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Cognitive map

A cognitive map is an internal mental representation of the spatial layout and relationships within an environment, allowing organisms to navigate, plan routes, and adapt to changes such as blocked paths or new goals.^[1] This concept enables flexible behavior beyond simple trial-and-error learning, as demonstrated in early experiments where rats could take shortcuts in mazes after initial exploration without immediate rewards.^[2] The term was coined by psychologist Edward C. Tolman in his seminal 1948 paper, which proposed that such maps form through latent learning, where knowledge is acquired and stored for later use.^[1] Tolman's foundational work involved maze studies with rats, revealing that they developed anticipatory spatial knowledge rather than mere stimulus-response associations.^[2] For instance, in maze experiments, rats took detours around barriers to reach food rewards, indicating an internalized understanding of the layout.^[1] This challenged behaviorist views dominant at the time and laid the groundwork for cognitive psychology, emphasizing purposive behavior guided by internal representations.^[3] In humans, cognitive maps support not only physical navigation but also memory formation and decision-making, with neural underpinnings in the hippocampus and entorhinal cortex.^[3] Place cells in the hippocampus fire in response to specific locations, while grid cells in the entorhinal cortex provide a coordinate-like framework for spatial encoding.^[3] These mechanisms allow individuals to mentally simulate paths and integrate landmarks, as seen in studies where people reorient themselves in novel environments using internalized maps.^[3] Beyond spatial domains, cognitive maps extend to abstract representations, such as social networks or conceptual knowledge structures, where the brain adapts similar hippocampal processes to navigate non-physical relationships.^[3] For example, activity patterns in the hippocampus during tasks involving social hierarchies mirror those in spatial navigation, suggesting a generalized mapping system for relational inference.^[3] This versatility underscores the cognitive map's role in broader adaptive behaviors, from urban planning to problem-solving in complex scenarios.

Definition and Overview

Core Concept

A cognitive map is an internal, abstract representation of the spatial layout of an environment, enabling organisms to anticipate outcomes and navigate without relying on direct sensory experience or trial-and-error.^[1] Introduced by psychologist Edward C. Tolman in his seminal 1948 paper, this concept posits that learning involves forming a mental model of environmental relationships, rather than mere associations between stimuli and responses.^[1] Key functions of cognitive maps include facilitating route planning, discovering shortcuts, and supporting flexible, goal-directed behavior in familiar settings.^[1] These representations can manifest as vector-based structures encoding metric distances and directions, or topological frameworks capturing connectivity between locations.^[4] For instance, a person might use a cognitive map of their city to identify an efficient shortcut between two points, while a rat in a laboratory maze employs it to reach a food reward via an altered path after initial exploration.^[1] This idea forms a cornerstone of Tolman's purposive behaviorism, outlined in his 1932 book, which emphasized cognitive processes and goal orientation in behavior over rigid stimulus-response mechanisms.^[5] By integrating environmental cues into a cohesive mental framework, cognitive maps allow for latent learning, where knowledge accrues without immediate reinforcement and manifests adaptively when needed.^[1] The concept of a mental map, prevalent in geography and urban planning, refers to an individual's subjective and often distorted perception of geographic areas, such as the layout of a city, emphasizing qualitative biases like overestimation of familiar routes or centrality of personal landmarks rather than precise navigational utility. This usage, as explored by Kevin Lynch, focuses on how people image urban environments through elements like paths, edges, districts, nodes, and landmarks, which can vary widely due to personal experience and cultural factors.^[6] In distinction, the psychological notion of a cognitive map entails a more structured internal representation of spatial relationships that supports flexible, goal-directed navigation, independent of such perceptual distortions. Cognitive maps also differ from schemas, which are broader knowledge structures that abstract common patterns across diverse experiences to facilitate general comprehension and prediction.^[7] Schemas, as defined in memory research, integrate episodic details into generalized frameworks applicable to non-spatial domains, such as social interactions or event sequences, without requiring environment-specific encoding.^[7] By contrast, cognitive maps prioritize detailed, allocentric encodings of spatial and relational elements within a particular setting, enabling precise inference like shortcut selection, and are typically dependent on hippocampal mechanisms rather than the neocortical storage of abstract schemas.^[7] Heuristics represent another related but distinct construct, involving simplified rule-of-thumb strategies for navigation, such as beacon homing toward visible landmarks or path integration via accumulated sensory cues, which approximate spatial problem-solving without forming a holistic representation.^[1] These approaches, akin to stimulus-response chains in behaviorist accounts, rely on trial-and-error or sequential habits that limit flexibility to learned routes.^[1] Cognitive maps, however, transcend such approximations by incorporating latent learning, where unobserved spatial knowledge is inferred to support novel behaviors, as evidenced in Tolman's rat maze experiments where animals bypassed barriers via untraveled paths.^[1] The terminology of "cognitive map" thus specifically denotes representations enabling latent learning and inferential capabilities, setting it apart from mere memorized paths or rote navigational sequences that lack true spatial generalization.^[1]

Historical Development

Origins and Early Theories

The origins of the cognitive map concept trace back to early 20th-century Gestalt psychology, which prioritized holistic perception and the organized structure of mental processes over fragmented sensory elements. Gestalt theorists argued that learning and problem-solving involve perceiving the whole configuration of a situation, rather than accumulating isolated associations. A key example is Wolfgang Köhler's research in the 1920s, where he observed chimpanzees achieving insight learning—sudden solutions to novel problems, such as stacking boxes to reach bananas—indicating that animals form integrated representations of their environment for adaptive behavior.^[8] Edward C. Tolman, influenced by Gestalt ideas, extended this holistic approach to spatial learning in rodents. In his 1948 paper "Cognitive maps in rats and men," Tolman proposed cognitive maps as internal, survey-like representations of spatial layouts that guide purposive behavior, allowing organisms to navigate flexibly toward goals without relying solely on chained stimulus-response (S-R) associations. He contrasted this with behaviorist models, asserting that cognitive maps account for phenomena like detours and novel path selection, where animals demonstrate knowledge of the overall environment rather than rote habits.^[9] Tolman's sign-gestalt theory, developed earlier, further bridged Gestalt principles with behavioral explanations by positing that environmental cues form meaningful expectancies in the learner's mind.^[10] This theoretical advancement occurred during the broader transition from strict behaviorism to the cognitive revolution in the 1940s and 1950s, as psychologists increasingly recognized internal mental representations as essential to explaining complex behaviors. Tolman's ideas were contextualized by growing interest in animal navigation within ecological settings and the nascent field of ethology, which explored instinctive spatial adaptations in natural environments, such as birds' migratory routes. Early debates surrounding cognitive maps centered on their origins, with Tolman emphasizing learned structures built through exploratory experience and latent learning, rather than purely innate mechanisms.^[11]^[12]

Seminal Experiments

One of the earliest demonstrations of latent learning, which provided foundational evidence for cognitive mapping, came from Blodgett's 1929 experiment using a six-unit alley maze with rats. In this study, rats in experimental groups explored the maze without food rewards for several days (three or six days, depending on the group), after which rewards were introduced; these groups exhibited a sudden and dramatic drop in errors—often halving their error rates immediately—compared to a continuously rewarded control group, suggesting that the rats had formed an internal representation of the maze layout during unrewarded exploration rather than relying solely on trial-and-error reinforcement. Building on this, Tolman and Honzik's 1930 experiments further validated latent learning using a more complex 14-unit multiple T-maze. Here, an experimental group of hungry rats ran the maze without rewards for the first 10 days, showing no improvement, but upon reward introduction on day 11, their errors plummeted from an average of 15 to under 4 per trial within sessions, while control groups (continuously unrewarded or rewarded) improved gradually; this abrupt performance shift indicated the acquisition of a cognitive map independent of immediate reinforcement. Tolman's seminal 1948 review synthesized these findings and presented additional rat experiments that directly tested for map-like representations, such as spatial orientation and shortcut-taking. In one key setup, rats trained on an elevated plus-maze to reach food from a consistent starting point were later tested with barriers removed or alternative paths opened; upon release into an open field with radiating arms pointing toward the trained food location, the rats preferentially chose the arm aligned with the goal (e.g., 42% selected the correct path versus 20% for others), demonstrating flexible navigation via an internalized spatial map rather than fixed responses.^[13] These experiments highlighted methodological innovations like T-mazes and plus-mazes to distinguish cognitive mapping from stimulus-response habits. In Tolman, Ritchie, and Kalish's 1946 studies using a plus-maze, rats trained under "place learning" conditions (navigating to a fixed spatial location using extra-maze cues) readily took novel shortcuts when barriers were removed, whereas "response learning" groups (trained on body turns regardless of location) failed to adapt, confirming the role of holistic environmental representations in inference-based navigation. Early human analogs to these rat studies emerged in the 1950s through sketching tasks, where participants drew maps of familiar room layouts or indoor environments from memory, revealing map-like distortions such as elongation of central paths and underrepresentation of peripheral details, indicative of abstracted spatial cognition similar to Tolman's findings.

Formation and Acquisition

Learning Mechanisms

Cognitive maps are constructed through a combination of self-motion cues and environmental landmarks, enabling organisms to estimate their position and navigate effectively. Path integration, also known as dead reckoning, serves as a primary mechanism for updating spatial position by continuously integrating velocity and acceleration signals from vestibular and proprioceptive inputs. This process allows animals to maintain an internal estimate of location even in the absence of external cues, as demonstrated in studies where rodents accurately return to a starting point after passive displacement. However, path integration is inherently prone to cumulative errors over time, necessitating complementary learning strategies to refine the map.^[14] Landmark-based learning involves associating salient environmental features, such as visual or olfactory cues, with specific locations to anchor the cognitive map. In this process, organisms distinguish between beacon homing—direct approach to a single prominent landmark—and true cognitive mapping, where multiple landmarks are integrated to form relational spatial representations. For instance, rats prioritize geometric configurations of landmarks over individual beacons when disoriented, indicating a modular system for encoding spatial geometry that supports flexible navigation. This learning enables correction of path integration errors by resetting the internal position estimate upon encountering familiar landmarks.^[15] Error correction in cognitive map formation relies on Bayesian updating, where new sensory inputs are weighted against prior estimates to revise the spatial representation probabilistically. This mechanism treats path integration outputs as noisy priors that are adjusted based on landmark reliability, minimizing overall uncertainty in position estimates. For example, when discrepancies arise between integrated self-motion and observed landmarks, the system recalibrates by downweighting unreliable cues, as modeled in human and animal navigation tasks. Such updating ensures map stability, with exploration behaviors facilitating repeated encounters that refine accuracy over time.^[15] Multimodal integration combines inputs from visual, vestibular, proprioceptive, and other sensory modalities to construct a robust cognitive map, with each cue contributing to position estimation according to its precision.^[15] During navigation, self-motion cues from path integration are fused with landmark information, often through weighted averaging that favors more reliable sources, enhancing overall map fidelity. Exploration plays a critical role in this integration, as active movement exposes the organism to diverse sensory data, allowing iterative refinement of the map through trial-and-error adjustments. This process supports adaptive navigation across varying environments, though its efficiency can vary developmentally.

Developmental Processes

In infancy, cognitive maps begin to emerge through reliance on egocentric spatial coding, where infants use body-centered cues to track object locations. The classic A-not-B task illustrates this nascent mapping ability, as infants aged 6-12 months often search incorrectly at the previous hiding location (A) despite seeing the object moved to a new one (B), reflecting limitations in updating spatial representations based on egocentric frames. Performance improves rapidly during this period, with average success rates rising from near zero at 6 months to over 80% by 12 months, indicating the development of basic spatial working memory and inhibition of outdated cues. This early phase lays the foundation for more flexible mapping, though still dominated by proximal, self-referenced information.^[16] During childhood, a key transition occurs around ages 4-6, shifting from predominantly egocentric to allocentric representations, where children begin using external landmarks and geometric properties independent of their viewpoint to form cognitive maps. This developmental milestone aligns with Piaget's preoperational to concrete operational stages, during which children overcome egocentrism to integrate distal cues for navigation, as seen in tasks like disorientation paradigms where 4- to 5-year-olds show rudimentary allocentric recall in virtual environments, improving to reliable shortcut-taking by age 6. By middle childhood (7-12 years), experience further refines these maps, enabling adult-like integration of metric and landmark information in larger spaces.^[17]^[18] In adulthood, cognitive maps continue to refine through accumulated experience, enhancing precision in route planning and environmental integration via hippocampal plasticity. However, aging brings declines in hippocampal function, such as reduced place cell stability and synaptic efficiency, leading to navigation deficits in the elderly, including slower learning of new layouts and reliance on less efficient egocentric strategies. Studies using virtual Morris water mazes demonstrate that older adults exhibit 20-30% lower accuracy in allocentric tasks compared to younger ones, correlating with hippocampal volume loss and contributing to real-world wayfinding challenges.^[19] Cultural and environmental factors influence map acquisition across the lifespan, with variations tied to exposure levels. For instance, individuals raised in rural or suburban settings develop superior spatial navigation skills into adulthood compared to those from urban grid-like environments, as low-entropy street networks in cities limit exposure to varied topography.^[20] This highlights how diverse environmental experiences shape the robustness of cognitive maps from childhood onward.

Neural Basis

Key Brain Regions and Cells

The hippocampus plays a central role in cognitive maps through place cells, which are neurons that fire selectively when an animal is in specific locations within an environment. These cells were first identified in the CA1 region of the rat hippocampus, where they exhibit stable firing fields corresponding to particular places, independent of the animal's orientation or sensory inputs.^[21] Place cells integrate spatial information to form a representation of the environment's layout, contributing to the formation of allocentric cognitive maps. Adjacent to the hippocampus, the entorhinal cortex provides essential metric structure to these maps via grid cells, which fire in a hexagonal lattice pattern across the environment, offering a coordinate system for distance and direction. Discovered in the medial entorhinal cortex (MEC) of rats, grid cells maintain their firing patterns during navigation, scaling and rotating with environmental changes to support path integration.^[22] Additionally, the entorhinal cortex contains boundary vector cells (also termed border cells), which activate near environmental edges or walls, encoding the distance and direction to boundaries to delineate the map's limits. Beyond these core structures, the parietal cortex facilitates the transformation from egocentric (body-centered) to allocentric (world-centered) representations, enabling the integration of sensory cues into a coherent spatial framework. Neurons in areas like the ventral intraparietal area exhibit flexible coding that switches between reference frames based on navigational demands, supporting efficient route planning.^[23] The prefrontal cortex, particularly its medial regions, contributes to goal-directed navigation by encoding prospective paths and decision-making within the cognitive map, allowing for flexible planning and adaptation to novel routes.^[24] Neuroimaging studies in humans, including functional MRI (fMRI) evidence from the 2010s and 2020s, have revealed analogous map-like activity during virtual navigation tasks. These demonstrate grid-cell-like representations in the entorhinal cortex and place-cell-like activity in the hippocampus, activated when participants navigate or imagine movement through virtual environments, mirroring rodent findings and extending the cognitive map framework to human spatial cognition.^[25]

Representational Theories

Representational theories of cognitive maps propose diverse formats for how spatial environments are internally encoded, emphasizing structures that support navigation, memory, and decision-making. These models range from rigid geometric layouts to flexible relational networks, often integrating multiple representational layers to handle varying levels of environmental complexity. Seminal frameworks highlight the brain's capacity to construct overlapping or parallel maps, balancing precision with adaptability. A key distinction in representational theories lies between topological and Euclidean maps. Topological maps represent the environment as graph-like structures, capturing connectivity, order, and qualitative relations among locations without specifying exact distances or angles, which enables efficient route following in uncertain or changing settings. In contrast, Euclidean maps encode precise metric information, such as distances and directions, akin to a scaled diagram, facilitating accurate shortcutting and distance estimation. This dichotomy allows cognitive maps to support both coarse-grained exploration and fine-tuned localization. Poucet's hierarchical model (1993) integrates these formats, positing that cognitive maps begin with topological information for basic spatial organization—such as adjacency and sequence—and layer on Euclidean details for enhanced precision, particularly in familiar environments.^[26] This structure is thought to emerge across brain regions, with topological elements providing a flexible skeleton for navigation that metric components refine. Kuipers (1978) similarly emphasized topological representations as foundational, derived from sensory experiences of paths and landmarks, evolving into more Euclidean forms through accumulated metric learning.^[27] Parallel map theory extends this by proposing multiple, dissociable mapping systems operating concurrently in the hippocampus. Jacobs and Schenk (2003) describe a bearing map, primarily in the dentate gyrus, which tracks directional bearings from self-motion cues to support path integration, and a sketch map in the CA fields, built from landmark positions to form a holistic layout; these parallel systems overlap to resolve ambiguities in spatial encoding.^[28] This theory accounts for how animals and humans combine idiothetic (internal) and allothetic (external) information without relying on a single unified representation. Vector-based models conceptualize cognitive maps as accumulations of directional vectors encoding paths, displacements, and orientations, serving as a basis for dead-reckoning and goal vector navigation. These approaches treat spatial knowledge as composable vector fields, where trajectories are represented by summed displacement vectors from starting points to targets, enabling flexible recombination for novel routes. Such models align with neural path integration mechanisms, where head-direction and speed signals generate vector updates to maintain positional estimates. Starting in the 2010s and continuing into the 2020s, computational models advanced representational theories by integrating cognitive maps with reinforcement learning (RL) for goal-directed mapping. These frameworks employ successor representations—matrices encoding predicted future states from current positions—to form dynamic cognitive maps that optimize planning by linking spatial structure to reward anticipation. For instance, model-based RL variants use cognitive maps to simulate trajectories and evaluate policies, allowing efficient generalization across environments. This integration posits cognitive maps as predictive tools that adapt via RL algorithms, supporting both spatial navigation and abstract inference in value-based decisions.

Evidence in Animals

Studies in Rodents

Early studies on cognitive maps in rodents drew heavily from Edward Tolman's experiments using maze paradigms, where rats demonstrated the ability to form internal representations of spatial layouts that allowed for flexible navigation. In landmark experiments conducted in the 1940s, Tolman and colleagues trained rats to navigate elevated mazes to reach food rewards, revealing latent learning: rats that explored without reinforcement could quickly adapt and take shortcuts when rewards were relocated or barriers removed, indicating an updated cognitive map rather than mere stimulus-response habits.^[13] Subsequent work through the 1970s extended this by showing remapping after environmental alterations, such as changing maze configurations or cue placements, where rats' path choices reflected reorganization of their spatial knowledge without retraining.^[1] Advancements in electrophysiological recordings provided direct neural evidence for cognitive maps via place cells in the hippocampus. In a seminal 1987 study, Muller and Kubie recorded from hippocampal neurons in rats as they foraged in controlled environments, finding that place cells remap—exhibiting entirely new firing patterns—when introduced to novel arenas, while maintaining stable spatial selectivity in familiar ones over repeated exposures.^[29] This remapping underscores the dynamic nature of cognitive maps, allowing rodents to distinguish distinct contexts and update representations efficiently. Grid cells in the medial entorhinal cortex further illuminate map dynamics. For instance, research demonstrated that grid cell modules—groups firing at different spatial scales—can realign through rotation or rescaling in response to subtle environmental changes, such as resizing enclosures, preserving overall metric structure while adapting to new contexts.^[30] Recent experiments have enabled precise control over rodent exploration, highlighting how self-initiated behaviors drive cognitive map formation in mice. In 2024 experiments, mice navigating environments formed allocentric maps based on self-motion cues alone, shortcutting to goals after learning, which illustrates the emergence of flexible spatial representations without physical landmarks.^[31]

Cross-Species Comparisons

Cognitive maps, as internal representations of spatial environments, exhibit variations across non-rodent species, reflecting adaptations to diverse ecological niches. In primates, such as monkeys, hippocampal activity supports navigation similar to that observed in rodents, with neurons firing based on spatial views rather than egocentric positions. For instance, studies in the 2000s and 2010s recorded place-like and view cells in the macaque hippocampus during virtual and real-world tasks, enabling flexible path planning and detour navigation.^[32]^[33]^[34] In birds, particularly food-caching species like Clark's nutcrackers, cognitive maps underpin precise cache recovery, allowing birds to remember thousands of locations over months. The hippocampus in these species is disproportionately enlarged compared to non-caching relatives, correlating with enhanced spatial memory for scattered food sites. Migratory birds, such as homing pigeons, also demonstrate hippocampal involvement in map-based navigation, integrating geomagnetic and visual cues for long-distance orientation. These abilities build on rodent-like hippocampal mechanisms but adapt to aerial and caching demands.^[35]^[36]^[37] In insects, exemplified by desert ants, rely primarily on path integration for navigation, using an internal odometer to track distance and direction via stride counting and celestial cues, rather than holistic cognitive maps. This system enables efficient homing over flat terrains but shows limitations in complex environments, where view-matching supplements integration without evidence of flexible remapping typical of vertebrate cognitive maps.^[38]^[39] Evolutionary trends in cognitive mapping correlate with brain region size and ecological pressures; species with high foraging demands, like food-storing birds and navigating primates, show expanded hippocampal volumes relative to body size, enhancing spatial memory for resource distribution. In contrast, territorial behaviors in insects favor simpler integration over expansive mapping, suggesting that cognitive map complexity scales with environmental variability and energy costs of larger brains.^[40]^[37]

Extensions and Applications

Humans employ cognitive maps during wayfinding to integrate spatial information from landmarks, routes, and environmental cues, enabling efficient navigation in familiar and novel settings. In the GPS era, reliance on satellite-based navigation systems has shifted behaviors away from internal cognitive map construction toward external guidance, potentially diminishing spatial knowledge acquisition. A systematic review and meta-analysis found that frequent GPS use negatively impacts environmental knowledge and sense of direction, though its effect on actual wayfinding performance remains limited.^[41] Studies indicate that GPS users often bypass developing allocentric representations, treating devices as substitutes for internal spatial cognition, which can lead to poorer route planning without technological aid.^[42] Sex differences influence navigation strategies linked to cognitive map utilization, with men typically favoring route-based, vector-oriented approaches that emphasize directional and distance metrics, while women more commonly adopt landmark-centric strategies focused on sequential cues and relational positions. Research on route-learning tasks shows that males exhibit fewer errors in configurational navigation, whereas females demonstrate superior recall of landmarks both on and off paths, suggesting differential reliance on cognitive map elements.^[43]^[44] These patterns persist across contexts, moderated by factors like motivation, but highlight how cognitive maps support varied behavioral adaptations in human spatial decision-making.^[45] Impairments in cognitive map formation underlie navigation deficits in neurological disorders such as Alzheimer's disease (AD) and topographical disorientation. In AD, early-stage patients display profound disruptions in allocentric navigation, relying excessively on egocentric cues due to hippocampal atrophy that degrades cognitive map integrity, manifesting as disorientation in familiar environments.^[46]^[47] Topographical disorientation, whether developmental or acquired, involves a selective inability to construct or retrieve cognitive maps, resulting in frequent getting lost even in highly familiar surroundings, as affected individuals fail to integrate landmarks into coherent spatial representations.70133-9/fulltext)^[48] Training interventions, including video game play, can enhance cognitive map development and navigation proficiency. Studies from the 2010s and later demonstrate that action video games improve visuospatial competencies, with gamers showing superior performance in virtual navigation tasks that require building and using cognitive maps for orientation and path integration.^[49]^[50] Participants trained on navigation-focused games exhibit more efficient strategies, such as reduced reliance on trial-and-error and better allocentric processing, indicating transferable benefits to real-world wayfinding. Cultural practices among indigenous navigators, such as Polynesians, exemplify advanced cognitive map applications through dead reckoning and environmental cue integration. Polynesian wayfinders construct mental maps incorporating stellar paths, ocean swells, and wind patterns to traverse vast Pacific distances without instruments, maintaining positional awareness via continuous path integration.^[51] This etak system, akin to dead reckoning, relies on imagined reference points to update cognitive maps dynamically, enabling precise landfalls over thousands of kilometers.^[52] Cognitive maps, originally conceptualized for spatial navigation, have been extended to represent abstract relational structures that support reasoning beyond physical environments. In these abstract maps, the brain employs geometric principles akin to spatial layouts to organize non-spatial information, such as event sequences in planning and decision-making. For instance, hippocampal and entorhinal representations encode relational distances between abstract concepts, enabling flexible inference and adaptive behavior in tasks requiring sequence prediction or conceptual navigation.^[53] This framework allows individuals to simulate future scenarios by traversing these internal maps, much like path integration in space, thereby facilitating goal-directed actions in complex, non-physical domains.^[54] Social cognitive maps further generalize this representational scheme to interpersonal dynamics, capturing networks of relationships and hierarchies as structured geometries. These maps encode social knowledge—such as alliances, status rankings, and influence patterns—in a manner analogous to spatial grids, with the medial temporal lobe supporting distance-based metrics between individuals or groups.^[55] For example, people infer unseen social connections by mentally navigating these maps, using proximity in relational space to predict behaviors or outcomes in group settings, similar to shortcutting in physical environments.^[56] Such representations promote efficient social navigation, allowing adaptation to evolving interpersonal contexts without exhaustive direct interactions.^[57] Recent advancements have integrated cognitive map principles into artificial intelligence for enhanced decision-making, where neural networks construct predictive maps from sensory data to model relational structures. These AI models, inspired by hippocampal mechanisms, generate internal simulations for planning in uncertain environments, improving efficiency in tasks like reinforcement learning or multi-step reasoning.^[58] Additionally, emerging research highlights curiosity's role in exploration and map refinement, driving agents to seek novel information that updates social and abstract representations; for instance, intrinsic motivation correlates with more accurate relational encoding during social interactions.^[59] These extensions underscore a key gap in traditional views: cognitive maps integrate with episodic memory to bind relational knowledge across experiences, forming cohesive schemas for long-term abstraction and retrieval.^[60]

Criticisms and Alternatives

Limitations of the Framework

The cognitive map framework has faced criticism for its overreliance on the "map" analogy, which suggests a precise, Euclidean representation of spatial layouts akin to a cartographic tool, yet empirical evidence indicates that such internal representations are often distorted and biased. Studies from the 1970s, including those examining urban environments, revealed systematic distortions in cognitive maps, such as alignment biases toward major landmarks or rotation heuristics that skew perceived distances and directions, rather than faithful metric accuracy.^[61] These findings, led by geographers like Reginald Golledge, underscore how cognitive representations prioritize functional anchors and hierarchies over geometric fidelity, limiting the analogy's applicability to real-world navigation.^[62] Empirical gaps further weaken the framework, as certain navigation behaviors—such as taking novel shortcuts—can be parsimoniously explained by serial order memory or chained response sequences without necessitating a holistic map. For example, animal studies purporting to demonstrate mapping have been critiqued for failing to rule out simpler associative mechanisms, where sequences of landmarks or paths are recalled linearly rather than reconfigured flexibly.^[63] This alternative accounts for observed efficiencies in route-following tasks, suggesting that the evidence for true map-like inference remains inconclusive.^[64] A core methodological challenge lies in inferring cognitive maps from behavioral data, which assumes unobservable internal structures and risks conflating external actions with specific representational formats. Critics highlight that behaviors like detours or reorientations could stem from multiple underlying processes, such as beacon-based guidance or habitual chaining, making direct attribution to maps speculative and unverifiable without invasive neural measures.^[63] This inferential leap has persisted as a limitation, complicating validation across species and contexts. Post-2000s advancements in connectionist models have intensified these critiques by showing that spatial cognition and navigation can arise from distributed neural network dynamics, bypassing the need for explicit, symbolic map structures central to the original theory. These models demonstrate emergent route planning and generalization through weighted connections and pattern completion, challenging the framework's emphasis on discrete, metric embeddings as essential for flexible behavior.^[65]

Competing Models

Serial order theories posit that spatial navigation can be achieved through the memorization of sequential routes or chains of landmarks and actions, obviating the need for a holistic, Euclidean cognitive map. These models emphasize taxon-like strategies where animals follow familiar paths or respond to successive cues in a linear fashion, allowing for efficient traversal without geometric inference. Bennett (1996) argued that classic evidence for cognitive maps, such as novel shortcutting in rodents and insects, can be parsimoniously explained by route-based mechanisms, including recognition of landmarks from novel angles or path integration for short deviations, rather than internal surveying. For example, in experiments with dogs and honeybees, apparent spatial inference behaviors were attributable to sequential landmark chaining rather than map-like representations.^[63] Associative models, rooted in behaviorist principles, account for spatial cognition via direct links between environmental cues and responses, without invoking internal spatial representations. Elemental associative learning associates single cues, such as a prominent landmark, directly with a goal location, enabling navigation through stimulus-response pairings. Configural variants extend this by treating combinations of cues as unique gestalts that trigger specific behaviors, still avoiding global maps. Buatois and Gerlai (2020) demonstrated elemental and configural associative learning in zebrafish using a spatial task akin to a plus-maze, where performance involves cue-goal associations and spatial memory potentially mediated by the hippocampal homolog (lateral pallium), paralleling associative processes in rodent spatial navigation tasks like the Morris water maze. Such models explain robust navigation in cue-rich environments through incremental strengthening of associations, as formalized in theories like Rescorla-Wagner.^[66] Connectionist approaches employ artificial neural networks to model spatial navigation, generating map-like behaviors through distributed, emergent representations rather than predefined structures. These models use recurrent networks to encode spatial adjacencies or temporal sequences from experience, with error-driven learning adjusting weights to simulate route planning and subgoal formation. Arleo et al. (1998) proposed a framework where recurrent associative nets store environmental knowledge as predictive patterns, reinjected to support imagination and flexible navigation, akin to hippocampal function without explicit grids or place codes. Integrated into AI from the 1990s to recent decades, such networks have replicated animal homing and human wayfinding in simulations, highlighting adaptability without rigid cognitive maps.^[67] Hybrid views conceptualize cognitive maps as emergent properties arising from the interplay of distinct subsystems, such as path integration for self-motion tracking and landmark cues for external anchoring, rather than a unified framework. Path integration provides continuous egocentric updates prone to drift, which landmarks correct via allocentric referencing, yielding flexible spatial knowledge through Bayesian-like integration. Wiener et al. (2024) showed in virtual reality tasks that humans optimally combine these systems for homing accuracy with few landmarks (up to three), but shift to alternation or PI dominance in cluttered scenes, where maps form dynamically from cue competition. This perspective reconciles multiple navigation modes, explaining individual variability and environmental adaptability without relying on a singular map construct.^[68]

References

[1]
[PDF] Cognitive Maps in Rats and Men
EDWARD C. TOLMAN. G had not exhibited any stimulus-response propensities to go more to the side which became later the side of the ap- propriate goal. There ...
[2]
COGNITIVE MAPS - American Psychological Association
Tolman (1948) noted that cognitive maps allow rats to use place hypotheses. What this means, as we have just seen, is that rats can use their cognitive maps ...
[3]
The cognitive map in humans: Spatial navigation and beyond - PMC
The 'cognitive map' hypothesis proposes that brain builds a unified representation of the spatial environment to support memory and guide future action.
[4]
What Is a Cognitive Map? Organizing Knowledge for Flexible Behavior
Oct 24, 2018 · For Tolman, this cognitive map was a systematic organization of knowledge that spanned all domains of behavior (Tolman, 1948). However, its ...Main Text · Generalizing Spatial... · Unifying Spatial And...
[5]
[PDF] purposive behavior - animals and men
Tolman to expect Purposive Behavior in Animals and Men to be superseded by 'better' and more modern theories and systems, concluded what we had always ...
[6]
https://mitpress.mit.edu/9780262620017/the-image-of-the-city/
[7]
https://doi.org/10.1038/s41583-022-00655-9
[8]
The mentality of apes. - APA PsycNet
The mentality of apes. Citation. Kohler, W. (1925). The mentality of apes. (E. Winter, Trans.). Harcourt, Brace.
[9]
https://doi.org/10.1037/h0061626
[10]
[PDF] EDWARD CHACE TOLMAN - Biographical Memoirs
That is, I was already becoming influenced by Gestalt psychology and conceived that a rat in running a maze must be learning a lay-out or pattern and not ...
[11]
[PDF] The cognitive revolution: a historical perspective - cs.Princeton
The behavioral revolution transformed experimental psychology in the US. Perception became discrimination, memory became learning, language became verbal beha- ...
[12]
[PDF] The Hippocampus as a Cognitive Map - Faculty
learned or constructed. This obviously only applies to psychological space. By innate or a priori is meant that the structure and function of those parts of.
[13]
COGNITIVE MAPS IN RATS AND MEN[1] Edward C. Tolman (1948)
Tolman (1948). First published in The Psychological Review, 55(4), 189-208. I shall devote the body of this paper to a description of experiments with rats.Missing: primary | Show results with:primary
[14]
The Need to Fuse Boundary Knowledge with Path Integration
We show firstly that it is theoretically implausible for path integration alone to maintain a stable positional representation beyond three minutes.Missing: seminal | Show results with:seminal
[15]
Visual influence on path integration in darkness indicates a ... - PNAS
Overall, our findings suggest that a combined multimodal representation guides large-scale navigation, consistent with a role for visual imagery or a cognitive ...Results · Discussion · Multimodal Representation...
[16]
Timing and Rate of A-Not-B Performance Gains and EEG Maturation
This study examined how timing (ie, relative maturity) and rate (ie, how quickly infants attain proficiency) of A-not-B performance were related to changes in ...
[17]
The Development of Human Navigation in Middle Childhood
Aug 18, 2022 · The main result of this review showed how middle childhood is a crucial period for the improvement and development of allocentric strategies, ...Missing: shift | Show results with:shift
[18]
https://doi.org/10.1111/desc.12496
[19]
The Aging Navigational System - PMC - NIH
The following sections will provide an in-depth discussion of the behavioral and neural mechanisms that cause specific navigational impairments in old age.
[20]
The hippocampus as a spatial map. Preliminary evidence from unit ...
The hippocampus as a spatial map. Preliminary evidence from unit activity in the freely-moving rat. Author links open overlay panelJ. O'Keefe, J. Dostrovsky.
[21]
Microstructure of a spatial map in the entorhinal cortex - Nature
Jun 19, 2005 · Grid cells have tessellating firing fields. To visualize the spatial structure of firing fields in dMEC neurons, we first tested the rats in a ...
[22]
Flexible egocentric and allocentric representations of heading ...
Mar 19, 2018 · We report a flexible switching between egocentric and allocentric reference frames in a subpopulation of VIP neurons, depending on gaze strategy.
[23]
https://www.pnas.org/doi/10.1073/pnas.1715625115
[24]
The effects of changes in the environment on the spatial ... - PubMed
Using the techniques set out in the preceding paper (Muller et al., 1987), we investigated the response of place cells to changes in the animal's environment.Missing: novel stability familiar
[25]
Modular Realignment of Entorhinal Grid Cell Activity as a Basis for ...
Jun 22, 2011 · ... cells in exploring rodents, can undergo large changes in relative location during remapping. ... spin glass model of path integration in rat ...
[26]
Shortcutting from self-motion signals reveals a cognitive map in mice
Nov 11, 2024 · Hippocampal neurons of rats randomly foraging in a real world or virtual reality ... (2023) Interactions between rodent visual and spatial ...
[27]
Moculus: an immersive virtual reality system for mice incorporating ...
Dec 12, 2024 · Early rodent systems used flat monitors with predefined optic flow, whereas later systems with panoramic projectors incorporated treadmill-based ...
[28]
Neural Correlates of Spatial Navigation in Primate Hippocampus - NIH
Nov 2, 2022 · We review empirical studies examining the neural correlates of spatial navigation in the primate (including human) hippocampus at the levels of local field ...
[29]
Spatial representations in the primate hippocampus, and their ...
Hippocampal spatial view neurons in primates respond to the place where a monkey is looking, with some modulation by place. In contrast, hippocampal neurons ...
[30]
Significance of visual scene‐based learning in the hippocampal ...
Dec 2, 2022 · Wirth et al. (2017) also recorded hippocampal neurons in nonhuman primates during virtual navigation (Wirth et al., 2017). They trained monkeys ...1 Introduction · 4 Hippocampal Functions In... · 10 Conclusion And Future...
[31]
Hippocampal specialization of food-storing birds. - PNAS
Feb 15, 1989 · Food-storing species of passerines have an enlarged hippocampal complex as a specialization associated with the use of a specialized memory capacity.
[32]
Neural representations of space in the hippocampus of a food ...
Jul 16, 2021 · Accurate cache retrieval requires the hippocampus, which is enlarged in food-caching birds (2, 3, 26). We designed miniature microdrives ...Missing: recovery | Show results with:recovery
[33]
Integrating ecology, psychology and neurobiology within a food ...
Adaptive specialization of memory and the hippocampus. It has been well established that almost all food-caching species use spatial memory, at least in part, ...
[34]
On the 'cognitive map debate' in insect navigation - ScienceDirect.com
It is true that in contrast to the map proponents we have intensively studied the various compass, odometer, path integration, and view matching strategies ...
[35]
Path integration in a three-dimensional world: the case of desert ants
Feb 4, 2020 · Desert ants use path integration to return from foraging excursions on a shortcut way to their nests. Intriguingly, when walking over hills, the ants ...
[36]
The Evolution of the Cognitive Map - Karger Publishers
Aug 29, 2003 · This cognitive map thus appears to underlie long-distance navigation. Its mediation by the hippocampal formation and its presence in birds and ...Missing: demands | Show results with:demands
[37]
GPS use and navigation ability: A systematic review and meta-analysis
Although evidence suggests that using GPS tools can have a negative impact on environmental knowledge and sense of direction but a limited effect on wayfinding.Missing: childhood rural<|control11|><|separator|>
[38]
How is GPS used? Understanding navigation system use and its ...
Mar 19, 2024 · ... cognitive maps, so they use GPS to replace internal spatial knowledge. In contrast, as the use of GPS for time and traffic estimates is ...
[39]
The Role of Gender and Familiarity in a Modified Version of ... - NIH
May 22, 2021 · For instance, men rely on global reference points and configurational strategies, while females more often use landmarks and procedural ...
[40]
Sex differences in route-learning - ScienceDirect.com
As expected, males made fewer errors and took fewer trials to reach criterion. Females remembered more landmarks both on and off the route than males, and ...
[41]
Motivation moderates gender differences in navigation performance
Sep 25, 2023 · Previous research suggests that men outperform women in navigation tasks and that men and women exhibit different navigation strategies. Here, ...
[42]
https://pmc.ncbi.nlm.nih.gov/articles/PMC10951145/
[43]
Different Profiles of Spatial Navigation Deficits In Alzheimer's ...
Background: Spatial navigation impairment is a promising cognitive marker of Alzheimer's disease (AD) that can reflect the underlying pathology.
[44]
Topographical Disorientation: Clinical and Theoretical Significance ...
Sep 19, 2019 · Individuals unable to form a cognitive map get lost more frequently than individuals who are able to form and use cognitive maps of the ...
[45]
The cognitive effects of playing video games with a navigational ...
Our study provides important evidence suggesting that the use of video game training programs influence spatial navigation and orientation abilities. Previous ...
[46]
Positive Effects of Videogame Use on Visuospatial Competencies
May 30, 2019 · Previous research show that training with videogames enables children and adolescents to improve their scores in visuospatial tests.
[47]
Wayfinding and Navigation - University of Hawaii at Manoa
The early Polynesian voyagers were some of the best wayfinders in history (Fig. 8.3). They were able to find their way across vast reaches of the Pacific ocean ...Missing: cognitive | Show results with:cognitive
[48]
A View from the Islands: Spatial Cognition in the Western Pacific - jstor
The most complex Micronesian navigation strategy described is a dead reckoning system which uses an imagined reference island (etak), a star compass point ...
[49]
Navigating cognition: Spatial codes for human thinking - Science
Nov 9, 2018 · Bellmund et al. review and combine concepts from cognitive science and philosophy with findings from neurophysiology of spatial navigation in ...
[50]
Constructing, Combining, and Inferring on Abstract Cognitive Maps
Sep 23, 2020 · These findings reveal how abstract and discrete relational structures are represented, are combined, and enable novel inferences in the human brain.
[51]
Transforming social perspectives with cognitive maps
Growing evidence suggests that cognitive maps represent relations between social knowledge similar to how spatial locations are represented in an environment.
[52]
Cognitive maps of social features enable flexible inference ... - PNAS
Sep 13, 2021 · Our results reveal that cognitive maps play a powerful role in shaping how people represent and reason about relationships in a social network.
[53]
Abstract cognitive maps of social network structure aid adaptive ...
We propose a theory of network representation that explains how people learn inferential cognitive maps of social relations from direct observation.
[54]
Automated construction of cognitive maps with visual predictive coding
Jul 18, 2024 · Here we demonstrate that predictive coding provides a natural and versatile neural network algorithm for constructing spatial maps using sensory data.
[55]
Curiosity shapes spatial exploration and cognitive map formation in ...
Dec 30, 2024 · These results provide new evidence for a link between curiosity and exploratory behaviour, and how curiosity might shape cognitive map formation.
[56]
Cognitive mapping and episodic memory emerge from simple ...
Aug 28, 2024 · Through cognitive mapping of objects organized in context, episodic memory provides the associative substrates for autonoetic consciousness ( ...
[57]
Systematic Distortions in Urban Cognitive Maps - jstor
We argue that systematic distortions are related to a rotation heuristic and to key reference points used by the subjects to code and access spatial information ...
[58]
Exploring the anchor-point hypothesis of spatial cognition
The anchor-point hypothesis of spatial cognition, according to which primary nodes or reference points anchor distinct regions in cognitive space.
[59]
Do Animals Have Cognitive Maps? | Journal of Experimental Biology
Jan 1, 1996 · According to the Tolman, O'Keefe and Nadel definition, a cognitive map would be indicated by the ability to short-cut from A to B, or vice versa ...Introduction · Choices And Constraints · The Cognitive Maps Of Tolman...<|separator|>
[60]
Do animals have cognitive maps? - PubMed
The author argues that no animal has been conclusively shown to have a cognitive map, and the term should be avoided due to simpler explanations.
[61]
Connectionism - Stanford Encyclopedia of Philosophy
May 18, 1997 · Connectionist models provide a new paradigm for understanding how information might be represented in the brain. A seductive but naive idea is ...
[62]
Elemental and Configural Associative Learning in Spatial Tasks
Spatial learning represents a unique subset of relational/configural learning in which the subject establishes its position relative to spatial cues, and using ...Missing: behaviorist | Show results with:behaviorist
[63]
Spatial and temporal cognitive mapping: a neural network approach
We describe how both spatial and temporal cognitive maps can be mechanistically implemented in terms of recurrent associative networks.
[64]
Not seeing the forest for the trees: combination of path integration ...
Two of the most important cues in navigation are the spatial arrangement of landmarks in the environment, and the continuous path integration of travelled ...