and comprehend language. It involves much more than just learning. Using this broad notion of development, we distinguish between different types of bias that could operate in the processes of linguistic innovation and transmission, which correspond to genetic mutation and biological reproduction, respectively. Having thus sharpened our conceptual toolbox, we then reanalyze two well-documented cases of linguistic change and show that, in both these cases, linguists have only considered Neo-Darwinian evolutionary explanations, falsely deploying an exclusive disjunction of selection and drift. We show that there is at least a third relevant alternative in these examples, namely developmental constraint or bias in the sense we explicate here. (shrink)

The INBIOSA project brings together a group of experts across many disciplines who believe that science requires a revolutionary transformative step in order to address many of the vexing challenges presented by the world. It is INBIOSA’s purpose to enable the focused collaboration of an interdisciplinary community of original thinkers. This paper sets out the case for support for this effort. The focus of the transformative research program proposal is biology-centric. We admit that biology to date has been (...) more fact-oriented and less theoretical than physics. However, the key leverageable idea is that careful extension of the science of living systems can be more effectively applied to some of our most vexing modern problems than the prevailing scheme, derived from abstractions in physics. While these have some universal application and demonstrate computational advantages, they are not theoretically mandated for the living. A new set of mathematical abstractions derived from biology can now be similarly extended. This is made possible by leveraging new formal tools to understand abstraction and enable computability. [The latter has a much expanded meaning in our context from the one known and used in computer science and biology today, that is "by rote algorithmic means", since it is not known if a living system is computable in this sense (Mossio et al., 2009).] Two major challenges constitute the effort. The first challenge is to design an original general system of abstractions within the biological domain. The initial issue is descriptive leading to the explanatory. There has not yet been a serious formal examination of the abstractions of the biological domain. What is used today is an amalgam; much is inherited from physics (via the bridging abstractions of chemistry) and there are many new abstractions from advances in mathematics (incentivized by the need for more capable computational analyses). Interspersed are abstractions, concepts and underlying assumptions “native” to biology and distinct from the mechanical language of physics and computation as we know them. A pressing agenda should be to single out the most concrete and at the same time the most fundamental process-units in biology and to recruit them into the descriptive domain. Therefore, the first challenge is to build a coherent formal system of abstractions and operations that is truly native to living systems. Nothing will be thrown away, but many common methods will be philosophically recast, just as in physics relativity subsumed and reinterpreted Newtonian mechanics. -/- This step is required because we need a comprehensible, formal system to apply in many domains. Emphasis should be placed on the distinction between multi-perspective analysis and synthesis and on what could be the basic terms or tools needed. The second challenge is relatively simple: the actual application of this set of biology-centric ways and means to cross-disciplinary problems. In its early stages, this will seem to be a “new science”. This White Paper sets out the case of continuing support of Information and Communication Technology (ICT) for transformative research in biology and information processing centered on paradigm changes in the epistemological, ontological, mathematical and computational bases of the science of living systems. Today, curiously, living systems cannot be said to be anything more than dissipative structures organized internally by genetic information. There is not anything substantially different from abiotic systems other than the empirical nature of their robustness. We believe that there are other new and unique properties and patterns comprehensible at this bio-logical level. The report lays out a fundamental set of approaches to articulate these properties and patterns, and is composed as follows. -/- Sections 1 through 4 (preamble, introduction, motivation and major biomathematical problems) are incipient. Section 5 describes the issues affecting Integral Biomathics and Section 6 -- the aspects of the Grand Challenge we face with this project. Section 7 contemplates the effort to formalize a General Theory of Living Systems (GTLS) from what we have today. The goal is to have a formal system, equivalent to that which exists in the physics community. Here we define how to perceive the role of time in biology. Section 8 describes the initial efforts to apply this general theory of living systems in many domains, with special emphasis on crossdisciplinary problems and multiple domains spanning both “hard” and “soft” sciences. The expected result is a coherent collection of integrated mathematical techniques. Section 9 discusses the first two test cases, project proposals, of our approach. They are designed to demonstrate the ability of our approach to address “wicked problems” which span across physics, chemistry, biology, societies and societal dynamics. The solutions require integrated measurable results at multiple levels known as “grand challenges” to existing methods. Finally, Section 10 adheres to an appeal for action, advocating the necessity for further long-term support of the INBIOSA program. -/- The report is concluded with preliminary non-exclusive list of challenging research themes to address, as well as required administrative actions. The efforts described in the ten sections of this White Paper will proceed concurrently. Collectively, they describe a program that can be managed and measured as it progresses. (shrink)

The concept of downward causation is frequently used in an explanatory capacity in biology to account for certain regularities and processes. Some philosophers, however, argue that downward causation is metaphysically incoherent, providing three main objections. Underlying these objections is the assumption that entities are connected by compositional hierarchies of levels of organization. In this paper, I introduce the notions of weak and strong compositional relations using examples from evolutionary developmental biology. I argue that downward causation becomes unproblematic (...) if we use features of interventionist theories of causation to explain the causal relations between levels. I show that an interventionist account of downward causation successfully responds to the three central objections to downward causation in the philosophical literature and I clarify the explanatory usefulness of the concept in biology. As such, this paper provides an epistemic solution to demystify downward causation in the context of scientific practice. The solution proposed is compatible with how biologists seem to use a concept of downward causation fruitfully in their work. (shrink)

The context for biological emergence is modular hierarchical structures; their existence is what enables functional complexity to arise. Because of the openness of organisms to their environment, complete initial data (position, momentum) of all particles making up their structure is insufficient to determine future outcomes, because unpredictable new matter, energy, and information impacts each organism from the exterior. Consequently, through Darwinian evolution, life has developed processes to handle this issue functionally on short time scales as well on longer developmental (...) timescales. Symbolism and technology are the transforming factors handling this issue at the social levels, which is where the most sophisticated outcomes of openness occur. Considering the cosmological context, the issue is, should the universe itself be regarded as an open system over time? I make the case that is indeed so, because radically new outcomes occur such as the existence of aircraft, iPads, and the internet, which could not plausibly have been encoded in some form of data on the Last Scattering Surface in the expanding universe. (shrink)

The aim of this paper is to propose a systematic classification of emotions which can also characterize their nature. The first challenge we address is the submission of clear criteria for a theory of emotions that determine which mental phenomena are emotions and which are not. We suggest that emotions as a subclass of mental states are determined by their functional roles. The second and main challenge is the presentation of a classification and theory of emotions that can account for (...) all existing varieties. We argue that we must classify emotions according to four developmental stages: 1. pre-emotions as unfocussed expressive emotion states, 2. basic emotions, 3. primary cognitive emotions, and 4. secondary cognitive emotions. We suggest four types of basic emotions (fear, anger, joy and sadness) which are systematically differentiated into a diversity of more complex emotions during emotional development. The classification distinguishes between basic and non-basic emotions and our multi-factorial account considers cognitive, experiential, physiological and behavioral parameters as relevant for constituting an emotion. However, each emotion type is constituted by a typical pattern according to which some features may be more significant than others. Emotions differ strongly where these patterns of features are concerned, while their essential functional roles are the same. We argue that emotions form a unified ontological category that is coherent and can be well defined by their characteristic functional roles. Our account of emotions is supported by data from developmental psychology, neurobiology, evolutionary biology and sociology. (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)

Denis Diderot’s natural philosophy is deeply and centrally ‘biologistic’: as it emerges between the 1740s and 1780s, thus right before the appearance of the term ‘biology’ as a way of designating a unified science of life (McLaughlin), his project is motivated by the desire both to understand the laws governing organic beings and to emphasize, more ‘philosophically’, the uniqueness of organic beings within the physical world as a whole. This is apparent both in the metaphysics of vital matter he (...) puts forth in works such as D’Alembert’s Dream (1769) and the more empirical concern with the mechanics of life in his manuscript Elements of Physiology, on which he worked during the last twenty years of his life. This ‘biologism’ obviously presents the interpreter of Diderot with some difficulties, notably as regards his materialism, given that contemporary forms of materialism have on the contrary strongly rejected notions of emergence, vitalism, teleology and any concepts appealing to unique, irreducible features of organisms. In response, some have described him as a ‘holist’ (Kaitaro) while others have emphasized his materialist, naturalist project (Bourdin, Wolfe). In what follows I examine a little-known aspect of Diderot’s articulation of his biological project: his statement in favour of epigenesis within the short but suggestive Encyclopédie article “Spinosiste.” Diderot was, of course, a partisan of epigenesis (the developmental-biological theory opposed to preformation, according to which beings develop by successive adjunction of layers of matter), but why include a statement in favour of a particular biological (or developmental) theory within an entry dealing with a philosopher, Spinoza, who does not seem to have been concerned at all with the specific properties of living beings, how they grow from embryonic to developed states, and so on? By trying to answer this question I also try and locate Diderot’s biological project in relation to what will become, in the years after his death, the project for a science called ‘biology’, with figures such as Treviranus and Lamarck. For it is not clear that the two can be easily correlated or causally linked: Diderot’s ‘epigenetic Spinozism’ is a different conceptual entity from what we find in histories of biology. (shrink)

The allure of perennial questions in biology: temporary excitement or substantive advance? Content Type Journal Article Pages 1-4 DOI 10.1007/s11016-011-9533-5 Authors Alan C. Love, Department of Philosophy, Minnesota Center for Philosophy of Science, University of Minnesota, 831 Heller Hall, 271 19th Ave. S, Minneapolis, MN 55455-0310, USA Journal Metascience Online ISSN 1467-9981 Print ISSN 0815-0796.

Theories of cultural evolution rest on the assumption that cultural inheritance is distinct from biological inheritance. Cultural and biological inheritance are two separate so-called channels of inheritance, two sub-systems of the sum total of developmental resources traveling in distinct ways between individual agents. This paper asks: what justifies this assumption? In reply, a philosophical account is offered that points at three related but distinct criteria that (taken together) make the distinction between cultural and biological inheritance not only precise but (...) also justify it as real, i.e. as ontologically adequate. These three criteria are: (a) the autonomy of cultural change, (b) the near-decomposability of culture, and (c) differences in temporal order between cultural and biological inheritance. (shrink)

The aim of this paper is to evaluate the level of gender bias in Aristotle’s Generation of Animals while exercising due care in the analysis of its arguments. I argue that while the GA theory is clearly sexist, the traditional interpretation fails to diagnose the problem correctly. The traditional interpretation focuses on three main sources of evidence: (1) Aristotle’s claim that the female is, as it were, a “disabled” (πεπηρωμένον) male; (2) the claim at GA IV.3, 767b6-8 that females are (...) a departure from the kind; and (3) Aristotle’s supposed claim at GA IV.3, 768a21-8 that the most ideal outcome of reproduction is a male offspring that perfectly resembles its father. I argue that each of these passages has either been misunderstood or misrepresented by commentators. In none of these places is Aristotle suggesting that females are imperfect members of the species or that they result from the failure to achieve some teleological goal. I defend the view that the GA does not see reproduction as occurring for the sake of producing males; rather, what sex an embryo happens to become is determined entirely by non-teleological forces operating through material necessity. This interpretation is consistent with Aristotle’s view in GA II.5 that females have the same soul as the male (741a7) as well as the argument in Metaphysics X.9 that sexual difference is not part of the species form but is an affection (πάθος) arising from the matter (1058b21-4). While the traditional interpretation has tended to exaggerate the level of sexism in Aristotle’s
developmental biology, the GA is by no means free of gender bias as some recent scholarship has claimed. In the final section of the paper I point to one passage where Aristotle clearly falls back on sexist assumptions in order to answer the difficult question, “Why are animals divided into sexes?”. I argue that this passage in particular poses a serious challenge
to anyone attempting to absolve Aristotle’s developmental biology of the charge of sexism. (shrink)

Organisms live not as discrete entities on which an independent environment acts, but as members of a reproductive lineage in an ongoing series of interactions between that lineage and a dynamic ecological niche. These interactions continuously shape both systems in a reciprocal manner, resulting in the emergence of reliably co-occurring configurations within and between both systems. The enactive approach to cognition describes this relationship as the structural coupling between an organism and its environment; similarly, Developmental Systems Theory emphasizes the (...) reciprocal nature of structurally coupled systems in its analysis of organisms as developmental processes embedded within a developmental system. Through an enactive-developmental systems framing, this paper identifies the organizational features of cognizing systems in order to motivate a picture of how organism and environment co-determine and co-construct one another. I argue that organisms can be characterized as self-organizing, operationally closed, plastic systems ecologically embedded within a developmental system. In virtue of this organizational makeup, organisms actively engage in the modulation and assessment of their coupling with their environment: cognitive strategies that entail contextualized responses to variations across the web of interactions that comprises the developmental system. (shrink)

According to neo-Aristotelian ethical naturalism, moral goodness is an instance of natural goodness, a kind of normativity supposedly already present in nature in the biological realm of non-human living things. Proponents of this view appeal to Michael Thompson’s conception of a life-form--the form of a living organism--to give an account of natural goodness. However, although neo-Aristotelians call themselves naturalists, they hardly ever consult the science of biology to defend their commitments regarding biological organisms. This has led many critics to (...) argue that the neo-Aristotelian account of natural normativity is out of touch with the findings of modern evolutionary biology. One line of response to this objection, presented by John Hacker-Wright and Micah Lott, claims that the neo-Aristotelian concept of a living organism has to be presupposed in evolutionary biology as long as organisms are the subjects of evolutionary explanation. In this paper, I examine this response by tracing the concept of organism in modern evolutionary biology. I first argue that the Modern Synthesis theory of evolution, which understands evolution as change in gene frequencies within a population, does not presuppose the relevant concept of organism. I then explore an alternative view of evolution that has emerged in the past twenty years from advances in evolutionary developmental biology. I argue that this so called ‘evo-devo’ approach makes room for an explanatory concept of organism that can be reconciled with the neo-Aristotelian view. Moreover, I argue that although the explanatory role of the concept of organism in evolutionary biology is still contentious, the well-established role of this concept in developmental biology can be used to defend the biological commitments of neo-Aristotelian naturalism. (shrink)

In recent years, the concept of evolvability has been gaining in prominence both within evolutionary developmental biology (evo-devo) and the broader field of evolutionary biology. Despite this, there remains considerable disagreement about what evolvability is. This article offers a solution to this problem. I argue that, in focusing too closely on the role played by evolvability as an explanandum in evo-devo, existing philosophical attempts to clarify the evolvability concept have been overly narrow. Within evolutionary biology more (...) broadly, evolvability offers a robust explanation for the evolutionary trajectories of populations. Evolvability is an abstract, robust, dispositional property of populations, which captures the joint causal influence of their internal features on the outcomes of evolution (as opposed to the causal influence of selection, which is often characterized as external). When considering the nature of the physical basis of this disposition, it becomes clear that the many existing definitions of evolvability at play within evo-devo should be understood as capturing only aspects of a much broader phenomenon. 1 Introduction2 The Problem of Evolvability3 The Theoretical Role of Evolvability in Evolutionary Biology3.1 The explanatory targets of evolutionary biology3.2 Selection-based explanations3.3 Lineage explanations3.4 Evolvability-based explanations3.5 What properties must evolvability have?4 What Evolvability Really Is4.1 Making sense of ft4.2 Making sense of x and b5 What of the Limbs? The Power of E6 Conclusion. (shrink)

Homology is a biological sameness relation that is purported to hold in the face of changes in form, composition, and function. In spite of the centrality and importance of homology, there is no consensus on how we should understand this concept. The two leading views of homology, the genealogical and developmental accounts, have significant shortcomings. We propose a new account, the hierarchical-dependency account of homology, which avoids these shortcomings. Furthermore, our account provides for continuity between special, general, and serial (...) homology. (shrink)

Is psychopathy born or made? Contemporary psychopathy research shows that there is much wrong with this question. It is increasingly accepted that the development of psychopathy is dependent on multiple causal factors interacting with one another. However, there remains the major theoretical challenge of understanding the relations between these multiple causal factors in the developmental process. In this paper, I argue that the conventional picture of gene-environment interactionism does not offer an adequate account of psychopathy development. Instead, I propose (...) that a theoretical framework from the philosophy of biology, namely developmental systems theory, can facilitate a better understanding of psychopathy development that captures the contingent and dynamic relations between multiple causal factors. Some practical implications of a developmental systems theory approach to psychopathy are also explored. (shrink)

Evolutionary adaptation has been suggested as the hallmark of life that best accounts for life’s creativity. However, current evolutionary approaches still fail to give an adequate account of it, even if they are able to explain both the origin of novelties and the proliferation of certain traits in a population. Although modern-synthesis Darwinism is today usually appraised as too narrow a position to cope with all the complexities of developmental and structural biology—not to say biosemiotic phenomena—, Darwinism need (...) not be if we separate metaphor from reality in natural selection in order to show the axiological complexity of this concept. This can shed light on the relationship between biosemiotics and biological evolution. (shrink)

Causal selection is the task of picking out, from a field of known causally relevant factors, some factors as elements of an explanation. The Causal Parity Thesis in the philosophy of biology challenges the usual ways of making such selections among different causes operating in a developing organism. The main target of this thesis is usually gene centrism, the doctrine that genes play some special role in ontogeny, which is often described in terms of information-bearing or programming. This paper (...) is concerned with the attempt of confronting the challenge coming from the Causal Parity Thesis by offering principles of causal selection that are spelled out in terms of an explicit philosophical account of causation, namely an interventionist account. I show that two such accounts that have been developed, although they contain important insights about causation in biology, nonetheless fail to provide an adequate reply to the Causal Parity challenge: Ken Waters's account of actual-difference making and Jim Woodward's account of causal specificity. A combination of the two also doesn't do the trick, nor does Laura Franklin-Hall's account of explanation (in this volume). We need additional conceptual resources. I argue that the resources we need consist in a special class of counterfactual conditionals, namely counterfactuals the antecedents of which describe biologically normal interventions. (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 functionality, 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 adaptations, 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 opportunistic 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 suggest. 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 implement 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 ‘‘engineering’’ 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 engineering 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)

The nature and role of the patient in biomedicine comprise issues central to bioethical inquiry. Given its developmental history grounded firmly in a backlash against 20th-century cases of egregious human subjects abuse, contemporary medical bioethics has come to rely on a fundamental assumption: the unit of care is the autonomous self-directing patient. In this article we examine first the structure of the feminist social critique of autonomy. Then we show that a parallel argument can be made against relational autonomy (...) as well, demonstrating how this second concept of autonomy fails to take sufficiently into account an array of biological determinants, particularly those from microbial biology. Finally, in light of this biological critique, we question whether or to what extent any relevant and meaningful view of autonomy can be recovered in the contemporary landscape of bioethics. (shrink)

The article presents a perspective on the scientific explanation of the subjectivity of conscious experience. It proposes plausible answers for two empirically valid questions: the ‘how’ question concerning the developmental mechanisms of subjectivity, and the ‘why’ question concerning its function. Biological individuation, which is acquired in several different stages, serves as a provisional description of how subjective perspectives may have evolved. To the extent that an individuated informational space seems the most efficient way for a given organism to select (...) biologically valuable information, subjectivity is deemed to constitute an adaptive response to informational overflow. One of the possible consequences of this view is that subjectivity might be (at least functionally) dissociated from consciousness, insofar as the former primarily facilitates selection, the latter action. (shrink)

Several authors have used the notion of causal specificity in order to defend non-parity about genetic causes (Waters 2007, Woodward 2010, Weber 2017, forthcoming). Non-parity in this context is the idea that DNA and some other biomolecules that are often described as information-bearers by biologists play a unique role in life processes, an idea that has been challenged by Developmental Systems Theory (e.g., Oyama 2000). Indeed, it has proven to be quite difficult to state clearly what the alleged special (...) role of genetic causes consists in. In this paper, I show that the set of biomolecules that are normally considered to be information-bearers (DNA, mRNA) can be shown to be the most specific causes of protein primary structure, provided that causal specificity is measured over a relevant space of biological possibilities, disregarding physical as well as logically possible states of the causal variables. (shrink)

The advent of contemporary evolutionary theory ushered in the eventual decline of Aristotelian Essentialism (Æ) – for it is widely assumed that essence does not, and cannot have any proper place in the age of evolution. This paper argues that this assumption is a mistake: if Æ can be suitably evolved, it need not face extinction. In it, I claim that if that theory’s fundamental ontology consists of dispositional properties, and if its characteristic metaphysical machinery is interpreted within the framework (...) of contemporary evolutionary developmental biology, an evolved essentialism is available. The reformulated theory of Æ offered in this paper not only fails to fall prey to the typical collection of criticisms, but is also independently both theoretically and empirically plausible. The paper contends that, properly understood, essence belongs in the age of evolution. (shrink)

The paper addresses the formation of striking patterns within originally near-homogenous tissue, the process prototypical for embryology, and represented in particularly purist form by cut sections of hydra regenerating, by internal reorganisation of the pre-existing tissue, a complete animal with head and foot. The essential requirements are autocatalytic, self-enhancing activation, combined with inhibitory or depletion effects of wider range – “lateral inhibition”. Not only de-novo-pattern formation, but also well known, striking features of developmental regulation such as induction, inhibition, and (...) proportion regulation can be explained on this basis. The theory provides a mathematical recipe for the construction of molecular models with criteria for the necessary non-linear interactions. It has since been widely applied to different developmental processes. (shrink)

This volume provides a broad overview of issues in the philosophy of behavioral biology, covering four main themes: genetic, developmental, evolutionary, and neurobiological explanations of behavior. It is both interdisciplinary and empirically informed in its approach, addressing philosophical issues that arise from recent scientific findings in biological research on human and non-human animal behavior. Accordingly, it includes papers by professional philosophers and philosophers of science, as well as practicing scientists. Much of the work in this volume builds on (...) presentations given at the international conference, “Biological Explanations of Behavior: Philosophical Perspectives”, held in 2008 at the Leibniz Universität Hannover in Germany. The volume is intended to be of interest to a broad range of audiences, which includes philosophers (e.g., philosophers of mind, philosophers of biology, and metaethicists), as well as practicing scientists, such as biologists or psychologists whose interests relate to biological explanations of behavior. (shrink)

Evolutionary developmental biology represents a paradigm shift in the understanding of the ontogenesis and evolutionary progression of the denizens of the natural world. Given the empirical successes of the evo-devo framework, and its now widespread acceptance, a timely and important task for the philosophy of biology is to critically discern the ontological commitments of that framework and assess whether and to what extent our current metaphysical models are able to accommodate them. In this paper, I argue that (...) one particular model is a natural fit: an ontology of dispositional properties coherently and adequately captures the crucial casual-cum-explanatory role that the fundamental elements of evo-devo play within that framework. (shrink)

On a book concerned with taking developmental biology seriously within the context of evolutionary theory.

Multicellular organisms contain numerous symbiotic microorganisms, collectively called microbiomes. Recently, microbiomic research has shown that these microorganisms are responsible for the proper functioning of many of the systems (digestive, immune, nervous, etc.) of multicellular organisms. This has inclined some scholars to argue that it is about time to reconceptualise the organism and to develop a concept that would place the greatest emphasis on the vital role of microorganisms in the life of plants and animals. We believe that, unfortunately, there is (...) a problem with this suggestion, since there is no such thing as a universal concept of the organism which could constitute a basis for all biological sciences. Rather, the opposite is true: numerous alternative definitions exist. Therefore, comprehending how microbiomics is changing our understanding of organisms may be a very complex matter. In this paper we will demonstrate that this pluralism proves that claims about a change in our understanding of organisms can be treated as both true and untrue. Mainly, we assert that the existing concepts differ substantially, and that only some of them have to be reconsidered in order to incorporate the discoveries of microbiomics, while others are already flexible enough to do so. Taking into account the plurality of conceptualisations within different branches of modern biology, we will conduct our discussion using the developmental and the cooperation–conflict concepts of the organism. Then we will explain our results by referring to the recent philosophical debate on the nature of the concept of the organism within biology. (shrink)

Griffiths et al. (2015) have proposed a quantitative measure of causal specificity and used it to assess various attempts to single out genetic causes as being causally more specific than other cellular mechanisms, for example, alternative splicing. Focusing in particular on developmental processes, they have identified a number of important challenges for this project. In this discussion note, I would like to show how these challenges can be met.

It is widely assumed that there is value in the biological tie between parent and child. An implication of this is that adoption is often considered a less desirable alternative to procreation. This paper offers a philosophical defence of adoptive parenthood as a valuable and authentic form of parenthood. While previous defences have suggested that society’s valorisation of the biological tie is unjustified, I argue herein that the conception of the biological tie that features in the normative discourse on parenthood (...) is too narrowly genocentric. Against this genocentric conception, recent work in the philosophy of biology has emphasised the roles of joint determination, dynamic construction, and extended inheritance in development, which I suggest can substantiate a more inclusive conception of the biological tie. Accordingly, I propose that adoptive parents form a rich variety of biological ties with their children, some of which are as heritable and formative as genetic relatedness. (shrink)

Empirical evidence from developmental psychology and anthropology points out that the human mind is predisposed to conceptualize the world in particular, species-specific ways. These cognitive predispositions lead to universal human commonsense views, often referred to as folk theories. Nevertheless, humans can transgress these views – i.e. they can contradict them with alternative descriptions, they perceive as more accurate – as exemplified in modern sciences. In this paper, I enquire about the cognitive faculties underlying such transgressions. I claim that there (...) are three faculties enabling us to part with these universal commonsense views of the world imposed by our nature. The first is our ability to represent representations – i.e. to form metarepresentations. The second is our ability to produce alternative representations both by explaining a familiar subject matter in terms of the principles governing different conceptual domains than the one that we are predisposed to apply to the subject matter and by directing our mind to new subject matters (for which we have no predisposed conceptual grasp), understanding them in terms of familiar domains. The third, finally, is our ability to give these representations an epistemic orientation. Key words: Human cognition, Domain-specific knowledge systems, Cognitive predispositions, Folk theories, Cognitive flexibility, Human – nonhuman distinction. (shrink)

Ontologies of living things are increasingly grounded on the concepts and practices of current life science. Biological development is a process, undergone by living things, which begins with a single cell and (in an important class of cases) ends with formation of a multicellular organism. The process of development is thus prima facie central for ideas about biological individuality and organismality. However, recent accounts of these concepts do not engage developmental biology. This paper aims to fill the gap, (...) proposing the lineage view of stem cells as an ontological framework for conceptualizing organismal development. This account is grounded on experimental practices of stem cell research, with emphasis on new techniques for generating biological organization in vitro. On the lineage view, a stem cell is the starting point of a cell lineage with a specific organismal source, time-interval of existence, and ‘tree topology’ of branch-points linking the stem to developmental termini. The concept of ‘enkapsis’ accommodates the cell-organism relation within the lineage view; this hierarchical notion is further explicated by considering the methods and results of stem cell experiments. Results of this examination include a (partial) characterization of stem cells’ developmental versatility, and the context-dependence of developmental processes involving stem cells. (shrink)

Molecular developmental biology has expanded our conceptions of gene actions, underpinning that embryonic development is not only governed by a set of specific genes, but as much by space–time conditions of its developing modules. Typically, formation of cellular spheres, their transformation into planar epithelia, followed by tube formations and laminations are modular steps leading to the development of nervous tissues. Thereby, actions of organising centres, morphogenetic movements, inductive events between epithelia, tissue polarity reversal, widening of epithelia, and all (...) these occurring orderly in space and time, are driving forces of emergent laminar neural tissues, e.g. the vertebrate retina. Analyses of self-organisational formation of retina-like 3D structures from dispersed cells under defined cell culture conditions demonstrate that not only particular genetic networks, but—at least as important—the applied culture conditions define phenotypes of emergent tissues. Such in vitro approaches allow assigning emerging tissue formation to ground-laying genetic networks separately from contributions by conditional constraints. (shrink)