The developmental trajectories of children's reorientation to global and local properties of environmental geometry.

The way in which organisms represent the shape of their environments during navigation has been debated in cognitive, comparative, and developmental psychology. While there is evidence that adult humans encode the entire boundary shape of an environment (a global-shape representation), there are also data demonstrating that organisms reorient using only segments of the boundary that signal a goal location (a local-shape representation). Developmental studies offer unique insights into this debate; however, most studies have used designs that cannot dissociate the type of boundary-shape representation that children use to guide reorientation. Thus, we examined the developmental trajectories of children's reorientation according to local and global boundary shape. Participants aged 6-12 years were trained to find a goal hidden in one corner of a virtual arena, after which they were required to reorient in a novel test arena. From 10.5 years, children performed above chance when the test arena permitted reorientation based only on local-shape (Experiment 2), or only global-shape (Experiment 3) information. Moreover, when these responses were placed into conflict, older children reoriented with respect to global-shape information (Experiment 4). These age-related findings were not due to older children being better able to reorient in virtual environments per se: when trained and tested within the same environment (Experiment 1), children performed above chance from 6 years. Together, our results suggest (a) the ability to reorient on the basis of global- and local-shape representations develops in parallel, and (b) shape-based information is weighted to determine which representation informs reorientation. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

The spatial layout of doorways and environmental boundaries shape the content of event memories.

Physical boundaries in our environment have been observed to define separate events in episodic memory. To date, however, there is little evidence that the spatial properties of boundaries exert any control over event memories. To examine this possibility, we conducted four experiments that took manipulations involving boundaries that have been demonstrated to influence spatial representations, and adapted them for use in an episodic object memory paradigm. Here, participants were given 15 min to freely explore an environment that contained 36 objects, equally dispersed among six discriminable buildings. In a subsequent test of object-location binding, participants were required to indicate where they remembered encountering the objects. In Experiment 1 the spatial properties of the building boundaries were identical; however, in Experiment 2 the boundaries were differentiated by their geometric shape and the location of the doorways in the buildings. In the test phases of these experiments, we observed a shift from a bias towards remembering the positions of objects within a building but not the building itself (Experiment 1), to a bias towards remembering which building an object was in but not the location within the building (Experiment 2). In Experiment 3, the buildings shared the same geometry but were differentiated by the locations of doorways, and we observed no significant differences between response types. Finally, in Experiment 4, the buildings were uniquely shaped but shared the same doorway location, and we observed a bias towards remembering the positions of objects within a building. In addition, exploratory analyses of non-spatial interference revealed more correct recall for objects housed in the first building a participant visited during exploration, compared to all other buildings. Together, our data indicates that the location of doorways in boundaries and, to a lesser extent, boundary geometries influence event models, and that a primacy effect can be observed in the recall of multiple object-location bindings.

Temporal and spatial contiguity are necessary for competition between events.

Over the last 50 years, cue competition phenomena have shaped theoretical developments in animal and human learning. However, recent failures to observe competition effects in standard conditioning procedures, as well as the lengthy and ongoing debate surrounding cue competition in the spatial learning literature, have cast doubts on the generality of these phenomena. In the present study, we manipulated temporal contiguity between simultaneously trained predictors and outcomes (Experiments 1-4), and spatial contiguity between landmarks and goals in spatial learning (Supplemental Experiments 1 and 2; Experiment 5). Across different parametric variations, we observed overshadowing when temporal and spatial contiguity were strong, but no overshadowing when contiguity was weak. Thus, across temporal and spatial domains, we observed that contiguity is necessary for competition to occur, and that competition between cues presented simultaneously during learning is absent when these cues were either spatially or temporally discontiguous from the outcome. Consequently, we advance a model in which the contiguity between events is accounted for and which explains these results and reconciles the previously contradictory findings observed in spatial learning. (PsycInfo Database Record (c) 2022 APA, all rights reserved).

The effects of spatial stability and cue type on spatial learning: Implications for theories of parallel memory systems.

Some theories of spatial learning predict that associative rules apply under only limited circumstances. For example, learning based on a boundary has been claimed to be immune to cue competition effects because boundary information is the basis for the formation of a cognitive map, whilst landmark learning does not involve cognitive mapping. This is referred to as the cue type hypothesis. However, it has also been claimed that cue stability is a prerequisite for the formation of a cognitive map, meaning that whichever cue type was perceived as stable would enter a cognitive map and thus be immune to cue competition, while unstable cues will be subject to cue competition, regardless of cue type. In experiments 1 and 2 we manipulated the stability of boundary and landmark cues when learning the location of two hidden goals. One goal location was constant with respect to the boundary, and the other constant with respect to the landmark cues. For both cue types, the presence of distal orientation cues provided directional information. For half the participants the landmark cues were unstable relative to the boundary and orientation cues, whereas for the remainder of the participants the boundary was unstable relative to landmarks and orientation cues. In a second stage of training, all cues remained stable so that both goal locations could be learned with respect to both landmark and boundary information. According to the cue type hypothesis, boundary information should block learning about landmarks regardless of cue stability. According to the cue stability hypothesis, however, landmarks should block learning about the boundary when the landmarks appear stable relative to the boundary. Regardless of cue type or stability the results showed reciprocal blocking, contrary to both formulations of incidental cognitive mapping. Experiment 3 established that the results of Experiments 1 and 2 could not be explained in terms of difficulty in learning certain locations with respect to different cue types. In a final experiment, following training in which both landmarks and boundary cues signalled two goal locations, a new goal location was established with respect to the landmark cues, before testing with the boundary, which had never been used to define the new goal location. The results of this novel test of the interaction between boundary and landmark cues indicated that new learning with respect to the landmark had a profound effect on navigation with respect to the boundary, counter to the predictions of incidental cognitive mapping of boundaries.

Individual differences in theta-band oscillations in a spatial memory network revealed by electroencephalography predict rapid place learning.

Spatial memory has been closely related to the medial temporal lobe and theta oscillations are thought to play a key role. However, it remains difficult to investigate medial temporal lobe activation related to spatial memory with non-invasive electrophysiological methods in humans. Here, we combined the virtual delayed-matching-to-place task, reverse-translated from the watermaze delayed-matching-to-place task in rats, with high-density electroencephalography recordings. Healthy young volunteers performed this computerised task in a virtual circular arena, which contained a hidden target whose location moved to a new place every four trials, allowing the assessment of rapid memory formation. Using behavioural measures as predictor variables for source reconstructed frequency-specific electroencephalography power, we found that inter-individual differences in 'search preference' during 'probe trials', a measure of one-trial place learning known from rodent studies to be particularly hippocampus-dependent, correlated predominantly with distinct theta-band oscillations (approximately 7 Hz), particularly in the right temporal lobe, the right striatum and inferior occipital cortex or cerebellum. This pattern was found during both encoding and retrieval/expression, but not in control analyses and could not be explained by motor confounds. Alpha-activity in sensorimotor and parietal cortex contralateral to the hand used for navigation also correlated (inversely) with search preference. This latter finding likely reflects movement-related factors associated with task performance, as well as a frequency difference in (ongoing) alpha-rhythm for high-performers versus low-performers that may contribute to these results indirectly. Relating inter-individual differences in ongoing brain activity to behaviour in a continuous rapid place-learning task that is suitable for a variety of populations, we could demonstrate that memory-related theta-band activity in temporal lobe can be measured with electroencephalography recordings. This approach holds great potential for further studies investigating the interactions within this network during encoding and retrieval, as well as neuromodulatory impacts and age-related changes.

Crossing boundaries: Global reorientation following transfer from the inside to the outside of an arena.

In 2 spatial navigation experiments, human participants were asked to find a hidden goal (a WiFi signal) that was located in 1 of the right-angled corners of a kite-shaped (Experiment 1) or a cross-shaped (Experiment 2) virtual environment. Goal location was defined solely with respect to the geometry of the environment. Following this training, in a test conducted in extinction, participants were placed onto the outside of the same environments and asked to locate the WiFi signal. The results of both experiments revealed that participants spent more time searching in regions on the outside of the environments that were closest to where the WiFi signal was located during training. These results are difficult to explain in terms of analyses of spatial navigation and reorientation that emphasize the role of local representational encoding or view matching. Instead, we suggest that these results are better understood in terms of a global representation of the shape of the environment. (PsycINFO Database Record (c) 2019 APA, all rights reserved).

Walking through doorways differentially affects recall and familiarity.

Previous research has reported that walking through a doorway to a new location makes memory for objects and events experienced in the previous location less accurate. This effect, termed the location updating effect, has been used to suggest that location changes are used to mark boundaries between events in memory: memories for objects encountered within the current event are more available than those from beyond an event boundary. Within a computer-generated memory task, participants navigated through virtual rooms, walking through doorways, and interacting with objects. The accuracy and their subjective experience of their memory for the objects (remember/know and confidence) were assessed. The findings showed that shifts in location decreased accurate responses associated with the subjective experience of remembering but not those associated with the experience of knowing, even when considering only the most confident responses in each condition. These findings demonstrate that a shift in location selectively impacts recollection and so contributes to our understanding of boundaries in event memory.

Thinking outside of the box II: Disrupting the cognitive map.

A number of influential spatial learning theories posit that organisms encode a viewpoint independent (i.e. allocentric) representation of the global boundary shape of their environment in order to support spatial reorientation and place learning. In contrast to the trial and error learning mechanisms that support domain-general processes, a representation of the global-shape of the environment is thought to be encoded automatically as part of a cognitive map, and without interference from other spatial cues. To date, however, this core theoretical assumption has not been appropriately examined. This is because previous attempts to address this question have failed to employ tasks that fully dissociate reorientation based on an allocentric representation of global-shape from egocentric reorientation strategies. Here, we address this issue in two experiments. Participants were trained to navigate to a hidden goal on one side of a virtual arena (e.g. the inside) before being required to find the same point on the alternative side (e.g. the outside). At test, performing the correct search behaviour requires an allocentric representation of the global boundary-shape. Using established associative learning procedures of overshadowing and blocking, we find that search behaviour at test is disrupted when participants were able to form landmark-goal associations during training. These results demonstrate that encoding of an allocentric representation of boundary information is susceptible to interference from landmark cues, and is not acquired through special means. Instead, the results suggest that allocentric representations of environmental boundaries are acquired through the same kind of error-correction mechanisms that support domain-general non-spatial learning.

A new human delayed-matching-to-place test in a virtual environment reverse-translated from the rodent watermaze paradigm: Characterization of performance measures and sex differences.

Watermaze tests of place learning and memory in rodents and corresponding reverse-translated human paradigms in real or virtual environments are key tools to study hippocampal function. In common variants, the animal or human participant has to find a hidden goal that remains in the same place over many trials, allowing for incremental learning of the place with reference to distal cues surrounding the circular, featureless maze. Although the hippocampus is involved in incremental place learning, rodent studies have shown that the delayed-matching-to-place (DMP) watermaze test is a more sensitive assay of hippocampal function. On the DMP test, the goal location changes every four trials, requiring the rapid updating of place memory. Here, we developed a virtual DMP test reverse-translated from the rat watermaze DMP paradigm. In two replications, participants showed 1-trial place learning, evidenced by marked latency and path length savings between Trials 1 and 2 to the same goal location, and by search preference for the vicinity of the goal when Trial 2 was run as probe trial (during which the goal was removed). The performance was remarkably similar to rats' performance on the watermaze DMP test. In both replications, male participants showed greater savings and search preferences compared to female participants. Male participants also showed better mental rotation performance, although mental rotation scores did not consistently correlate with DMP performance measures, pointing to distinct neurocognitive mechanisms. The remarkable similarity between rodent and human DMP performance suggests similar underlying neuro-psychological mechanisms, including hippocampus dependence. The new virtual DMP test may, therefore, provide a sensitive tool to probe human hippocampal function.

Thinking outside of the box: Transfer of shape-based reorientation across the boundary of an arena.

The way in which human and non-human animals represent the shape of their environments remains a contentious issue. According to local theories of shape learning, organisms encode the local geometric features of the environment that signal a goal location. In contrast, global theories of shape learning suggest that organisms encode the overall shape of the environment. There is, however, a surprising lack of evidence to support this latter claim, despite the fact that common behaviours seem to require encoding of the global-shape of an environment. We tested one such behaviour in 5 experiments, in which human participants were trained to navigate to a hidden goal on one side of a virtual arena (e.g. the inside) before being required to find the same point on the alternative side (e.g. the outside). Participants navigated to the appropriate goal location, both when inside and outside the virtual arena, but only when the shape of the arena remained the same between training and test (Experiments 1a and 1b). When the arena shape was transformed between these stages, participants were lost (Experiments 2a and 2b). When training and testing was conducted on the outside of two different-shaped arenas that shared local geometric cues participants once again explored the appropriate goal location (Experiment 3). These results provide core evidence that humans encode a global representation of the overall shape of the environments in, or around, which they navigate.

Blocking spatial navigation across environments that have a different shape.

According to the geometric module hypothesis, organisms encode a global representation of the space in which they navigate, and this representation is not prone to interference from other cues. A number of studies, however, have shown that both human and non-human animals can navigate on the basis of local geometric cues provided by the shape of an environment. According to the model of spatial learning proposed by Miller and Shettleworth (2007, 2008), geometric cues compete for associative strength in the same manner as non-geometric cues do. The experiments reported here were designed to test if humans learn about local geometric cues in a manner consistent with the Miller-Shettleworth model. Experiment 1 replicated previous findings that humans transfer navigational behavior, based on local geometric cues, from a rectangle-shaped environment to a kite-shaped environment, and vice versa. In Experiments 2 and 3, it was observed that learning about non-geometric cues blocked, and were blocked by, learning about local geometric cues. The reciprocal blocking observed is consistent with associative theories of spatial learning; however, it is difficult to explain the observed effects with theories of global-shape encoding in their current form.

The developmental trajectory of intramaze and extramaze landmark biases in spatial navigation: An unexpected journey.

Adults learning to navigate to a hidden goal within an enclosed space have been found to prefer information provided by the distal cues of an environment, as opposed to proximal landmarks within the environment. Studies with children, however, have shown that 5- or 7-year-olds do not display any preference toward distal or proximal cues during navigation. This suggests that a bias toward learning about distal cues occurs somewhere between the age of 7 years and adulthood. We recruited 5- to 11-year-old children and an adult sample to explore the developmental profile of this putative change. Across a series of 3 experiments, participants were required to navigate to a hidden goal in a virtual environment, the location of which was signaled by both extramaze and intramaze landmark cues. During testing, these cues were placed into conflict to assess the search preferences of participants. Consistent with previously reported findings, adults were biased toward using extramaze information. However, analysis of the data from children, which incorporated age as a continuous variable, suggested that older children in our sample were, in fact, biased toward using the intramaze landmark in our task. These findings suggest the bias toward using distal cues in spatial navigation, frequently displayed by adults, may be a comparatively late developing trait, and one that could supersede an initial developmental preference for proximal landmarks.

Learned predictiveness training modulates biases towards using boundary or landmark cues during navigation.

A number of navigational theories state that learning about landmark information should not interfere with learning about shape information provided by the boundary walls of an environment. A common test of such theories has been to assess whether landmark information will overshadow, or restrict, learning about shape information. Whilst a number of studies have shown that landmarks are not able to overshadow learning about shape information, some have shown that landmarks can, in fact, overshadow learning about shape information. Given the continued importance of theories that grant the shape information that is provided by the boundary of an environment a special status during learning, the experiments presented here were designed to assess whether the relative salience of shape and landmark information could account for the discrepant results of overshadowing studies. In Experiment 1, participants were first trained that either the landmarks within an arena (landmark-relevant), or the shape information provided by the boundary walls of an arena (shape-relevant), were relevant to finding a hidden goal. In a subsequent stage, when novel landmark and shape information were made relevant to finding the hidden goal, landmarks dominated behaviour for those given landmark-relevant training, whereas shape information dominated behaviour for those given shape-relevant training. Experiment 2, which was conducted without prior relevance training, revealed that the landmark cues, unconditionally, dominated behaviour in our task. The results of the present experiments, and the conflicting results from previous overshadowing experiments, are explained in terms of associative models that incorporate an attention variant.

Shape shifting: Local landmarks interfere with navigation by, and recognition of, global shape.

An influential theory of spatial navigation states that the boundary shape of an environment is preferentially encoded over and above other spatial cues, such that it is impervious to interference from alternative sources of information. We explored this claim with 3 intradimensional-extradimensional shift experiments, designed to examine the interaction of landmark and geometric features of the environment in a virtual navigation task. In Experiments 1 and 2, participants were first required to find a hidden goal using information provided by the shape of the arena or landmarks integrated into the arena boundary (Experiment 1) or within the arena itself (Experiment 2). Participants were then transferred to a different-shaped arena that contained novel landmarks and were again required to find a hidden goal. In both experiments, participants who were navigating on the basis of cues that were from the same dimension that was previously relevant (intradimensional shift) learned to find the goal significantly faster than participants who were navigating on the basis of cues that were from a dimension that was previously irrelevant (extradimensional shift). This suggests that shape information does not hold special status when learning about an environment. Experiment 3 replicated Experiment 2 and also assessed participants' recognition of the global shape of the navigated arenas. Recognition was attenuated when landmarks were relevant to navigation throughout the experiment. The results of these experiments are discussed in terms of associative and non-associative theories of spatial learning.

Evidence for spatial navigational impairments in hydrocephalus patients without spina bifida.

The cognitive sequelae of hydrocephalus have mostly been explored with standardised clinical tasks. The aim of the present research was determine whether impairments on these abstract tasks extend to everyday spatial and navigational behaviour. Patients with hydrocephalus, but without spina bifida, were compared to a control group on tests of searching behaviour, landmark memory, route learning, and path integration. Participants with hydrocephalus displayed reduced sensitivity to spatial cueing, less accurate route-learning, and significantly less accurate spatial updating. These data represent an important empirical demonstration of spatial navigational impairments due to hydrocephalus outside of the context of spina bifida. We discuss some of the cognitive, neural, and individual differences factors that might contribute to this particular pattern of impairments.

Spatial navigational impairments in hydrocephalus.

Whilst much is known about the neuropathological consequences of hydrocephalus, there have been comparatively few studies of the cognitive impairments associated with it. Studies using standardised tests of cognitive function have identified a general pattern of impairments, with patients exhibiting particular difficulty on tests of spatial memory and executive function. A strong prediction is that these deficits are likely to affect daily wayfinding behaviour, and we report a study of spatial and navigational abilities in a group of patients with hydrocephalus but without spina bifida. Participants completed a range of experimental tasks assessing spatial cueing behaviour, landmark memory and route-learning, and idiothetic path integration. This patient group was compared to a control sample matched on verbal, spatial, and intelligence measures, and hydrocephalus was found to be associated with relative impairments in each of the tasks. Patients exhibited reduced sensitivity to spatial cueing, less accurate route-learning, poorer memory for landmark objects, and less accurate spatial updating (with particular impairments in the calculation of heading). Overall, these data represent the first empirical demonstration of navigational impairments in hydrocephalus, and we suggest some of the cognitive, neural, and individual differences factors that may contribute to the pattern of performance reported.