The relationship between object-based spatial ability and virtual navigation performance.

Spatial navigation is a multi-faceted behaviour drawing on many different aspects of cognition. Visuospatial abilities, such as mental rotation and visuospatial working memory, in particular, may be key factors. A range of tests have been developed to assess visuospatial processing and memory, but how such tests relate to navigation ability remains unclear. This understanding is important to advance tests of navigation for disease monitoring in various disorders (e.g., Alzheimer's disease) where spatial impairment is an early symptom. Here, we report the use of an established mobile gaming app, Sea Hero Quest (SHQ), as a measure of navigation ability in a sample of young, predominantly female university students (N = 78; 20; female = 74.3%; mean age = 20.33 years). We used three separate tests of navigation embedded in SHQ: wayfinding, path integration and spatial memory in a radial arm maze. In the same participants, we also collected measures of mental rotation (Mental Rotation Test), visuospatial processing (Design Organization Test) and visuospatial working memory (Digital Corsi). We found few strong correlations across our measures. Being good at wayfinding in a virtual navigation test does not mean an individual will also be good at path integration, have a superior memory in a radial arm maze, or rate themself as having a strong sense of direction. However, we observed that participants who were good in the wayfinding task of SHQ tended to perform well on the three visuospatial tasks examined here, and to also use a landmark strategy in the radial maze task. These findings help clarify the associations between different abilities involved in spatial navigation.

Shorter self-reported sleep duration is associated with worse virtual spatial navigation performance in men.

Sleep has been shown to impact navigation ability. However, it remains unclear how different sleep-related variables may be independently associated with spatial navigation performance, and as to whether gender may play a role in these associations. We used a mobile video game app, Sea Hero Quest (SHQ), to measure wayfinding ability in US-based participants. Wayfinding performance on SHQ has been shown to correlate with real-world wayfinding. Participants were asked to report their sleep duration, quality, daytime sleepiness and nap frequency and duration on a typical night (n = 766, 335 men, 431 women, mean age = 26.5 years, range = 18-59 years). A multiple linear regression was used to identify which self-reported sleep variables were independently associated with wayfinding performance. Shorter self-reported sleep durations were significantly associated with worse wayfinding performance in men only. Other self-reported sleep variables showed non-significant trends of association with wayfinding performance. When removing non-typical sleepers (< 6 or > 9 h of sleep on a typical night), the significant association between sleep duration and spatial navigation performance in men was no longer present. These findings from U.S.-based participants suggest that a longer self-reported sleep duration may be an important contributor to successful navigation ability in men.

Wayfinding across ocean and tundra: what traditional cultures teach us about navigation.

Research on human navigation by psychologists and neuroscientists has come mainly from a limited range of environments and participants inhabiting western countries. By contrast, numerous anthropological accounts illustrate the diverse ways in which cultures adapt to their surrounding environment to navigate. Here, we provide an overview of these studies and relate them to cognitive science research. The diversity of cues in traditional navigation is much higher and multimodal compared with navigation experiments in the laboratory. It typically involves an integrated system of methods, drawing on a detailed understanding of the environmental cues, specific tools, and forms part of a broader cultural system. We highlight recent methodological developments for measuring navigation skill and modelling behaviour that will aid future research into how culture and environment shape human navigation.

A neural circuit for spatial orientation derived from brain lesions.

There is disagreement regarding the major components of the brain network supporting spatial cognition. To address this issue, we applied a lesion mapping approach to the clinical phenomenon of topographical disorientation. Topographical disorientation is the inability to maintain accurate knowledge about the physical environment and use it for navigation. A review of published topographical disorientation cases identified 65 different lesion sites. Our lesion mapping analysis yielded a topographical disorientation brain map encompassing the classic regions of the navigation network: medial parietal, medial temporal, and temporo-parietal cortices. We also identified a ventromedial region of the prefrontal cortex, which has been absent from prior descriptions of this network. Moreover, we revealed that the regions mapped are correlated with the Default Mode Network sub-network C. Taken together, this study provides causal evidence for the distribution of the spatial cognitive system, demarking the major components and identifying novel regions.

Examining individual learning patterns using generalised linear mixed models.

Everyone learns differently, but individual performance is often ignored in favour of a group-level analysis. Using data from four different experiments, we show that generalised linear mixed models (GLMMs) and extensions can be used to examine individual learning patterns. Producing ellipsoids and cluster analyses based on predicted random effects, individual learning patterns can be identified, clustered and used for comparisons across various experimental conditions or groups. This analysis can handle a range of datasets including discrete, continuous, censored and non-censored, as well as different experimental conditions, sample sizes and trial numbers. Using this approach, we show that learning a face-named paired associative task produced individuals that can learn quickly, with the performance of some remaining high, but with a drop-off in others, whereas other individuals show poor performance throughout the learning period. We see this more clearly in a virtual navigation spatial learning task (NavWell). Two prominent clusters of learning emerged, one showing individuals who produced a rapid learning and another showing a slow and gradual learning pattern. Using data from another spatial learning task (Sea Hero Quest), we show that individuals' performance generally reflects their age category, but not always. Overall, using this analytical approach may help practitioners in education and medicine to identify those individuals who might need extra help and attention. In addition, identifying learning patterns may enable further investigation of the underlying neural, biological, environmental and other factors associated with these individuals.

Goal-seeking compresses neural codes for space in the human hippocampus and orbitofrontal cortex.

Humans can navigate flexibly to meet their goals. Here, we asked how the neural representation of allocentric space is distorted by goal-directed behavior. Participants navigated an agent to two successive goal locations in a grid world environment comprising four interlinked rooms, with a contextual cue indicating the conditional dependence of one goal location on another. Examining the neural geometry by which room and context were encoded in fMRI signals, we found that map-like representations of the environment emerged in both hippocampus and neocortex. Cognitive maps in hippocampus and orbitofrontal cortices were compressed so that locations cued as goals were coded together in neural state space, and these distortions predicted successful learning. This effect was captured by a computational model in which current and prospective locations are jointly encoded in a place code, providing a theory of how goals warp the neural representation of space in macroscopic neural signals.

Wayfinding and path integration deficits detected using a virtual reality mobile app in patients with traumatic brain injury.

The ability to navigate is supported by a wide network of brain areas which are particularly vulnerable to disruption brain injury, including traumatic brain injury (TBI). Wayfinding and the ability to orient back to the direction you have recently come (path integration) may likely be impacted in daily life but have so far not been tested with patients with TBI. Here, we assessed spatial navigation in thirty-eight participants, fifteen of whom had a history of TBI, and twenty-three control participants. Self-estimated spatial navigation ability was assessed using the Santa Barbara Sense of Direction (SBSOD) scale. No significant difference between TBI patients and a control group was identified. Rather, results indicated that both participant groups demonstrated 'good' self-inferred spatial navigational ability on the SBSOD scale. Objective navigation ability was tested via the virtual mobile app test Sea Hero Quest (SHQ), which has been shown to predict real-world navigation difficulties and assesses (a) wayfinding across several environments and (b) path integration. Compared to a sub-sample of 13 control participants, a matched subsample of 10 TBI patients demonstrated generally poorer performance on all wayfinding environments tested. Further analysis revealed that TBI participants consistently spent a shorter duration viewing a map prior to navigating to goals. Patients showed mixed performance on the path integration task, with poor performance evident when proximal cues were absent. Our results provide preliminary evidence that TBI impacts both wayfinding and, to some extent, path integration. The findings suggest long-lasting clinical difficulties experienced in TBI patients affect both wayfinding and to some degree path integration ability.

Hydrocephalus: A neuropsychological and theoretical primer.

Hydrocephalus is a common neurological condition, the hallmark feature of which is an excess in production, or accumulation, of cerebrospinal fluid in the ventricles. Although it is associated with diffuse damage to paraventricular brain areas, patients are broadly typified by a particular pattern of cognitive impairments that include deficits in working memory, attention, and spatial abilities. There have, however, been relatively few neuropsychological accounts of the condition. Moreover, theories of the relationship between aetiology and impairment appear to have emerged in isolation of each other, and proffer fundamentally different accounts. In this primer, we aim to provide a comprehensive and contemporary overview of hydrocephalus for the neuropsychologist, covering cognitive sequelae and theoretical interpretations of their origins. We review clinical and neuropsychological assays of cognitive profiles, along with the few studies that have addressed more integrative behaviours. In particular, we explore the distinction between congenital or early-onset hydrocephalus with a normal-pressure variant that can be acquired later in life. The relationship between these two populations is a singularly interesting one in neuropsychology since it can allow for the examination of typical and atypical developmental trajectories, and their interaction with chronic and acute impairment, within the same broad neurological condition. We reflect on the ramifications of this for our subject and suggest avenues for future research.

Landmark-dependent Navigation Strategy Declines across the Human Life-Span: Evidence from Over 37,000 Participants.

Humans show a remarkable capacity to navigate various environments using different navigation strategies, and we know that strategy changes across the life span. However, this observation has been based on studies of small sample sizes. To this end, we used a mobile app-based video game (Sea Hero Quest) to test virtual navigation strategies and memory performance within a distinct radial arm maze level in over 37,000 participants. Players were presented with six pathways (three open and three closed) and were required to navigate to the three open pathways to collect a target. Next, all six pathways were made available and the player was required to visit the pathways that were previously unavailable. Both reference memory and working memory errors were calculated. Crucially, at the end of the level, the player was asked a multiple-choice question about how they found the targets (i.e., a counting-dependent strategy vs. a landmark-dependent strategy). As predicted from previous laboratory studies, we found the use of landmarks declined linearly with age. Those using landmark-based strategies also performed better on reference memory than those using a counting-based strategy. These results extend previous observations in the laboratory showing a decreased use of landmark-dependent strategies with age.

Humans account for cognitive costs when finding shortcuts: An information-theoretic analysis of navigation.

When faced with navigating back somewhere we have been before we might either retrace our steps or seek a shorter path. Both choices have costs. Here, we ask whether it is possible to characterize formally the choice of navigational plans as a bounded rational process that trades off the quality of the plan (e.g., its length) and the cognitive cost required to find and implement it. We analyze the navigation strategies of two groups of people that are firstly trained to follow a "default policy" taking a route in a virtual maze and then asked to navigate to various known goal destinations, either in the way they want ("Go To Goal") or by taking novel shortcuts ("Take Shortcut"). We address these wayfinding problems using InfoRL: an information-theoretic approach that formalizes the cognitive cost of devising a navigational plan, as the informational cost to deviate from a well-learned route (the "default policy"). In InfoRL, optimality refers to finding the best trade-off between route length and the amount of control information required to find it. We report five main findings. First, the navigational strategies automatically identified by InfoRL correspond closely to different routes (optimal or suboptimal) in the virtual reality map, which were annotated by hand in previous research. Second, people deliberate more in places where the value of investing cognitive resources (i.e., relevant goal information) is greater. Third, compared to the group of people who receive the "Go To Goal" instruction, those who receive the "Take Shortcut" instruction find shorter but less optimal solutions, reflecting the intrinsic difficulty of finding optimal shortcuts. Fourth, those who receive the "Go To Goal" instruction modulate flexibly their cognitive resources, depending on the benefits of finding the shortcut. Finally, we found a surprising amount of variability in the choice of navigational strategies and resource investment across participants. Taken together, these results illustrate the benefits of using InfoRL to address navigational planning problems from a bounded rational perspective.

From cognitive maps to spatial schemas.

A schema refers to a structured body of prior knowledge that captures common patterns across related experiences. Schemas have been studied separately in the realms of episodic memory and spatial navigation across different species and have been grounded in theories of memory consolidation, but there has been little attempt to integrate our understanding across domains, particularly in humans. We propose that experiences during navigation with many similarly structured environments give rise to the formation of spatial schemas (for example, the expected layout of modern cities) that share properties with but are distinct from cognitive maps (for example, the memory of a modern city) and event schemas (such as expected events in a modern city) at both cognitive and neural levels. We describe earlier theoretical frameworks and empirical findings relevant to spatial schemas, along with more targeted investigations of spatial schemas in human and non-human animals. Consideration of architecture and urban analytics, including the influence of scale and regionalization, on different properties of spatial schemas may provide a powerful approach to advance our understanding of spatial schemas.

Explaining World-Wide Variation in Navigation Ability from Millions of People: Citizen Science Project Sea Hero Quest.

Navigation ability varies widely across humans. Prior studies have reported that being younger and a male has an advantage for navigation ability. However, these studies have generally involved small numbers of participants from a handful of western countries. Here, we review findings from our project Sea Hero Quest, which used a video game for mobile and tablet devices to test 3.9 million people on their navigation ability, sampling across every nation-state and from 18 to 99 years of age. Results revealed that the task has good ecological validity and across all countries sufficiently sampled (N = 63), age is linked to a near-linear decline in navigation ability from the early 20s. All countries showed a male advantage, but this varied considerably and could be partly predicted by gender inequality. We found that those who reported growing up in a city were on average worse at navigating than those who grew up outside cities and that navigation performance helped identify those at greater genetic risk of Alzheimer's disease. We discuss the advantages and challenges of using a mobile app to study cognition and the future avenues for understanding individual differences in navigation ability arising from this research.

Predictive maps in rats and humans for spatial navigation.

Much of our understanding of navigation comes from the study of individual species, often with specific tasks tailored to those species. Here, we provide a novel experimental and analytic framework integrating across humans, rats, and simulated reinforcement learning (RL) agents to interrogate the dynamics of behavior during spatial navigation. We developed a novel open-field navigation task ("Tartarus maze") requiring dynamic adaptation (shortcuts and detours) to frequently changing obstructions on the path to a hidden goal. Humans and rats were remarkably similar in their trajectories. Both species showed the greatest similarity to RL agents utilizing a "successor representation," which creates a predictive map. Humans also displayed trajectory features similar to model-based RL agents, which implemented an optimal tree-search planning procedure. Our results help refine models seeking to explain mammalian navigation in dynamic environments and highlight the utility of modeling the behavior of different species to uncover the shared mechanisms that support behavior.

Extending neural systems for navigation to hunting behavior.

For decades, a central question in neuroscience has been: How does the brain support navigation? Recent research on navigation has explored how brain regions support the capacity to adapt to changes in the environment and track the distance and direction to goal locations. Here, we provide a brief review of this literature and speculate how these neural systems may be involved in another, parallel behavior-hunting. Hunting shares many of the same challenges as navigation. Like navigation, hunting requires the hunter to orient towards a goal while minimizing their distance from it while traveling. Likewise, hunting may require the accommodation of detours to locate prey or the exploitation of shortcuts for a quicker capture. Recent research suggests that neurons in the periaqueductal gray, hypothalamus, and dorsal anterior cingulate play key roles in such hunting behavior. In this review, we speculate on how these regions may operate functionally with other key brain regions involved in navigation, such as the hippocampus, to support hunting. Additionally, we posit that hunting in a group presents an additional set of challenges, where success relies on multicentric tracking and prediction of prey position as well as the position of co-hunters.

Spatial goal coding in the hippocampal formation.

The mammalian hippocampal formation contains several distinct populations of neurons involved in representing self-position and orientation. These neurons, which include place, grid, head direction, and boundary cells, are thought to collectively instantiate cognitive maps supporting flexible navigation. However, to flexibly navigate, it is necessary to also maintain internal representations of goal locations, such that goal-directed routes can be planned and executed. Although it has remained unclear how the mammalian brain represents goal locations, multiple neural candidates have recently been uncovered during different phases of navigation. For example, during planning, sequential activation of spatial cells may enable simulation of future routes toward the goal. During travel, modulation of spatial cells by the prospective route, or by distance and direction to the goal, may allow maintenance of route and goal-location information, supporting navigation on an ongoing basis. As the goal is approached, an increased activation of spatial cells may enable the goal location to become distinctly represented within cognitive maps, aiding goal localization. Lastly, after arrival at the goal, sequential activation of spatial cells may represent the just-taken route, enabling route learning and evaluation. Here, we review and synthesize these and other evidence for goal coding in mammalian brains, relate the experimental findings to predictions from computational models, and discuss outstanding questions and future challenges.

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.

The Versatile Wayfinder: Prefrontal Contributions to Spatial Navigation.

The prefrontal cortex (PFC) supports decision-making, goal tracking, and planning. Spatial navigation is a behavior that taxes these cognitive processes, yet the role of the PFC in models of navigation has been largely overlooked. In humans, activity in dorsolateral PFC (dlPFC) and ventrolateral PFC (vlPFC) during detours, reveal a role in inhibition and replanning. Dorsal anterior cingulate cortex (dACC) is implicated in planning and spontaneous internally-generated changes of route. Orbitofrontal cortex (OFC) integrates representations of the environment with the value of actions, providing a 'map' of possible decisions. In rodents, medial frontal areas interact with hippocampus during spatial decisions and switching between navigation strategies. In reviewing these advances, we provide a framework for how different prefrontal regions may contribute to different stages of navigation.

Hippocampal place cells encode global location but not connectivity in a complex space.

Flexible navigation relies on a cognitive map of space, thought to be implemented by hippocampal place cells: neurons that exhibit location-specific firing. In connected environments, optimal navigation requires keeping track of one's location and of the available connections between subspaces. We examined whether the dorsal CA1 place cells of rats encode environmental connectivity in four geometrically identical boxes arranged in a square. Rats moved between boxes by pushing saloon-type doors that could be locked in one or both directions. Although rats demonstrated knowledge of environmental connectivity, their place cells did not respond to connectivity changes, nor did they represent doorways differently from other locations. Place cells coded location in a global reference frame, with a different map for each box and minimal repetitive fields despite the repetitive geometry. These results suggest that CA1 place cells provide a spatial map that does not explicitly include connectivity.