From cognitive maps to spatial schemas
Open accessAbstract
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.
AI snapshot
Written from the paper’s full text.
TL;DR
The review proposes that spatial schemas are superordinate, generalizable spatial representations formed by integrating spatial gists across multiple similar environments, distinct from environment-specific cognitive maps and event schemas. It synthesizes rodent, human lesion/neuroimaging, and urban-analytics evidence to outline cognitive and neural mechanisms, with HPC–neocortical interactions and extra-HPC hubs such as mPFC, RSC, ERC, and PPC implicated.
Key findings
- Spatial schemas are proposed as extra-HPC, environment-independent representations that support navigation in novel environments sharing geometric/visuospatial properties; they are distinct from cognitive maps and spatial gists, which remain environment-specific and can be kept distinct via pattern separation.
- Rodent studies: rats with HPC lesions more than 24 hours after paired-associate learning retained original and novel paired associates, but lesions before 24 hours disrupted acquisition; in the village maze, HPC-lesioned rats with extensive pre-lesion experience learned novel rewarded locations as quickly as controls, whereas inexperienced HPC-lesioned rats took much longer, and performance depended on stable room geometry rather than distal cues.
- Human evidence: K.C., an individual with bilateral HPC damage, produced accurate sketch maps of his hometown’s overall layout and judged relative distances/directions between major landmarks, but was impaired for minor landmarks and perceptual details; an fMRI analogue showed retrieval activity shifting from HPC to vmPFC after 24 hours and to vlPFC by 90 days, with accelerated consolidation for congruent novel associates from days 91–105.
- Sea Hero Quest data from about 3.9 million people worldwide found that those who grew up in grid-like cities were generally worse navigators than those from disorganized cities or outside cities, but had a slight advantage in grid-like virtual environments; street network entropy values cited include Prague 3.6 and Chicago 2.5.
Methods
- Narrative review/conceptual synthesis; no new primary data were collected. It integrates prior theoretical frameworks and empirical findings from rodent spatial navigation, human lesion/amnesia, fMRI, virtual navigation, and urban analytics/space syntax.
- Evidence discussed includes rodent paradigms (event arena, village maze, HexMaze), human cases (e.g., K.C.), neuroimaging studies, and large-scale virtual navigation data (Sea Hero Quest, ~3.9 million participants, 38 countries). Main analysis is qualitative integration, not meta-analysis.
Limitations
- Stated: the concept of spatial schema is still emerging; schema operationalizations are heterogeneous (statistical rule learning, associative inference, paired-associate learning, prior knowledge retrieval), and specific experimental paradigms for spatial schemas are needed.
- Stated: systematic measurement of quantitative differences in real-world spatial organization is a major challenge; future work must determine where and how detailed and schematic spatial representations coexist and whether new schemas can be acquired/updated after HPC lesions.
- Evident: as a review, it does not directly test the proposed neurocognitive framework; many neural mechanisms and distinctions are proposed or extrapolated from event-schema and animal work, and some cited evidence is indirect.
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Claims the paper makes
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- Premisecitation
Similar parts of one environment or multiple environments are represented using generalizable spatial codes.
“similar parts of one environment or multiple environments seem to be represented using generalizable spatial codes [14][15][16][17][18][19] , which refer to a set of abstract spatial relations that can be used flexibly to facilitate navigation.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [14][15][16][17][18][19]
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- Premisecitation
The HPC can store unique cognitive maps of different environments or of altered versions of the same environment.
“The HPC can store unique cognitive maps of different environments or of altered versions of the same environment 2,[10][11][12][13] .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites 2,[10][11][12][13]
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- Interpretationcitation
Spatial schemas would be used to guide navigation by enhancing expectations in predictive coding.
“In both cases, spatial schemas would be used to guide navigation by enhancing expectations in predictive coding [G] 28 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [28] de Lange, F. P., Heilbron, M. & Kok, P. How do expectations shape perception? Trends Cogn. Sci. 22, 764–779 (2018). source
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- Interpretationnone given
Experiences during navigation with many similarly structured environments give rise to the formation of spatial schemas.
“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.”
Quote in the paperNot a substantial claim - Premisecitation
In participants navigating novel virtual reality environments, segments that overlapped with familiar environments learned the day before produced proactive interference.
“A more recent study similarly showed proactive interference in participants who were navigating novel virtual reality environments that contained segments that overlapped with familiar environments learned the day before.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [37] He, Q., Beveridge, E. H., Starnes, J., Goodroe, S. C. & Brown, T. I. Environmental overlap and individual encoding strategy modulate memory interference in spatial navigation. Cognition 207, 104508 (2021). source
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- Interpretationcitation
The relationship between schemas and distinct elements is bidirectional: by pointing to elements that are in common, schemas bring into focus what is distinct.
“We view this as a bidirectional process: by pointing to elements that are in common, schemas bring into focus what is distinct 67 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [67] Bein, O., Reggev, N. & Maril, A. Prior knowledge promotes hippocampal separation but cortical assimilation in the left inferior frontal gyrus. Nat. Comm. 11, 4590 (2020). source
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- Premisecitation
New paired associates consistent with learned associations were acquired in only a single trial in rats.
“New paired associates that were consistent with learned associations were acquired in only a single trial.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [53] Tse, D. et al. Schemas and memory consolidation. Science 316, 76–82 (2007). This study in rodents with hippocampal lesions provides evidence for spatial schemas and the involvement of the hippocampus in rapid learning. source
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- Interpretationcitation
The observed learning gains in these studies were attributed to the formation of a spatial schema.
“The observed learning gains in these studies were attributed to the formation of a spatial schema 53,54 (for a recent theoretical framework based on these findings, see ref. 36 ).”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [53] Tse, D. et al. Schemas and memory consolidation. Science 316, 76–82 (2007). This study in rodents with hippocampal lesions provides evidence for spatial schemas and the involvement of the hippocampus in rapid learning. source
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- cites [54] Tse, D. et al. Schema-dependent gene activation and memory encoding in neocortex. Science 333, 891–895 (2011). source
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- Premisecitation
Experienced London taxi drivers had considerable difficulty learning a novel version of London that differed from their previously learned version, unlike non-expert controls.
“Performance on spatial memory tests, such as drawing the layout of the novel environment in a sketch map and recognizing scenes from photographs, indicated that, unlike non-expert controls, the taxi drivers had considerable difficulty learning a novel version of London that differed from their previously learned version of London 68 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [68] Woollett, K. & Maguire, E. A. The effect of navigational expertise on wayfinding in new environments. J. Environ. Psychol. 30, 565–573 (2010). source
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- Interpretationcitation
Spatial schemas do not depend on the HPC for their maintenance or retrieval.
“extrapolating from event schemas 22 , which do not rely on the HPC, we believe spatial schemas do not depend on the HPC for their maintenance or retrieval.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [22] Gilboa, A. & Marlatte, H. Neurobiology of schemas and schema-mediated memory. Trends Cogn. Sci. 21, 618–631 (2017). source
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- Interpretationcitation
The interfering effects of the original environment suggest that a cognitive map or gist of the environment had formed.
“The interfering effects of the original environment might suggest that a cognitive map or gist of the environment had formed, as is suggested by additional evidence that proactive interference increases as a function of cognitive mapping and self-reported sense of direction 37 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [37] He, Q., Beveridge, E. H., Starnes, J., Goodroe, S. C. & Brown, T. I. Environmental overlap and individual encoding strategy modulate memory interference in spatial navigation. Cognition 207, 104508 (2021). source
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- Premisecitation
Upregulated expression of immediate early genes in neocortex, including mPFC and RSC, was associated with more rapid acquisition of novel paired associates in nonlesioned rats exposed to original paired associates.
“In a follow-up study, upregulated expression of immediate early genes in several regions of neocortex, including the medial prefrontal cortex (mPFC) and retrosplenial cortex (RSC), was associated with more rapid acquisition of novel paired associates in (nonlesioned) rats that were exposed to the original paired associates 54 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [54] Tse, D. et al. Schema-dependent gene activation and memory encoding in neocortex. Science 333, 891–895 (2011). source
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- Premisecitation
HPC lesions disrupted initial acquisition if applied before 24 hours.
“However, HPC lesions did disrupt initial acquisition if they were applied before the representation had an opportunity to undergo rapid consolidation; that is, before 24 hours.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [53] Tse, D. et al. Schemas and memory consolidation. Science 316, 76–82 (2007). This study in rodents with hippocampal lesions provides evidence for spatial schemas and the involvement of the hippocampus in rapid learning. source
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- Interpretationcitation
Prior knowledge can interfere with the acquisition of spatial knowledge for schema-incongruent elements of a novel environment.
“These findings provide indirect evidence that prior knowledge can interfere with the acquisition of spatial knowledge for schema-incongruent elements of a novel environment.”
Quote in the paperSubstantial claim - Premisecitation
HPC lesions more than 24 hours after paired-associate learning did not disrupt recall of original or novel paired associates.
“HPC lesions more than 24 hours after paired-associate learning did not disrupt the recall of original or novel paired associates.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [53] Tse, D. et al. Schemas and memory consolidation. Science 316, 76–82 (2007). This study in rodents with hippocampal lesions provides evidence for spatial schemas and the involvement of the hippocampus in rapid learning. source
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- Premisecitation
An acquired cognitive map of a particular environment interferes with learning a novel environment that possesses features inconsistent with previous knowledge.
“Evidence suggests that an acquired cognitive map of a particular environment interferes with learning a novel environment that possesses features that are inconsistent with previous knowledge.”
Quote in the paperSubstantial claim - Premisecitation
In the HexMaze, the passage of time, rather than frequency of training, is responsible for accumulation of spatial knowledge into a long-term representation.
“The more recently developed 'HexMaze', a large-scale, complex, gangway maze experienced by mice for different durations, has been used to show that the passage of time, rather than the frequency of training, is responsible for the accumulation of spatial knowledge into a long-term representation 69 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [69] Alonso, A. et al. The HexMaze: a previous knowledge task on map learning for mice. eNeuro 8, ENEURO.0554–20.2021 (2021). source
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- Interpretationreasoning
Rapid updating in experienced HPC-lesioned rats is consistent with embedding novel paths into a schema of the environment, rather than completely new learning.
“Such rapid updating as seen in the experienced HPC-lesioned rats is consistent with the embedding of the novel paths to a goal into a schema of the environment, rather than with completely new learning of novel paths.”
Quote in the paperSubstantial claim - Premisecitation
Human HPC representations for context-item associations accumulated over multiple learning trials generalize across events that share similar item-context associations.
“Human HPC representations for context-item associations that are accumulated over multiple learning trials also seem to generalize across events that share similar item-context associations 79 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [79] Libby, L. A., Reagh, Z. M., Bouffard, N. R., Ragland, J. D. & Ranganath, C. The hippocampus generalizes across memories that share item and context information. J. Cogn. Neurosci. 31, 24–35 (2019). source
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- Premisecitation
Once knowledge was semanticized, vmPFC was active during retrieval of subsequently learned novel associates (day 91 to 105) congruent with previously learned associates.
“Once knowledge was semanticized, the vmPFC was found to be active during retrieval of subsequently learned novel associates (from day 91 to 105) that were congruent with previously learned associates.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [72] Sommer, T. The emergence of knowledge and how it supports the memory for novel related information. Cereb. Cortex 27, 1906–1921 (2017).
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- Premisecitation
Schematic aspects of remote spatial memories are those features that survive HPC damage and continue to support navigation.
“Schematic aspects of remote spatial memories are those features that survive HPC damage and continue to support navigation 27,80 .”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [27] Spiers, H. J. & Maguire, E. A. The neuroscience of remote spatial memory: a tale of two cities. Neuroscience 149, 7–27 (2007). source
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- cites [80] Rosenbaum, R. S. et al. Remote spatial memory in an amnesic person with extensive bilateral hippocampal lesions. Nat. Neurosci. 3, 1044–1048 (2000). source
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- Interpretationreasoning
These findings are consistent with a fine-to-coarse detail gradient for encoding information along the posterior-to-anterior axis of the HPC.
“These findings are consistent with a fine-to-coarse detail gradient for encoding information along the posterior-to-anterior axis of the HPC [76][77][78] (Fig. 1).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [76][77][78]
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- Premisecitation
In consolidation, memories are transformed from highly specific instances to more generalized representations, losing specificity.
“It is important to keep in mind that, in this consolidation process, memories are transformed from highly specific instances to more generalized representations, losing specificity in the process 91 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [91] Ngo, C. T., Benear, S. L., Popal, H., Olson, I. R. & Newcombe, N. S. Contingency of semantic generalization on episodic specificity varies across development. Curr. Biol. 31, 2690–2697.e5 (2021). source
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- Interpretationcitation
Schematized spatial memories primarily include geometric properties rather than surface properties.
“Schematized spatial memories primarily include geometric properties [G], rather than surface properties [G] 27,70,80,83 .”
Quote in the paperSubstantial claim0 of 4 sources checked- cites [27] Spiers, H. J. & Maguire, E. A. The neuroscience of remote spatial memory: a tale of two cities. Neuroscience 149, 7–27 (2007). source
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- cites [70] Winocur, G., Moscovitch, M., Fogel, S., Rosenbaum, R. S. & Sekeres, M. Preserved spatial memory after hippocampal lesions: effects of extensive experience in a complex environment. Nat. Neurosci. 8, 273–275 (2005). source
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- cites [80] Rosenbaum, R. S. et al. Remote spatial memory in an amnesic person with extensive bilateral hippocampal lesions. Nat. Neurosci. 3, 1044–1048 (2000). source
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- cites [83] Rosenbaum, R. S., Ziegler, M., Winocur, G., Grady, C. L. & Moscovitch, M. “I have often walked down this street before”: fMRI studies on the hippocampus and other structures during mental navigation of an old environment. Hippocampus 14, 826–835 (2004). source
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- Interpretationreasoning
The village maze is an experimental animal model of the use of pre-existing spatial schemas in humans.
“The village maze is a rare example of an animal task, designed on the basis of findings in K.C., that serves as an experimental animal model of the use of pre-existing spatial schemas in humans (Fig. 2b).”
Quote in the paperSubstantial claim - Interpretationcitation
The process of generalization probably relies on the formation and use of gist-based representations in the HPC rather than on the formation and use of schemas per se.
“The process of generalization, by contrast, probably relies on the formation and use of gist-based representations in the HPC rather than on the formation and use of schemas per se.”
Quote in the paperSubstantial claim - Interpretationcitation
Generalized representations formed by rodents in the reviewed studies seem to rely on gist-like representations rather than schemas.
“For example, the generalized representations formed by rodents in the studies reviewed above seem to rely on gist-like representations rather than schemas 18,53,54 .”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [18] McKenzie, S. et al. Hippocampal representation of related and opposing memories develop within distinct, hierarchically organized neural schemas. Neuron 83, 202–215 (2014). source
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- cites [53] Tse, D. et al. Schemas and memory consolidation. Science 316, 76–82 (2007). This study in rodents with hippocampal lesions provides evidence for spatial schemas and the involvement of the hippocampus in rapid learning. source
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- cites [54] Tse, D. et al. Schema-dependent gene activation and memory encoding in neocortex. Science 333, 891–895 (2011). source
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- Interpretationcitation
The village maze provided converging evidence that semantic-like spatial representations or spatial schemas exist independently of the HPC.
“It provided converging evidence that semantic-like spatial representations or spatial schemas exist independently of the HPC 70 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [70] Winocur, G., Moscovitch, M., Fogel, S., Rosenbaum, R. S. & Sekeres, M. Preserved spatial memory after hippocampal lesions: effects of extensive experience in a complex environment. Nat. Neurosci. 8, 273–275 (2005). source
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- Premisecitation
K.C. and other individuals with hippocampal amnesia were still able to navigate familiar environments using only a schematic form of their spatial representation.
“Notably, K.C. and other individuals with hippocampal amnesia were still able to navigate familiar environments using only a schematic form of their spatial representation 81,82 .”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [81] Herdman, K. A., Calarco, N., Moscovitch, M., Hirshhorn, M. & Rosenbaum, R. S. Impoverished descriptions of familiar routes in three cases of hippocampal/medial temporal lobe amnesia. Cortex 71, 248–263 (2015). This patient-lesion study provides converging evidence that the hippocampus is needed to support remote spatial memory for visuospatial details but not the gist of environments learned long ago. source
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- cites [82] Rosenbaum, R. S., Cassidy, B. N. & Herdman, K. A. Patterns of preserved and impaired spatial memory in a case of developmental amnesia. Front. Hum. Neurosci. 9, 196 (2015). source
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- Premisecitation
In rats with pre-lesion experience in the complex village maze, allocentric navigation abilities are preserved after HPC lesions.
“As described above, in rats with pre-lesion experience in the complex village maze, allocentric navigation abilities are preserved after HPC lesions 70 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [70] Winocur, G., Moscovitch, M., Fogel, S., Rosenbaum, R. S. & Sekeres, M. Preserved spatial memory after hippocampal lesions: effects of extensive experience in a complex environment. Nat. Neurosci. 8, 273–275 (2005). source
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- Interpretationreasoning
Gist-like representations in the HPC may feed into schematic ones elsewhere in the brain, such as the mPFC in rodents and vmPFC in humans.
“Gist-like representations in the HPC may feed into schematic ones elsewhere in the brain, such as the mPFC in rodents and vmPFC in humans (Box 1).”
Quote in the paperNot a substantial claim - Interpretationcitation
A plausible physiological mechanism by which schemas develop is through the integration of repeated memories with overlapping elements during HPC replay during slow-wave sleep, through coordination of neural network oscillations that facilitate HPC-neocortical interactions.
“A plausible physiological mechanism by which schemas develop is through the integration of repeated memories with overlapping elements during HPC replay [G] during slow-wave sleep (SWS) 89,[95][96][97][98] , through the coordination of neural network oscillations that facilitate HPCneocortical interactions 99,100 .”
Quote in the paperSubstantial claim0 of 3 sources checked- cites 89,[95][96][97][98]
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- cites [99] Maingret, N., Girardeau, G., Todorova, R., Goutierre, M. & Zugaro, M. Hippocampo-cortical coupling mediates memory consolidation during sleep. Nat. Neurosci. 19, 959–964 (2016). source
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- cites [100] Ohki, T. & Takei, Y. Neural mechanisms of mental schema: a triplet of delta, low beta/spindle and ripple oscillations. Eur. J. Neurosci. 48, 2416–2430 (2018). source
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- Premisecitation
The receptive fields of place cells in the rodent HPC are sensitive to specific locations in space.
“Analogous to the selectivity of concept cells, the receptive fields of place cells in the rodent HPC are sensitive to specific locations in space 60,61 .”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites [60] O’Keefe, J. & Dostrovsky, J. The hippocampus as a spatial map. Preliminary evidence from unit activity in the freely-moving rat. Brain Res. 34, 171–175 (1971). source
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- cites [61] Ekstrom, A. D. et al. Cellular networks underlying human spatial navigation. Nature 425, 184 (2003). source
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- Interpretationcitation
This interplay is similar to that between event schemas and representations of event details, which co-exist in both the HPC and neocortex.
“This interplay is similar to that between event schemas and representations of event details, which co-exist in both the HPC and neocortex 94 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [94] Gilboa, A. & Moscovitch, M. No consolidation without representation: correspondence between neural and psychological representations in recent and remote memory. Neuron 109, 2239–2255 (2021). source
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- Premisecitation
Sleep enhances specific instances via HPC circuitry.
“On one hand, sleep enhances specific instances via HPC circuitry 101,102 .”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites [101] Lee, A. K. & Wilson, M. A. Memory of sequential experience in the hippocampus during slow wave sleep. Neuron 36, 1183–1194 (2002). source
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- cites [102] Wilson, M. A. & McNaughton, B. L. Reactivation of hippocampal ensemble memories during sleep. Science 265, 676–679 (1994). source
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- Interpretationcitation
The extraction of the meaning of stimuli beyond their superficial sensory properties may be the first step in capturing the essence of a schema, possibly mediated by gist representations.
“The extraction of the meaning of stimuli beyond their superficial sensory properties may be the first step in capturing the essence of a schema 14,20,22 , possibly mediated by gist representations.”
Quote in the paperNot a substantial claim0 of 3 sources checked- cites [14] Baraduc, P., Duhamel, J. R. & Wirth, S. Schema cells in the macaque hippocampus. Science 363, 635–639 (2019). Recordings from hippocampal neurons in the macaque monkey during virtual navigation show the existence of cells that generalize parts of similar environments across environments. source
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- cites [20] Robin, J. & Moscovitch, M. Details, gist and schema: hippocampal–neocortical interactions underlying recent and remote episodic and spatial memory. Curr. Opin. Behav. Sci. 17, 114–123 (2017). source
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- cites [22] Gilboa, A. & Marlatte, H. Neurobiology of schemas and schema-mediated memory. Trends Cogn. Sci. 21, 618–631 (2017). source
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- Premisecitation
Geometric resemblance guides the generalization of spatial learning of locations from one environment to another.
“Geometric resemblance guides the generalization of spatial learning of locations from one environment to another [14][15][16][17][18]53,107 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [14][15][16][17][18]53,107
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- Interpretationreasoning
The latter links value-based judgements to specific spatial features, enabling rewards to guide the development of spatial schemas, particularly early in their formation.
“The latter links value-based judgements to specific spatial features, enabling rewards to guide the development of spatial schemas, particularly early in their formation.”
Quote in the paperNot a substantial claim - Premisecitation
Zheng et al. showed that the mPFC represents a common spatial structure across environments that have shared and unique elements.
“In line with these ideas, Zheng et al. showed that the mPFC represents a common spatial structure across environments that have shared and unique elements 38 (see also Box 1).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [38] Zheng, L., Gao, Z., McAvan, A. S., Isham, E. A. & Ekstrom, A. D. Partially overlapping spatial environments trigger reinstatement in hippocampus and schema representations in prefrontal cortex. Nat. Commun. 12, 6231 (2021). source
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- Premisecitation
Multivoxel pattern similarity analyses in a recent fMRI study showed that HPC responses to items that share a similar context are more likely to occur in overlapping regions than responses to items that do not share a context.
“Multivoxel pattern similarity analyses in a recent fMRI study in which participants learned item-context associations showed that HPC responses to items that share a similar context, a likely requisite for schema formation, are more likely to occur in overlapping regions than responses to items that do not share a context 79 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [79] Libby, L. A., Reagh, Z. M., Bouffard, N. R., Ragland, J. D. & Ranganath, C. The hippocampus generalizes across memories that share item and context information. J. Cogn. Neurosci. 31, 24–35 (2019). source
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- Premisecitation
When geometric or surface features are modified, the firing patterns of HPC place cells can change in a process known as remapping.
“When geometric or surface features are modified, the firing patterns of HPC place cells can change in a process known as remapping 11,113 .”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [11] Moser, E. I., Moser, M.-B. & McNaughton, B. L. Spatial representation in the hippocampal formation: a history. Nat. Neurosci. 20, 1448 (2017). source
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- cites [113] Kubie, J. L., Levy, E. R. J. & Fenton, A. A. Is hippocampal remapping the physiological basis for context? Hippocampus https://doi.org/10.1002/hipo.23160 (2019). source
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- Premisecitation
Cells in the macaque HPC, termed 'schema cells,' encode task representations that encompass spatial and cognitive dimensions, dissociated from immediate visual properties of the environment.
“Cells in the macaque HPC, termed 'schema cells,' have been found to encode task representations that encompass spatial and cognitive dimensions, dissociated from immediate visual properties of the environment 14 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [14] Baraduc, P., Duhamel, J. R. & Wirth, S. Schema cells in the macaque hippocampus. Science 363, 635–639 (2019). Recordings from hippocampal neurons in the macaque monkey during virtual navigation show the existence of cells that generalize parts of similar environments across environments. source
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- Premisecitation
Remapping is thought to underlie the ability of the HPC to store multiple unique representations of similar environments or altered states of the same environment.
“Remapping is thought to underlie the ability of the HPC to store multiple unique representations of similar environments or altered states of the same environment 12 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [12] Alme, C. B. et al. Place cells in the hippocampus: eleven maps for eleven rooms. Proc. Natl Acad. Sci. USA 111, 18428–18435 (2014). source
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- Interpretationcitation
Overlapping item-context associations may serve as the building blocks for schemas, or for gists that combine to form schemas.
“Overlapping item-context associations may serve as the building blocks for schemas, or for gists that combine to form schemas.”
Quote in the paperSubstantial claim - Premisecitation
Pattern recognition is a neural mechanism for gist extraction and schema formation in mice and humans.
“Pattern recognition is a neural mechanism for gist extraction and schema formation in mice 1 and humans 119,120 .”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [1] Richards, B. A. et al. Patterns across multiple memories are identified over time. Nat. Neurosci. 17, 981–986 (2014). source
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- cites [119] Antony, J. et al. Spatial gist extraction during human memory consolidation. J. Exp. Psychol. Learn. Mem. Cogn. 48, 929–941 (2022). Distributional learning during spatial navigation is examined in three behavioural experiments, demonstrating the extraction of spatial gist representations that decrease over time. source
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- Premisecitation
Rodents with HPC lesions and extensive pre-lesion experience in an environment are able to use the overall layout of the learned environment, or its spatial gist.
“This ditinction is similar to that found in rodents with HPC lesions and extensive pre-lesion experience in an environment: they are able to use the overall layout of the learned environment, or its spatial gist, and take detours to reach goal locations, although often not the most optimal ones 70,125,126 .”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [70] Winocur, G., Moscovitch, M., Fogel, S., Rosenbaum, R. S. & Sekeres, M. Preserved spatial memory after hippocampal lesions: effects of extensive experience in a complex environment. Nat. Neurosci. 8, 273–275 (2005). source
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- cites [125] Winocur, G., Moscovitch, M., Rosenbaum, R. S. & Sekeres, M. An investigation of the effects of hippocampal lesions in rats on pre- and postoperatively acquired spatial memory in a complex environment. Hippocampus 20, 1350–1365 (2010). source
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- cites [126] Spiers, H. J. & Gilbert, S. J. Solving the detour problem in navigation: a model of prefrontal and hippocampal interactions. Front. Hum. Neurosci. 9, 125 (2015). source
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- Premisecitation
In a water-maze paradigm, mice exposed to platform locations moved across trials according to selections from a normal distribution acquired a generalized representation of the spatial distribution.
“In a water-maze paradigm, mice exposed to platform locations that were moved across trials according to selections from a normal distribution were able to acquire a generalized representation of the spatial distribution that was disrupted by pharmacological inhibition of the mPFC 1 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [1] Richards, B. A. et al. Patterns across multiple memories are identified over time. Nat. Neurosci. 17, 981–986 (2014). source
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- Premisecitation
In individuals with bilateral HPC damage, memory for details or surface properties is lost.
“whereas memory for details or surface properties is lost [80][81][82]123,124 .”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [80] Rosenbaum, R. S. et al. Remote spatial memory in an amnesic person with extensive bilateral hippocampal lesions. Nat. Neurosci. 3, 1044–1048 (2000). source
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- cites [81] Herdman, K. A., Calarco, N., Moscovitch, M., Hirshhorn, M. & Rosenbaum, R. S. Impoverished descriptions of familiar routes in three cases of hippocampal/medial temporal lobe amnesia. Cortex 71, 248–263 (2015). This patient-lesion study provides converging evidence that the hippocampus is needed to support remote spatial memory for visuospatial details but not the gist of environments learned long ago. source
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- cites [82] Rosenbaum, R. S., Cassidy, B. N. & Herdman, K. A. Patterns of preserved and impaired spatial memory in a case of developmental amnesia. Front. Hum. Neurosci. 9, 196 (2015). source
Not checked in the free run.
- Interpretationreasoning
The mPFC is involved in monitoring the degree of consistency between existing schemas and novel information.
“Given the known role of the mPFC in monitoring the contents of memory retrieval 22 , this finding suggests that the mPFC is involved in monitoring the degree of consistency between existing schemas and novel information 1,53,121 .”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [1] Richards, B. A. et al. Patterns across multiple memories are identified over time. Nat. Neurosci. 17, 981–986 (2014). source
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- cites [53] Tse, D. et al. Schemas and memory consolidation. Science 316, 76–82 (2007). This study in rodents with hippocampal lesions provides evidence for spatial schemas and the involvement of the hippocampus in rapid learning. source
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- cites [121] Wang, S. H., Tse, D. & Morris, R. G. Anterior cingulate cortex in schema assimilation and expression. Learn. Mem. 19, 315–318 (2012). source
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- Interpretationcitation
Geometric resemblance could help drive spatial schema construction via HPC remapping.
“Geometric resemblance could help drive spatial schema construction via HPC remapping 115 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [115] Hartley, T., Burgess, N., Lever, C., Cacucci, F. & O’Keefe, J. Modeling place fields in terms of the cortical inputs to the hippocampus. Hippocampus 10, 369–379 (2000). source
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- Premisecitation
The generalized representation of the spatial distribution in mice was disrupted by pharmacological inhibition of the mPFC.
“that was disrupted by pharmacological inhibition of the mPFC 1 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [1] Richards, B. A. et al. Patterns across multiple memories are identified over time. Nat. Neurosci. 17, 981–986 (2014). source
Not checked in the free run.
- Premisecitation
In individuals with bilateral HPC damage, the gist or overall layout of a familiar environment learned before the onset of HPC damage is preserved.
“Neuropsychological evidence in individuals with bilateral HPC damage, such as K.C. described above 80 , indicates that the gist or overall layout of a familiar environment learned before the onset of HPC damage is preserved 3,[80][81][82][122][123][124] , whereas memory for details or surface properties is lost [80][81][82]123,124 .”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [3] Robin, J., Rivest, J., Rosenbaum, R. S. & Moscovitch, M. Remote spatial and autobiographical memory in cases of episodic amnesia and topographical disorientation. Cortex 119, 237–257 (2019). source
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- cites [80] Rosenbaum, R. S. et al. Remote spatial memory in an amnesic person with extensive bilateral hippocampal lesions. Nat. Neurosci. 3, 1044–1048 (2000). source
Not checked in the free run.
- cites [81] Herdman, K. A., Calarco, N., Moscovitch, M., Hirshhorn, M. & Rosenbaum, R. S. Impoverished descriptions of familiar routes in three cases of hippocampal/medial temporal lobe amnesia. Cortex 71, 248–263 (2015). This patient-lesion study provides converging evidence that the hippocampus is needed to support remote spatial memory for visuospatial details but not the gist of environments learned long ago. source
Not checked in the free run.
- Premisecitation
Angles of intersections are remembered as being closer to 90 degrees.
“and angles of intersections are remembered as being closer to 90 degrees 138 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [138] Waller, D. E. & Nadel, L. E. Handbook of Spatial Cognition (American Psychological Association, 2013). source
Not checked in the free run.
- Premisecitation
Rodents with HPC lesions and extensive pre-lesion experience take detours to reach goal locations, although often not the most optimal ones.
“they are able to use the overall layout of the learned environment, or its spatial gist, and take detours to reach goal locations, although often not the most optimal ones 70,125,126 .”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [70] Winocur, G., Moscovitch, M., Fogel, S., Rosenbaum, R. S. & Sekeres, M. Preserved spatial memory after hippocampal lesions: effects of extensive experience in a complex environment. Nat. Neurosci. 8, 273–275 (2005). source
Not checked in the free run.
- cites [125] Winocur, G., Moscovitch, M., Rosenbaum, R. S. & Sekeres, M. An investigation of the effects of hippocampal lesions in rats on pre- and postoperatively acquired spatial memory in a complex environment. Hippocampus 20, 1350–1365 (2010). source
Not checked in the free run.
- cites [126] Spiers, H. J. & Gilbert, S. J. Solving the detour problem in navigation: a model of prefrontal and hippocampal interactions. Front. Hum. Neurosci. 9, 125 (2015). source
Not checked in the free run.
- Interpretationnone given
Sensitivity to region boundaries may reflect a neurophysiological mechanism by which variations in different visuospatial forms in the environment are identified.
“Sensitivity to region boundaries may reflect a neurophysiological mechanism by which variations in different visuospatial forms in the environment are identified.”
Quote in the paperNot a substantial claim - Premisecitation
In a virtual maze task, landmarks belonging to a semantic category were clustered to form implicit semantic regions that exerted an influence on navigation decisions.
“In a virtual maze task, landmarks belonging to a semantic category (vehicles, animals or buildings) were clustered to form implicit semantic regions that exerted an influence on navigation decisions.”
Quote in the paperSubstantial claim - Premisecitation
A city's spatial structure is characterized by multiple subregions connected via distribution networks.
“Recent findings indicate that a city's spatial structure is characterized by multiple subregions (such as urban zoning areas) connected via distribution networks (such as streets) 40,43,134,135 .”
Quote in the paperNot a substantial claim0 of 3 sources checked- cites [40] Lee, M., Barbosa, H., Youn, H., Holme, P. & Ghoshal, G. Morphology of travel routes and the organization of cities. Nat. Commun. 8, 2229 (2017). source
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- cites [43] Roth, C., Kang, S. M., Batty, M. & Barthelemy, M. Structure of urban movements: polycentric activity and entangled hierarchical flows. PLoS One 6, e15923 (2011). source
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- cites [134] Schirmer, P. M. & Axhausen, K. W. A multiscale classification of urban morphology. J. Transp. Land Use 9, 101–130 (2016).
Not checked in the free run.
- Interpretationcitation
Heuristic-based decisions in relation to space are based on parcellating the environment into semantic regions.
“These findings suggest that semantic cues influence spatial judgements, and that heuristic-based decisions in relation to space are based on parcellating the environment into semantic regions.”
Quote in the paperSubstantial claim - Premisecitation
Individuals prefer to take routes that cross fewer semantic regions, even if they are the same length as routes that contain more regions.
“Specifically, individuals prefer to take routes that cross fewer semantic regions, even if they are the same length as routes that contain more regions 29 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [29] Noack, H., Schick, W., Mallot, H. & Born, J. Sleep enhances knowledge of routes and regions in spatial environments. Learn. Mem. 24, 140–144 (2017). source
Not checked in the free run.
- Interpretationnone given
A collection of multiple gists makes up a spatial schema.
“A collection of multiple gists makes up a spatial schema, which we argue is probably supported by the mPFC, possibly together with the HPC and a network of posterior regions (Fig. 1).”
Quote in the paperSubstantial claim - Premisecitation
Familiar locations are drawn larger on maps than are unfamiliar locations.
“and familiar locations are drawn larger on maps than are unfamiliar locations 141 (Box 1).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [141] Jafarpour, A. & Spiers, H. Familiarity expands space and contracts time. Hippocampus 27, 12–16 (2017). source
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- Interpretationcitation
The role of geometry in spatial schemas extends to complex human settlements.
“The role of geometry in spatial schemas extends to complex human settlements 129 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [129] Batty, M. Cities and Complexity: Understanding Cities with Cellular Automata, Agent-Based Models, and Fractals (The MIT Press, 2007).
Not checked in the free run.
- Interpretationreasoning
A spatial schema is probably supported by the mPFC, possibly together with the HPC and a network of posterior regions.
“A collection of multiple gists makes up a spatial schema, which we argue is probably supported by the mPFC, possibly together with the HPC and a network of posterior regions (Fig. 1).”
Quote in the paperSubstantial claim - Premisecitation
Real-world spaces are segmented by physical barriers that yield various substructures or regions.
“Real-world spaces are segmented by physical barriers that yield various substructures or regions 41,[130][131][132][133] .”
Quote in the paperNot a substantial claim0 of 3 sources checked- cites [41] Lämmer, S., Gehlsen, B. & Helbing, D. Scaling laws in the spatial structure of urban road networks. Phys. A Stat. Mech. Appl. 363, 89–95 (2006). source
Not checked in the free run.
- cites [130] Kim, M. & Maguire, E. A. Hippocampus, retrosplenial and parahippocampal cortices encode multicompartment 3D space in a hierarchical manner. Cereb. Cortex 28, 1898–1909 (2018). source
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- cites [131] He, Q. & Brown, T. I. Environmental barriers disrupt grid-like representations in humans during navigation. Curr. Biol. 29, 2718–2722.e13 (2019). source
Not checked in the free run.
- Interpretationreasoning
Regions and boundaries are determined not only by the physical structure of the built environment, but also by the way we process the environment in relation to stored knowledge in semantic memory.
“However, regions and boundaries are determined not only by the physical structure of the built environment, but also by the way we process the environment in relation to stored knowledge in semantic memory.”
Quote in the paperSubstantial claim - Premisecitation
Distances between two locations are overestimated if they are separated by a barrier or boundary.
“For example, distances between two locations are overestimated if they are separated by a barrier or boundary (such as state borders), and angles of intersections are remembered as being closer to 90 degrees 138 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [138] Waller, D. E. & Nadel, L. E. Handbook of Spatial Cognition (American Psychological Association, 2013). source
Not checked in the free run.
- Interpretationcitation
Semantic cues influence spatial judgements.
“These findings suggest that semantic cues influence spatial judgements, and that heuristic-based decisions in relation to space are based on parcellating the environment into semantic regions.”
Quote in the paperSubstantial claim - Premisecitation
Distances from less familiar places to more familiar places are often underestimated compared with estimates of distances from more familiar to less familiar places.
“Familiarity with an environment produces other biases: distances from less familiar places to more familiar places are often underestimated compared with estimates of distances from more familiar to less familiar places 136,140 , and familiar locations are drawn larger on maps than are unfamiliar locations 141 (Box 1).”
Quote in the paperNot a substantial claim0 of 2 sources checked - Interpretationcitation
The involvement of grid cells in the extraction of a gist, and ultimately a schema, is supported by fMRI evidence showing that barriers that segment an environment modulate activity in several brain regions, including human ERC.
“As discussed below, the involvement of grid cells in the extraction of a gist, and ultimately a schema, is supported by fMRI evidence showing that the presence of barriers that segment an environment modulates activity in several brain regions, including human ERC 131 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [131] He, Q. & Brown, T. I. Environmental barriers disrupt grid-like representations in humans during navigation. Curr. Biol. 29, 2718–2722.e13 (2019). source
Not checked in the free run.
- Interpretationcitation
Neural mechanisms exist outside of the HPC to represent similar substructures of an environment that may give rise to spatial schemas.
“Neural mechanisms exist outside of the HPC to represent similar substructures of an environment that may give rise to spatial schemas.”
Quote in the paperSubstantial claim - Interpretationcitation
The repetition-activation patterns in RSC reflect similarity between metric distances travelled in different directions from specified spatial boundaries, akin to a local code for the space traversed.
“Although this finding supports the importance of a global code in capturing similar spatial contexts, the repetition-activation patterns reflect similarity between metric distances travelled in different directions from specified spatial boundaries, akin to a local code for the space traversed 151 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [151] Alexander, A. S. & Nitz, D. A. Retrosplenial cortex maps the conjunction of internal and external spaces. Nat. Neurosci. 18, 1143–1151 (2015). source
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- Premisecitation
Consolidation mechanisms, such as sleep, seem to facilitate the explicit knowledge and retrieval of semantic regions and their boundaries.
“Consolidation mechanisms, such as sleep, seem to facilitate the explicit knowledge and retrieval of semantic regions and their boundaries 29 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [29] Noack, H., Schick, W., Mallot, H. & Born, J. Sleep enhances knowledge of routes and regions in spatial environments. Learn. Mem. 24, 140–144 (2017). source
Not checked in the free run.
- Premisecitation
RSC neurons encode common features of routes with repeating segments, periodically firing across segments of routes that share a similar shape.
“For example, RSC neurons encode common features of routes with repeating segments, periodically firing across segments of routes that share a similar shape 151 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [151] Alexander, A. S. & Nitz, D. A. Retrosplenial cortex maps the conjunction of internal and external spaces. Nat. Neurosci. 18, 1143–1151 (2015). source
Not checked in the free run.
- Premisecitation
Sleep contributes greatly to the development of spatial schemas in mice navigating environments with repeating connectivity patterns.
“A study in mice navigating environments with repeating connectivity patterns further shows that sleep contributes greatly to the development of spatial schemas 69 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [69] Alonso, A. et al. The HexMaze: a previous knowledge task on map learning for mice. eNeuro 8, ENEURO.0554–20.2021 (2021). source
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- Premisecitation
Interacting networks of brain regions including specialized hubs help to support navigation and spatial memory.
“Interacting networks of brain regions including specialized 'hubs' help to support navigation and spatial memory 61,[142][143][144] (Fig. 1).”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [61] Ekstrom, A. D. et al. Cellular networks underlying human spatial navigation. Nature 425, 184 (2003). source
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- cites [142] Schedlbauer, A. M., Copara, M. S., Watrous, A. J. & Ekstrom, A. D. Multiple interacting brain areas underlie successful spatiotemporal memory retrieval in humans. Sci. Rep. 4, 6431 (2014). source
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- cites [143] Watrous, A. J., Tandon, N., Conner, C. R., Pieters, T. & Ekstrom, A. D. Frequency-specific network connectivity increases underlie accurate spatiotemporal memory retrieval. Nat. Neurosci. 16, 349–356 (2013). source
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- Premisecitation
In participants who learned object locations in geometrically similar but visually distinct virtual buildings, the representation of facing direction was generalized across environments with similar geometry, as indicated by fMRI activity pattern similarity in the RSC across multiple trials.
“In participants who learned the location of objects in geometrically similar but visually distinct (virtual) buildings 152 , the representation of facing direction (for example, south-west) was generalized across environments with similar geometry, as indicated by fMRI activity pattern similarity in the RSC across multiple trials.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [152] Marchette, S. A., Vass, L. K., Ryan, J. & Epstein, R. A. Anchoring the neural compass: coding of local spatial reference frames in human medial parietal lobe. Nat. Neurosci. 17, 1598–1606 (2014). source
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- Premisecitation
The RSC has been implicated in the instantiation and reinstatement of event schemas in a neuroanatomical model of schemas.
“The RSC has also been implicated in the instantiation and reinstatement of event schemas in a neuroanatomical model of schemas 22 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [22] Gilboa, A. & Marlatte, H. Neurobiology of schemas and schema-mediated memory. Trends Cogn. Sci. 21, 618–631 (2017). source
Not checked in the free run.
- Interpretationcitation
The ERC is another candidate for a hub involved in the formation of spatial schemas.
“The ERC is another candidate for a hub involved in the formation of spatial schemas.”
Quote in the paperNot a substantial claim - Interpretationcitation
The structure of the environment has a strong impact on how the brain organizes its representation of space.
“indicating that the structure of the environment has a strong impact on how the brain organizes its representation of space.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [131] He, Q. & Brown, T. I. Environmental barriers disrupt grid-like representations in humans during navigation. Curr. Biol. 29, 2718–2722.e13 (2019). source
Not checked in the free run.
- Premisecitation
The posterior HPC showed abstraction over different views of a room across the building.
“The posterior HPC showed abstraction over different views of a room across the building, suggesting that the posterior HPC might help build a more global map of the building.”
Quote in the paperSubstantial claim - Premisecitation
Similar findings have been reported in a virtual outdoor environment segmented into subspaces by a natural barrier (a river).
“Similar findings have been reported in a virtual outdoor environment segmented into subspaces by a natural barrier (a river) 39 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [39] Peer, M. & Epstein, R. A. The human brain uses spatial schemas to represent segmented environments. Curr. Biol. 31, 4677–4688.e8 (2021). Collectively, refs 37–39 investigate the cognitive and neural precursors to, and properties of, schema formation in humans, including the effects of overlap with previously learned environments, segmentation, and the roles of the hippocampus, retrosplenial cortex, and medial prefrontal cortex. source
Not checked in the free run.
- Interpretationown data
The RSC provides a long-term store of the spatial structure of the environment, such as the distance between locations in an integrated spatial map of a segmented environment.
“This is consistent with the RSC providing a long-term store of the spatial structure of the environment, such as the distance between locations in an integrated spatial map of a segmented environment 39 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [39] Peer, M. & Epstein, R. A. The human brain uses spatial schemas to represent segmented environments. Curr. Biol. 31, 4677–4688.e8 (2021). Collectively, refs 37–39 investigate the cognitive and neural precursors to, and properties of, schema formation in humans, including the effects of overlap with previously learned environments, segmentation, and the roles of the hippocampus, retrosplenial cortex, and medial prefrontal cortex. source
Not checked in the free run.
- Premisecitation
Human ERC grid-cell-like activity shifts from 6-fold rotational symmetry in an open arena to 4-fold rotational symmetry in an environment compartmentalized by barriers.
“Recent fMRI research has revealed that, like rodent grid cells, human ERC grid-cell-like activity shifts from 6-fold rotational symmetry in an open arena to 4-fold rotational symmetry in an environment compartmentalized by barriers 131 indicating that the structure of the environment has a strong impact on how the brain organizes its representation of space.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [131] He, Q. & Brown, T. I. Environmental barriers disrupt grid-like representations in humans during navigation. Curr. Biol. 29, 2718–2722.e13 (2019). source
Not checked in the free run.
- Premisecitation
In a follow-up study, when asked to navigate to a target object, participants were more likely to navigate to an equivalent location in an incorrect building rather than navigate to an incorrect location in the correct building.
“In a follow-up study 19 , when asked to navigate to a target object, participants were more likely to navigate to an equivalent location in an incorrect building rather than navigate to an incorrect location in the correct building.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [19] Marchette, S. A., Ryan, J. & Epstein, R. A. Schematic representations of local environmental space guide goal-directed navigation. Cognition 158, 68–80 (2017). source
Not checked in the free run.
- Interpretationreasoning
These regions are likely involved in integrating statistical regularities of visual percepts with high-level semantic concepts about spatial environments to aiding hierarchical structuring of the knowledge.
“Thus, it is likely that these regions are involved in integrating statistical regularities of visual percepts with high-level semantic concepts about spatial environments to aiding hierarchical structuring of the knowledge.”
Quote in the paperSubstantial claim - Interpretationnone given
Spatial schemas are a type of extra-HPC spatial representation.
“Above, we have presented behavioural and neurobiological evidence to suggest that spatial schemas are a type of extra-HPC spatial representation.”
Quote in the paperSubstantial claim - Premisecitation
Local rules often define the topological relations of containment and boundary.
“Local rules often define the topological relations of containment and boundary 165 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [165] Penn, A. & Turner, J. S. Can we identify general architectural principles that impact the collective behaviour of both human and animal systems? Philos. Trans. R. Soc. B Biol. Sci. 373, 20180253 (2018). source
Not checked in the free run.
- Interpretationreasoning
The PFC and PPC (particularly the angular gyrus) may sit atop such a functional cortical hierarchy and integrate low-level features with high-level categorical concepts to facilitate decision making.
“The PFC and PPC (particularly the angular gyrus) may sit atop such a functional cortical hierarchy and integrate low-level features with high-level categorical concepts to facilitate decision making.”
Quote in the paperNot a substantial claim - Premisecitation
If a small square cell is adjoined by other squares on three sides, and the process is repeated, a 'courtyard complex' scheme is generated.
“For example, if a small square cell is adjoined by other squares on three sides, and the process is repeated, a 'courtyard complex' scheme is generated 166 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [166] Hillier, B. & Julienne, H. The Social Logic of Space (Cambridge University Press, 1989).
Not checked in the free run.
- Interpretationreasoning
The posterior HPC might help build a more global map of the building.
“suggesting that the posterior HPC might help build a more global map of the building.”
Quote in the paperSubstantial claim - Premisecitation
fMRI data from participants navigating city streets presented in film footage contrasts with the posterior HPC abstraction finding.
“This contrasts with fMRI data from participants navigating city streets presented in film footage 158 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [158] Javadi, A. H. et al. Hippocampal and prefrontal processing of network topology to simulate the future. Nat. Commun. 8, 14652 (2017). source
Not checked in the free run.
- Premisecitation
The global form of 'beady-ring' settlements of Les Yves in southern France results from a recursive addition of spatial units according to a local rule.
“For example, the global form of 'beady-ring' settlements of Les Yves in southern France results from a recursive addition of spatial units according to a local rule 165 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [165] Penn, A. & Turner, J. S. Can we identify general architectural principles that impact the collective behaviour of both human and animal systems? Philos. Trans. R. Soc. B Biol. Sci. 373, 20180253 (2018). source
Not checked in the free run.
- Premisereasoning
These local rules give rise to the global form of real human settlements.
“These local rules give rise to the global form of real human settlements.”
Quote in the paperSubstantial claim - Premisecitation
Different classes of spatial configurations are associated with differences in the connectedness of regions of space and thereby influence the movement and interactions of individuals in space.
“Different classes of spatial configurations are associated with differences in the connectedness of regions of space and thereby influence the movement and interactions of individuals in space 168,169 .”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites [168] Haq, S. & Zimring, C. Just down the road a piece: the development of topological knowledge of building layouts. Environ. Behav. 35, 132–160 (2003). source
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- cites [169] Hillier, B. Space is the Machine: A Configurational Theory of Architecture (Space Syntax, 2007).
Not checked in the free run.
- Premisecitation
Virtual landmarks may help in navigating electronic space (such as the world-wide web) by influencing the frequency of visits and connectivity between related webpages.
“Similarly, virtual landmarks may help in navigating electronic space (such as the world-wide web) by influencing the frequency of visits and connectivity between related webpages 171 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [171] Sorrows, M. E. & Hirtle, S. C. The nature of landmarks for real and electronic spaces. Spatial Information Theory. Cognitive and Computational Foundations of Geographic Information Science, Vol. 1661, 37–50 (Springer, 1999). source
Not checked in the free run.
- Premisecitation
Variations in local rules lead to different probabilities for different global spatial forms.
“These investigations have shown that variations in local rules lead to different probabilities for different global spatial forms 166 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [166] Hillier, B. & Julienne, H. The Social Logic of Space (Cambridge University Press, 1989).
Not checked in the free run.
- Premisecitation
In a more recent computational study, the original sketch map of Boston consisting of elements collectively described by its residents was recreated using graph-network indices such as betweenness centrality, incidence angle of vertices and modularity optimization.
“In a more recent computational study, the original sketch map of Boston consisting of elements collectively described by its residents, was recreated using graph-network indices such as betweenness centrality [G], incidence angle of vertices and modularity optimization 34 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [34] Filomena, G., Verstegen, J. & Manley, E. A computational approach to ‘The Image of the City’. Cities 89, 14–25 (2019). The authors apply a mathematical analysis to the layout of cities to identify hidden structures in the street layouts for regions. source
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- Premisecitation
Graph-theoretic measures of street networks have been used to operationalize the essential urban elements that characterize a city form, as theorized by Kevin Lynch.
“Graph-theoretic measures of street networks have also been used to operationalize the essential urban elements that characterize a city form, as theorized by the urban theorist Kevin Lynch 33,34 .”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites [33] Lynch, K. The Image of the City. Vol. 11 (MIT Press, 1960).
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- cites [34] Filomena, G., Verstegen, J. & Manley, E. A computational approach to ‘The Image of the City’. Cities 89, 14–25 (2019). The authors apply a mathematical analysis to the layout of cities to identify hidden structures in the street layouts for regions. source
Not checked in the free run.
- Premisecitation
Industrial cities can be classified using characteristic spatial patterns and common principles, such as an organization centred around nodes of economic activity.
“Urban data analytics further show that industrial cities can be classified using characteristic spatial patterns and common principles, such as an organization centred around nodes of economic activity 42,43,129,177 .”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [42] Strano, E. et al. Urban street networks, a comparative analysis of ten European cities. Environ. Plan. B Plan. Des. 40, 1071–1086 (2013). source
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- cites [43] Roth, C., Kang, S. M., Batty, M. & Barthelemy, M. Structure of urban movements: polycentric activity and entangled hierarchical flows. PLoS One 6, e15923 (2011). source
Not checked in the free run.
- cites [129] Batty, M. Cities and Complexity: Understanding Cities with Cellular Automata, Agent-Based Models, and Fractals (The MIT Press, 2007).
Not checked in the free run.
- Interpretationreasoning
The different functional responses of the anterior and posterior HPC may reflect the role of these brain regions in simulating future routes through the street network.
“The different functional responses of the anterior and posterior HPC may reflect the role of these brain regions in simulating future routes through the street network.”
Quote in the paperSubstantial claim - Premisereasoning
A city street network can be represented as a graph in which street segments or axial lines are the units rather than the nodes.
“A city street network can thus be represented as a graph in which street segments or axial lines are the units rather than the nodes.”
Quote in the paperSubstantial claim - Interpretationreasoning
Such responses probably reflect the activity of neurons that, during exploration, learn the important, highly connected streets that maximally enable navigation across the network of city streets.
“Such responses probably reflect the activity of neurons that, during exploration, learn the important, highly connected streets that maximally enable navigation across the network of city streets.”
Quote in the paperSubstantial claim - Premisecitation
Space syntax-derived segment and axial maps can be analysed as various types of graphs to derive spatial measures, such as connectivity, integration and choice.
“Space syntax-derived segment and axial maps can then be analysed as various types of graphs to derive spatial measures, such as connectivity (measured using the graph-theoretic index of degree centrality), integration (closeness centrality) and choice (betweenness centrality) 175 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [175] Emo, B. Seeing the axial line: evidence from wayfinding experiments. Behav. Sci. 4, 167–180 (2014). source
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- Premisecitation
The mobility patterns of individuals towards and away from nodes of economic activity in cities are consistent over time.
“Interestingly, the mobility patterns of individuals towards and away from nodes of economic activity in cities are consistent over time.”
Quote in the paperSubstantial claim - Interpretationcitation
Early-life experiences with environments lead to the formation of spatial schemas that support particular navigational approaches that prepare us to variable extents for navigating different novel terrains.
“These findings suggest that our early-life experiences with environments lead to the formation of spatial schemas that support particular navigational approaches that prepare us to variable extents for navigating different novel terrains 187 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [187] Barhorst-Cates, E. M., Meneghetti, C., Zhao, Y., Pazzaglia, F. & Creem-Regehr, S. H. Effects of home environment structure on navigation preference and performance: a comparison in Veneto, Italy and Utah, USA. J. Environ. Psychol. 74, 101580 (2021). source
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- Premisecitation
Analysis of daily travel patterns of individuals using origin-destination matrices indicates stable movement patterns between functionally distinct regions in different cities.
“Analysis of daily travel patterns of individuals using origin-destination matrices indicates stable movement patterns between functionally distinct regions (such as residential and commercial regions) in different cities 177 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [177] Schneider, C. M., Belik, V., Couronne, T., Smoreda, Z. & Gonzalez, M. C. Unravelling daily human mobility motifs. J. R. Soc. Interface 10, 20130246 (2013). source
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- Premisecitation
The topological relations of containment and boundary resemble the gestalt grouping rules that organize visual percepts at a neuronal and behavioural level.
“For example, the topological relations of containment and boundary resemble the gestalt grouping rules that organize visual percepts at a neuronal and behavioural level [180][181][182][183] .”
Quote in the paperNot a substantial claim0 of 3 sources checked- cites [180] Elder, J. H. & Goldberg, R. M. Ecological statistics of Gestalt laws for the perceptual organization of contours. J. Vis. 2, 324–353 (2002). source
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- cites [181] Wannig, A., Stanisor, L. & Roelfsema, P. R. Automatic spread of attentional response modulation along Gestalt criteria in primary visual cortex. Nat. Neurosci. 14, 1243 (2011). source
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- cites [182] Wertheimer, M. in A Source Book of Gestalt Psychology (ed Willis. E) (Humanities Press, 1950).
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- Interpretationcitation
The observation that local rules drive global form in human settlements suggests a link between gestalt notions of grouping and configuration.
“The observation that local rules drive global form in human settlements 166 suggests a link between gestalt [G] notions of grouping (or assembling parts into a unit) and configuration (or positioning the units in space) 180 .”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites [166] Hillier, B. & Julienne, H. The Social Logic of Space (Cambridge University Press, 1989).
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- cites [180] Elder, J. H. & Goldberg, R. M. Ecological statistics of Gestalt laws for the perceptual organization of contours. J. Vis. 2, 324–353 (2002). source
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- Premisecitation
People who grew up in 'griddy' cities had a slight advantage at navigating grid-like virtual environments in Sea Hero Quest.
“However, people who grew up in 'griddy' cities had a slight advantage at navigating grid-like virtual environments in Sea Hero Quest (Fig. 4c), suggesting that experience of gridded cities does not have a purely negative effect on navigation skill 117 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [117] Coutrot, A. et al. Entropy of city street networks linked to future spatial navigation ability. Nature 604, 104–110 (2022). This study reports evidence that growing up in griddy cities adapts navigators to navigate better in grid-like environments but less so in more disorganized environments. source
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- Interpretationcitation
Experience of gridded cities does not have a purely negative effect on navigation skill.
“suggesting that experience of gridded cities does not have a purely negative effect on navigation skill 117 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [117] Coutrot, A. et al. Entropy of city street networks linked to future spatial navigation ability. Nature 604, 104–110 (2022). This study reports evidence that growing up in griddy cities adapts navigators to navigate better in grid-like environments but less so in more disorganized environments. source
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- Interpretationreasoning
Spatial representations may reflect the accumulation of 'spatial gists' of an environment type that are extracted from multiple experiences navigating similar or different environments.
“It follows that spatial representations may also reflect the accumulation of 'spatial gists' of an environment type (for example, a grid-like city) that are extracted from multiple experiences navigating similar or different environments.”
Quote in the paperSubstantial claim - Interpretationnone given
A hippocampus (HPC)-based representation of an environment is highly linked with detailed perceptual features of episodic events.
“A hippocampus (HPC)-based representation of an environment is highly linked with detailed perceptual features of episodic events (panels c,f).”
Quote in the paperNot a substantial claim - Interpretationnone given
The interaction of hubs with networks of regions eventually gives rise to spatial schemas.
“it is the interaction of hubs with networks of regions that eventually gives rise to spatial schemas.”
Quote in the paperNot a substantial claim- Rests on an assumption: The various posterior regions play specialized roles in representing components of surface properties and their relations and/or coding of these properties within local or global scales.
- Interpretationnone given
Geometric resemblance is a key feature of environments that gives rise to generalizable spatial representations.
“We have presented evidence from animal and human studies that suggests that geometric resemblance is a key feature of environments that gives rise to generalizable spatial representations.”
Quote in the paperSubstantial claim - Premisecitation
The update to the flavor-place associations can be learned following a single exposure.
“This update can be learned following a single exposure 36 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [36] Alonso, A., van der Meij, J., Tse, D. & Genzel, L. Naïve to expert: considering the role of previous knowledge in memory. Brain Neurosci. Adv. 4, 2398212820948686 (2020). source
Not checked in the free run.
- Interpretationreasoning
The integration of overlapping features of similar environments results in prototypical spatial forms that act as templates for wayfinding behaviour in novel contexts.
“We propose that the integration of overlapping features of similar environments results in prototypical spatial forms that act as templates for wayfinding behaviour in novel contexts.”
Quote in the paperSubstantial claim - Interpretationcitation
Geometric cues, rather than surface properties, seem to facilitate allocentric navigation in a highly familiar environment, despite HPC lesions.
“Therefore, geometric cues, rather than surface properties, seem to facilitate allocentric navigation in a highly familiar environment, despite HPC lesions 70 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [70] Winocur, G., Moscovitch, M., Fogel, S., Rosenbaum, R. S. & Sekeres, M. Preserved spatial memory after hippocampal lesions: effects of extensive experience in a complex environment. Nat. Neurosci. 8, 273–275 (2005). source
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- Premisecitation
Street networks for the centres of the 10 most populous cities in Argentina and Romania differ most in terms of the organized versus disorganized orientations of their streets.
“Street networks for the centres of the 10 most populous cities in Argentina and Romania; two countries that differ most in terms of the organized versus disorganized orientations of their streets 115 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [115] Hartley, T., Burgess, N., Lever, C., Cacucci, F. & O’Keefe, J. Modeling place fields in terms of the cortical inputs to the hippocampus. Hippocampus 10, 369–379 (2000). source
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- Interpretationcitation
HPC-lesioned rats used stable room geometry rather than distal or surface cues to navigate to correct goal locations.
“These findings indicate that HPC-lesioned rats used stable room geometry rather than distal or surface cues to navigate to correct goal locations.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [70] Winocur, G., Moscovitch, M., Fogel, S., Rosenbaum, R. S. & Sekeres, M. Preserved spatial memory after hippocampal lesions: effects of extensive experience in a complex environment. Nat. Neurosci. 8, 273–275 (2005). source
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- Premisecitation
There is an association between street network entropy (SNE) and the effect size for the negative impact of growing up in a city compared to non-city participants.
“Association between SNE and the effect size for the negative impact of growing up in a city compared to non-city participants, and also the identification of two clusters of countries (with high and low SNE).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [117] Coutrot, A. et al. Entropy of city street networks linked to future spatial navigation ability. Nature 604, 104–110 (2022). This study reports evidence that growing up in griddy cities adapts navigators to navigate better in grid-like environments but less so in more disorganized environments. source
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- Premisecitation
The difficulty that an individual with vmPFC damage (L.G.) had in navigating in a familiar environment was found to relate to L.G.'s inability to maintain a goal in mind, rather than to a loss of schema representations per se.
“In one such study, the difficulty that an individual with vmPFC damage, who was known as L.G., had in navigating in a familiar environment was found to relate to L.G.'s inability to maintain a goal in mind, rather than to a loss of schema representations per se 199 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [199] Rosenbaum, R. S., Winocur, G., Grady, C. L., Ziegler, M. & Moscovitch, M. Memory for familiar environments learned in the remote past: fMRI studies of healthy people and an amnesic person with extensive bilateral hippocampal lesions. Hippocampus 17, 1241–1251 (2007). source
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- Premisecitation
There is evidence of grid-like coding in mPFC similar to that observed in entorhinal cortex.
“This possibility is further suggested by evidence of grid-like coding in mPFC similar to that observed in entorhinal cortex 193 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [193] Amodio, D. M. & Frith, C. D. Meeting of minds: the medial frontal cortex and social cognition. Nat. Rev. Neurosci. 7, 268–277 (2006). source
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- Premisecitation
A neural network model of brain regions underlying spatial schemas, consisting of an HPC-supported indexing stream and a neocortex-dominant representation stream, replicates early experiments in rodents.
“A neural network model of brain regions underlying spatial schemas, consisting of an HPC-supported indexing stream and a neocortex-dominant representation stream 201 , replicates early experiments in rodents 53,54 .”
Quote in the paperNot a substantial claim0 of 3 sources checked- cites [201] Goh, S., Choi, M., Lee, K. & Kim, K.-M. How complexity emerges in urban systems: Theory of urban morphology. Phys. Rev. E 93, 052309 (2016). source
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- cites [53] Tse, D. et al. Schemas and memory consolidation. Science 316, 76–82 (2007). This study in rodents with hippocampal lesions provides evidence for spatial schemas and the involvement of the hippocampus in rapid learning. source
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- cites [54] Tse, D. et al. Schema-dependent gene activation and memory encoding in neocortex. Science 333, 891–895 (2011). source
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- Premisecitation
There is some functional MRI evidence of increased activation of the dorsomedial prefrontal cortex in highly familiar environments that had been navigated extensively over many years, even following bilateral hippocampal lesions.
“There is some functional MRI evidence of increased activation of the dorsomedial prefrontal cortex in highly familiar environments that had been navigated extensively over many years 127,154 , even following bilateral hippocampal lesions 198 , but the precise conditions in which this region is activated is unclear.”
Quote in the paperNot a substantial claim0 of 3 sources checked- cites [127] Rosenbaum, R. S. et al. fMRI studies of remote spatial memory in an amnesic person. Brain Cogn. 54, 170–172 (2004).
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- cites [154] Spiers, H. J., & Maguire, E. A. (2006). Thoughts, behaviour, and brain dynamics during navigation in the real world. Neuroimage, 31(4), 1826–1840. source
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- cites [198] Ross, R. S. & Eichenbaum, H. Dynamics of hippocampal and cortical activation during consolidation of a nonspatial memory. J. Neurosci. 26, 4852–4859 (2006). source
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- Premisecitation
Systematic measurement of quantitative differences in the spatial organization of real-world environments is a major challenge.
“However, systematic measurement of quantitative differences in the spatial organization of real-world environments, which often have varied layout patterns and route morphologies, is a major challenge [40][41][42]44,132,176,200 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [40][41][42]44,132,176,200
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- Interpretationcitation
Computational approaches have provided converging evidence of a core network of brain regions that enable schemas, as identified in patient-lesion and fMRI studies.
“Computational approaches have also provided converging evidence of a core network of brain regions that enable schemas, as identified in patient-lesion and fMRI studies.”
Quote in the paperNot a substantial claim - Interpretationcitation
The vmPFC may support a flexible representation of the world, providing structure to make sense of an ongoing stream of sensory input to enable relational inferences.
“Similar to a cognitive map, the vmFPC may support a flexible representation of the world, providing structure to make sense of an ongoing stream of sensory input to enable relational inferences 191 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [191] Kan, I. P., Rosenbaum, R. S. & Verfaellie, M. Schema processing across the lifespan: from theory to applications. Cogn. Neuropsychol. 37, 1–7 (2020). source
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- Premisecitation
A deep neural network that mimics the computational mechanisms in the primate posterior parietal cortex suggests that high-dimensional sensory inputs from scenes (global properties) are projected onto low-dimensional inputs (local properties) according to a learned spatial structure, akin to a spatial schema in this region.
“A deep neural network that mimics the computational mechanisms in the primate posterior parietal cortex suggests that high-dimensional sensory inputs from scenes (that is, global properties) are projected onto low-dimensional inputs (that is, local properties) according to a learned spatial structure, akin to a spatial schema in this region 160 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [160] Schick, W., Halfmann, M., Hardiess, G. & Mallot, H. A. Language cues in the formation of hierarchical representation of space. Cogn. Process. 15, S63–S64 (2014).
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- Premisecitation
Navigation planning follows a hierarchical representation of space that consists of a series of contexts that, in humans, is mediated by the dorsomedial PFC.
“Studies of navigation support the idea that navigation planning follows a hierarchical representation of space that consist of a series of contexts that, in humans, is mediated by the dorsomedial PFC 128,133,159,204 .”
Quote in the paperNot a substantial claim0 of 4 sources checked- cites [128] Hirshhorn, M., Grady, C., Rosenbaum, R. S., Winocur, G. & Moscovitch, M. Brain regions involved in the retrieval of spatial and episodic details associated with a familiar environment: an fMRI study. Neuropsychologia 50, 3094–3106 (2012). source
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- cites [133] Schick, W., Halfmann, M., Hardiess, G. & Mallot, H. A. Language cues in the formation of hierarchical representation of space. Cogn. Process. 16, S78–S78 (2015). source
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- cites [159] Balaguer, J., Spiers, H., Hassabis, D. & Summerfield, C. Neural mechanisms of hierarchical planning in a virtual subway network. Neuron 90, 893–903 (2016). source
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- cites [204] Kriegeskorte, N. & Douglas, P. K. Cognitive computational neuroscience. Nat. Neurosci. 21, 1148–1160 (2018). source
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- Premisecitation
The regionalization of space provides an efficient heuristic for planning actions in spatial navigation.
“The regionalization of space provides an efficient heuristic for planning actions in spatial navigation 29,131,159,205,206 , and computational models are particularly useful in understanding the correlations between physical regions and substructures, and their representations in the brain.”
Quote in the paperNot a substantial claim0 of 5 sources checked- cites [29] Noack, H., Schick, W., Mallot, H. & Born, J. Sleep enhances knowledge of routes and regions in spatial environments. Learn. Mem. 24, 140–144 (2017). source
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- cites [131] He, Q. & Brown, T. I. Environmental barriers disrupt grid-like representations in humans during navigation. Curr. Biol. 29, 2718–2722.e13 (2019). source
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- cites [159] Balaguer, J., Spiers, H., Hassabis, D. & Summerfield, C. Neural mechanisms of hierarchical planning in a virtual subway network. Neuron 90, 893–903 (2016). source
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- cites [205] McNamee, D., Wolpert, D. & Lengyel, M. in Advances in Neural Information Processing Systems 29 (NIPS, 2016).
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- cites [206] Wiener, J. M. & Mallot, H. A. ‘Fine-to-coarse’ route planning and navigation in regionalized environments. Spat. Cogn. Comput. 3, 331–358 (2003). source
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- Interpretationcitation
Spatial schemas are stable neural representations of environmental regularities, similar to how statistical regularities form priors or expectations in visual perception.
“Spatial schemas are stable neural representations of such environmental regularities, similar to how statistical regularities form priors or expectations in visual perception 28,47,211,212 .”
Quote in the paperNot a substantial claim0 of 4 sources checked- cites [28] de Lange, F. P., Heilbron, M. & Kok, P. How do expectations shape perception? Trends Cogn. Sci. 22, 764–779 (2018). source
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- cites [47] Mack, M. L., Love, B. C. & Preston, A. R. Dynamic updating of hippocampal object representations reflects new conceptual knowledge. Proc. Natl Acad. Sci. USA 113, 13203–13208 (2016). source
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- cites [211] Jiang, S., Ferreira, J. & González, M. C. Clustering daily patterns of human activities in the city. Data Min. Knowl. Discov. 25, 478–510 (2012). source
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- cites [212] Girshick, A. R., Landy, M. S. & Simoncelli, E. P. Cardinal rules: visual orientation perception reflects knowledge of environmental statistics. Nat. Neurosci. 14, 926–932 (2011). source
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- Premisecitation
Mobility patterns of 90 percent of the population in different countries can be captured by repeating 'motifs' of individuals' movements over time.
“Moreover, mobility patterns of 90 percent of the population in different countries can be captured by repeating 'motifs' of individuals' movements over time 177 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [177] Schneider, C. M., Belik, V., Couronne, T., Smoreda, Z. & Gonzalez, M. C. Unravelling daily human mobility motifs. J. R. Soc. Interface 10, 20130246 (2013). source
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- Premisecitation
The pattern of an individual's movements around a city seems to be highly consistent over an observational period of 6 months.
“The pattern of an individual's movements around a city seems to be highly consistent over an observational period of 6 months 177 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [177] Schneider, C. M., Belik, V., Couronne, T., Smoreda, Z. & Gonzalez, M. C. Unravelling daily human mobility motifs. J. R. Soc. Interface 10, 20130246 (2013). source
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- May combine two claims. 62
- Classified as probably not a substantial claim. 54
May combine two claims.
FindingcitationIn our recent research, the RSC was found to code distance to goals in well-known, real-world environments, but not in recently learned environments.
“In our recent research, the RSC was found to code distance to goals in well-known, real-world environments, but not in recently learned environments 153 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [153] Patai, E. Z. et al. Hippocampal and retrosplenial goal distance coding after long-term consolidation of a real-world environment. Cereb. Cortex 29, 2748–2758 (2019). This fMRI study provides evidence that the retrosplenial cortex codes spatial information in familiar environments in line with extraction of spatial knowledge to cortical regions. source
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Probably not a substantial claim: may not pass S4 (based). May combine two claims.
Interpretationnone givenThere are two aspects to spatial schemas: a psychological aspect relating to using previously formed schemas as reference templates for behaviour, and a neural aspect relating to schema formation via HPC-neocortical interaction.
“There are two aspects to spatial schemas: one that is psychological and relates to using previously formed schemas as reference templates for behaviour, and the other that is also neural and relates to schema formation via HPC-neocortical interaction.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S2 (checkable).
PremisecitationA spatial gist is a summary of the general layout extracted from experience navigating in that same environment, stripped of its perceptual details.
“Like a cognitive map, a spatial gist [G] is composed of key elements specific to a particular environment, but it may be viewed as a summary of the general layout extracted from experience navigating in that same environment, stripped of its perceptual details 20 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [20] Robin, J. & Moscovitch, M. Details, gist and schema: hippocampal–neocortical interactions underlying recent and remote episodic and spatial memory. Curr. Opin. Behav. Sci. 17, 114–123 (2017). source
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Probably not a substantial claim: may not pass S2 (checkable), S4 (based).
InterpretationreasoningA spatial schema is a superordinate spatial representation that supports optimal decision-making in multiple environments that share common geometric and visuospatial properties.
“we argue that a spatial schema is a superordinate spatial representation that supports optimal decision-making in multiple environments that share common geometric and visuospatial properties.”
Quote in the paperNot a substantial claimMay combine two claims.
InterpretationcitationA spatial schema is separate from an event schema, reflecting learned recurring patterns of spatial relations among key elements of an environment, and facilitated through navigation in that environment.
“Here we consider the possibility of a spatial schema as separate from an event schema [G], instead reflecting learned recurring patterns of spatial relations among key elements of an environment 14,18,28 , and facilitated through the act of navigating in that environment [29][30][31][32] .”
Quote in the paperSubstantial claim0 of 4 sources checked- cites [14] Baraduc, P., Duhamel, J. R. & Wirth, S. Schema cells in the macaque hippocampus. Science 363, 635–639 (2019). Recordings from hippocampal neurons in the macaque monkey during virtual navigation show the existence of cells that generalize parts of similar environments across environments. source
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- cites [18] McKenzie, S. et al. Hippocampal representation of related and opposing memories develop within distinct, hierarchically organized neural schemas. Neuron 83, 202–215 (2014). source
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- cites [28] de Lange, F. P., Heilbron, M. & Kok, P. How do expectations shape perception? Trends Cogn. Sci. 22, 764–779 (2018). source
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- cites [29][30][31][32]
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May combine two claims.
InterpretationreasoningSchemas may draw on multiple gists and are not specific to any particular environment or location.
“In this way, schemas may draw on multiple gists and, thus, are not specific to any particular environment or location.”
Quote in the paperSubstantial claimProbably not a substantial claim: may not pass S4 (based).
InterpretationreasoningThe integration of multiple gists could give rise to spatial schemas that are independent of a specific environment and the HPC.
“The integration of multiple gists, in turn, could give rise to spatial schemas [G] that are independent of a specific environment and the HPC.”
Quote in the paperNot a substantial claimClassified as probably not a substantial claim.
PremisecitationSpatial navigation is guided by an internal representation of spatial relations in a specific environment, referred to as a cognitive map.
“Spatial navigation is thought to be guided by the internal representation of spatial relations in a specific environment, referred to as a cognitive map [G] 2,9 .”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [2] Epstein, R. A., Patai, E. Z., Julian, J. B. & Spiers, H. J. The cognitive map in humans: spatial navigation and beyond. Nat. Neurosci. 20, 1504–1513 (2017). source
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- cites [9] O’Keefe, J. & Nadel, L. The Hippocampus as a Cognitive Map (Clarendon Press, 1978).
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May combine two claims.
PremisecitationPattern separation keeps cognitive maps and gists distinct from one another, but they can work in concert with schemas to aid in navigation.
“A process known as pattern separation [G] 65,66 keeps cognitive maps and gists distinct from one another, but they can work in concert with schemas to aid in navigation 38 .”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [65] Rolls, E. T. On pattern separation in the primate, including human, hippocampus. Trends Cogn. Sci. 25, 920–922 (2021). source
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- cites [66] Yassa, M. A. & Stark, C. E. Pattern separation in the hippocampus. Trends Neurosci. 34, 515–525 (2011). source
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- cites [38] Zheng, L., Gao, Z., McAvan, A. S., Isham, E. A. & Ekstrom, A. D. Partially overlapping spatial environments trigger reinstatement in hippocampus and schema representations in prefrontal cortex. Nat. Commun. 12, 6231 (2021). source
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Probably not a substantial claim: may not pass S4 (based). May combine two claims.
Interpretationnone givenA spatial schema might be reinstated when one encounters similar elements in a novel environment, or when one must assimilate previously encountered elements with novel elements when a portion of a familiar environment is substantially altered.
“A spatial schema might be reinstated when one encounters similar elements in a novel environment, or when one must assimilate previously encountered elements with novel elements when a portion of a familiar environment is substantially altered.”
Quote in the paperNot a substantial claimMay combine two claims.
InterpretationreasoningGists and the cognitive maps that relate to them are not folded into schemas; they need to remain distinct so that not all environments are viewed as alike.
“Gists, and the cognitive maps that relate to them, however, are not folded into schemas. They need to remain distinct from one another so that not all environments are viewed as alike -as would be the case if one only relied on schemas.”
Quote in the paperSubstantial claimMay combine two claims.
Interpretationnone givenSpatial schemas share properties with, but are distinct from, cognitive maps and event schemas at both cognitive and neural levels.
“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.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S2 (checkable), S4 (based).
InterpretationreasoningA spatial schema is a generalizable spatial representation of the category of an environment, formed by integrating overlapping neural representations across similar environments.
“As such, a spatial schema could be thought of as a generalizable spatial representation of the category of an environment, formed by integrating overlapping neural representations across similar environments.”
Quote in the paperNot a substantial claimClassified as probably not a substantial claim.
InterpretationreasoningA schema might facilitate learning of a novel environment that resembles a familiar one.
“A schema, by contrast, might facilitate learning of a novel environment that resembles a familiar one.”
Quote in the paperSubstantial claimProbably not a substantial claim: may not pass S2 (checkable), S4 (based).
Interpretationnone givenThe difference between spatial gists and spatial schemas should be revealed at a neuroanatomical level.
“The difference between spatial gists and spatial schemas should be revealed at a neuroanatomical level.”
Quote in the paperNot a substantial claimMay combine two claims.
InterpretationcitationSpatial gists may be mediated by the anterior HPC, although lesion evidence suggests they do not depend on it, at least in humans.
“Spatial gists may be mediated by the anterior HPC (Fig. 1), although there is lesion evidence reviewed below to suggest that they do not depend on it, at least in humans.”
Quote in the paperNot a substantial claimMay combine two claims.
PremisecitationRats with HPC lesions but extensive experience with the village maze acquired associations as quickly as controls, whereas inexperienced HPC-lesioned rats took much longer.
“Rats with HPC lesions but extensive experience with the village acquired those associations as quickly as controls did, whereas inexperienced HPC-lesioned rats took much longer.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [70] Winocur, G., Moscovitch, M., Fogel, S., Rosenbaum, R. S. & Sekeres, M. Preserved spatial memory after hippocampal lesions: effects of extensive experience in a complex environment. Nat. Neurosci. 8, 273–275 (2005). source
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May combine two claims.
PremisecitationK.C. was impaired in remembering minor landmarks and perceptual details of familiar environments, despite largely intact visuoperceptual processing abilities and semantic knowledge of those same places.
“K.C. was impaired, however, in remembering minor landmarks and perceptual details of familiar environments, despite having largely intact visuoperceptual processing abilities and semantic knowledge of those same places.”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [27] Spiers, H. J. & Maguire, E. A. The neuroscience of remote spatial memory: a tale of two cities. Neuroscience 149, 7–27 (2007). source
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- cites [81] Herdman, K. A., Calarco, N., Moscovitch, M., Hirshhorn, M. & Rosenbaum, R. S. Impoverished descriptions of familiar routes in three cases of hippocampal/medial temporal lobe amnesia. Cortex 71, 248–263 (2015). This patient-lesion study provides converging evidence that the hippocampus is needed to support remote spatial memory for visuospatial details but not the gist of environments learned long ago. source
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- cites [82] Rosenbaum, R. S., Cassidy, B. N. & Herdman, K. A. Patterns of preserved and impaired spatial memory in a case of developmental amnesia. Front. Hum. Neurosci. 9, 196 (2015). source
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PremisecitationRepresentational similarity analyses indicate that representations of learned associations, particularly previously established in memory, are integrated in anterior HPC and posterior mPFC, but remain distinct in posterior HPC and anterior mPFC.
“Representational similarity analyses [G] indicate that the representations of learned associations, particularly those that are previously established in memory, are integrated in anterior HPC and posterior mPFC, but remain distinct in the posterior HPC and anterior mPFC 49 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [49] Schlichting, M. L., Mumford, J. A. & Preston, A. R. Learning-related representational changes reveal dissociable integration and separation signatures in the hippocampus and prefrontal cortex. Nat. Commun. 6, 8151 (2015). source
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InterpretationreasoningThis was accompanied by a more rapid shift of activity from HPC to vlPFC, reflecting reduced forgetting and accelerated systems consolidation.
“This was accompanied by a more rapid shift of activity from HPC to vlPFC, reflecting reduced forgetting and accelerated systems consolidation.”
Quote in the paperSubstantial claimMay combine two claims.
PremisecitationfMRI data showed an activity shift from HPC to vmPFC for retrieval of learned associates after 24 hours, and with overlearning by 90 days, retrieval responses shifted to vlPFC, representing semanticization.
“functional MRI (fMRI) data showed an activity shift from HPC to ventromedial prefrontal cortex (vmPFC) for retrieval of learned associates after 24 hours, but that with overlearning (by 90 days), retrieval responses shifted to ventrolateral PFC (vlPFC), representing a process of semanticization.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [72] Sommer, T. The emergence of knowledge and how it supports the memory for novel related information. Cereb. Cortex 27, 1906–1921 (2017).
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PremisecitationK.C., an individual with bilateral HPC damage, could produce accurate sketch maps of the overall layout of his hometown and judge relative distances and directions between its major landmarks.
“This was demonstrated in the case K.C., an individual with bilateral HPC damage, who could produce accurate sketch maps of the overall layout of his hometown (Fig. 3) and judge relative distances and directions between its major landmarks 27,81,82 .”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [27] Spiers, H. J. & Maguire, E. A. The neuroscience of remote spatial memory: a tale of two cities. Neuroscience 149, 7–27 (2007). source
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- cites [81] Herdman, K. A., Calarco, N., Moscovitch, M., Hirshhorn, M. & Rosenbaum, R. S. Impoverished descriptions of familiar routes in three cases of hippocampal/medial temporal lobe amnesia. Cortex 71, 248–263 (2015). This patient-lesion study provides converging evidence that the hippocampus is needed to support remote spatial memory for visuospatial details but not the gist of environments learned long ago. source
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- cites [82] Rosenbaum, R. S., Cassidy, B. N. & Herdman, K. A. Patterns of preserved and impaired spatial memory in a case of developmental amnesia. Front. Hum. Neurosci. 9, 196 (2015). source
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May combine two claims.
InterpretationcitationNavigation ability in these animals was impaired after manipulating distal cues, indicating that stable room geometry determined success.
“Navigation ability in these animals was impaired after manipulating distal cues (for example, pictures on walls surrounding the maze), however, indicating that stable room geometry determined success.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [70] Winocur, G., Moscovitch, M., Fogel, S., Rosenbaum, R. S. & Sekeres, M. Preserved spatial memory after hippocampal lesions: effects of extensive experience in a complex environment. Nat. Neurosci. 8, 273–275 (2005). source
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Probably not a substantial claim: may not pass S4 (based).
Interpretationnone givenA spatial schema reflects the integration of spatial gists across multiple environments that have shared features, identified via statistical abstraction.
“A spatial schema reflects the integration of spatial gists across multiple environments that have shared features, identified via statistical abstraction.”
Quote in the paperNot a substantial claimMay combine two claims.
PremisecitationIn rodents and humans, sleep enhances gist extraction from memory, HPC spatial representations, and explicit knowledge of regions in space.
“In rodents and humans, sleep has been found to enhance gist extraction from memory 89 , HPC spatial representations 101 and explicit knowledge of regions in space 29 -mechanisms that have been attributed to schema learning 22,63 .”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [89] Lewis, P. A. & Durrant, S. J. Overlapping memory replay during sleep builds cognitive schemata. Trends Cogn. Sci. 15, 343–351 (2011). source
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- cites [101] Lee, A. K. & Wilson, M. A. Memory of sequential experience in the hippocampus during slow wave sleep. Neuron 36, 1183–1194 (2002). source
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- cites [29] Noack, H., Schick, W., Mallot, H. & Born, J. Sleep enhances knowledge of routes and regions in spatial environments. Learn. Mem. 24, 140–144 (2017). source
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PremisecitationHuman concept cells seem to encode a stable representation of a specific person or landmark, regardless of size or viewing angle, providing sparse codes for visually invariant abstractions related to personal events.
“Human 'concept cells' seem to encode a stable representation of a specific person or landmark, regardless of size or viewing angle 104,105 , providing sparse codes [G] for visually invariant abstractions related to personal events 106 .”
Quote in the paperNot a substantial claim0 of 3 sources checked- cites [104] Quiroga, R. Q., Reddy, L., Kreiman, G., Koch, C. & Fried, I. Invariant visual representation by single neurons in the human brain. Nature 435, 1102–1107 (2005). source
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- cites [105] Quiroga, R. Q. Neural representations across species. Science 363, 1388–1389 (2019). source
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- cites [106] Quiroga, R. Concept cells: the building blocks of declarative memory functions. Nat. Rev. Neurosci. 13, 587–597 (2012). source
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PremisecitationSleep enables extraction of regularities across instances, loss of contextual information, and retention of gist, as seen in extra-hippocampal regions.
“On the other hand, sleep enables the extraction of regularities across instances, the loss of contextual information and the retention of gist, as seen in representations in extra-hippocampal regions 89,97 .”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [89] Lewis, P. A. & Durrant, S. J. Overlapping memory replay during sleep builds cognitive schemata. Trends Cogn. Sci. 15, 343–351 (2011). source
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- cites [97] Stickgold, R. & Walker, M. P. Sleep-dependent memory triage: evolving generalization through selective processing. Nat. Neurosci. 16, 139–145 (2013). source
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PremisecitationSome spatial memories retain their detailed HPC representations in perpetuity, along with their schematic representation, and these representations interact with each other, with one or the other dominating depending on conditions including time since acquisition, task demands and retrieval cues.
“However, some spatial memories retain their detailed HPC representations in perpetuity, along with their schematic representation, and these representations interact with each other, with one or the other dominating depending on various conditions, including time since acquisition, task demands and retrieval cues 92,93 .”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [92] Winocur, G. & Moscovitch, M. Memory transformation and systems consolidation. J. Int. Neuropsychol. Soc. 17, 766–780 (2011). source
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- cites [93] Sekeres, M. J., Winocur, G. & Moscovitch, M. The hippocampus and related neocortical structures in memory transformation. Neurosci. Lett. 680, 39–53 (2018). source
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InterpretationcitationDisruption of replay of HPC neuronal sequences in rodents during SWS severely impairs retrieval of spatial representations, suggesting that replay mechanisms are needed for consolidation of spatial memories.
“Disruption of replay of HPC neuronal sequences in rodents during SWS 101,102 severely impairs the retrieval of spatial representations 103 , suggesting that replay mechanisms are needed for the consolidation of spatial memories.”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [101] Lee, A. K. & Wilson, M. A. Memory of sequential experience in the hippocampus during slow wave sleep. Neuron 36, 1183–1194 (2002). source
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- cites [102] Wilson, M. A. & McNaughton, B. L. Reactivation of hippocampal ensemble memories during sleep. Science 265, 676–679 (1994). source
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- cites [103] Girardeau, G., Benchenane, K., Wiener, S. I., Buzsaki, G. & Zugaro, M. B. Selective suppression of hippocampal ripples impairs spatial memory. Nat. Neurosci. 12, 1222–1223 (2009). source
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InterpretationcitationGeometric modifications are accompanied by more substantial changes in the firing patterns of place cells than are surface modifications, suggesting that HPC remapping actively supports the role of geometry in the stability of spatial representations.
“Geometric modifications are accompanied by more substantial changes in the firing patterns of place cells than are surface modifications 15,113,114 , suggesting that HPC remapping actively supports the role of geometry in the stability of spatial representations 115 .”
Quote in the paperSubstantial claim0 of 4 sources checked- cites [15] Spiers, H., Hayman, R. M., Jovalekic, A., Marozzi, E. & Jeffery, K. J. Place field repetition and purely local remapping in a multicompartment environment. Cereb. Cortex 25, 10–25 (2015). source
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- cites [113] Kubie, J. L., Levy, E. R. J. & Fenton, A. A. Is hippocampal remapping the physiological basis for context? Hippocampus https://doi.org/10.1002/hipo.23160 (2019). source
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- cites [114] Latuske, P., Kornienko, O., Kohler, L. & Allen, K. Hippocampal remapping and its entorhinal origin. Front. Behav. Neurosci. 11, 253 (2017). source
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- cites [115] Hartley, T., Burgess, N., Lever, C., Cacucci, F. & O’Keefe, J. Modeling place fields in terms of the cortical inputs to the hippocampus. Hippocampus 10, 369–379 (2000). source
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InterpretationreasoningThe HPC, guided by schemas, could extract the particular gist of the environment without attention to specific details.
“In this way, the HPC, guided by schemas, could extract the particular gist of the environment without attention to specific details.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationRemapping has been proposed to be conceived as the HPC performing 'hidden state inference', whereby sensory information is integrated with stored representations to create a 'belief' about current occupancy in the environment.
“Such remapping has been proposed to be conceived as the HPC performing 'hidden state inference', whereby sensory information is integrated with stored representations to create a 'belief' about current occupancy in the environment 116 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [116] Sanders, H., Wilson, M. A. & Gershman, S. J. Hippocampal remapping as hidden state inference. eLife 9, e51140 (2020). source
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PremisecitationThe increasing similarity between firing patterns of schema cells was not driven by visual similarity between environments, but by their common spatial geometry, and by past and future actions within the task structure.
“The increasing similarity between firing patterns of schema cells was not driven by visual similarity between environments, as indicated by point-ofgaze maps 14,118 , but by their common spatial geometry 14 , and by past and future actions within the task structure 14,105 .”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [14] Baraduc, P., Duhamel, J. R. & Wirth, S. Schema cells in the macaque hippocampus. Science 363, 635–639 (2019). Recordings from hippocampal neurons in the macaque monkey during virtual navigation show the existence of cells that generalize parts of similar environments across environments. source
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- cites [118] Wirth, S., Baraduc, P., Plante, A., Pinede, S. & Duhamel, J. R. Gaze-informed, task-situated representation of space in primate hippocampus during virtual navigation. PLoS Biol. 15, e2001045 (2017). source
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- cites [105] Quiroga, R. Q. Neural representations across species. Science 363, 1388–1389 (2019). source
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InterpretationcitationGeometric resemblance might explain findings that people are less likely to detect changes in surface properties than geometric properties of highly familiar and extensively experienced landmarks and scenes; if such changes are detected, they are at a global or coarse level.
“Geometric resemblance might explain findings that people are less likely to detect changes in surface properties than geometric properties of highly familiar and extensively experienced landmarks and scenes, whether tested directly by recognition or indirectly by eye-movements; if such changes are detected, the changes are at a global or coarse level 112 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [112] Rosenbaum, R. S. et al. Dramatic changes to well-known places go unnoticed. Preprint at PsyArXiv https://doi.org/10.31234/osf.io/ypg96 (2020). source
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Probably not a substantial claim: may not pass S3 (load-bearing), S4 (based). May combine two claims.
InterpretationreasoningThis process of biasing stored representations is akin to how schemas supported by the mPFC facilitate pattern separation to disambiguate overlapping representations: HPC cell remapping by modifying spatial firing patterns may enable the differentiation of similar experiences or locations.
“This process of biasing stored representations is akin to how schemas supported by the mPFC facilitate pattern separation to disambiguate overlapping representations: HPC cell remapping by modifying spatial firing patterns may enable the differentiation of similar experiences or locations.”
Quote in the paperNot a substantial claimMay combine two claims.
InterpretationcitationStatistical pattern recognition underlies spatial gist extraction and navigation search patterns in humans.
“A recent study that used a similar distributional learning paradigm during spatial navigation suggests that statistical pattern recognition also underlies spatial gist extraction and navigation search patterns in humans 119 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [119] Antony, J. et al. Spatial gist extraction during human memory consolidation. J. Exp. Psychol. Learn. Mem. Cogn. 48, 929–941 (2022). Distributional learning during spatial navigation is examined in three behavioural experiments, demonstrating the extraction of spatial gist representations that decrease over time. source
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Probably not a substantial claim: may not pass S3 (load-bearing), S4 (based).
Interpretationnone givenSchemas themselves may be better represented by cell assemblies outside of the HPC than by single schema cells, at least in humans.
“However, schemas themselves may be better represented by cell assemblies outside of the HPC than by single schema cells, at least in humans.”
Quote in the paperNot a substantial claimMay combine two claims.
InterpretationcitationAlthough the anterior HPC may help to generate and support a spatial gist, anterior HPC is not necessarily required to maintain or retrieve spatial gists.
“Human and rodent lesion studies suggest that although the anterior HPC may help to generate and support a spatial gist, anterior HPC is not necessarily required to maintain or retrieve spatial gists.”
Quote in the paperSubstantial claimMay combine two claims.
InterpretationcitationThe influence of stored knowledge on segmentation of environments into subregions is suggested by studies showing that human spatial representations are prone to systematic biases and may be hierarchically organized, similar to semantic and event knowledge.
“The influence of stored knowledge on segmentation of environments into subregions is suggested by studies showing that human spatial representations are prone to systematic biases 136,137 and may be hierarchically organized, similar to semantic and event knowledge 133,138,139 .”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [136] Tversky, B. Distortions in cognitive maps. Geoforum 23, 131–138 (1992). source
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- cites [137] Poucet, B. Spatial cognitive maps in animals: new hypotheses on their structure and neural mechanisms. Psychol. Rev. 100, 163–182 (1993). source
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- cites [133] Schick, W., Halfmann, M., Hardiess, G. & Mallot, H. A. Language cues in the formation of hierarchical representation of space. Cogn. Process. 16, S78–S78 (2015). source
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InterpretationcitationThe posterior HPC supports detailed cognitive maps, whereas a network of extra-HPC regions including PHC, RSC and PPC supports cognitive maps transformed over repeated experiences into a spatial gist.
“Thus, whereas the (posterior) HPC seems to support detailed cognitive maps, a network of extra-HPC regions, including the parahippocampal cortex (PHC), RSC and PPC, seems to support cognitive maps that have been transformed over repeated experiences navigating in a particular environment to form a 'spatial gist' -a summary of a specific environment 25,27,127,128 .”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [25] Frankland, P. W. & Bontempi, B. The organization of recent and remote memories. Nat. Rev. Neurosci. 6, 119–130 (2005). source
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- cites [27] Spiers, H. J. & Maguire, E. A. The neuroscience of remote spatial memory: a tale of two cities. Neuroscience 149, 7–27 (2007). source
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- cites [127] Rosenbaum, R. S. et al. fMRI studies of remote spatial memory in an amnesic person. Brain Cogn. 54, 170–172 (2004).
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InterpretationcitationMany of the same extra-HPC regions involved in the initial acquisition of spatial knowledge for large-scale environments are also involved in the retrieval of schematized spatial memory and event schemas, suggesting that the core elements of spatial schemas may be laid down at acquisition.
“fMRI studies indicate that many of the same extra-HPC regions involved in the initial acquisition of spatial knowledge for large-scale environments are also involved in the retrieval of schematized spatial memory 27,83,128 and event schemas 22 , suggesting that the core elements of spatial schemas may be laid down at acquisition 92,[147][148][149][150] .”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [27] Spiers, H. J. & Maguire, E. A. The neuroscience of remote spatial memory: a tale of two cities. Neuroscience 149, 7–27 (2007). source
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- cites [83] Rosenbaum, R. S., Ziegler, M., Winocur, G., Grady, C. L. & Moscovitch, M. “I have often walked down this street before”: fMRI studies on the hippocampus and other structures during mental navigation of an old environment. Hippocampus 14, 826–835 (2004). source
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- cites [128] Hirshhorn, M., Grady, C., Rosenbaum, R. S., Winocur, G. & Moscovitch, M. Brain regions involved in the retrieval of spatial and episodic details associated with a familiar environment: an fMRI study. Neuropsychologia 50, 3094–3106 (2012). source
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InterpretationcitationConsolidation mechanisms might act on place cells in the HPC and on grid cells in the ERC, which show repeating patterns of activity in an arrangement comparable to equilateral triangles that tile the environment.
“These consolidation mechanisms might act on place cells in the HPC and on grid cells in the entorhinal cortex (ERC), which show repeating patterns of activity in an arrangement comparable to equilateral triangles that tile the environment 11 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [11] Moser, E. I., Moser, M.-B. & McNaughton, B. L. Spatial representation in the hippocampal formation: a history. Nat. Neurosci. 20, 1448 (2017). source
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InterpretationcitationThe ERC may work with the HPC and occipital place area, which represent local geometry common across segments, and with the RSC, which integrates segments into a superordinate map, to give rise to a gist in the service of schema formation.
“The ERC may work with the HPC and occipital place area, which represent local geometry common across segments, and with the RSC, which integrates segments into a superordinate map, to give rise to a gist in the service of schema formation 39 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [39] Peer, M. & Epstein, R. A. The human brain uses spatial schemas to represent segmented environments. Curr. Biol. 31, 4677–4688.e8 (2021). Collectively, refs 37–39 investigate the cognitive and neural precursors to, and properties of, schema formation in humans, including the effects of overlap with previously learned environments, segmentation, and the roles of the hippocampus, retrosplenial cortex, and medial prefrontal cortex. source
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PremisecitationAccording to one model, the HPC and RSC are hubs of two largely overlapping networks that preferentially support allocentric and egocentric spatial representations, respectively.
“According to one model 145 , the HPC and RSC are hubs of two largely overlapping networks that preferentially support allocentric and egocentric spatial representations, respectively (see also ref. 146 ).”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites [145] Ekstrom, A. D., Huffman, D. J. & Starrett, M. Interacting networks of brain regions underlie human spatial navigation: a review and novel synthesis of the literature. J. Neurophysiol. 118, 3328–3344 (2017). source
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- cites [146] Bicanski, A. & Burgess, N. A neural-level model of spatial memory and imagery. eLife 7, e33752 (2018). source
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PremisecitationRodent HPC cells show repeated firing in multicompartment environments, with increased clustering of activity when the animal is near open doorways.
“Rodent HPC cells show repeated firing in multicompartment environments, with increased clustering of activity when the animal is near open doorways 15,16 .”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [15] Spiers, H., Hayman, R. M., Jovalekic, A., Marozzi, E. & Jeffery, K. J. Place field repetition and purely local remapping in a multicompartment environment. Cereb. Cortex 25, 10–25 (2015). source
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- cites [16] Derdikman, D. et al. Fragmentation of grid cell maps in a multicompartment environment. Nat. Neurosci. 12, 1325–1332 (2009). source
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Interpretationnone givenRegionalization using physical barriers or metric distance travelled are both determined by the global properties of the environment.
“In either case, regionalization using physical barriers or metric distance travelled are both determined by the global properties of the environment.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationSeveral brain regions have been implicated in neuroanatomical models of navigation, including the HPC, RSC, ERC, PHC, PPC, mPFC and inferior temporal cortex (ITC).
“Several brain regions have been implicated in neuroanatomical models of navigation, including the HPC, RSC, ERC, PHC, PPC, mPFC and inferior temporal cortex (ITC) 145 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [145] Ekstrom, A. D., Huffman, D. J. & Starrett, M. Interacting networks of brain regions underlie human spatial navigation: a review and novel synthesis of the literature. J. Neurophysiol. 118, 3328–3344 (2017). source
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PremisecitationThe RSC integrates egocentric and allocentric spatial frames of reference and generalizes local codes across geometrically similar environments.
“The RSC has also been shown to integrate egocentric and allocentric spatial frames of reference 146,152 and to generalize local codes (that is, codes for specific regions, such as the southwest corner of a specific room) across geometrically similar environments (in this example, to the south-west corner of various rooms) 19,39,152 .”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [146] Bicanski, A. & Burgess, N. A neural-level model of spatial memory and imagery. eLife 7, e33752 (2018). source
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- cites [152] Marchette, S. A., Vass, L. K., Ryan, J. & Epstein, R. A. Anchoring the neural compass: coding of local spatial reference frames in human medial parietal lobe. Nat. Neurosci. 17, 1598–1606 (2014). source
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- cites [19] Marchette, S. A., Ryan, J. & Epstein, R. A. Schematic representations of local environmental space guide goal-directed navigation. Cognition 158, 68–80 (2017). source
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PremisecitationfMRI repetition suppression effects in several brain regions, including the RSC, HPC and PHC, in participants navigating to target landmarks in rooms within a virtual multi-level building revealed that the repetition suppression effect was similar across rooms that had the same layout, suggesting generalization of neural representations.
“fMRI repetition suppression [G] effects in several brain regions, including the RSC, HPC and PHC in participants navigating to target landmarks in rooms within a virtual multi-level building revealed that the repetition suppression effect was similar across rooms that had the same layout, suggesting generalization of neural representations 130 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [130] Kim, M. & Maguire, E. A. Hippocampus, retrosplenial and parahippocampal cortices encode multicompartment 3D space in a hierarchical manner. Cereb. Cortex 28, 1898–1909 (2018). source
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PremisecitationThe ERC achieves this outcome by learning the transition structure of an environment and the regularities that occur within it, offering a more generalized code that is optimal for spatial schema instantiation.
“The ERC achieves this outcome by learning the transition structure of an environment and the regularities that occur within it, offering a more generalized code that is optimal for spatial schema instantiation 157 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [157] Behrens, T. E. J. et al. What is a cognitive map? Organizing knowledge for flexible behavior. Neuron 100, 490–509 (2018). source
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Classified as probably not a substantial claim.
InterpretationcitationSuch a representation might take a schematic form, such that similar distances across different environments are processed similarly or by the same neural populations in the RSC.
“Such a representation might take a schematic form, such that similar distances across different environments are processed similarly or by the same neural populations in the RSC 153,154 .”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [153] Patai, E. Z. et al. Hippocampal and retrosplenial goal distance coding after long-term consolidation of a real-world environment. Cereb. Cortex 29, 2748–2758 (2019). This fMRI study provides evidence that the retrosplenial cortex codes spatial information in familiar environments in line with extraction of spatial knowledge to cortical regions. source
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- cites [154] Spiers, H. J., & Maguire, E. A. (2006). Thoughts, behaviour, and brain dynamics during navigation in the real world. Neuroimage, 31(4), 1826–1840. source
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PremisecitationGrid cells in the ERC are posited to provide a structural scaffold for episodic events and spatial representations, with ERC acting as a mediator between HPC and neocortex by providing a geometric code for the global properties of the environment.
“Grid cells in the ERC are posited to provide a structural scaffold for episodic events and spatial representations 7 , with ERC acting as a mediator between HPC and neocortex by providing a geometric code for the global properties of the environment 155,156 .”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [7] Bellmund, J. L. S., Gardenfors, P., Moser, E. I. & Doeller, C. F. Navigating cognition: spatial codes for human thinking. Science 362, eaat6766 (2018). source
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- cites [155] Nau, M., Julian, J. B. & Doeller, C. F. How the brain’s navigation system shapes our visual experience. Trends Cogn. Sci. 22, 810–825 (2018). source
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- cites [156] Nau, M., Navarro Schroder, T., Bellmund, J. L. S. & Doeller, C. F. Hexadirectional coding of visual space in human entorhinal cortex. Nat. Neurosci. 21, 188–190 (2018). source
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Interpretationnone givenSchema instantiation involves pattern recognition and the regionalization of previously encountered spaces to extract a common geometric structure.
“We suggest that schema instantiation involves pattern recognition and the regionalization of previously encountered spaces to extract a common geometric structure.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
InterpretationcitationOne possible explanation for the different engagement of the posterior and anterior HPC in generalization relates to spatial scale, with the posterior HPC encoding room or junction spaces, and the anterior HPC encoding large-scale (outdoor) spaces.
“One possible explanation for the different engagement of the posterior and anterior HPC in generalization relates to spatial scale, with the posterior HPC encoding room or junction spaces, and the anterior HPC encoding large-scale (outdoor) spaces 76 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [76] Poppenk, J., Evensmoen, H. R., Moscovitch, M. & Nadel, L. Long-axis specialization of the human hippocampus. Trends Cogn. Sci. 17, 230–240 (2013). source
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PremisecitationOne approach to understanding how the built environment is organized and related to human behaviour is through the field of space syntax.
“One approach to understanding how the built environment is organized and related to human behaviour is through the field of space syntax 166 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [166] Hillier, B. & Julienne, H. The Social Logic of Space (Cambridge University Press, 1989).
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PremisecitationIn participants navigating city streets presented in film footage, the anterior HPC provided a more global code (tracking global street connections), while the posterior HPC provided a more local code (tracking local street connections).
“In this context, the anterior HPC provided a more global code (tracking global street connections), while the posterior HPC provided a more local code (tracking local street connections) 158 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [158] Javadi, A. H. et al. Hippocampal and prefrontal processing of network topology to simulate the future. Nat. Commun. 8, 14652 (2017). source
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PremisecitationRepresentations of navigation plans, the regionalization of physical environments and the neural representations of physical environments are all typically hierarchical, whereby a high-level structure encompasses multiple mid-level or low-level features.
“Representations of navigation plans 159 , the regionalization of physical environments and the neural representations of physical environments 29,160 are all typically hierarchical -whereby a high-level structure encompasses multiple mid-level or low-level features.”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [159] Balaguer, J., Spiers, H., Hassabis, D. & Summerfield, C. Neural mechanisms of hierarchical planning in a virtual subway network. Neuron 90, 893–903 (2016). source
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- cites [29] Noack, H., Schick, W., Mallot, H. & Born, J. Sleep enhances knowledge of routes and regions in spatial environments. Learn. Mem. 24, 140–144 (2017). source
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- cites [160] Schick, W., Halfmann, M., Hardiess, G. & Mallot, H. A. Language cues in the formation of hierarchical representation of space. Cogn. Process. 15, S63–S64 (2014).
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PremisecitationBuildings are similar to many objects in that they are constructed by assembling smaller parts into a whole, whereas buildings differ from many objects in that buildings create volumes of space through the arrangement of objects, such as walls.
“Buildings are similar to many objects, in that they are constructed by assembling smaller parts into a whole, whereas buildings differ from many objects in that buildings create volumes of space through the arrangement of objects, such as walls 166,167 .”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites [166] Hillier, B. & Julienne, H. The Social Logic of Space (Cambridge University Press, 1989).
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- cites [167] Pallasmaa, J. The Eyes of the Skin: Architecture and the Senses (John Wiley & Sons, 2012).
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PremisecitationThe structure (or geometry) of an artifact is determined by the function it is meant to serve.
“'Form follows function' is an influential design principle often followed by architects; the structure (or geometry) of an artifact is determined by the function it is meant to serve [162][163][164] .”
Quote in the paperNot a substantial claim0 of 3 sources checked- cites [162] Koolhaas, R. The Generic City (Sikkens Foundation, 1995).
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- cites [163] Kristan, W. B. & Katz, P. Form and function in systems neuroscience. Curr. Biol. 16, R828–R831 (2006). source
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- cites [164] Hillier, B., Penn, A., Hanson, J., Grajewski, T. & Xu, J. Natural movement: or, configuration and attraction in urban pedestrian movement. Environ. Plan. B Plan. Des. 20, 29–66 (1993). source
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PremisecitationBuildings establish social importance and 'meaning' through embellishments and modifications to their surface properties and structural features.
“Buildings also establish social importance and 'meaning' through embellishments and modifications to their surface properties and structural features 165 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [165] Penn, A. & Turner, J. S. Can we identify general architectural principles that impact the collective behaviour of both human and animal systems? Philos. Trans. R. Soc. B Biol. Sci. 373, 20180253 (2018). source
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Premisenone givenSpace syntax aims to identify the series of construction steps or local rules that result in a global spatial pattern.
“Space syntax aims to identify the series of construction steps or local rules that result in a global spatial pattern.”
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PremisecitationA building regulates the flow of air and individuals across its boundaries, similar to a living system.
“For example, a building regulates the flow of air and individuals across its boundaries, similar to a living system 165 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [165] Penn, A. & Turner, J. S. Can we identify general architectural principles that impact the collective behaviour of both human and animal systems? Philos. Trans. R. Soc. B Biol. Sci. 373, 20180253 (2018). source
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InterpretationcitationA posterior region, such as AG, may bind the contents and the structure of experience at a higher level of abstraction than does the HPC and is probably an important area for relating spatial gists to spatial schema.
“A posterior region, such as AG, may bind the contents and the structure of experience at a higher level of abstraction than does the HPC 161 and, thus, is probably an important area for relating spatial gists to spatial schema (summarised in Fig. 1).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [161] Summerfield, C., Luyckx, F. & Sheahan, H. Structure learning and the parietal cortex. Prog. Neurobiol. 184, 101717 (2020). source
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Premisenone givenRegions implicated in scene-processing, such as the parahippocampal place area and RSC, have a role in the integration of common local properties such as surface features.
“Regions implicated in scene-processing, such as the parahippocampal place area and RSC, have a role in the integration of common local properties such as surface features.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
Premisenone givenSpace syntax theorists have identified other elementary formulae besides the courtyard complex and the beady-ring templates, such as central spaces, blocks and ring streets.
“Space syntax theorists have identified other elementary formulae besides the courtyard complex and the beady-ring templates, such as central spaces, blocks and ring streets.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationThrough qualitative analysis of verbal descriptions of Boston residents, Lynch found that the presence of these essential elements in their descriptions was associated with the ease with which navigators understand the spatial coherence of their environment.
“Through qualitative analysis of verbal descriptions of Boston residents, Lynch found that the presence of these essential elements in their descriptions was associated with the ease with which navigators understand the spatial coherence of their environment 33 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [33] Lynch, K. The Image of the City. Vol. 11 (MIT Press, 1960).
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PremisecitationFive elements - landmarks, paths, nodes, edges and districts - have been identified as building blocks of a common understanding of the space that is shared amongst people with extensive experience of the region.
“Five elements -landmarks, paths, nodes, edges and districts -have been identified as building blocks of a common understanding of the space that is shared amongst people with extensive experience of the region.”
Quote in the paperNot a substantial claimMay combine two claims.
PremisecitationFindings from spatial cognition studies in humans show that semantic knowledge and language cues exert a powerful influence on human spatial judgements and may contribute to spatial schemas.
“As mentioned in the section on regionalization of space, findings from spatial cognition studies in humans show that semantic knowledge and language cues exert a powerful influence on human spatial judgements 29,138 and may contribute to spatial schemas.”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites [29] Noack, H., Schick, W., Mallot, H. & Born, J. Sleep enhances knowledge of routes and regions in spatial environments. Learn. Mem. 24, 140–144 (2017). source
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- cites [138] Waller, D. E. & Nadel, L. E. Handbook of Spatial Cognition (American Psychological Association, 2013). source
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PremisecitationSpatial forms have been proposed to be a 'fossil' that reflects the social and psychological characteristics of a group of people that share common linguistic and cultural values.
“Spatial forms have also been proposed to be a 'fossil' the reflects the social and psychological characteristics of a group of people that share common linguistic and cultural values 166,170 .”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites [166] Hillier, B. & Julienne, H. The Social Logic of Space (Cambridge University Press, 1989).
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- cites [170] Levi-Strauss, C. Structural Anthropology Translated from the French by Jacobson, C. & Grundfest Schoepf, B. (Basic Books, 1963).
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PremisecitationWhen navigating recently learned city streets in London, UK, activity in the anterior HPC increases when entering streets with greater global connections (higher integration) and decreases when entering streets that are less globally connected.
“When navigating recently learned city streets in London, UK, activity in the anterior HPC increases when entering streets with greater global connections (that is, with higher integration) and decreases when entering streets that are less globally connected 176 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [176] Bettencourt, L. & West, G. A unified theory of urban living. Nature 467, 912 (2010). source
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PremisecitationIndustrial cities in different regions of the world exhibit scale-invariant structural characteristics (for example, road networks), whereas the number of nodes and their configuration depends on infrastructure development, size and other sociocultural factors.
“Industrial cities in different regions of the world are suggested to exhibit scale-invariant structural characteristics (for example, road networks) 40,129 , whereas the number of nodes and their configuration depends on infrastructure development, size and other sociocultural factors 40 .”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites [40] Lee, M., Barbosa, H., Youn, H., Holme, P. & Ghoshal, G. Morphology of travel routes and the organization of cities. Nat. Commun. 8, 2229 (2017). source
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- cites [129] Batty, M. Cities and Complexity: Understanding Cities with Cellular Automata, Agent-Based Models, and Fractals (The MIT Press, 2007).
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PremisecitationA city street network is transformed into a network of lines using its topological properties.
“First, a city street network is transformed into a network of lines using its topological properties 34,[173][174][175] .”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [34] Filomena, G., Verstegen, J. & Manley, E. A computational approach to ‘The Image of the City’. Cities 89, 14–25 (2019). The authors apply a mathematical analysis to the layout of cities to identify hidden structures in the street layouts for regions. source
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- cites [173] Manley, E., Addison, J. & Cheng, T. Shortest path or anchor-based route choice: a large-scale empirical analysis of minicab routing in London. J. Transp. Geogr. 43, 123–139 (2015). source
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- cites [174] Manley, E., Orr, S. W. & Cheng, T. A heuristic model of bounded route choice in urban areas. Transp. Res. C. Emerg. Technol. 56, 195–209 (2015). source
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PremisecitationSpace syntax has been used to study prototypical spatial patterns on a global scale (for example, across different street grids).
“Space syntax has been used to study prototypical spatial patterns on a global scale (for example, across different street grids) 166,172 .”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites [166] Hillier, B. & Julienne, H. The Social Logic of Space (Cambridge University Press, 1989).
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- cites [172] Peponis, J., Zimring, C. & Choi, Y. K. Finding the building in wayfinding. Environ. Behav. 22, 555–590 (1990). source
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Premisenone givenThe posterior HPC is responsive on entering streets and reacts to connectivity, showing higher activity when entering a road with many connections than a road with dead-end.
“The posterior HPC is also responsive on entering streets, but rather than signalling a global code, it reacts to the connectivity (for example, showing higher activity when entering a road with many connections than a road with dead-end).”
Quote in the paperSubstantial claimProbably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
Premisenone givenA long straight main road can be represented as a single axial line or as a large set of connected street segments.
“For example, a long straight main road can be represented as a single axial line [G] or as a large set of connected street segments [G].”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationThe connectivity of a space in view is reflected in the activity of the occipital place area.
“Recent work has shown that the connectivity of a space in view is reflected in the activity of the occipital place area 18 , but how this connectivity extends to active navigation is not known.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [18] McKenzie, S. et al. Hippocampal representation of related and opposing memories develop within distinct, hierarchically organized neural schemas. Neuron 83, 202–215 (2014). source
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InterpretationcitationThese statistical commonalities of mobility patterns characterize the underlying structure of a city and navigational behaviours, and provide support for the notion of a spatial schema for an environment with key elements that can be generalized to other environments that belong to a superordinate category.
“Given that these statistical commonalities of mobility patterns characterize the underlying structure of a city 41,43 and navigational behaviours 34,117,178,179 , they provide support for the notion of a spatial schema for an environment with key elements that can be generalized to other environments that belong to a superordinate category 159 .”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [41] Lämmer, S., Gehlsen, B. & Helbing, D. Scaling laws in the spatial structure of urban road networks. Phys. A Stat. Mech. Appl. 363, 89–95 (2006). source
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- cites [43] Roth, C., Kang, S. M., Batty, M. & Barthelemy, M. Structure of urban movements: polycentric activity and entangled hierarchical flows. PLoS One 6, e15923 (2011). source
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- cites [34] Filomena, G., Verstegen, J. & Manley, E. A computational approach to ‘The Image of the City’. Cities 89, 14–25 (2019). The authors apply a mathematical analysis to the layout of cities to identify hidden structures in the street layouts for regions. source
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
Premisenone givenMeasures that involve egocentric information have been derived within the space syntax framework to characterize spatial relationships in scenes (vista spaces).
“Alongside the allocentric graph-theoretic indices such as integration and choice, measures that involve egocentric information have been derived within the from space syntax framework to characterize spatial relationships in scenes (also known as vista spaces).”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationViewshed analysis (which uses 2D polygons known as isovists to represent the visible area from a given vantage point) has been used to characterize local scenes.
“For example, viewshed analysis (which uses 2D polygons known as isovists to represent the visible area from a given vantage point) have been used to characterize local scenes 186 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [186] Bonner, M. F. & Epstein, R. A. Computational mechanisms underlying cortical responses to the affordance properties of visual scenes. PLoS Comput. Biol. 14, e1006111 (2018). source
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Interpretationnone givenThe distinction between geometric and surface properties as two forms of spatial representation parallels the dissociable gist and detailed aspects of remote spatial memories.
“We have discussed the distinction between geometric and surface properties as two forms of spatial representation that parallel the dissociable gist and detailed aspects of remote spatial memories.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S2 (checkable), S4 (based).
Interpretationnone givenA spatial schema is a consolidated distribution of spatial gists for a given prototype.
“A spatial schema is a consolidated distribution of spatial gists for a given prototype (panels a,d).”
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PremisecitationWith extensive practice across 3 months, rats show improved performance, indicated by increased digging time in the correct location and fewer errors when choosing the location in which to dig first.
“With extensive practice across 3 months, the rats show improved performance, indicated by increased digging time in the correct location and fewer errors when choosing the location in which to dig first.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [36] Alonso, A., van der Meij, J., Tse, D. & Genzel, L. Naïve to expert: considering the role of previous knowledge in memory. Brain Neurosci. Adv. 4, 2398212820948686 (2020). source
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Interpretationnone givenThe RSC serves as a hub for translating information between two orthogonal frames: detailed or local cues versus coarse or global cues, and egocentric versus allocentric.
“The RSC serves as a hub for translating information between two orthogonal frames: detailed or local cues versus coarse or global cues, and egocentric versus allocentric.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S4 (based).
Interpretationnone givenEnvironment geometry input is received by the entorhinal cortex (ERC).
“Environment geometry input is received by the entorhinal cortex (ERC).”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S4 (based). May combine two claims.
Interpretationnone givenStatistical learning of spatial context is integrated in the angular gyrus (AG) and mPFC into spatial schemas and is continuously updated through reciprocal connections with HPC and other posterior regions.
“Statistical learning of spatial context is integrated in the angular gyrus (AG) and mPFC into spatial schemas and is continuously updated through reciprocal connections with HPC and other posterior regions.”
Quote in the paperNot a substantial claimMay combine two claims.
PremisecitationHPC-lesioned rats were severely impaired in room change and village rotation conditions, but performed well in the cue distortion condition.
“HPC-lesioned rats were severely impaired in room change and village rotation conditions, but performed well in the cue distortion condition.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [70] Winocur, G., Moscovitch, M., Fogel, S., Rosenbaum, R. S. & Sekeres, M. Preserved spatial memory after hippocampal lesions: effects of extensive experience in a complex environment. Nat. Neurosci. 8, 273–275 (2005). source
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Interpretationnone givenSchemas are dynamic knowledge structures formed by extracting generalizable common features of particular detailed episodic events.
“Schemas are dynamic knowledge structures formed by extracting generalizable common features of particular detailed episodic events.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S4 (based).
Interpretationnone givenA network of posterior regions, including parahippocampal cortex (PHC) and posterior parietal cortex (PPC), is involved in representing perceptual features and spatial relations.
“A network of posterior regions, including parahippocampal cortex (PHC) and posterior parietal cortex (PPC), is involved in representing perceptual features and spatial relations.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S2 (checkable), S4 (based).
InterpretationreasoningSchemas may be built through specific expectations on the basis of a store of many repeated cognitive maps, culminating in a meta-map prediction of what lies ahead in the environment in which we are navigating.
“Schemas may be built through specific expectations on the basis of a store of many repeated cognitive maps, culminating in a meta-map prediction of what lies ahead in the environment in which we are navigating.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S4 (based).
Interpretationnone givenThe medial prefrontal cortex (mPFC) facilitates schema development by linking spatial features with goal-relevant actions and behaviours.
“The medial prefrontal cortex (mPFC) facilitates schema development by linking spatial features with goal-relevant actions and behaviours.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S2 (checkable), S4 (based). May combine two claims.
Interpretationnone givenA cognitive map can vary in precision from a highly detailed map to a gist, whereas a schema is a knowledge structure that is extracted from multiple gist-like cognitive maps in a way that is similar to how schemas form from events.
“A cognitive map can vary in precision from a highly detailed map to a gist, whereas a schema is a knowledge structure that is extracted from multiple gist-like cognitive maps in a way that is similar to how schemas form from events.”
Quote in the paperNot a substantial claimMay combine two claims.
PremisecitationArgentina and the USA have the largest gap in performance between those who grew up inside cities versus those who grew up outside cities, whereas Romania lies at the other end of the scale.
“Argentina and the USA have the largest gap in performance between those who grew up inside cites versus those who grew up outside cities, whereas Romania lies at the other end of the scale.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [117] Coutrot, A. et al. Entropy of city street networks linked to future spatial navigation ability. Nature 604, 104–110 (2022). This study reports evidence that growing up in griddy cities adapts navigators to navigate better in grid-like environments but less so in more disorganized environments. source
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PremisecitationAlthough the overall spatial configuration of K.C.'s sketch map was intact, he included fewer streets and landmarks than did control participants.
“Although the overall spatial configuration of K.C.'s sketch map was intact, he included fewer streets and landmarks than did control participants.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [213] Kok, P., Mostert, P. & de Lange, F. P. Prior expectations induce prestimulus sensory templates. Proc. Natl Acad. Sci. USA 114, 10473–10478 (2017). source
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PremisecitationSketch maps drawn by other individuals with amnesia relating to bilateral hippocampal damage are similarly accurate in terms of overall layout yet similarly impoverished in detail.
“Sketch maps drawn by other individuals with amnesia relating to bilateral hippocampal damage are similarly accurate in terms of overall layout yet similarly impoverished in detail 81,82 .”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [81] Herdman, K. A., Calarco, N., Moscovitch, M., Hirshhorn, M. & Rosenbaum, R. S. Impoverished descriptions of familiar routes in three cases of hippocampal/medial temporal lobe amnesia. Cortex 71, 248–263 (2015). This patient-lesion study provides converging evidence that the hippocampus is needed to support remote spatial memory for visuospatial details but not the gist of environments learned long ago. source
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- cites [82] Rosenbaum, R. S., Cassidy, B. N. & Herdman, K. A. Patterns of preserved and impaired spatial memory in a case of developmental amnesia. Front. Hum. Neurosci. 9, 196 (2015). source
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PremisecitationParticipants who reported growing up in cities from the low-SNE countries were most impacted by environments in Sea Hero Quest with a high SNE, and marginally better at navigating gridlike environments with a low SNE.
“Participants who reported growing up in cities from the low-SNE countries were most impacted by environments in Sea Hero Quest with a high SNE, and marginally better at navigating gridlike environments with a low SNE.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [117] Coutrot, A. et al. Entropy of city street networks linked to future spatial navigation ability. Nature 604, 104–110 (2022). This study reports evidence that growing up in griddy cities adapts navigators to navigate better in grid-like environments but less so in more disorganized environments. source
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InterpretationreasoningThe hippocampus is needed for detailed spatial representations, similar to its role in episodic memory.
“This pattern of performance suggests that the hippocampus is needed for detailed spatial representations, similar to its role in episodic memory [4][5][6]20 .”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [4][5][6]20
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PremisecitationNeural network models are powerful tools for understanding pattern recognition mechanisms that underlie visual perception, whereas Bayesian generative models explain how abstract prior knowledge is used to make inferences about familiar and novel stimuli.
“Neural network models are powerful tools for understanding pattern recognition mechanisms that underlie visual perception, whereas Bayesian generative models explain how abstract prior knowledge is used to make inferences about familiar and novel stimuli 28,[201][202][203] .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites 28,[201][202][203]
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PremisecitationL.G.'s knowledge of the routes and of the locations of landmarks along them were intact; he needed only to be reminded of the goal for the representation to be expressed appropriately during navigation.
“L.G.'s knowledge of the routes and of the locations of landmarks along them were intact; he needed only to be reminded of the goal for the representation to be expressed appropriately during navigation.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [199] Rosenbaum, R. S., Winocur, G., Grady, C. L., Ziegler, M. & Moscovitch, M. Memory for familiar environments learned in the remote past: fMRI studies of healthy people and an amnesic person with extensive bilateral hippocampal lesions. Hippocampus 17, 1241–1251 (2007). source
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PremisecitationThe ventromedial prefrontal cortex (vmPFC) features prominently in the literature on event schemas in humans.
“The ventromedial prefrontal cortex (vmPFC) features prominently in the literature on event schemas in humans 22,190 .”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites [22] Gilboa, A. & Marlatte, H. Neurobiology of schemas and schema-mediated memory. Trends Cogn. Sci. 21, 618–631 (2017). source
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- cites [190] Rosenbaum, R. S. et al. The case of KC: contributions of a memory-impaired person to memory theory. Neuropsychologia 43, 989–1021 (2005). source
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PremisecitationFamiliar places are drawn larger than unfamiliar places.
“This value attribution may be reflected in the finding that familiar places are drawn larger than unfamiliar places 141 , as well-known places are most likely to be valued or to contain reward.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [141] Jafarpour, A. & Spiers, H. Familiarity expands space and contracts time. Hippocampus 27, 12–16 (2017). source
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PremisecitationFormal artificial intelligence models provide flexibility in planning and account for a range of biological processes at neural and behavioural levels.
“Formal artificial intelligence models provide flexibility in planning and account for a range of biological processes at neural and behavioural levels 204 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [204] Kriegeskorte, N. & Douglas, P. K. Cognitive computational neuroscience. Nat. Neurosci. 21, 1148–1160 (2018). source
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PremisecitationStudies that focus on mathematical analyses of topology have helped us gain insight into spatial patterns that characterize different types of environments.
“Studies that focus on mathematical analyses of topology have helped us gain insight into spatial patterns that characterize different types of environments 43,135,208 .”
Quote in the paperNot a substantial claim0 of 3 sources checked- cites [43] Roth, C., Kang, S. M., Batty, M. & Barthelemy, M. Structure of urban movements: polycentric activity and entangled hierarchical flows. PLoS One 6, e15923 (2011). source
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- cites [135] Louail, T. et al. Uncovering the spatial structure of mobility networks. Nat. Commun. 6, 6007 (2015). source
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- cites [208] de Cothi, W. J. et al. Predictive maps in rats and humans for spatial navigation. Curr. Biol. 32, 3676–3689.e5 (2022). source
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PremisecitationSections of the urban fabric of Mumbai can be deformed using topological transformations into the suburban neighbourhoods of Las Vegas or into the regular grid structure of New York city blocks.
“For example, sections of the urban fabric of Mumbai can be deformed using topological transformations into the suburban neighbourhoods of Las Vegas or into the regular grid structure of New York city blocks 208 .”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [208] de Cothi, W. J. et al. Predictive maps in rats and humans for spatial navigation. Curr. Biol. 32, 3676–3689.e5 (2022). source
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PremisecitationA dominant model of modern city form is the clustering of economic activities in urban hubs, known as polycentricity.
“For example, a dominant model of modern city form is the clustering of economic activities in urban hubs, known as polycentricity 43,209 .”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites [43] Roth, C., Kang, S. M., Batty, M. & Barthelemy, M. Structure of urban movements: polycentric activity and entangled hierarchical flows. PLoS One 6, e15923 (2011). source
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- cites [209] Brelsford, C., Martin, T., Hand, J. & Bettencourt, L. M. A. Toward cities without slums: Topology and the spatial evolution of neighborhoods. Sci. Adv. 4, eaar4644 (2018). source
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PremisecitationThe patterns of individuals' movement to and from urban hubs across cities can be collected using GIS tools and phone-usage data.
“The patterns of individuals' movement to and from urban hubs across cities can be collected using GIS (geographic information system) tools and phone-usage data 40,43,177,210 .”
Quote in the paperNot a substantial claim0 of 4 sources checked- cites [40] Lee, M., Barbosa, H., Youn, H., Holme, P. & Ghoshal, G. Morphology of travel routes and the organization of cities. Nat. Commun. 8, 2229 (2017). source
Not checked in the free run.
- cites [43] Roth, C., Kang, S. M., Batty, M. & Barthelemy, M. Structure of urban movements: polycentric activity and entangled hierarchical flows. PLoS One 6, e15923 (2011). source
Not checked in the free run.
- cites [177] Schneider, C. M., Belik, V., Couronne, T., Smoreda, Z. & Gonzalez, M. C. Unravelling daily human mobility motifs. J. R. Soc. Interface 10, 20130246 (2013). source
Not checked in the free run.
- cites [210] Berry, B. J. & Garrison, W. L. The functional bases of the central place hierarchy. Econ. Geogr. 34, 145–154 (1958). source
Not checked in the free run.
The paper relies on this without support.
The various posterior regions play specialized roles in representing components of surface properties and their relations and/or coding of these properties within local or global scales.
“although the various posterior regions play specialized roles in representing components of surface properties and their relations and/or coding of these properties within local or global scales”
Claims other papers make about this one
Each is checked against this paper’s own text: does it say what the citing paper says it does?
Memory of navigational episodes emerges from the interaction of multiple parallel spatial representations, each supported by a widespread hippocampal-neocortical network.
“Taken together, our findings suggest that memory of navigational episodes emerges from the interaction of multiple parallel spatial representations, each supported by a widespread hippocampal–neocortical network2,5,38.”
From Long-term Memory Reorganization of Navigational Episodes (2026) · cites this paper as [5]
The ‘Memory consolidation, generalization and hippocampal replay’ section says detailed and schematic spatial representations can coexist and interact. The ‘Networks and hubs for spatial schemas’ section describes interacting brain networks that support navigation and spatial memory. But the paper does not establish that each representation is supported by a widespread hippocampal–neocortical network; its Introduction says spatial schemas can be independent of the hippocampus. No page numbers are available in our copy. (LLM, conf=0.83)
Experimental — accuracy not yet measuredFor natural environments, memory of spatial navigational episodes mostly occurs against the backdrop of semantic knowledge and patterns extracted from previous experiences that shape expectations and guide behaviour.
“For natural environments, memory of spatial navigational episodes mostly occurs against the backdrop of semantic knowledge and patterns extracted from previous experiences that shape our expectations and guide actual behaviour2,5.”
From Long-term Memory Reorganization of Navigational Episodes (2026) · cites this paper as [5]
Memory of egocentric and allocentric representational modes relies on at least partially dissociable neural substrates.
“Studies suggest that memory of these representational modes relies on at least partially dissociable neural substrates5,11,12.”
From Long-term Memory Reorganization of Navigational Episodes (2026) · cites this paper as [5]
Long-term spatial memory of navigational episodes may shift from a detailed cognitive map to a more gist-like spatial representation.
“Instead, several details fade, whereas other aspects remain salient, thus suggesting that long-term spatial memory of navigational episodes may shift from a detailed cognitive map to a more gist-like spatial representation5.”
From Long-term Memory Reorganization of Navigational Episodes (2026) · cites this paper as [5]
Cognitive maps, spatial gist and spatial schema are not three successive steps of abstraction of an initial experience but rather three classes of spatial representations with distinct relationships to navigational episodes that exist in parallel and mutually influence each other.
“In this framework, cognitive maps, spatial gist and spatial schema are not three successive steps of abstraction of an initial experience but rather three classes of spatial representations with distinct relationships to navigational episodes that exist in parallel and mutually influence each other5.”
From Long-term Memory Reorganization of Navigational Episodes (2026) · cites this paper as [5]
Visual stimuli may have guided behaviour by prompting the retrieval of schematic representations of previously visited environments that share spatial features with the Berlin Zoo.
“Thus, visual stimuli may have guided behaviour by prompting the retrieval of schematic representations of previously visited environments that share spatial features with the Berlin Zoo5.”
From Long-term Memory Reorganization of Navigational Episodes (2026) · cites this paper as [5]
Allocentric representations may be conceived as abstracted templates that may ultimately contribute to an overall spatial 'zoo'-schema that is independent of a distinct navigational episode.
“They may thus be conceived as abstracted templates that may ultimately contribute to an overall spatial ‘zoo’-schema that is independent of a distinct navigational episode5.”
From Long-term Memory Reorganization of Navigational Episodes (2026) · cites this paper as [5]
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Percentages, sums and reported differences re-computed from the paper’s own numbers.
Each contradiction between two passages listed here was confirmed by an independent check.
Conflicting results for altered distal cues
MajorThe main text says that manipulating distal cues impaired navigation in experienced hippocampal-lesioned rats. The figure caption says those rats performed well in the cue-distortion condition.
“Navigation ability in these animals was impaired after manipulating distal cues (for example, pictures on walls surrounding the maze), however, indicating that stable room geometry determined success.”
“HPC-lesioned rats were severely impaired in room change and village rotation conditions, but performed well in the cue distortion condition.”
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- Authors
- Farzanfar, D., Spiers, H. J., Moscovitch, M., & Rosenbaum, R. S.
- Affiliations
- University of Toronto (CA)
- University College London (GB)
- York University (CA)
- Venue
- Nature Reviews Neuroscience
- Published
- 2022-11-21
- Volume
- 24
- Issue
- 2
- Pages
- 63-79
- ISSN
- 1471-003X (ISSN-L)
- 1471-0048
- OpenAlex
- W4309646944
- Cited by
- 115
- References
- 198
- Citations by year
- 2026: 25 · 2025: 33 · 2024: 33 · 2023: 24
- FWCI
- 1.89
- Type
- journal article
- Language
- en
- Open access
- green
- discovery.ucl.ac.uk
- Licence
- Locations
- Nature reviews. Neuroscience (publishedVersion)
- PubMed (publishedVersion)
- UCL Discovery (University College London) (submittedVersion)
- Funders
- Canada First Research Excellence Fund
- Canadian Institutes of Health Research
- Topics
- Physical Sciences › Engineering › Automotive Engineering › Spatial Cognition and Navigation
- Life Sciences › Neuroscience › Cognitive Neuroscience › Memory and Neural Mechanisms
- Physical Sciences › Engineering › Building and Construction › Urban Design and Spatial Analysis
- Keywords
- spatial schema, cognitive maps, event schemas, spatial memory, episodic memory, spatial navigation, memory consolidation, urban layout, urban analytics, scale, regionalization, non-human animals
- MeSH terms
- Memory Consolidation, Animals, Cognition, Humans
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