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 Biology 3.1 The explanatory targets of evolutionary biology 3.2 Selection-based explanations 3.3 Lineage explanations 3.4 Evolvability-based explanations 3.5 What properties must evolvability have?4 What Evolvability Really Is 4.1 Making sense of ft 4.2 Making sense of x and b5 What of the Limbs? The Power of E6 Conclusion. (shrink)

In the last decade, robustness has been extensively mentioned and discussed in biology as well as in the philosophy of the life sciences. Nevertheless, from both fields, someone has affirmed that this debate has resulted in more semantic confusion than in semantic clearness. Starting from this claim, we wish to offer a sort of prima facie map of the different usages of the term. In this manner we would intend to predispose a sort of “semantic platform” which could be exploited (...) by those who wish to discuss or simply use it. We do this by starting from a core distinction between the robustness of representations, which is a philosophy of science issue, and the representations of robustness, which instead pertains to science. We illustrate our proposal with examples from biology, physics and mathematics. (shrink)

Complexity and integration are longstanding widely debated issues in philosophy of science and recent contributions have largely focused on biology and biomedicine. This paper specifically considers some methodological novelties in cancer research, motivated by various features of tumours as complex diseases, and shows how they encourage some rethinking of philosophical discourses on those topics. In particular, we discuss the integrative-cluster approach, and analyse its potential in the epistemology of cancer. We suggest that, far from being the solution to tame cancer (...) complexity, this approach offers a philosophically interesting new manner of considering integration, and show how it can help addressing the apparent contrast between a pluralistic and a unitary account. (shrink)

Complexity and integration are longstanding widely debated issues in philosophy of science and recent contributions have largely focused on biology and biomedicine. This paper specifically considers some methodological novelties in cancer research, motivated by various features of tumours as complex diseases, and shows how they encourage some rethinking of philosophical discourses on those topics. In particular, we discuss the integrative-cluster approach, and analyse its potential in the epistemology of cancer. We suggest that, far from being the solution to tame cancer (...) complexity, this approach offers a philosophically interesting new manner of considering integration, and show how it can help addressing the apparent contrast between a pluralistic and a unitary account. (shrink)

Complexity and integration are longstanding widely debated issues in philosophy of science and recent contributions have largely focused on biology and biomedicine. This paper specifically considers some methodological novelties in cancer research, motivated by various features of tumours as complex diseases, and shows how they encourage some rethinking of philosophical discourses on those topics. In particular, we discuss the integrative-cluster approach, and analyse its potential in the epistemology of cancer. We suggest that, far from being the solution to tame cancer (...) complexity, this approach offers a philosophically interesting new manner of considering integration, and show how it can help addressing the apparent contrast between a pluralistic and a unitary account. (shrink)

Artificial life (also known as “ALife”) is a broad, interdisciplinary endeavor that studies life and life-like processes through simulation and synthesis. The goals of this activity include modelling and even creating life and life-like systems, as well as developing practical applications using intuitions and methods taken from living systems. Artificial life both illuminates traditional philosophical questions and raises new philosophical questions. Since both artificial life and philosophy investigate the essential nature of certain fundamental aspects of reality like life and adaptation, (...) artificial life offers philosophy a new perspective on these phenomena. This chapter provides an introduction to current research in artificial life and explains its philosophical implications. (shrink)

Research on diseases such as cancer reveals that primary mechanisms, which have been the focus of study by the new mechanists in philosophy of science, are often subject to control by other mechanisms. Cancer cells employ the same primary mechanisms as healthy cells but control them differently. I use cancer research to highlight just how widespread control is in individual cells. To provide a framework for understanding control, I reconceptualize mechanisms as imposing constraints on flows of free energy, with control (...) mechanisms operating on flexible constraints in primary mechanisms. I argue that control mechanisms themselves often form complex, integrated networks. (shrink)

Systems biology provides alternatives to the strategies to developing mechanistic explanations traditionally pursued in cell and molecular biology and much discussed in accounts of mechanistic explanation. Rather than starting by identifying a mechanism for a given phenomenon and decomposing it, systems biologists often start by developing cell-wide networks of detected connections between proteins or genes and construe clusters of highly interactive components as potential mechanisms. Using inference strategies such as ‘guilt-by-association’, researchers advance hypotheses about functions performed of these mechanisms. I (...) examine several examples of research on budding yeast, first on what are taken to be enduring networks and subsequently on networks that change as cells perform different activities or respond to different external conditions. (shrink)

Immunology is central to contemporary biology and medicine, but it also provides novel philosophical insights. Its most significant contribution to philosophy concerns the understanding of biological individuality: what a biological individual is, what makes it unique, how its boundaries are established and what ensures its identity through time. Immunology also offers answers to some of the most interesting philosophical questions. What is the definition of life? How are bodily systems delineated? How do the mind and the body interact? In this (...) Element, Thomas Pradeu considers the ways in which immunology can shed light on these and other important philosophical issues. This title is also available as Open Access on Cambridge Core. (shrink)

Intent and mitigating circumstances play a central role in moral and legal assessments in large-scale industrialized societies. Al- though these features of moral assessment are widely assumed to be universal, to date, they have only been studied in a narrow range of societies. We show that there is substantial cross-cultural variation among eight traditional small-scale societies (ranging from hunter-gatherer to pastoralist to horticulturalist) and two Western societies (one urban, one rural) in the extent to which intent and mitigating circumstances influence (...) moral judgments. Although participants in all societies took such factors into account to some degree, they did so to very different extents, varying in both the types of considerations taken into account and the types of violations to which such considerations were applied. The particular patterns of assessment characteristic of large-scale industrialized societies may thus reflect relatively recently culturally evolved norms rather than inherent features of human moral judgment. (shrink)

Complex systems research is becoming ever more important in both the natural and social sciences. It is commonly implied that there is such a thing as a complex system, different examples of which are studied across many disciplines. However, there is no concise definition of a complex system, let alone a definition on which all scientists agree. We review various attempts to characterize a complex system, and consider a core set of features that are widely associated with complex systems in (...) the literature and by those in the field. We argue that some of these features are neither necessary nor sufficient for complexity, and that some of them are too vague or confused to be of any analytical use. In order to bring mathematical rigour to the issue we then review some standard measures of complexity from the scientific literature, and offer a taxonomy for them, before arguing that the one that best captures the qualitative notion of the order produced by complex systems is that of the Statistical Complexity. Finally, we offer our own list of necessary conditions as a characterization of complexity. These conditions are qualitative and may not be jointly sufficient for complexity. We close with some suggestions for future work. (shrink)

This book offers a philosophy for error-prone humans trying to understand messy systems in the real world.

This book, based on Jaegwon Kim's 1996 Townsend Lectures, presents the philosopher's current views on a variety of issues in the metaphysics of the mind...

Though most contemporary philosophers and scientists accept a physicalist view of mind, the recent surge of interest in the problem of consciousness has put the mind /body problem back into play. The physicalists' lack of success in dispelling the air of residual mystery that surrounds the question of how consciousness might be physically explained has led to a proliferation of options. Some offer alternative formulations of physicalism, but others forgo physicalism in favour of views that are more dualistic or that (...) bring in mentalistic features at the ground- floor level of reality as in pan-proto-psychism. My aim here is to give an overview of the recent philosophic discussion to serve as a map in locating issues and options. I will not offer a comprehensive survey of the debate or mark every important variant to be found in the recent literature. I will mark the principal features of the philosophic landscape that one might use as general orientation points in navigating the terrain. I will focus in particular on three central and interrelated ideas: those of emergence, reduction, and nonreductive physicalism. The third of these, which has emerged as more or less the majority view among current philosophers of mind, combines a pluralist view about the diversity of what needs to be explained by science with an underlying metaphysical commitment to the physical as the ultimate basis of all that is real. The view has been challenged from both left and right, on one side from dualists and on the other from hard core reductive materialists. Despite their differences, those critics agree in finding nonreductive physicalism an unacceptable and perhaps even incoherent position. They agree as well in treating reducibility as the essential criterion for physicality; they differ only about whether the criterion can be met. Reductive physicalists argue that it can, and dualists deny it. (shrink)

This paper explores the fundamental ideas that have motivated the idea of emergence and the movement of emergentism. The concept of reduction, which lies at the heart of the emergence idea is explicated, and it is shown how the thesis that emergent properties are irreducible gives a unified account of emergence. The paper goes on to discuss two fundamental unresolved issues for emergentism. The first is that of giving a “positive” characterization of emergence; the second is to give a coherent (...) explanation of how “downward” causation, a central component of emergentism, is able to avoid the problem of overdetermination. (shrink)

The increasing application of network models to interpret biological systems raises a number of important methodological and epistemological questions. What novel insights can network analysis provide in biology? Are network approaches an extension of or in conflict with mechanistic research strategies? When and how can network and mechanistic approaches interact in productive ways? In this paper we address these questions by focusing on how biological networks are represented and analyzed in a diverse class of case studies. Our examples span from (...) the investigation of organizational properties of biological networks using tools from graph theory to the application of dynamical systems theory to understand the behavior of complex biological systems. We show how network approaches support and extend traditional mechanistic strategies but also offer novel strategies for dealing with biological complexity. (shrink)

The functional complexity, or the number of functions, of organisms hasfigured prominently in certain theoretical and empirical work inevolutionary biology. Large-scale trends in functional complexity andcorrelations between functional complexity and other variables, such assize, have been proposed. However, the notion of number of functions hasalso been operationally intractable, in that no method has been developedfor counting functions in an organism in a systematic and reliable way.Thus, studies have had to rely on the largely unsupported assumption thatnumber of functions can be (...) measured indirectly, by using number ofmorphological, physiological, and behavioral parts as a proxy. Here, amodel is developed that supports this assumption. Specifically, the modelpredicts that few parts will have many functions overlapping in them, andtherefore the variance in number of functions per part will be low. If so,then number of parts is expected to be well correlated with number offunctions, and we can use part counts as proxies for function counts incomparative studies of organisms, even when part counts are low. Alsodiscussed briefly is a strategy for identifying certain kinds of parts inorganisms in a systematic way. (shrink)

The consensus among evolutionists seems to be that the morphological complexity of organisms increases in evolution, although almost no empirical evidence for such a trend exists. Most studies of complexity have been theoretical, and the few empirical studies have not, with the exception of certain recent ones, been especially rigorous; reviews are presented of both the theoretical and empirical literature. The paucity of evidence raises the question of what sustains the consensus, and a number of suggestions are offered, including the (...) possibility that certain cultural and/or perceptual biases are at work. In addition, a shift in emphasis from theoretical to empirical inquiry is recommended for the study of complexity, and guidelines for future empirical studies are proposed. (shrink)

The world is complex, but acknowledging its complexity requires an appreciation for the many roles context plays in shaping natural phenomena. In _Unsimple Truths, _Sandra Mitchell argues that the long-standing scientific and philosophical deference to reductive explanations founded on simple universal laws, linear causal models, and predict-and-act strategies fails to accommodate the kinds of knowledge that many contemporary sciences are providing about the world. She advocates, instead, for a new understanding that represents the rich, variegated, interdependent fabric of many levels (...) and kinds of explanation that are integrated with one another to ground effective prediction and action. Mitchell draws from diverse fields including psychiatry, social insect biology, and studies of climate change to defend “integrative pluralism”—a theory of scientific practices that makes sense of how many natural and social sciences represent the multi-level, multi-component, dynamic structures they study. She explains how we must, in light of the now-acknowledged complexity and contingency of biological and social systems, revise how we conceptualize the world, how we investigate the world, and how we act in the world. Ultimately _Unsimple Truths _argues that the very idea of what should count as legitimate science itself should change. (shrink)

Concerns with the use of engineering approaches in biology have recently been raised. I examine two related challenges to biological research that I call the synchronic and diachronic underdetermination problem. The former refers to challenges associated with the inference of design principles underlying system capacities when the synchronic relations between lower-level processes and higher-level systems capacities are degenerate. The diachronic underdetermination problem regards the problem of reverse engineering a system where the non-linear relations between system capacities and lower-level mechanisms are (...) changing over time. Braun and Marom argue that recent insights to biological complexity leave the aim of reverse engineering hopeless - in principle as well as in practice. While I support their call for systemic approaches to capture the dynamic nature of living systems, I take issue with the conflation of reverse engineering with naïve reductionism. I clarify how the notion of design principles can be more broadly conceived and argue that reverse engineering is compatible with a dynamic view of organisms. It may even help to facilitate an integrated account that bridges the gap between mechanistic and systems approaches. (shrink)

This article defends the notion of downward causation, relating it to a notion of conditional independence.

This paper argues that besides mechanistic explanations, there is a kind of explanation that relies upon “topological” properties of systems in order to derive the explanandum as a consequence, and which does not consider mechanisms or causal processes. I first investigate topological explanations in the case of ecological research on the stability of ecosystems. Then I contrast them with mechanistic explanations, thereby distinguishing the kind of realization they involve from the realization relations entailed by mechanistic explanations, and explain how both (...) kinds of explanations may be articulated in practice. The second section, expanding on the case of ecological stability, considers the phenomenon of robustness at all levels of the biological hierarchy in order to show that topological explanations are indeed pervasive there. Reasons are suggested for this, in which “neutral network” explanations are singled out as a form of topological explanation that spans across many levels. Finally, I appeal to the distinction of explanatory regimes to cast light on a controversy in philosophy of biology, the issue of contingence in evolution, which is shown to essentially involve issues about realization. (shrink)

Philosophical accounts of emergence have been explicated in terms of logical relationships between statements (derivation) or static properties (function and realization). Jaegwon Kim is a modern proponent. A property is emergent if it is not explainable by (or reducible to) the properties of lower level components. This approach, I will argue, is unable to make sense of the kinds of emergence that are widespread in scientific explanations of complex systems. The standard philosophical notion of emergence posits the wrong dichotomies, confuses (...) compositional physicalism with explanatory physicalism, and is unable to represent the type of dynamic processes (self-organizing feedback) that both generate emergent properties and express downward causation. (shrink)

In recent cancer research, strong and apparently conflicting epistemological stances have been advocated by different research teams in a mist of an ever-growing body of knowledge ignited by ever-more perplexing and non-conclusive experimental facts: in the past few years, an 'organicist' approach investigating cancer development at the tissue level has challenged the established and so-called 'reductionist' approach focusing on disentangling the genetic and molecular circuitry of carcinogenesis. This article reviews the ways in which 'organicism' and 'reductionism' are used and opposed (...) in this context, with an aim at clarifying the debate. Methodological, epistemological and ontological implications of both approaches are discussed. We argue that the 'organicist/reductionist' opposition in the present case of carcinogenesis is more a matter of diverging heuristics than a claim about theoretical or ontological (ir)reducibility. As a matter of fact, except for the downward causation claim, which we question, we argue that the organicist arguments are compatible with the reductionist approach. Moreover, we speculate that both approaches, which currently focus on specific entities i.e., genes versus tissues, will need to shift their conceptual frameworks to studying complex arrays of relationships potentially ranging over several levels of entities, as is the case with 'systems biology'. (shrink)

The clonal evolution model and the cancer stem cell model are two independent models of cancers, yet recent data shows intersections between the two models. This article explores the impacts of the CSC model on the CE model. I show that CSC restriction, which depends on CSC frequency in cancer cell populations and on the probability of dedifferentiation of cancer non-stem cells into CSCs, can favor or impede some patterns of evolution and some processes of evolution. Taking CSC restriction into (...) account for the CE model thus has implications for the way in which we understand the patterns and processes of evolution, and can also provide new leads for therapeutic interventions. (shrink)

The local microenvironment, or niche, of a cancer cell plays important roles in cancer development. A major component of the niche is the extracellular matrix, a complex network of macromolecules with distinctive physical, biochemical, and biomechanical properties. Although tightly controlled during embryonic development and organ homeostasis, the ECM is commonly deregulated and becomes disorganized in diseases such as cancer. Abnormal ECM affects cancer progression by directly promoting cellular transformation and metastasis. Importantly, however, ECM anomalies also deregulate behavior of stromal cells, (...) facilitate tumor-associated angiogenesis and inflammation, and thus lead to generation of a tumorigenic microenvironment. Understanding how ECM composition and topography are maintained and how their deregulation influences cancer progression may help develop new therapeutic interventions by targeting the tumor niche. © 2012 Lu et al. (shrink)

This fine collection of essays by a leading philosopher of science presents a defence of integrative pluralism as the best description for the complexity of scientific inquiry today. The tendency of some scientists to unify science by reducing all theories to a few fundamental laws of the most basic particles that populate our universe is ill-suited to the biological sciences, which study multi-component, multi-level, evolved complex systems. This integrative pluralism is the most efficient way to understand the different and complex (...) processes - historical and interactive - that generate biological phenomena. This book will be of interest to students and professionals in the philosophy of science. (shrink)

This paper argues that scale-dependence of physical and biological processes offers resistance to reductionism and has implications that support a specific kind of downward causation. I demonstrate how insights from multiscale modeling can provide a concrete mathematical interpretation of downward causation as boundary conditions for models used to represent processes at lower scales. The autonomy and role of macroscale parameters and higher-level constraints are illustrated through examples of multiscale modeling in physics, developmental biology, and systems biology. Drawing on these examples, (...) I defend the explanatory importance of constraining relations for understanding the behavior of biological systems. (shrink)

Complexity arises from interaction dynamics, but its forms are co-determined by the operative constraints within which the dynamics are expressed. The basic interaction dynamics underlying complex systems is mostly well understood. The formation and operation of constraints is often not, and oftener under appreciated. The attempt to reduce constraints to basic interaction fails in key cases. The overall aim of this paper is to highlight the key role played by constraints in shaping the field of complex systems. Following an introduction (...) to constraints, the paper develops the roles of constraints in specifying forms of complexity and illustrates the roles of constraints in formulating the fundamental challenges to understanding posed by complex systems. (shrink)

The mechanistic perspective has dominated biological disciplines such as biochemistry, physiology, cell and molecular biology, and neuroscience, especially during the 20th century. The primary strategy is reductionist: organisms are to be decomposed into component parts and operations at multiple levels. Researchers adopting this perspective have generated an enormous body of information about the mechanisms of life at scales ranging from the whole organism down to genetic and other molecular operations.

1 The Zero-Force Evolutionary Law 2 Randomness, Hierarchy, and Constraint 3 Diversity 4 Complexity 5 Evidence, Predictions, and Tests 6 Philosophical Foundations 7 Implications.

Due to the variation, contingency and complexity of living systems, biology is often taken to be a science without fundamental theories, laws or general principles. I revisit this question in light of the quest for design principles in systems biology and show that different views can be reconciled if we distinguish between different types of generality. The philosophical literature has primarily focused on generality of specific models or explanations, or on the heuristic role of abstraction. This paper takes a different (...) approach in emphasizing a theory-constituting role of general principles. Design principles signify general dependency-relations between structures and functions, given a set of formally defined constraints. I contend that design principles increase our understanding of living systems by relating specific models to general types. The categorization of types is based on a delineation of the scope of biological possibilities, which serves to identify and define the generic features of classes of systems. To characterize the basis for general principles through generic abstraction and reasoning about possibility spaces, I coin the term constraint-based generality. I show that constraint-based generality is distinct from other types of generality in biology, and argue that general principles play a unifying role that does not entail theory reduction. (shrink)

Cancer research has been at the forefront of biomedical activity in recent decades, and advances in molecular biology have provided a growing amount of information on the mechanisms involved in the etiopathogenesis of tumors. Nevertheless, despite these advances, the complexity of cancer is more evident, especially as different levels of phenomena are considered to explain the heterogeneity of the neoplastic process. A synthetic analysis of advances in cancer research illustrates these changes. In attempting to overcome the limits of epistemological reductionism, (...) there is a move from a view of a genetic basis for cancer to a more comprehensive perspective aiming to integrate a large amount of information and to understand the neoplastic phenomenon at higher levels of biological complexity through a new interpretative framework. (shrink)