Navigating to goal locations in a known environment (wayfinding) can be accomplished by different strategies, notably by taking habitual, well-learned routes (response strategy) or by inferring novel paths, such as shortcuts, from spatial knowledge of the environment's layout (place strategy). Human and animal neuroscience studies reveal that these strategies reflect different brain systems, with response strategies relying more on activation of the striatum and place strategies associated with activation of the hippocampus. In addition to individual differences in strategy, recent behavioral (...) studies show sex differences such that men use place strategies more than women, and age differences such that older adults use more response strategies than younger adults. This paper takes a comprehensive multilevel approach to understanding these differences, characterizing wayfinding as a complex information processing task. This analysis reveals factors that affect navigation strategy, including availability of the relevant type of environmental knowledge, momentary access to this knowledge, trade-offs between physical and mental effort in different navigation contexts, and risk taking. We consider how strategies are influenced by the computational demands of a navigation task and by factors that affect the neural circuits underlying navigation. We also discuss limitations of laboratory studies to date and outline priorities for future research, including relating wayfinding strategies to independent measures of spatial knowledge, and studying wayfinding strategies in naturalistic environments. (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)

Event concepts are unstructured atomic concepts that apply to event types. A paradigm example of such an event type would be that of diaper changing, and so a putative example of an atomic event concept would be DADDY'S-CHANGING-MY-DIAPER.1 I will defend two claims about such concepts. First, the conceptual claim that it is in principle possible to possess a concept such as DADDY'S-CHANGING-MY-DIAPER without possessing the concept DIAPER. Second, the empirical claim that we actually possess such concepts and that they (...) play an important role in our cognitive lives. The argument for the empirical claim has the form of an inference to the best explanation and is aimed at those who are already willing to attribute concepts and beliefs to infants and nonhuman animals. Many animals and prelinguistic infants seem capable of re-identifying event-types in the world, and they seem to store information about things happening at particular times and places. My account offers a plausible model of how such organisms are able to do this without attributing linguistically structured mental states to them. And although language allows adults to form linguistically structured mental representations of the world, there is no good reason to think that such structured representations necessarily replace the unstructured ones. There is also no good reason for a philosopher who is willing to explain the behavior of an organism by appealing to atomic concepts of individuals or kinds to not use a similar form of explanation when explaining the organism's capacity to recognize events. -/- We can form empirical concepts of individuals, kinds, properties, event-types, and states of affairs, among other things, and I assume that such concepts function like what François Recanati calls ‘mental files’ or what Ruth Millikan calls ‘substance concepts’ (Recanati 2012; Millikan 1999, 2000, 2017). To possess such a concept one must have a reliable capacity to re-identify the object in question, but this capacity of re-identification does not fix the reference of the concept. Such concepts allow us to collect and utilize useful information about things that we re-encounter in our environment. We can distinguish between a perception-action system and a perception-belief system, and I will argue that empirical concepts, including atomic event concepts, can play a role in both systems. The perception-action system involves the application of concepts in the service of (often skilled) action. We can think of the concept as a mental file containing motor-plans that can be activated once the individual recognizes that they are in a certain situation. In this way, recognizing something (whether an object or an event) as a token of a type, plays a role in guiding immediate action. The perception-belief system, in contrast, allows for the formation of beliefs that can play a role in deliberation and planning and in the formation of expectations. I distinguish between two particular types of belief which I call where-beliefs and when-beliefs, and I argue that we can model the formation of such perceptual beliefs in nonlinguistic animals and human infants in terms of the formation of a link between an empirical concept and a position on a cognitive map. According to the account offered, seemingly complex beliefs, such as a baby's belief that Daddy changed her diaper in the kitchen earlier, will not be linguistically structured. If we think that prelinguistic infants possess such concepts and are able to form such beliefs, it is likely that adults do too. The ability to form such beliefs does not require the capacity for public language, and we can model them in nonlinguistic terms; thus, we have no good reason to think of such beliefs as propositional attitudes. Of course, we can use sentences to refer to such beliefs, and thus it is possible to think of such beliefs as somehow relations to propositions. But it is not clear to me what is gained by this as we have a perfectly good way to think about the structure of such beliefs that does not involve any appeal to language. (shrink)

In this commentary on Bastin et al., we suggest that spatial context plays a critical role in the encoding and retrieval of events. Specifically, the translation process between the viewpoint-independent content of a memory and the viewpoint-dependent stimuli activating the retrieval plays a critical role in spatial memory recollection. This perspective also provides an explanatory model for pathological disturbances such as Alzheimer's disease.

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)

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)

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)

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)

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)

We propose that rapid eye movement (REM) and slow-wave sleep contribute differently to the formation of episodic memories. REM sleep is important for building up invariant object representations that eventually recur to gamma-band oscillations in the neocortex. In contrast, slow-wave sleep is more directly involved in the consolidation of episodic memories through replay of sequential neural activity in hippocampal place cells.

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)

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)

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)

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)

A model of the neuropsychology of anxiety is proposed. The model is based in the first instance upon an analysis of the behavioural effects of the antianxiety drugs in animals. From such psychopharmacologi-cal experiments the concept of a “behavioural inhibition system” has been developed. This system responds to novel stimuli or to those associated with punishment or nonreward by inhibiting ongoing behaviour and increasing arousal and attention to the environment. It is activity in the BIS that constitutes anxiety and that (...) is reduced by antianxiety drugs. The effects of the antianxiety drugs in the brain also suggest hypotheses concerning the neural substrate of anxiety. Although the benzodiazepines and barbiturates facilitate the effects of γ-aminobutyrate, this is insufficient to explain their highly specific behavioural effects. Because of similarities between the behavioural effects of certain lesions and those of the antianxiety drugs, it is proposed that these drugs reduce anxiety by impairing the functioning of a widespread neural system including the septo-hippocampal system, the Papez circuit, the prefrontal cortex, and ascending monoaminergic and cholinergic pathways which innervate these forebrain structures. Analysis of the functions of this system suggests that it acts as a comparator: it compares predicted to actual sensory events and activates the outputs of the BIS when there is a mismatch or when the predicted event is aversive. Suggestions are made as to the functions of particular pathways within this overall brain system. The resulting theory is applied to the symptoms and treatment of anxiety in man, its relations to depression, and the personality of individuals who are susceptible to anxiety or depression. (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)

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)