Woodward's long awaited book is an attempt to construct a comprehensive account of causation explanation that applies to a wide variety of causal and explanatory claims in different areas of science and everyday life. The book engages some of the relevant literature from other disciplines, as Woodward weaves together examples, counterexamples, criticisms, defenses, objections, and replies into a convincing defense of the core of his theory, which is that we can analyze causation by appeal to the notion of manipulation.

Traditionally a scientific theory is viewed as based on universal laws of nature that serve as axioms for logical deduction. In analyzing the logical structure of evolutionary biology, Elisabeth Lloyd argues that the semantic account is more appropriate and powerful. This book will be of interest to biologists and philosophers alike.

The architects of punctuated equilibrium and species selection as well as more recent workers (Vrba) have narrowed the original formulation of species selection and made it dependent upon so-called emergent characters. One criticism of this narrow version is the dearth of emergent characters with a consequent diminution in the robustness of species selection as an important evolutionary process. We argue that monomorphic species characters may at times be the focus of selection and that under these circumstances selection at the organism (...) level is by-passed due to the absence of critical variance. Selection therefore shifts to the species level where variability reemerges in a clade. The absence of critical variance among organisms prevents effect macroevolution from operating. If species-wide properties are important in macroevolutionary processes, as we contend, systematists should pay more attention to their elucidation. (shrink)

Once upon a time in evolutionary theory, everything happened for the best. Predators killed only the old or the sick. Pecking orders and other dominance hierarchies minimized wasteful conflict within the group. Male displays ensured that only the best and the fittest had mates. In the culmination of this tradition, Wynne-Edwards argued that many species have mechanisms that ensure groups do not over-exploit their resource base. The “central function” of territoriality in birds and other higher animals is “of limiting the (...) numbers of occupants per unit area of habitat”. Species with dominance hierarchies, species with lekking breeding systems, and species with communal breeding regulate their populations. These social mechanisms have population regulation as their “underlying primary function”. Wynne-Edwards argued that these mechanisms evolve through group selection. Populations without such mechanisms are apt to go extinct by eroding their own resource base. (shrink)

Elliott Sober and Ken Waters both raise interesting and difficult challenges for various aspects of the position I set out in Evolution and the Levels of the Selection. I am grateful to them for their penetrating criticisms of my work, and find myself in agreement with many of their points.

In models of multi-level selection, the property of Darwinian fitness is attributed to entities at more than one level of the biological hierarchy, e.g. individuals and groups. However, the relation between individual and group fitness is a controversial matter. Theorists disagree about whether group fitness should always, or ever, be defined as total (or average) individual fitness. This paper tries to shed light on the issue by drawing on work in social choice theory, and pursuing an analogy between fitness and (...) utility. Social choice theorists have long been interested in the relation between individual and social utility, and have identified conditions under which social utility equals total (or average) individual utility. These ideas are used to shed light on the biological problem. (shrink)

The fitness of any evolutionary unit can be understood in terms of its two basic components: fecundity (reproduction) and viability (survival). Trade-offs between these fitness components drive the evolution of life-history traits in extant multicellular organisms. We argue that these trade-offs gain special significance during the transition from unicellular to multicellular life. In particular, the evolution of germ–soma specialization and the emergence of individuality at the cell group (or organism) level are also consequences of trade-offs between the two basic fitness (...) components, or so we argue using a multilevel selection approach. During the origin of multicellularity, we study how the group trade-offs between viability and fecundity are initially determined by the cell level trade-offs, but as the transition proceeds, the fitness trade-offs at the group level depart from those at the cell level. We predict that these trade-offs begin with concave curvature in single-celled organisms but become increasingly convex as group size increases in multicellular organisms. We argue that the increasingly convex curvature of the trade-off function is driven by the cost of reproduction which increases as group size increases. We consider aspects of the biology of the volvocine green algae – which contain both unicellular and multicellular members – to illustrate the principles and conclusions discussed. (shrink)

Most models of generational succession in sexually reproducing populations necessarily move back and forth between genic and genotypic spaces. We show that transitions between and within these spaces are usually hidden by unstated assumptions about processes in these spaces. We also examine a widely endorsed claim regarding the mathematical equivalence of kin-, group-, individual-, and allelic-selection models made by Lee Dugatkin and Kern Reeve. We show that the claimed mathematical equivalence of the models does not hold.

The concept of fitness, central to population genetics and to the synthetic theory of evolution, is discussed. After a historical introduction on the origin of this concept, the current meaning of it in population genetics is examined: a cause of the selective process and its quantification. Several difficulties arise for its exact definition. Three adequacy criteria for such a definition are formulated. It is shown that it is impossible to formulate an adequate definition of fitness respecting these criteria. The propensity (...) definition of fitness is presented and rejected. Finally it is argued that fitness is a conceptual device, a useful tool, only for descriptive purposes of selective processes, changing from case to case, and thus devoid of any substantial physical counterpart. Any attempt to its reification is an apport to the metaphysical load evolutionary theory has inherited from Natural Theology. (shrink)

Recent years have seen a renewed debate over the importance of groupselection, especially as it relates to the evolution of altruism. Onefeature of this debate has been disagreement over which kinds ofprocesses should be described in terms of selection at multiple levels,within and between groups. Adapting some earlier discussions, we presenta mathematical framework that can be used to explore the exactrelationships between evolutionary models that do, and those that donot, explicitly recognize biological groups as fitness-bearing entities.We show a fundamental set (...) of mathematical equivalences between these twokinds of models, one of which applies a form of multi-level selectiontheory and the other being a form of ``individualism.'' However, we alsoargue that each type of model can have heuristic advantages over theother. Indeed, it can be positively useful to engage in a kind ofback-and-forth switching between two different perspectives on theevolutionary role of groups. So the position we defend is a``gestalt-switching pluralism.''. (shrink)

A central subject is the main strand around which the fabric of an historical narrative is woven. Such a subject must possess both spatial and temporal continuity. It is integrated into an historical entity through the relationship between those properties which make it an individual, and their interaction with the historical event. Scientific theory is useful in the reconstruction of past events and the definition of the central subject. Ideas used as central subjects present the problem of finding internal principles (...) of integration which will make the idea continuous over time. The purpose of narratives is to explain an event by integrating it into an organized whole. (shrink)

Judea Pearl has been at the forefront of research in the burgeoning field of causal modeling, and Causality is the culmination of his work over the last dozen or so years. For philosophers of science with a serious interest in causal modeling, Causality is simply mandatory reading. Chapter 2, in particular, addresses many of the issues familiar from works such as Causation, Prediction and Search by Peter Spirtes, Clark Glymour, and Richard Scheines. But philosophers with a more general interest in (...) causation will also profit from reading Pearl’s book, especially the material in chapters 7, 9, and 10, which is self-contained and less technical than other parts of the book. The present review is aimed primarily at readers of the second type. (shrink)

Hierarchical expansions of the theory of natural selection exist in two distinct bodies of thought in evolutionary biology, the group selection and the species selection traditions. Both traditions share the point of view that the principles of natural selection apply at levels of biological organization above the level of the individual organism. This leads them both to considermultilevel selection situations, where selection is occurring simultaneously at more than one level. Impeding unification of the theoretical approaches of the multilevel selection traditions (...) are the different goals of investigators in the different subdisciplines and the different types of data potentially available for analysis. We identify two alternative approaches to multilevel situations, which we termmultilevel selection [1] andmultilevel selection [2]. Of interest in the former case are the effects of group membership onindividual fitnesses, and in the latter the tendencies for the groups themselves to go extinct or to found new groups (i.e., group fitnesses). We argue that: neither represents the entire multilevel selection process; both are aspects of any multilevel selection situation; and both are legitimate approaches, suitable for answering different questions. Using this formalism, we show that: multilevel selection [2] does not require emergent group properties in order to provide an explanatory mechanism of evolutionary change; multilevel selection [1] is usually more appropriate for neontological group selection studies; and species selection is most fruitfully considered from the point of view of multilevel selection [2]. Finally we argue that the effect hypothesis of macroevolution, requiring, in selection among species, both the absence of group effects on organismic fitness (multilevel selection [1]), and the direct determination of species fitnesses by those of organisms, is untestable with paleontological data. Furthermore, the conditions for the effect hypothesis to hold are extremely restrictive and unlikely to apply to the vast majority of situations encountered in nature. (shrink)

In this paper I argue against the claim, recently put forward by some philosophers of biology and evolutionary biologists, that there can be two or more ontologically distinct levels of selection. I show by comparing the fitness of individuals with that of collectives of individuals in the same environment and over the same period of time – as required to decide if one or more levels of selection is acting in a population – that the selection of collectives is a (...) by-product of selection at the individual level; thus, talking about two or more levels of selection represents merely a different perspective on one and the same process. (shrink)

For evolution by natural selection to occur it is classically admitted that the three ingredients of variation, difference in fitness and heredity are necessary and sufficient. In this paper, I show using simple individual-based models, that evolution by natural selection can occur in populations of entities in which neither heredity nor reproduction are present. Furthermore, I demonstrate by complexifying these models that both reproduction and heredity are predictable Darwinian products (i.e. complex adaptations) of populations initially lacking these two properties but (...) in which new variation is introduced via mutations. Later on, I show that replicators are not necessary for evolution by natural selection, but rather the ultimate product of such processes of adaptation. Finally, I assess the value of these models in three relevant domains for Darwinian evolution. (shrink)

Following Wallace’s suggestion, Darwin framed his theory using Spencer’s expression “survival of the fittest”. Since then, fitness occupies a significant place in the conventional understanding of Darwinism, even though the explicit meaning of the term ‘fitness’ is rarely stated. In this paper I examine some of the different roles that fitness has played in the development of the theory. Whereas the meaning of fitness was originally understood in ecological terms, it took a statistical turn in terms of reproductive success throughout (...) the 20th Century. This has lead to the ever-increasing importance of sexually reproducing organisms and the populations they compose in evolutionary explanations. I will argue that, moving forward, evolutionary theory should look back at its ecological roots in order to be more inclusive in the type of systems it examines. Many biological systems can only be satisfactorily accounted for by offering a non-reproductive account of fitness. This argument will be made by examining biological systems with very small or transient population structures. I argue this has significant consequences for how we define Darwinism, increasing the significance of survival over that of reproduction. (shrink)

Ecological fitness has been suggested to provide a unifying definition of fitness. However, a metric for this notion of fitness was in most cases unavailable except by proxy with differential reproductive success. In this article, I show how differential persistence of lineages can be used as a way to assess ecological fitness. This view is inspired by a better understanding of the evolution of some clonal plants, colonial organisms, and ecosystems. Differential persistence shows the limitation of an ensemblist noncausal understanding (...) of evolution. Causal explanations are necessary to understand the evolution by natural selection of these biological systems. †To contact the author write to: Department of Philosophy, University of Montreal, P.O. Box 6128, Station Centre‐Ville, Montreal, Quebec, H3C 3J7 Canada; e‐mail: [email protected]. (shrink)

The central point of this essay is to demonstrate the incommensurability of ‘Darwinian fitness’ with the numeric values associated with reproductive rates used in population genetics. While sometimes both are called ‘fitness’, they are distinct concepts coming from distinct explanatory schemes. Further, we try to outline a possible answer to the following question: from the natural properties of organisms and a knowledge of their environment, can we construct an algorithm for a particular kind of organismic life-history pattern that itself will (...) allow us to predict whether a type in the population will increase or decrease relative to other types? Introduction Darwinian fitness Reproductive fitness and genetical models of evolution The models of reproductive fitness 4.1 The Standard Viability Model 4.2 Frequency-dependent selection 4.3 Fertility models 4.4 Overlapping generations Fitness as outcome 5.1 Fitness as actual increase in type 5.2 Fitness as expected increase in type 5.2.1 Expected increase within a generation 5.2.2 Expected increase between generations 5.2.3 Postponed reproductive fitness effects The book-keeping problem Conclusion. (shrink)

The central point of this essay is to demonstrate the incommensurability of ‘Darwinian fitness’ with the numeric values associated with reproductive rates used in population genetics. While sometimes both are called ‘fitness’, they are distinct concepts coming from distinct explanatory schemes. Further, we try to outline a possible answer to the following question: from the natural properties of organisms and a knowledge of their environment, can we construct an algorithm for a particular kind of organismic life-history pattern that itself will (...) allow us to predict whether a type in the population will increase or decrease relative to other types? 1. Introduction2. Darwinian fitness3. Reproductive fitness and genetical models of evolution4. The models of reproductive fitness4.1 The Standard Viability Model4.2 Frequency-dependent selection4.3 Fertility models4.4 Overlapping generations5. Fitness as outcome5.1 Fitness as actual increase in type5.2 Fitness as expected increase in type5.2.1 Expected increase within a generation5.2.2 Expected increase between generations5.2.3 Postponed reproductive fitness effects6. The book-keeping problem7. Conclusion. (shrink)

It has been argued that biological fitness cannot be defined as expected number of offspring in all contexts. Some authors argue that fitness therefore merely satisfies a common schema or that no unified mathematical characterization of fitness is possible. I argue that comparative fitness must be relativized to an evolutionary effect; thus relativized, fitness can be given a unitary mathematical characterization in terms of probabilities of producing offspring and other effects. Such fitnesses will sometimes be defined in terms of probabilities (...) of effects occurring over the long term, but these probabilities nevertheless concern effects occurring over the short term. †To contact the author, please write to: Department of Philosophy, University of Alabama at Birmingham, HB 414A, 900 13th Street South, Birmingham, AL 35294‐1260; e‐mail: [email protected]. (shrink)

Causality offers the first comprehensive coverage of causal analysis in many sciences, including recent advances using graphical methods. Pearl presents a unified account of the probabilistic, manipulative, counterfactual and structural approaches to causation, and devises simple mathematical tools for analyzing the relationships between causal connections, statistical associations, actions and observations. The book will open the way for including causal analysis in the standard curriculum of statistics, artificial intelligence, business, epidemiology, social science and economics.

The book presents a new way of understanding Darwinism and evolution by natural selection, combining work in biology, philosophy, and other fields.

Manipulablity theories of causation, according to which causes are to be regarded as handles or devices for manipulating effects, have considerable intuitive appeal and are popular among social scientists and statisticians. This article surveys several prominent versions of such theories advocated by philosophers, and the many difficulties they face. Philosophical statements of the manipulationist approach are generally reductionist in aspiration and assign a central role to human action. These contrast with recent discussions employing a broadly manipulationist framework for understanding causation, (...) such as those due to the computer scientist Judea Pearl and others, which are non-reductionist and rely instead on the notion of an intervention. This is simply an appropriately exogenous causal process; it has no essential connection with human action. This interventionist framework manages to avoid at least some of these difficulties faced by traditional philosophical versions of the manipulability theory and helps to clarify the content of causal claims. (shrink)

this requirement for adaptations. Emergent characters are always potential adaptations. Not all selection processes produce adaptations, however. The key issue, in delineating a selection process, is the relationship between a character and fitness. The emergent character approach is more restrictive than alternative schemas that delineate selection..

The concept of fitness began its career in biology long before evolutionary theory was mathematized. Fitness was used to describe an organism’s vigor, or the degree to which organisms “fit” into their environments. An organism’s success in avoiding predators and in building a nest obviously contribute to its fitness and to the fitness of its offspring, but the peacock’s gaudy tail seemed to be in an entirely different line of work. Fitness, as a term in ordinary language (as in “physical (...) fitness”) and in its original biological meaning, applied to the survival of an organism and its offspring, not to sheer reproductive output (Paul ////; Cronin 1991). Darwin’s separation of natural from sexual selection may sound odd from a modern perspective, but it made sense from this earlier point of view. (shrink)