Interplay of wayfinding strategies in route repetition and route retracing.

This study explores the interplay of navigation strategies in route repetition (repeating a recently travelled route) and route retracing (returning to the start location of a recently travelled route). Specifically, we investigated how sequence knowledge contributes to route repetition and retracing. In the learning phase, participants passively transported along a route. In the test phase, they were then asked to repeat or retrace the route. Decision points were either presented in an order coherent with the learning phase (from start to destination in route repetition, or from destination to start in route retracing), or in a randomised order. As expected, participants performed better in route repetition than in route retracing. Performance declined when intersections were presented in a randomised order indicating that sequence knowledge contributed to route repetition and route retracing. Presenting intersections in an order coherent with learning boosted performance specifically on the first part of the route during route repetition. This effect was not observed during route retracing. These results show that sequence knowledge is utilised differently during route repetition and retracing. We argue that participants use a "sequence of turns" strategy alongside associating landmarks with direction changes during route repetition, and that it is unlikely that route retracing relies on the same type of sequence knowledge. Instead, we believe route retracing utilises knowledge about the sequence in which decision points are encountered. Overall, the findings highlight a complex interplay of different strategies in route repetition and retracing, shedding light on how navigators utilise sequence knowledge for effective navigation.

Biases in object location estimation: The role of rotations and translation.

Spatial memory studies often employ static images depicting a scene, an array of objects, or environmental features from one perspective and then following a perspective-shift-prompt memory either of the scene or objects within the scene. The current study investigated a previously reported systematic bias in spatial memory where, following a perspective shift from encoding to recall, participants indicated the location of an object farther to the direction of the shift. In Experiment 1, we aimed to replicate this bias by asking participants to encode the location of an object in a virtual room and then indicate it from memory following a perspective shift induced by camera translation and rotation. In Experiment 2, we decoupled the influence of camera translations and rotations and examined whether adding additional objects to the virtual room would reduce the bias. Overall, our results indicate that camera translations result in greater systematic bias than camera rotations. We propose that the accurate representation of camera translations requires more demanding mental computations than camera rotations, leading to greater uncertainty regarding the location of an object in memory. This uncertainty causes people to rely on an egocentric anchor, thereby giving rise to the systematic bias in the direction of camera translation.

The Relative Contribution of Executive Functions and Aging on Attentional Control During Road Crossing.

As we age, many physical, perceptual and cognitive abilities decline, which can critically impact our day-to-day lives. However, the decline of many abilities is concurrent; thus, it is challenging to disentangle the relative contributions of different abilities in the performance deterioration in realistic tasks, such as road crossing, with age. Research into road crossing has shown that aging and a decline in executive functioning (EFs) is associated with altered information sampling and less safe crossing decisions compared to younger adults. However, in these studies declines in age and EFs were confounded. Therefore, it is impossible to disentangle whether age-related declines in EFs impact on visual sampling and road-crossing performance, or whether visual exploration, and road-crossing performance, are impacted by aging independently of a decline in EFs. In this study, we recruited older adults with maintained EFs to isolate the impacts of aging independently of a decline EFs on road crossing abilities. We recorded eye movements of younger adults and older adults while they watched videos of road traffic and were asked to decide when they could cross the road. Overall, our results show that older adults with maintained EFs sample visual information and make similar road crossing decisions to younger adults. Our findings also reveal that both environmental constraints and EF abilities interact with aging to influence how the road-crossing task is performed. Our findings suggest that older pedestrians' safety, and independence in day-to-day life, can be improved through a limitation of scene complexity and a preservation of EF abilities.

Different Profiles of Spatial Navigation Deficits In Alzheimer's Disease Biomarker-Positive Versus Biomarker-Negative Older Adults With Amnestic Mild Cognitive Impairment.

Background: Spatial navigation impairment is a promising cognitive marker of Alzheimer's disease (AD) that can reflect the underlying pathology. Objectives: We assessed spatial navigation performance in AD biomarker positive older adults with amnestic mild cognitive impairment (AD aMCI) vs. those AD biomarker negative (non-AD aMCI), and examined associations between navigation performance, MRI measures of brain atrophy, and cerebrospinal fluid (CSF) biomarkers. Methods: A total of 122 participants with AD aMCI (n = 33), non-AD aMCI (n = 31), mild AD dementia (n = 28), and 30 cognitively normal older adults (CN) underwent cognitive assessment, brain MRI (n = 100 had high-quality images for volumetric analysis) and three virtual navigation tasks focused on route learning (body-centered navigation), wayfinding (world-centered navigation) and perspective taking/wayfinding. Cognitively impaired participants underwent CSF biomarker assessment [amyloid-ß1-42, total tau, and phosphorylated tau181 (p-tau181)] and amyloid PET imaging (n = 47 and n = 45, respectively), with a subset having both (n = 19). Results: In route learning, AD aMCI performed worse than non-AD aMCI (p < 0.001), who performed similarly to CN. In wayfinding, aMCI participants performed worse than CN (both p ≤ 0.009) and AD aMCI performed worse than non-AD aMCI in the second task session (p = 0.032). In perspective taking/wayfinding, aMCI participants performed worse than CN (both p ≤ 0.001). AD aMCI and non-AD aMCI did not differ in conventional cognitive tests. Route learning was associated with parietal thickness and amyloid-ß1-42, wayfinding was associated with posterior medial temporal lobe (MTL) volume and p-tau181 and perspective taking/wayfinding was correlated with MRI measures of several brain regions and all CSF biomarkers. Conclusion: AD biomarker positive and negative older adults with aMCI had different profiles of spatial navigation deficits that were associated with posterior MTL and parietal atrophy and reflected AD pathology.

The role of memory and perspective shifts in systematic biases during object location estimation.

In the current study, we investigated whether the introduction of perspective shifts in a spatial memory task results in systematic biases in object location estimations. To do so, we asked participants to first encode the position of an object in a virtual room and then to report its position from memory or perception following a perspective shift. Overall, our results showed that participants made systematic errors in estimating object positions in the same direction as the perspective shift. Notably, this bias was present in both memory and perception conditions. We propose that the observed systematic bias was driven by difficulties in understanding the perspective shifts that led participants to use an egocentric representation of object positions as an anchor when estimating the object location following a perspective shift.

Age-related changes in visual encoding strategy preferences during a spatial memory task.

Ageing is associated with declines in spatial memory, however, the source of these deficits remains unclear. Here we used eye-tracking to investigate age-related differences in spatial encoding strategies and the cognitive processes underlying the age-related deficits in spatial memory tasks. To do so we asked young and older participants to encode the locations of objects in a virtual room shown as a picture on a computer screen. The availability and utility of room-based landmarks were manipulated by removing landmarks, presenting identical landmarks rendering them uninformative, or by presenting unique landmarks that could be used to encode object locations. In the test phase, participants viewed a second picture of the same room taken from the same (0°) or a different perspective (30°) and judged whether the objects occupied the same or different locations in the room. We found that the introduction of a perspective shift and swapping of objects between encoding and testing impaired performance in both age groups. Furthermore, our results revealed that although older adults performed the task as well as younger participants, they relied on different visual encoding strategies to solve the task. Specifically, gaze analysis revealed that older adults showed a greater preference towards a more categorical encoding strategy in which they formed relationships between objects and landmarks.

London taxi drivers: A review of neurocognitive studies and an exploration of how they build their cognitive map of London.

Licensed London taxi drivers have been found to show changes in the gray matter density of their hippocampus over the course of training and decades of navigation in London (UK). This has been linked to their learning and using of the "Knowledge of London," the names and layout of over 26,000 streets and thousands of points of interest in London. Here we review past behavioral and neuroimaging studies of London taxi drivers, covering the structural differences in hippocampal gray matter density and brain dynamics associated with navigating London. We examine the process by which they learn the layout of London, detailing the key learning steps: systematic study of maps, travel on selected overlapping routes, the mental visualization of places and the optimal use of subgoals. Our analysis provides the first map of the street network covered by the routes used to learn the network, allowing insight into where there are gaps in this network. The methods described could be widely applied to aid spatial learning in the general population and may provide insights for artificial intelligence systems to efficiently learn new environments.

Spatial Navigation and Visuospatial Strategies in Typical and Atypical Aging.

Age-related spatial navigation decline is more pronounced in patients with mild cognitive impairment (MCI) and Alzheimer's disease (AD) dementia. We used a realistic-looking virtual navigation test suite to analyze different aspects of visuospatial processing in typical and atypical aging. A total of 219 older adults were recruited from the Czech Brain Aging Study cohort. Cognitively normal older adults (CN; n = 78), patients with amnestic MCI (n = 75), and those with mild AD dementia (n = 66) underwent three navigational tasks, cognitive assessment, and brain MRI. Route learning and wayfinding/perspective-taking tasks distinguished the groups as performance and learning declined and specific visuospatial strategies were less utilized with increasing cognitive impairment. Increased perspective shift and utilization of non-specific strategies were associated with worse task performance across the groups. Primacy and recency effects were observed across the groups in the route learning and the wayfinding/perspective-taking task, respectively. In addition, a primacy effect was present in the wayfinding/perspective-taking task in the CN older adults. More effective spatial navigation was associated with better memory and executive functions. The results demonstrate that a realistic and ecologically valid spatial navigation test suite can reveal different aspects of visuospatial processing in typical and atypical aging.

Ageing- and dementia-friendly design: theory and evidence from cognitive psychology, neuropsychology and environmental psychology can contribute to design guidelines that minimise spatial disorientation.

Many older people, both with and without dementia, eventually move from their familiar home environments into unfamiliar surroundings, such as sheltered housing or care homes. Age-related declines in wayfinding skills can make it difficult to learn to navigate in these new, unfamiliar environments. To facilitate the transition to their new accommodation, it is therefore important to develop retirement complexes and care homes specifically designed to reduce the wayfinding difficulties of older people and those with Alzheimer's disease (AD). Residential complexes that are designed to support spatial orientation and that compensate for impaired navigation abilities would make it easier for people with dementia to adapt to their new living environment. This would improve the independence, quality of life and well-being of residents, and reduce the caregivers' workload. Based on these premises, this opinion paper considers how evidence from cognitive psychology, neuropsychology and environmental psychology can contribute to ageing- and dementia-friendly design with a view to minimising spatial disorientation. After an introduction of the cognitive mechanisms and processes involved in spatial navigation, and the changes that occur in typical and atypical ageing, research from the field of environmental psychology is considered, highlighting design factors likely to facilitate (or impair) indoor wayfinding in complex buildings. Finally, psychological theories and design knowledge are combined to suggest ageing- and dementia-friendly design guidelines that aim to minimise spatial disorientation by focusing on residual navigation skills.

Perspective taking and systematic biases in object location memory.

The aim of the current study was to develop a novel task that allows for the quick assessment of spatial memory precision with minimal technical and training requirements. In this task, participants memorized the position of an object in a virtual room and then judged from a different perspective, whether the object has moved to the left or to the right. Results revealed that participants exhibited a systematic bias in their responses that we termed the reversed congruency effect. Specifically, they performed worse when the camera and the object moved in the same direction than when they moved in opposite directions. Notably, participants responded correctly in almost 100% of the incongruent trials, regardless of the distance by which the object was displaced. In Experiment 2, we showed that this effect cannot be explained by the movement of the object on the screen, but that it relates to the perspective shift and the movement of the object in the virtual world. We also showed that the presence of additional objects in the environment reduces the reversed congruency effect such that it no longer predicts performance. In Experiment 3, we showed that the reversed congruency effect is greater in older adults, suggesting that the quality of spatial memory and perspective-taking abilities are critical. Overall, our results suggest that this effect is driven by difficulties in the precise encoding of object locations in the environment and in understanding how perspective shifts affect the projected positions of the objects in the two-dimensional image.

From repeating routes to planning novel routes: the impact of landmarks and ageing on route integration and cognitive mapping.

The integration of intersecting routes is an important process for the formation of cognitive maps and thus successful navigation. Here we present a novel task to study route integration and the effects that landmark information and cognitive ageing have on this process. We created two virtual environments, each comprising five places and one central intersection but with different landmark settings: in the Identical Landmark environment, the intersection contained visually monotonic features whereas the intersection contained visually distinctive features in the Different Landmarks environment. In both environments young and older participants were presented with two short routes that both traversed through the shared intersection. To test route integration, participants were asked to either repeat the learning routes, to navigate the routes from the destination to the starting place or to plan novel routes. As expected, results demonstrate better performance when repeating or retracing routes than when planning novel routes. Performance was better in younger than older participants and in the Different Landmark environment which does not require detailed knowledge of the spatial configuration of all places in the environment. A subgroup of the older participants who performed lower on a screening test for cognitive impairments could not successfully complete the experiment or did not reach the required performance criterion. These results demonstrate that strategically placed landmarks support the integration of route knowledge into spatial representations that allow for goal-dependent flexible navigation behaviour and that earliest signs of atypical cognitive ageing affect this process of route integration.

Age-related differences in visual encoding and response strategies contribute to spatial memory deficits.

Successful navigation requires memorising and recognising the locations of objects across different perspectives. Although these abilities rely on hippocampal functioning, which is susceptible to degeneration in older adults, little is known about the effects of ageing on encoding and response strategies that are used to recognise spatial configurations. To investigate this, we asked young and older participants to encode the locations of objects in a virtual room shown as a picture on a computer screen. Participants were then shown a second picture of the same room taken from the same (0°) or a different perspective (45° or 135°) and had to judge whether the objects occupied the same or different locations. Overall, older adults had greater difficulty with the task than younger adults although the introduction of a perspective shift between encoding and testing impaired performance in both age groups. Diffusion modelling revealed that older adults adopted a more conservative response strategy, while the analysis of gaze patterns showed an age-related shift in visual-encoding strategies with older adults attending to more information when memorising the positions of objects in space. Overall, results suggest that ageing is associated with declines in spatial processing abilities, with older individuals shifting towards a more conservative decision style and relying more on encoding target object positions using room-based cues compared to younger adults, who focus more on encoding the spatial relationships among object clusters.

The impact of cognitive aging on route learning rate and the acquisition of landmark knowledge.

Aging is accompanied by changes in general cognitive functioning which may impact the learning rate of older adults; however, this is often not controlled for in cognitive aging studies. We investigated the contribution of differences in learning rates to age-related differences in landmark knowledge acquired from route learning. In Experiment 1 we used a standard learning procedure in which participants received a fixed amount of exposure to a route. Consistent with previous research, we found age-related deficits in associative cue and landmark sequence knowledge. Experiment 2 controlled for differences in learning rates by using a flexible exposure learning procedure. Specifically, participants were trained to a performance criterion during route learning before being tested on the content of their route knowledge. While older adults took longer to learn the route than younger adults, the age-related differences in associative cue knowledge were abolished. The deficit in landmark sequence knowledge, however, remained. Experiment 3 replicated these results and introduced a test situation in which a deficit in landmark sequence knowledge yielded an increased likelihood of disorientation in older adults. The findings of this study suggest that age-related deficits in landmark associative cue knowledge are attenuated by controlling for learning rates. In contrast, landmark sequence knowledge deficits persist and are best explained by changes in the learning strategy of older adults to acquire task essential associative cue knowledge at the expense of supplementary sequence knowledge.

A novel virtual-reality-based route-learning test suite: Assessing the effects of cognitive aging on navigation.

Most research groups studying human navigational behavior with virtual environment (VE) technology develop their own tasks and protocols. This makes it difficult to compare results between groups and to create normative data sets for any specific navigational task. Such norms, however, are prerequisites for the use of navigation assessments as diagnostic tools-for example, to support the early and differential diagnosis of atypical aging. Here we start addressing these problems by presenting and evaluating a new navigation test suite that we make freely available to other researchers (https://osf.io/mx52y/). Specifically, we designed three navigational tasks, which are adaptations of earlier published tasks used to study the effects of typical and atypical aging on navigation: a route-repetition task that can be solved using egocentric navigation strategies, and route-retracing and directional-approach tasks that both require allocentric spatial processing. Despite introducing a number of changes to the original tasks to make them look more realistic and ecologically valid, and therefore easy to explain to people unfamiliar with a VE or who have cognitive impairments, we replicated the findings from the original studies. Specifically, we found general age-related declines in navigation performance and additional specific difficulties in tasks that required allocentric processes. These findings demonstrate that our new tasks have task demands similar to those of the original tasks, and are thus suited to be used more widely.

Evidence for age-related deficits in object-location binding during place recognition.

Deciding whether a place is the same or different than places encountered previously is a common task in daily navigation which requires to develop knowledge about the locations of objects (object-location binding) and to recognize places from different perspectives. These abilities rely on hippocampal functioning which is susceptible to increasing age. Thus, the question of the present study is how they both together impact on place recognition in aging. Forty people aged 20-29, 44 aged 60-69, and 32 aged 70-79 were presented with places consisting of four different objects during the encoding phase. In the test phase, they were then presented with a second place and had to decide whether it was the same or different. Test places were presented from different perspectives (0°, 30°, 60°) and with different object conditions (same, a swap of two objects, a substitution with a novel object). The sensitivity for detecting changes (d') decreased from 20-29 to 60-69 and to 70-79 years old, and with increasing perspective shifts. Importantly, older adults were less sensitive to object swapping than to object substitution, while young participants did not show any difference. Overall, these results suggest specific age-related difficulties in object-location binding in the context of place recognition.

Investigating the Effect of the Environment on Prey Detection Ability in Humans.

Visual search experiments used in the field of psychology may be applied to investigate the relationship between environments and prey detection rates that could influence hunting behaviours in ancient humans. Two lab-based experiments were designed to examine the effects of differing virtual environments, representing Marine Isotope Stage 3 (MIS3) in Europe, on participants' ability to locate prey. The results show that prey detection performance is highly influenced by vegetation structure, both in terms of the biome type (wooded vs. grassland environments) and the density of the vegetation (trees in wooded and shrubs in grassland environments). However, the density of vegetation has a greater relative effect in grassland than in wooded biomes. Closer examination of the transition between biomes (relative percentages of trees vs. shrubs) at the same vegetative density shows a non-linear relationship between prey detection performance and the relative tree to shrub percentages. Changes in the distribution of biomes occurred throughout the Quaternary. The composition of those biomes will have likely affected hominin hunting behaviours because of their intermediary effects on prey detection performance. This may, therefore, have played a role in the turn-overs of hunter-gatherer hominin populations during MIS3 and at other times in the Quaternary.

The contribution of visual attention and declining verbal memory abilities to age-related route learning deficits.

Our ability to learn unfamiliar routes declines in typical and atypical ageing. The reasons for this decline, however, are not well understood. Here we used eye-tracking to investigate how ageing affects people's ability to attend to navigationally relevant information and to select unique objects as landmarks. We created short routes through a virtual environment, each comprised of four intersections with two objects each, and we systematically manipulated the saliency and uniqueness of these objects. While salient objects might be easier to memorise than non-salient objects, they cannot be used as reliable landmarks if they appear more than once along the route. As cognitive ageing affects executive functions and control of attention, we hypothesised that the process of selecting navigationally relevant objects as landmarks might be affected as well. The behavioural data showed that younger participants outperformed the older participants and the eye-movement data revealed some systematic differences between age groups. Specifically, older adults spent less time looking at the unique, and therefore navigationally relevant, landmark objects. Both young and older participants, however, effectively directed gaze towards the unique and away from the non-unique objects, even if these were more salient. These findings highlight specific age-related differences in the control of attention that could contribute to declining route learning abilities in older age. Interestingly, route-learning performance in the older age group was more variable than in the young age group with some older adults showing performance similar to the young group. These individual differences in route learning performance were strongly associated with verbal and episodic memory abilities.

Virtual navigation tested on a mobile app is predictive of real-world wayfinding navigation performance.

Virtual reality environments presented on tablets and smartphones have potential to aid the early diagnosis of conditions such as Alzheimer's dementia by quantifying impairments in navigation performance. However, it is unclear whether performance on mobile devices can predict navigation errors in the real world. We compared the performance of 49 participants (25 females, 18-35 years old) at wayfinding and path integration tasks designed in our mobile app 'Sea Hero Quest' with their performance at similar tasks in a real-world environment. We first performed this experiment in the streets of London (UK) and replicated it in Paris (France). In both cities, we found a significant correlation between virtual and real-world wayfinding performance and a male advantage in both environments, although smaller in the real world (Cohen's d in the game = 0.89, in the real world = 0.59). Results in London and Paris were highly similar, and controlling for familiarity with video games did not change the results. The strength of the correlation between real world and virtual environment increased with the difficulty of the virtual wayfinding task, indicating that Sea Hero Quest does not merely capture video gaming skills. The fact that the Sea Hero Quest wayfinding task has real-world ecological validity constitutes a step toward controllable, sensitive, safe, low-cost, and easy to administer digital cognitive assessment of navigation ability.

Global Determinants of Navigation Ability.

Human spatial ability is modulated by a number of factors, including age [1-3] and gender [4, 5]. Although a few studies showed that culture influences cognitive strategies [6-13], the interaction between these factors has never been globally assessed as this requires testing millions of people of all ages across many different countries in the world. Since countries vary in their geographical and cultural properties, we predicted that these variations give rise to an organized spatial distribution of cognition at a planetary-wide scale. To test this hypothesis, we developed a mobile-app-based cognitive task, measuring non-verbal spatial navigation ability in more than 2.5 million people and sampling populations in every nation state. We focused on spatial navigation due to its universal requirement across cultures. Using a clustering approach, we find that navigation ability is clustered into five distinct, yet geographically related, groups of countries. Specifically, the economic wealth of a nation was predictive of the average navigation ability of its inhabitants, and gender inequality was predictive of the size of performance difference between males and females. Thus, cognitive abilities, at least for spatial navigation, are clustered according to economic wealth and gender inequalities globally, which has significant implications for cross-cultural studies and multi-center clinical trials using cognitive testing.

Spatial navigation from same and different directions: The role of executive functions, memory and attention in adults with autism spectrum disorder.

To resolve some of the inconsistencies in existing research into spatial navigation in autism spectrum disorder (ASD), we tested two large age- and ability-matched groups of ASD and typically developing (TD) participants for their spatial navigation abilities in a route learning task, which has been shown to shed light on the strategies participants employ when navigating complex environments. Participants studied a route through a virtual maze by watching a short video of a first-person perspective navigating a maze. The maze included four four-way intersections that were each marked with two unique landmarks in two corners of the intersection. At test, static images of the intersections, either as seen during the video or as approached from a different direction, were presented and participants had to indicate in which direction they would need to travel (straight, left, or right) in order to follow the originally studied route. On both types of test trials, the ASD group performed worse and their difficulties were related to reduced cognitive flexibility. Eye-movement data and follow-up item-memory tests suggested that navigation difficulties may have been related to differences in attention during encoding and less spontaneous use of landmarks as cues for navigation. Spatial navigation performance was best predicted by memory for landmarks as well as by executive functions. The results are discussed in relation to theories of underlying navigation-related brain regions. More research is needed to disentangle the influence of executive functions, memory and attention on spatial navigation. Autism Res 2018, 11: 798-810. © 2018 International Society for Autism Research, Wiley Periodicals, Inc. LAY SUMMARY: Navigating an environment is difficult for people with ASD independent of whether they are travelling in the same or in a different direction from that which they originally studied. The present study suggests that flexibility in alternating travel directions, difficulties in remembering landmarks as well as reduced attention to landmarks while learning a route play a role in the navigation difficulties in ASD. Guidance at route learning might help autistic individuals to improve their ability to navigate in their environments.