Paul Edmund Griffiths

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.

Integrating the study of human diversity into the human evolutionary sciences requires substantial revision of traditional conceptions of a shared human nature. This process may be made more difficult by entrenched, 'folkbiological' modes of thought. Earlier work by the authors suggests that biologically naive subjects hold an implicit theory according to which some traits are expressions of an animal's inner nature while others are imposed by its environment. In this paper, we report further studies that extend and refine our account of this aspect of folkbiology. We examine biologically naive subjects' judgments about whether traits of an animal are 'innate', 'in its DNA' or 'part of its nature'. Subjects do not understand these three descriptions to be equivalent. Both innate and in its DNA have the connotation that the trait is species-typical. This poses an obstacle to the assimilation of the biology of polymorphic and plastic traits by biologically naive audiences. Researchers themselves may not be immune to the continuing pull of folkbiological modes of thought.

I defend the view that many biological categories are defined by homology against a series of arguments designed to show that all biological categories are defined, at least in part, by selected function. I show that categories of homology are `abnormality inclusive'—something often alleged to be unique to selected function categories. I show that classifications by selected function are logically dependent on classifications by homology, but not vice-versa. Finally, I reject the view that biologists must use considerations of selected function to abstract away from variation and pathology to form a canonical description of a class of biological systems.

We describe an approach to measuring biological information where ‘information’ is understood in the sense found in Francis Crick’s foundational contributions to molecular biology. Genes contain information in this sense, but so do epigenetic factors, as many biologists have recognized. The term ‘epigenetic’ is ambiguous, and we introduce a distinction between epigenetic and exogenetic inheritance to clarify one aspect of this ambiguity. These three heredity systems play complementary roles in supplying information for development. We then consider the evolutionary significance of the three inheritance systems. Whilst the genetic inheritance system was the key innovation in the evolution of heredity, in modern organisms the three systems each play important and complementary roles in heredity and evolution. Our focus in the earlier part of the paper is on ‘proximate biology’, where information is a substantial causal factor that causes organisms to develop and causes offspring to resemble their parents. But much philosophical work has focused on information in ‘ultimate biology’. Ultimate information is a way of talking about the evolutionary design of the mechanisms of development and inheritance. We conclude by clarifying the relationship between the two. Ultimate information is not a causal factor that acts in development or heredity, but it can help to explain the evolution of proximate information, which is.

The idea that development is the expression of information accumulated during evolution and that heredity is the transmission of this information is surprisingly hard to cash out in strict, scientific terms. This paper seeks to do so using the sense of information introduced by Francis Crick in his sequence hypothesis and central dogma of molecular biology. It focuses on Crick's idea of precise determination. This is analysed using an information-theoretic measure of causal specificity. This allows us to reconstruct some of Crick's claims about information in transcription and translation. Crick's approach to information has natural extensions to non-coding regions of DNA, to epigenetic marks, and to the genetic or environmental upstream causes of those epigenetic marks. Epigenetic information cannot be reduced to genetic information. The existence of biological information in epigenetic and exogenetic factors is relevant to evolution as well as to development.

It is now widely accepted that a scientifically credible conception of human nature must reject the folkbiological idea of a fixed, inner essence that makes us human. We argue here that to understand human nature is to understand the plastic process of human development and the diversity it produces. Drawing on the framework of developmental systems theory and the idea of developmental niche construction we argue that human nature is not embodied in only one input to development, such as the genome, and that it should not be confined to universal or typical human characteristics. Both similarities and certain classes of differences are explained by a human developmental system that reaches well out into the 'environment'. We point to a significant overlap between our account and the ‘Life History Trait Cluster’ account of Grant Ramsey, and defend the developmental systems account against the accusation that trying to encompass developmental plasticity and human diversity leads to an unmanageably complex account of human nature.

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.

Experimental philosophy of science gathers empirical data on how key scientific concepts are understood by particular scientific communities. In this paper we briefly describe two recent studies in experimental philosophy of biology, one investigating the concept of the gene, the other the concept of innateness. The use of experimental methods reveals facts about these concepts that would not be accessible using the traditional method of intuitions about possible cases. It also contributes to the study of conceptual change in science, which we understand as the result of a form of conceptual ecology, in which concepts become adapted to specific epistemic niches.

The Royal Institute of Philosophy’s London lecture series for 2004–2005 offers a useful snapshot of the current state of philosophy of biology in Britain. With one or two exceptions the papers are not simply current research articles. The authors map out questions they feel need more research, analyse ongoing debates, or outline the program of their own previously published work. This presumably reflects the fact that the papers are based closely on public lectures. It also makes for surprisingly easy and enjoyable reading in the context of such a volume.

There remains a division between the work of philosophers who draw on the sciences of the mind to understand emotion and those who see the philosophy of emotion as more self-sufficient. This article examines this methodological division before reviewing some of the debates that have figured in the philosophical literature of the last decade: whether emotion is a single kind of thing, whether there are discrete categories of emotion, and whether emotion is a form of perception. These questions have been addressed by both sides of the methodological divide and the integration of these two approaches would have clear benefits

The proposal that the concept of innateness expresses a 'folk biological' theory of the 'inner natures' of organisms was tested by examining the response of biologically naive participants to a series of realistic scenarios concerning the development of birdsong. Our results explain the intuitive appeal of existing philosophical analyses of the innateness concept. They simultaneously explain why these analyses are subject to compelling counterexamples. We argue that this explanation undermines the appeal of these analyses, whether understood as analyses of the vernacular concept or as explications of that concept for the purposes of science.

The current state of knowledge in psychology, cognitive neuroscience and behavioral ecology allows a fairly robust characterization of at least some, so-called ?basic emotions? - short-lived emotional responses with homologues in other vertebrates. Philosophers, however are understandably more focused on the complex emotion episodes that figure in folk-psychological narratives about mental life, episodes such as the evolving jealousy and anger of a person in an unraveling sexual relationship. One of the most pressing issues for the philosophy of emotion is the relationship between basic emotions and these complex emotion episodes. In this paper, I add to the list of existing, not necessarily incompatible, proposals concerning the relationship between basic emotions and complex emotions. I analyze the writings of ?transactional? psychologists of emotion, particularly those who see their work as a contribution to behavioral ecology, and offer a view of the basic emotion that focuses as much on their interpersonal functions as on their intrapersonal functions. Locating basic emotions and their evolutionary development in a context of processes of social interaction, I suggest, provides a way to integrate our knowledge of basic emotions into an understanding of the larger emotional episodes that have more obvious implications for philosophical disciplines such as moral psychology

The 'developmental systems' perspective in biology is intended to replace the idea of a genetic program. This new perspective is strongly convergent with recent work in psychology on situated/embodied cognition and on the role of external 'scaffolding' in cognitive development. Cognitive processes, including those which can be explained in evolutionary terms, are not 'inherited' or produced in accordance with an inherited program. Instead, they are constructed in each generation through the interaction of a range of developmental resources. The attractors which emerge during development and explain robust and/or widespread outcomes are themselves constructed during the process. At no stage is there an explanatory stopping point where some resources control or program the rest of the developmental cascade. 'Human nature' is a description of how things generally turn out, not an explanation of why they turn out that way. Finally, we suggest that what is distinctive about human development is its degree of reliance on external scaffolding.

The Developmental Systems approach to evolution is defended against the alternative extended replicator approach of Sterelny, Smith and Dickison (1996). A precise definition is provided of the spatial and temporal boundaries of the life-cycle that DST claims is the unit of evolution. Pacé Sterelny et al., the extended replicator theory is not a bulwark against excessive holism. Everything which DST claims is replicated in evolution can be shown to be an extended replicator on Sterelny et al.s definition. Reasons are given for scepticism about the heuristic value claimed for the extended replicator concept. For every competitive, individualistic insight the replicator theorist has a cooperative, systematic blindspot.

In this chapter we examine the relationship between biological information, the key biological concept of specificity, and recent philosophical work on causation. We begin by showing how talk of information in the molecular biosciences grew out of efforts to understand the sources of biological specificity. We then introduce the idea of ‘causal specificity’ from recent work on causation in philosophy, and our own, information theoretic measure of causal specificity. Biological specificity, we argue, is simple the causal specificity of certain biological processes. This, we suggest, means that causal relationships in biology are ‘informational’ relationships simply when they are highly specific relationships. Biological information can be identified with the storage, transmission and exercise of biological specificity. It has been argued that causal relationships should not be regarded as informational relationship unless they are ‘arbitrary’. We argue that, whilst arbitrariness is an important feature of many causal relationships in living systems, it should not be used in this way to delimit biological information. Finally, we argue that biological specificity, and hence biological information, is not confined to nucleic acids but distributed among a wide range of entities and processes.

The emerging discipline of evolutionary developmental biology has opened up many new lines of investigation into morphological evolution. Here I explore how two of the core theoretical concepts in ‘evo-devo’ – modularity and homology – apply to evolutionary psychology. I distinguish three sorts of module – developmental, functional and mental modules and argue that mental modules need only be ‘virtual’ functional modules. Evolutionary psychologists have argued that separate mental modules are solutions to separate evolutionary problems. I argue that the structure of developmental modules in an organism helps determine what counts as a separate evolutionary problem for that organism. I suggest that homology as an organizing principle for research in evolutionary psychology, has been severely neglected in favor of analogy (adaptive function). I consider some arguments suggesting that determining homology is less epistemically demanding than determining adaptive function and argue that psychological categories defined by homology are, in fact, more suitable objects of psychological – and particularly neuropsychological – investigation than categories defined by analogy.

It is unreasonable to assume that our pre-scientific emotion vocabulary embodies all and only those distinctions required for a scientific psychology of emotion. The psychoevolutionary approach to emotion yields an alternative classification of certain emotion phenomena. The new categories are based on a set of evolved adaptive responses, or affect-programs, which are found in all cultures. The triggering of these responses involves a modular system of stimulus appraisal, whose evoluations may conflict with those of higher-level cognitive processes. Whilst the structure of the adaptive responses is innate, the contents of the system which triggers them are largely learnt. The circuits subserving the adaptive responses are probably located in the limbic system. This theory of emotion is directly applicable only to a small sub-domain of the traditional realm of emotion. It can be used, however, to explain the grouping of various other phenomena under the heading of emotion, and to explain various characteristic failings of the pre-scientific conception of emotion.