We examine two different descriptions of the behavioral functions of the hippocampal system. One emphasizes spatially organized behaviors, especially those using cognitive maps. The other emphasizes memory, particularly working memory, a short-term memory that requires iexible stimulus-response associations and is highly susceptible to interference. The predictive value of the spatial and memory descriptions were evaluated by testing rats with damage to the hippocampal system in a series of experiments, independently manipulating the spatial and memory characteristics of a behavioral task. No (...) dissociations were found when the spatial characteristics of the stimuli to be remembered were changed; lesions produced a similar deficit in both spatial and nonspatial test procedures, indicating that the hippocampus was similarly involved regardless of the spatial nature of the task. In contrast, a marked dissociation was found when the memory requirements were altered. Rats with lesions were able to perform accurately in tasks that could be solved exclusively on the basis of reference memory. They performed at chance levels and showed no signs of recovery even with extensive postoperative training in tasks that required working memory. In one experiment all the characteristics of the reference memory and working memory procedures were identical except the type of memory required. Consequently, the behavioral dissociation cannot be explained by differences in attention, motivation, response inhibition, or the type of stimuli to be remembered. As a result of these experiments we propose that the hippocampus is selectively involved in behaviors that require working memory, irrespective of the type of material that is to be processed by that memory. (shrink)

Many philosophers and psychologists have attempted to elucidate the nature of mental representation by appealing to notions like isomorphism or abstract structural resemblance. The ‘structural representations’ that these theorists champion are said to count as representations by virtue of functioning as internal models of distal systems. In his 2007 book, Representation Reconsidered, William Ramsey endorses the structural conception of mental representation, but uses it to develop a novel argument against representationalism, the widespread view that cognition essentially involves the manipulation of (...) mental representations. Ramsey argues that although theories within the ‘classical’ tradition of cognitive science once posited structural representations, these theories are being superseded by newer theories, within the tradition of connectionism and cognitive neuroscience, which rarely if ever appeal to structural representations. Instead, these theories seem to be explaining cognition by invoking so-called ‘receptor representations’, which, Ramsey claims, aren’t genuine representations at all—despite being called representations, these mechanisms function more as triggers or causal relays than as genuine stand-ins for distal systems. I argue that when the notions of structural and receptor representation are properly explicated, there turns out to be no distinction between them. There only appears to be a distinction between receptor and structural representations because the latter are tacitly conflated with the ‘mental models’ ostensibly involved in offline cognitive processes such as episodic memory and mental imagery. While structural representations might count as genuine representations, they aren’t distinctively mental representations, for they can be found in all sorts of non-intentional systems such as plants. Thus to explain the kinds of offline cognitive capacities that have motivated talk of mental models, we must develop richer conceptions of mental representation than those provided by the notions of structural and receptor representation. (shrink)

Theories of spatial cognition are derived from many sources. Psychologists are concerned with determining the features of the mind which, in combination with external inputs, produce our spatialized experience. A review of philosophical and other approaches has convinced us that the brain must come equipped to impose a three-dimensional Euclidean framework on experience – our analysis suggests that object re-identification may require such a framework. We identify this absolute, nonegocentric, spatial framework with a specific neural system centered in the hippocampus.A (...) consideration of the kinds of behaviours in which such a spatial mapping system would be important is followed by an analysis of the anatomy and physiology of this system, with special emphasis on the place-coded neurons recorded in the hippocampus of freely moving rats. A tentative physiological model for the hippocampal cognitive map is proposed. A review of lesion studies, in tasks as diverse as discrimination learning, avoidance, and extinction, shows that the cognitive map notion can adequately explain much of the data.The model is extended to humans by the assumption that spatial maps are built in one hemisphere, semantic maps in the other. The latter provide a semantic deep structure within which discourse comprehension and production can be achieved. Evidence from the study of amnesic patients, briefly reviewed, is consistent with this extension. (shrink)

It is traditionally believed that cerebral activation (the presence of low voltage fast electrical activity in the neocortex and rhythmical slow activity in the hippocampus) is correlated with arousal, while deactivation (the presence of large amplitude irregular slow waves or spindles in both the neocortex and the hippocampus) is correlated with sleep or coma. However, since there are many exceptions, these generalizations have only limited validity. Activated patterns occur in normal sleep (active or paradoxical sleep) and during states of anesthesia (...) and coma. Deactivated patterns occur, at times, during normal waking, or during behavior in awake animals treated with atropinic drugs. Also, the fact that patterns characteristic of sleep, arousal, and waking behavior continue in decorticate animals indicates that reticulo-cortical mechanisms are not essential for these aspects of behavior. (shrink)

This paper argues that a set of questioning attitudes are among the foundations of human and animal minds. While both verbal questioning and states of curiosity are generally explained in terms of metacognitive desires for knowledge or true belief, I argue that each is better explained by a prelinguistic sui generis type of mental attitude of questioning. I review a range of considerations in support of such a proposal and improve on previous characterizations of the nature of these attitudes. I (...) then broaden their explanatory scope to include a number of forms of exploratory search. The paper has three main goals: to characterize the nature of the questioning attitudes, outlining their causal role and type of content; to argue that they are fundamental components of the mind, being widespread among animals and not constructed during ontogeny out of other attitudes; and to suggest that they explain a great deal more behavior than one might think. (shrink)

Colloquially, episodic memory is described as “the memory of personally experienced events”. Even though episodic memory has been studied in psychology and neuroscience for about six decades, there is still great uncertainty as to what episodic memory is. Here we ask how episodic memory should be characterized in order to be validated as a natural kind. We propose to conceive of episodic memory as a knowledge-like state that is identified with an experientially based mnemonic representation of an episode that allows (...) for a mnemonic simulation thereof. We call our analysis the Sequence Analysis of Episodic Memory since episodes will be analyzed in terms of sequences of events. Our philosophical analysis of episodic memory is driven and supported by experimental results from psychology and neuroscience. We discuss selected experimental results that provide exemplary evidence for uniform causal mechanisms underlying the properties of episodic memory and argue that episodic memory is a natural kind. The argumentation proceeds along three cornerstones: First, psychological evidence suggests that a violation of any of the proposed conditions for episodic memory amounts to a deficiency of episodic memory and no form of memory or cognitive process but episodic memory fulfills them. Second, empirical results support a claim that the principal anatomical substrate of episodic memory is the hippocampus. Finally, we can pin down causal mechanisms onto neural activities in the hippocampus to explain the psychological states and processes constituting episodic memory. (shrink)

While neuroscientists often characterize brain activity as representational, many philosophers have construed these accounts as just theorists’ glosses on the mechanism. Moreover, philosophical discussions commonly focus on finished accounts of explanation, not research in progress. I adopt a different perspective, considering how characterizations of neural activity as representational contributes to the development of mechanistic accounts, guiding the investigations neuroscientists pursue as they work from an initial proposal to a more detailed understanding of a mechanism. I develop one illustrative example involving (...) research on the information-processing mechanisms mammals employ in navigating their environments. This research was galvanized by the discovery in the 1970s of place cells in the hippocampus. This discovery prompted research in what the activity of these cells represents and how place representations figure in navigation. It also led to the discovery of a host of other types of neurons—grid cells, head-direction cells, boundary cells—that carry other types of spatial information and interact with place cells in the mechanism underlying spatial navigation. As I will try to make clear, the research is explicitly devoted to identifying representations and determining how they are constructed and used in an information processing mechanism. Construals of neural activity as representations are not mere glosses but are characterizations to which neuroscientists are committed in the development of their explanatory accounts. (shrink)

The dominant neuroscientific theory of spatial memory is, like many theories in neuroscience, a multilevel description of a mechanism. The theory links the activities of molecules, cells, brain regions, and whole organisms into an integrated sketch of an explanation for the ability of organisms to navigate novel environments. Here I develop a taxonomy of interlevel experimental strategies for integrating the levels in such multilevel mechanisms. These experimental strategies include activation strategies, interference strategies, and additive strategies. These strategies are mutually reinforcing, (...) providing a kind of interlevel and intratheoretic robustness that has not previously been recognized. (shrink)

We explore the contribution made by oscillatory, synchronous neural activity to representation in the brain. We closely examine six prominent examples of brain function in which neural oscillations play a central role, and identify two levels of involvement that these oscillations take in the emergence of representations: enabling (when oscillations help to establish a communication channel between sender and receiver, or are causally involved in triggering a representation) and properly representational (when oscillations are a constitutive part of the representation). We (...) show that even an idealized informational sender–receiver account of representation makes the representational status of oscillations a non-trivial matter, which depends on rather minute empirical details. (shrink)

The dominant neuroscientific theory of spatial memory is, like many theories in neuroscience, a multilevel description of a mechanism. The theory links the activities of molecules, cells, brain regions, and whole organisms into an integrated sketch of an explanation for the ability of organisms to navigate novel environments. Here I develop a taxonomy of interlevel experimental strategies for integrating the levels in such multilevel mechanisms. These experimental strategies include activation strategies, interference strategies, and additive strategies. These strategies are mutually reinforcing, (...) providing a kind of interlevel and intratheoretic robustness that has not previously been recognized. (shrink)

The Morris water maze has been put forward in the philosophy of neuroscience as an example of an experimental arrangement that may be used to delineate the cognitive faculty of spatial memory (e.g., Craver and Darden, Theory and method in the neurosciences, University of Pittsburgh Press, Pittsburgh, 2001; Craver, Explaining the brain: Mechanisms and the mosaic unity of neuroscience, Oxford University Press, Oxford, 2007). However, in the experimental and review literature on the water maze throughout the history of its use, (...) we encounter numerous responses to the question of “what” phenomenon it circumscribes ranging from cognitive functions (e.g., “spatial learning”, “spatial navigation”), to representational changes (e.g., “cognitive map formation”) to terms that appear to refer exclusively to observable changes in behavior (e.g., “water maze performance”). To date philosophical analyses of the water maze have not been directed at sorting out what phenomenon the device delineates nor the sources of the different answers to the question of what. I undertake both of these tasks in this paper. I begin with an analysis of Morris’s first published research study using the water maze and demonstrate that he emerged from it with an experimental learning paradigm that at best circumscribed a discrete set of observable changes in behavior. However, it delineated neither a discrete set of representational changes nor a discrete cognitive function. I cite this in combination with a reductionist-oriented research agenda in cellular and molecular neurobiology dating back to the 1980s as two sources of the lack of consistency across the history of the experimental and review literature as to what is under study in the water maze. (shrink)

Similarity-based cognition is commonplace. It occurs whenever an agent or system exploits the similarities that hold between two or more items—e.g., events, processes, objects, and so on—in order to perform some cognitive task. This kind of cognition is of special interest to cognitive neuroscientists. This paper explicates how similarity-based cognition can be understood through the lens of radical enactivism and why doing so has advantages over its representationalist rival, which posits the existence of structural representations or S-representations. Specifically, it is (...) argued that there are problems both with accounting for the content of S-representations and with understanding how neurally-based structural similarities can work as representations in guiding intelligent behavior. Finally, with these clarifications in place, it is revealed how radical enactivism can commit to an account of similarity-based cognition in its understanding of neurodynamics. (shrink)

Our perception of where touch occurs on our skin shapes our interactions with the world. Most accounts of cutaneous localisation emphasise spatial transformations from a skin-based reference frame into body-centred and external egocentric coordinates. We investigated another possible method of tactile localisation based on an intrinsic perception of ‘skin space’. The arrangement of cutaneous receptive fields (RFs) could allow one to track a stimulus as it moves across the skin, similarly to the way animals navigate using path integration. We applied (...) curved tactile motions to the hands of human volunteers. Participants identified the location midway between the start and end points of each motion path. Their bisection judgements were systematically biased towards the integrated motion path, consistent with the characteristic inward error that occurs in navigation by path integration. We thus showed that integration of continuous sensory inputs across several tactile RFs provides an intrinsic mechanism for spatial perception. (shrink)

Following Gilbert Ryle’s arguments, many philosophers took it for granted that someone knows how to do something just in case they have the ability to do it. Within the last couple decades, new intellectualists have challenged this longstanding anti-intellectualist assumption. Their central contention is that mere abilities aren’t on the same rational, epistemic level as know how. My goal is to intellectualize know how without over-intellectualizing it. Intelligent behavior is characteristically flexible or responsive to novelty, and the distinctive feature of (...) creatures who exhibit flexible behavior is their capacity to learn. As it turns out, Ryle already identified a core characteristic of learning shared widely across the animal kingdom from the lowly rat to the top athlete. Taking my cues from Ryle, I argue that know how is successful performance resulting from self-regulated abilities. To regulate an ability is to be disposed to adjust to error and respond to feedback. While regulating sophisticated human abilities often requires propositional knowledge, in many simpler cases it does not. I focus on the navigational know how of rats and honeybees. Although they possess know how, whether they possess propositional knowledge is at best an open question. (shrink)

One question of the bounds of cognition is that of which things have it. A scientifically relevant debate on this question must explain the persistent and selective use of psychological predicates to report findings throughout biology: for example, that neurons prefer, fruit flies and plants decide, and bacteria communicate linguistically. This paper argues that these claims should enjoy default literal interpretation. An epistemic consequence is that these findings can contribute directly to understanding the nature of psychological capacities.

In this essay, we examine the use of resting state fMRI data for psychological inferences. We argue that resting state studies hold the paired promises of discovering novel functional brain networks, and of avoiding some of the limitations of task-based fMRI. However, we argue that the very features of experimental design that enable resting state fMRI to support exploratory science also generate a novel confound. We argue that seemingly key features of resting state functional connectivity networks may be artefacts resulting (...) from sampling a ‘mixture distribution’ of diverse brain networks active at different times during the scan. We explore the consequences of this ‘mixture view’ for attempts to theorize about the cognitive or psychological functions of resting state networks, as well as the value of exploratory experiments. (shrink)

Cognitive science is unusual in that cognitive scientists have dramatic disagreements about the extension of their object of study, cognition. This paper defends a novel analysis of the scientific concept of cognition: that cognition is the sensitive management of an agent’s behavior. This analysis is “modular,” so that its extension varies depending on how one interprets certain of its constituent terms. I argue that these variations correspond to extant disagreements between cognitive scientists. This correspondence is evidence that the proposed analysis (...) models the contemporary understanding of cognition among scientists, without artificially resolving questions that are currently considered open. (shrink)

This paper offers a modeling account of episodic representation. I argue that the episodic system constructsmental models: representations that preserve the spatiotemporal structure of represented domains. In prototypical cases, these domains are events: occurrences taken by subjects to have characteristic structures, dynamics and relatively determinate beginnings and ends. Due to their simplicity and manipulability, mental event models can be used in a variety of cognitive contexts: in remembering the personal past, but also in future-oriented and counterfactual imagination. As structural representations, (...) they allow surrogative reasoning, supporting inferences about their constituents which can be used in reasoning about the represented events. (shrink)

Theories of superior collicular and hippocampal function have remarkable similarities. Both structures have been repeatedly implicated in spatial and attentional behaviour and in inhibitory control of locomotion. Moreover, they share certain electrophysiological properties in their single unit responses and in the synchronous appearance and disappearance of slow wave activity. Both are phylogenetically old and the colliculus projects strongly to brainstem nuclei instrumental in the generation of theta rhythm in the hippocampal EECOn the other hand, close inspection of behavioural and electrophysiological (...) data reveals disparities. In particular, hippocampal processing mainly concerns stimulus ambiguity, contextual significance, and spatial relations or other subtle, higher order characteristics. This requires the use of largely preprocessed sensory information and mediation of poststimulus investigation. Although collicular activity must also be integrated with that of “higher” centres, its primary role in attention is more “peripheral” and specific in controlling orienting/localisation via eye and body movements toward egocentrically labelled spatial positions. In addition, the colliculus may exert a nonspecific influence in alerting higher centres to the imminence of information potentially worthy of focal attention. Nevertheless, it is noteworthy that collicular and hippocampal lesions produce deficits on similar tasks, although the type of deficit is usually different in each case. Functional overlap between hippocampus and colliculus is virtually certain vis-à-vis stimulus sampling, for example in the acquisition of information via vibrissal movements and visual scanning. In addition, insofar as stimulus significance is a factor in collicular orienting mechanisms, the hippocampus — cingulate – cortex — colliculus pathway may play a significant role, modulating collicular responsiveness and thus ensuring an attentional strategy appropriate to current requirements. A tentative “reciprocal loop” model is proposed which bridges physiological and behavioural levels of analysis and which would account for the observed degree and nature of functional overlap between the superior colliculus and hippocampus. (shrink)

“The physics of representation” aims to define the word “representation” as used in the neurosciences, argue that such representations as described in neuroscience are related to and usefully illuminated by the representations generated by modern neural networks, and establish that these entities are “representations in good standing”. We suggest that Poldrack succeeds in, exposes some tensions between the broad use of the term in neuroscience and the narrower class of entities that he identifies in the end, and between the meaning (...) of “representation” in neuroscience and in psychology in, and fails in. This results in some hard choices: give up on the broad scope of the term in neuroscience or continue to embrace the broad, psychologically inflected sense of the term, and deny the entities generated by neural nets are representations in the relevant sense. (shrink)

Do animals require rich internal representations, such as cognitive maps, to navigate complex environments? Some researchers believe so, as they argue that sensory information is “too poor” to account for animals’ wayfinding abilities. However, this assumption is debatable, as James J. Gibson showed. Gibson proposed that wayfinding involves detecting information about environmental structure over time and used the concepts of “vistas” and “transitions” to explain terrestrial navigation. While these concepts may not apply universally to animal navigation, they highlight the importance (...) of exploiting stable environmental structures for wayfinding. By searching for species-relative environmental structures, we may gain insight into the navigational abilities of different nonhuman animals, while recognizing the unique evolutionary histories and ecological contexts that have shaped these abilities. (shrink)

This article argues that rapid eye movement (REM) dreaming is elaborative encoding for episodic memories. Elaborative encoding in REM can, at least partially, be understood through ancient art of memory (AAOM) principles: visualization, bizarre association, organization, narration, embodiment, and location. These principles render recent memories more distinctive through novel and meaningful association with emotionally salient, remote memories. The AAOM optimizes memory performance, suggesting that its principles may predict aspects of how episodic memory is configured in the brain. Integration and segregation (...) are fundamental organizing principles in the cerebral cortex. Episodic memory networks interconnect profusely within the cortex, creating omnidirectional “landmark” junctions. Memories may be integrated at junctions but segregated along connecting network paths that meet at junctions. Episodic junctions may be instantiated during non–rapid eye movement (NREM) sleep after hippocampal associational function during REM dreams. Hippocampal association involves relating, binding, and integrating episodic memories into a mnemonic compositional whole. This often bizarre, composite image has not been present to the senses; it is not “real” because it hyperassociates several memories. During REM sleep, on the phenomenological level, this composite image is experienced as a dream scene. A dream scene may be instantiated as omnidirectional neocortical junction and retained by the hippocampus as an index. On episodic memory retrieval, an external stimulus (or an internal representation) is matched by the hippocampus against its indices. One or more indices then reference the relevant neocortical junctions from which episodic memories can be retrieved. Episodic junctions reach a processing (rather than conscious) level during normal wake to enable retrieval. If this hypothesis is correct, the stuff of dreams is the stuff of memory. (shrink)

Robots are being extensively used for the purpose of discovering and testing empirical hypotheses about biological sensorimotor mechanisms. We examine here methodological problems that have to be addressed in order to design and perform “good” experiments with these machine models. These problems notably concern the mapping of biological mechanism descriptions into robotic mechanism descriptions; the distinction between theoretically unconstrained “implementation details” and robotic features that carry a modeling weight; the role of preliminary calibration experiments; the monitoring of experimental environments for (...) disturbing factors that affect both modeling features and theoretically unconstrained implementation details of robots. Various assumptions that are gradually introduced in the process of setting up and performing these robotic experiments become integral parts of the background hypotheses that are needed to bring experimental observations to bear on biological mechanism descriptions. (shrink)

Following Gilbert Ryle’s arguments, many philosophers took it for granted that someone knows how to do something just in case they have the ability to do it. Within the last couple decades, new intellectualists have challenged this longstanding anti-intellectualist assumption. Their central contention is that mere abilities aren’t on the same rational, epistemic level as know how. My goal is to intellectualize know how without over-intellectualizing it. Intelligent behavior is characteristically flexible or responsive to novelty, and the distinctive feature of (...) creatures who exhibit flexible behavior is their capacity to learn. As it turns out, Ryle already identified a core characteristic of learning shared widely across the animal kingdom from the lowly rat to the top athlete. Taking my cues from Ryle, I argue that know how is successful performance resulting from self-regulated abilities. To regulate an ability is to be disposed to adjust to error and respond to feedback. While regulating sophisticated human abilities often requires propositional knowledge, in many simpler cases it does not. I focus on the navigational know how of rats and honeybees. Although they possess know how, whether they possess propositional knowledge is at best an open question. (shrink)

We argue that a map is meaningless unless we have a process for using it. Thus, in this paper, we not only offer the world graph as a representation of relationships among situations the animal has encountered and may encounter again, but we also offer algorithms for how the information encoded in the world graph may be used by the animal in determining its behavior. Each node of the graph encodes a recognizable situation in the animal's world, but a given (...) place may well be encoded in a number of different nodes. Nodes not only require algorithms for the recognition of the situation; they store information about drive reduction associated with the encoded situation. We note that the use of graphs as a basis for exploring some search space is well known in artificial intelligence (AI), but we stress the importance of the animal's exploration of its environment for growing the graph, as distinct from the mathematically described potential nodes frequent in AI search spaces. To explore a number of hypotheses about the way information in the world graph is used to guide the animal's movement, we recall a number of classical experiments on maze exploration by animals, and use them to argue for the nonlocal hypothesis (selection of a path does not depend only upon information about the immediate environment of the animal) and the competing nodes hypothesis (more than one drive may enter into the determination of the animal's behavior at any time). An important feature of the model is that it yields exploration and latent learning without the introduction of an exploratory drive. We also note that the performance exhibited by the model appears to be state-dependent when the animal operates under high drive levels. The drive-interaction matrix is offered as a subject for future research.We complement the presentation of the world graph model, its drive dynamics, and how these are constrained by experiments on maze behavior, with a brief analysis of maps in the brain. We distinguish egocentric maps–which we relate to the many visual systems–from allocentric maps. We offer a somewhat unconventional view of short-term and long-term memory. We examine cooperative computation in somatotopically organized networks, relating this to visually guided behavior in the frog, and to the interaction of colliculus and cortex in the control of eye movements. We examine, but do not advocate, the hypothesis that the hippocampus is a cognitive map. We do stress that if it is a cognitive map, it must be seen as a chart of the local neighborhood, rather than the whole atlas; and we note that the cognitive map hypothesis would lead one to expect the region to exhibit activation of place cells before the animal leaves the previous place. (shrink)

What role does the concept of representation play in the contexts of experimentation and explanation in cognitive neurobiology? In this article, a distinction is drawn between minimal and substantive roles for representation. It is argued by appeal to a case study that representation currently plays a role in cognitive neurobiology somewhere in between minimal and substantive and that this is problematic given the ultimate explanatory goals of cognitive neurobiological research. It is suggested that what is needed is for representation to (...) instead play a more substantive role. (shrink)

In this paper, I evaluate recently defended mechanistic accounts of the unity of neuroscience from a metaphysical point of view. Considering the mechanistic framework in general , I argue that explanations of this kind are essentially reductive . The reductive character of mechanistic explanations provides a sufficiency criterion, according to which the mechanism underlying a certain phenomenon is sufficient for the latter. Thus, the concept of supervenience can be used in order to describe the relation between mechanisms and phenomena . (...) Against this background, I show that the mechanistic framework is subject to the causal exclusion problem and faces the classical metaphysical options when it comes to the relations obtaining between different levels of mechanisms . Finally, an attempt to improve the metaphysics of mechanisms is made and further difficulties are pointed out. (shrink)