Nicholas Shea

There is increasing evidence for epigenetically mediated transgenerational inheritance across taxa. However, the evolutionary implications of such alternative mechanisms of inheritance remain unclear. Herein, we show that epigenetic mechanisms can serve two fundamentally different functions in transgenerational inheritance: (i) selection-based effects, which carry adaptive information in virtue of selection over many generations of reliable transmission; and (ii) detection-based effects, which are a transgenerational form of adaptive phenotypic plasticity. The two functions interact differently with a third form of epigenetic information transmission, namely information about cell state transmitted for somatic cell heredity in multicellular organisms. Selection-based epigenetic information is more likely to conflict with somatic cell inheritance than is detection-based epigenetic information. Consequently, the evolutionary implications of epigenetic mechanisms are different for unicellular and multicellular organisms, which underscores the conceptual and empirical importance of distinguishing between these two different forms of transgenerational epigenetic effect.

The New Thinking contained in this volume rejects an Evolutionary Psychology that is committed to innate domain-specific psychological mechanisms: gene-based adaptations that are unlearnt, developmentally fixed and culturally universal. But the New Thinking does not simply deny the importance of innate psychological traits. The problem runs deeper: the concept of innateness is not suited to distinguishing between the two positions. That points to a more serious problem with the concept of innateness as it is applied to human psychological phenotypes. This paper argues that the features of recent human evolution highlighted by the New Thinking imply that the concept of inherited representation, set out here, is a better tool for theorising about human cognitive evolution.

One of the great outstanding problems in materialist philosophy of mind is the problem of how there can be space in the material world for intentionality. In the 1980s Ruth Millikan formulated a detailed theory according to which representations are physical particulars and their contents are complex relational properties of those particulars which can be specified in terms of respectable properties drawn from the natural sciences. In particular, she relied on the biological concept of the function of a trait, and the existence of historical conditions which enter into an evolutionary explanation of the operation of that trait. The present article is an introduction to this influential theory of intentionality.

Parents influence the development of their offspring in many ways beyond the transmission of DNA. This includes transfer of epigenetic states, nutrients, antibodies and hormones, and behavioural interactions after birth. While the evolutionary consequences of such nongenetic inheritance are increasingly well understood, less is known about how inheritance mechanisms evolve. Here, we present a simple but versatile model to explore the adaptive evolution of non-genetic inheritance. Our model is based on a switch mechanism that produces alternative phenotypes in response to different inputs, including genes and non-genetic factors transmitted from parents and the environment experienced during development. This framework shows how genetic and non-genetic inheritance mechanisms and environmental conditions can act as cues by carrying correlational information about future selective conditions. Differential use of these cues is manifested as different degrees of genetic, parental or environmental morph determination. We use this framework to evaluate the conditions favouring non-genetic inheritance, as opposed to genetic determination of phenotype or within-generation plasticity, by applying it to two putative examples of adaptive non-genetic inheritance: maternal effects on seed germination in plants and transgenerational phase shift in desert locusts. Our simulation models show how the adaptive value of non-genetic inheritance depends on its mechanism, the pace of environmental change, and life history characteristics.

Can findings from psychology and cognitive neuroscience about the neural mechanisms involved in decision-making can tell us anything useful about the commonly-understood mental phenomenon of making voluntary choices? Two philosophical objections are considered. First, that the neural data is subpersonal, and so cannot enter into illuminating explanations of personal level phenomena like voluntary action. Secondly, that mental properties are multiply realized in the brain in such a way as to make them insusceptible to neuroscientific study. The paper argues that both objections would be weakened by the discovery of empirical generalisations connecting subpersonal properties with the personal level. It gives three case studies that furnish evidence to that effect. It argues that the existence of such interrelations are consistent with a plausible construal of the personal-subpersonal distinction. Furthermore, there is no reason to suppose that the notion subpersonal representation relied on in cognitive neuroscience illicitly imports personal-level phenomena like consciousness or normativity, or is otherwise explanatorily problematic.

Recent theoretical work has identified a tightly-constrained sense in which genes carry representational content. Representational properties of the genome are founded in the transmission of DNA over phylogenetic time and its role in natural selection. However, genetic representation is not just relevant to questions of selection and evolution. This paper goes beyond existing treatments and argues for the heterodox view that information generated by a process of selection over phylogenetic time can be read in ontogenetic time, in the course of individual development. Recent results in evolutionary biology, drawn both from modelling work, and from experimental and observational data, support a role for genetic representation in explaining individual ontogeny: both genetic representations and environmental information are read by the mechanisms of development, in an individual, so as to lead to adaptive phenotypes. Furthermore, in some cases there appears to have been selection between individuals that rely to different degrees on the two sources of information. Thus, the theory of representation in inheritance systems like the genome is much more than just a coherent reconstruction of information talk in biology. Genetic representation is a property with considerable explanatory utility.

Developmental Systems Theory (DST) emphasises the importance of non-genetic factors in development and their relevance to evolution. A common, deflationary reaction is that it has long been appreciated that non-genetic factors are causally indispensable. This paper argues that DST can be reformulated to make a more substantive claim: that the special role played by genes is also played by some (but not all) non-genetic resources. That special role is to transmit inherited representations, in the sense of Shea (2007: Biology and Philosophy, 22, 313-331). Formulating DST as the claim that there are non-genetic inherited representations turns it into a striking, empirically-testable hypothesis, driving the sort of investigations that are only now beginning to appear in the scientific literature. DST’s characteristic rejection of a gene vs. environment dichotomy is preserved, but without dissolving all potentially explanatory distinctions into an interactionist causal soup, as some have alleged

The explosion of scientific results about epigenetic and other parental effects appears bewilderingly diverse. An important distinction helps to bring order to the data. Firstly, parents can detect adaptively-relevant information and transmit it to their offspring who rely on it to set a plastic phenotype adaptively. Secondly, adaptively-relevant information may be generated by a process of selection on a reliably transmitted parental effect. The distinction is particularly valuable in revealing two quite different ways in which human cultural transmission may operate.

Consciousness in experimental subjects is typically inferred from reports and other forms of voluntary behaviour. A wealth of everyday experience confirms that healthy subjects do not ordinarily behave in these ways unless they are conscious. Investigation of consciousness in vegetative state patients has been based on the search for neural evidence that such broad functional capacities are preserved in some vegetative state patients. We call this the standard approach. To date, the results of the standard approach have suggested that some vegetative state patients might indeed be conscious, although they fall short of being demonstrative. The fact that some vegetative state patients show evidence of consciousness according to the standard approach is remarkable, for the standard approach to consciousness is rather conservative, and leaves open the pressing question of how to ascertain whether patients who fail such tests are conscious or not. We argue for a cluster-based ‘natural kind’ methodology that is adequate to that task, both as a replacement for the approach that currently informs research into the presence or absence of consciousness in vegetative state patients and as a methodology for the science of consciousness more generally. 1. Introduction2. The Vegetative State3. The Standard Approach4. The Natural Kind Methodology5. Is Consciousness a Special Case? 5.1 Is consciousness a natural kind?5.2 A special obstacle?6. Conclusion.

Block’s well-known distinction between phenomenal consciousness and access consciousness has generated a large philosophical literature about putative conceptual connections between the two. The scientific literature about whether they come apart in any actual cases is rather smaller. Empirical evidence gathered to date has not settled the issue. Some put this down to a fundamental methodological obstacle to the empirical study of the relation between phenomenal consciousness and access consciousness. Block (2007) has drawn attention to the methodological puzzle and attempted to answer it. While the evidence Block points to is relevant and important, this paper puts forward a more systematic framework for addressing the puzzle. To give it a label, the approach is to study phenomenal consciousness as a natural kind. The approach allows consciousness studies to move beyond the initial means of identifying instances of the kind, like verbal report, and to find its underlying nature. It is wellrecognised that facts about an underlying kind may allow identification of instances of the kind that do not match the initial means of identification (cp. non-liquid samples of water). This paper shows that the same method can be deployed to investigate phenomenal consciousness independently of access consciousness

Just how far can externalism go? In this exciting new book Ruth Millikan explores a radically externalist treatment of empirical concepts (Millikan 2000). For the last thirty years philosophy of mind’s ties to meaning internalism have been loosened. The theory of content has swung uncomfortably on its moorings in a fickle current, straining against opposing ties to mind and world. In this book Millikan casts conceptual content adrift from the thinker: what determines the content of a concept is not cognitively accessible. She has only the stanchion of the world to hold her theory fast. She hopes that the tide will turn, and the theory of meaning will come stably to rest downstream of this anchor. This book is a bold exploration of how that might be achieved.

This paper advocates explicitness about the type of entity to be considered as content- bearing in connectionist systems; it makes a positive proposal about how vehicles of content should be individuated; and it deploys that proposal to argue in favour of representation in connectionist systems. The proposal is that the vehicles of content in some connectionist systems are clusters in the state space of a hidden layer. Attributing content to such vehicles is required to vindicate the standard explanation for some classificatory networks’ ability to generalise to novel samples their correct classification of the samples on which they were trained

A step towards a theory of consciousness would be to characterise the effect of consciousness on information processing. One set of results suggests that the effect of consciousness is to interfere with computations that are optimally performed non-consciously. Another set of results suggests that conscious, system 2 processing is the home of norm-compliant computation. This is contrasted with system 1 processing, thought to be typically unconscious, which operates with useful but error-prone heuristics. These results can be reconciled by separating out two different distinctions: between conscious and non-conscious representations, on the one hand, and between automatic and deliberate processes, on the other. This pair of distinctions is used to illuminate some existing experimental results and to resolve the puzzle about whether consciousness helps or hinders accurate information processing. This way of resolving the puzzle shows the importance of another category, which we label ‘type 0 cognition’, characterised by automatic computational processes operating on non-conscious representations.

This paper sets out a view about the explanatory role of representational content and advocates one approach to naturalising content – to giving a naturalistic account of what makes an entity a representation and in virtue of what it has the content it does. It argues for pluralism about the metaphysics of content and suggests that a good strategy is to ask the content question with respect to a variety of predictively successful information processing models in experimental psychology and cognitive neuroscience; and hence that data from psychology and cognitive neuroscience should play a greater role in theorising about the nature of content. Finally, the contours of the view are illustrated by drawing out and defending a surprising consequence: that individuation of vehicles of content is partly externalist.

An interesting feature of some sets of representations is that their structure mirrors the structure of the items they represent. Founding an account of representational content on isomorphism, homomorphism or structural resemblance has proven elusive, however, largely because these relations are too liberal when the candidate structure over representational vehicles is unconstrained. Furthermore, in many cases where there is a clear isomorphism, it is not relied on in the way the representations are used. That points to a potential resolution: that an isomorphism must be used, hence usable, if it is to be an ingredient in a theory of content. This paper argues that the class of exploitable isomorphisms can indeed play a content-constituting role

The success of a piece of behaviour is often explained by its being caused by a true representation (similarly, failure falsity). In some simple organisms, success is just survival and reproduction. Scientists explain why a piece of behaviour helped the organism to survive and reproduce by adverting to the behaviour’s having been caused by a true representation. That usage should, if possible, be vindicated by an adequate naturalistic theory of content. Teleosemantics cannot do so, when it is applied to simple representing systems (Godfrey-Smith 1996). Here it is argued that the teleosemantic approach to content should therefore be modified, not abandoned, at least for simple representing systems. The new ‘infotel-semantics’ adds an input condition to the output condition offered by teleosemantics, recognising that it is constitutive of content in a simple representing system that the tokening of a representation should correlate probabilistically with the obtaining of its specific evolutionary success condition

Humans can think about their conscious experiences using a special class of ?phenomenal? concepts. Psychophysical identity statements formulated using phenomenal concepts appear to be contingent. Kripke argued that this intuited contingency could not be explained away, in contrast to ordinary theoretical identities where it can. If the contingency is real, property dualism follows. Physicalists have attempted to answer this challenge by pointing to special features of phenomenal concepts that explain the intuition of contingency. However no physicalist account of their distinguishing features has proven to be satisfactory. Leading accounts rely on there being a phenomenological difference between tokening a physical-functional concept and tokening a phenomenal concept. This paper shows that existing psychological data undermine that claim. The paper goes on to suggest that the recalcitrance of the intuition of contingency may instead by explained by the limited means people typically have for applying their phenomenal concepts. Ways of testing that suggestion empirically are proposed

Fodor’s radical concept nativism flowed from his view that hypothesis testing is the only route to concept acquisition. Many have successfully objected to the overly-narrow restriction to learning by hypothesis testing. Existing representations can be connected to a new representational vehicle so as to constitute a sustaining mechanism for a new representation, without the new representation thereby being constituted by or structured out of the old. This paper argues that there is also a deeper objection. Connectionism shows that a more fundamental assumption underpinning the debate can also be rejected: the assumption that the development of a new representation must be explained in content-involving terms if innateness is to be avoided. Fodor has argued that connectionism offers no new resources to explain concept acquisition: unless it is merely an uninteresting claim about neural implementation, connectionism’s defining commitment to distributed representations reduces to the claim that some representations are structured out of others (which is the old, problematic research programme). Examination of examples of representational development in connectionist networks shows, however, that some such models explain the development of new representational capacities in non-representational terms. They illustrate the possibility of representational development that is not explicable-by-content. Connectionist representations can be distributed in an important sense, which is incompatible with the assumption of explanation-by-content: they can be distributed over non-representational resources that account for their development. Rejecting the assumption of explanation-by-content thereby opens up a more radical way of rejecting Fodor’s argument for radical concept nativism.

Millikan’s theory of content purports to rely heavily on the existence of isomorphisms between a system of representations and the things in the world which they represent — “the mapping requirement for being intentional signs” (Millikan 2004, p. 106). This paper asks whether those isomorphisms are doing any substantive explanatory work. Millikan’s isomorphism requirement is deployed for two main purposes. First, she claims that the existence of an isomorphism is the basic representing relation, with teleology playing a subsidiary role — to account for misrepresentation (the possibility of error). Second, Millikan relies on an isomorphism requirement in order to guarantee that a system of representations displays a kind of productivity. This seemingly strong reliance on isomorphism has prompted the objection that isomorphism is too liberal to be the basic representing relation: there are isomorphisms between any system of putative representations and any set, of the same cardinality, of items putatively represented. This paper argues that all the work in fixing content is in fact done by the teleology. Deploying Millikan’s teleology-based conditions to ascribe contents will ensure that there is an isomorphism between representations and the things they represent, but the isomorphism ‘requirement’ is playing no substantive explanatory role in Millikan’s account of content determination. So an objection to her theory based on the liberality of isomorphism is misplaced. The second role for isomorphism is to account for productivity. If some kind of productivity is indeed necessary for representation, then functional isomorphism will again be too liberal a constraint to account for that feature. The paper suggests an alternative way of specifying the relation between a system of representations and that which they represent which is capable of playing an explanatory role in accounting for Millikan’s type of productivity. In short, the liberality of isomorphism is no objection to Millikan’s teleosemantics, since the isomorphism ‘requirement’ need play no independent substantive role in Millikan’s account of representation.

The concept of innateness is used to make inferences between various better-understood properties, like developmental canalization, evolutionary adaptation, heritability, species-typicality, and so on (‘innateness-related properties’). This article uses a recently-developed account of the representational content carried by inheritance systems like the genome to explain why innateness-related properties cluster together, especially in non-human organisms. Although inferences between innateness-related properties are deductively invalid, and lead to false conclusions in many actual cases, where some aspect of a phenotypic trait develops in reliance on a genetic representation it will tend, better than chance, to have many of the innateness-related properties. The account also shows why inferences between innateness-related properties sometimes fail and argues that such inferences are especially misleading when applied to human psychology and behaviour because human psychological development is especially reliant on non-genetic inherited representations

Emerging evidence suggests that the long-established distinction between habit-based and goal-directed decision-making mechanisms can also be sustained in humans. Although the habit-based system has been extensively studied in humans, the goal-directed system is less well characterized. This review brings to that task the distinction between conceptual and nonconceptual representational mechanisms. Conceptual representations are structured out of semantic consituents - the use of which requires an ability to perform some language-like syntactic processing. Decision making - as investigated by neuroscience and psychology - is normally studied in isolation from questions about concepts as studied in philosophy and cognitive psychology. We ask what role concepts play in the "goal-directed" decision-making systems. We argue that one fruitful way of studying this system in humans is to investigate the extent to which it deploys conceptual representations

The distinction between top-down and bottom-up effects is widely relied on in experimental psychology. However, there is an important problem with the way it is normally defined. Top-down effects are effects of previously-stored information on processing the current input. But on the face of it that includes the information that is implicit in the operation of any psychological process – in its dispositions to transition from some types of representational state to others. This paper suggests a way to distinguish information stored in that way from the kind of influence of prior information that psychologists are concerned to classify as a top-down effect. So-drawn, the distinction is not just of service to theoretical psychology. Asking about the extent of top-down processing is one way to pose some of the questions at issue in philosophical debates about cognitive penetrability – about the extent of the influence of cognitive states on perception. The existence of a theoretically-useful perception-cognition distinction has come under pressure, but even if it has to be abandoned, some of the concerns addressed in the cognitive penetrability literature can be recaptured by asking about the extent of top-down influences on any given psychological process. That formulation is more general, since it can be applied to any psychological process, not just those that are paradigmatically perceptual.

Recent work combining cognitive neuroscience with computational modelling suggests that distributed patterns of neural firing may represent probability distributions. This paper asks: what makes it the case that distributed patterns of firing, as well as carrying information about (correlating with) probability distributions over worldly parameters, represent such distributions? In examples of probabilistic population coding, it is the way information is used in downstream processing so as to lead to successful behaviour. In these cases content depends on factors beyond bare information, contra Brian Skyrms’ (2010) view that representational content can be fully characterised in information-theoretic terms.