From cognitive maps to spatial schemas

In the paper

“For example, a dominant model of modern city form is the clustering of economic activities in urban hubs, known as polycentricity 43,209 .”

What the sources say

• Supported

  The Introduction says contemporary cities display patterns of polycentricity and describes the clustering of economic activity in many centers as an important feature of the urban landscape. That supports the citing sentence’s characterization of polycentricity as a model of modern city form. (LLM, conf=0.90)

  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
• ?Could not verify

  The Introduction discusses varied urban forms, and the Results examine street networks and city-block topology. Neither those passages nor searches for polycentricity, economic hubs, and employment centers address the clustering of economic activity in urban hubs. (LLM, conf=0.72)

  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). sourceExperimental — accuracy not yet measured

In the paper

“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 .”

What the sources say

• Not checked yetcites [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

In the paper

“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.”

In the paper

“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 .”

What the sources say

• Not checked yetcites [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

In the paper

“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.”

In the paper

“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] .”

What the sources say

• Not checked yetcites [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
• Not checked yetcites [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
• Not checked yetcites [28] de Lange, F. P., Heilbron, M. & Kok, P. How do expectations shape perception? Trends Cogn. Sci. 22, 764–779 (2018). source
• Not checked yetcites [29][30][31][32]

In the paper

“In this way, schemas may draw on multiple gists and, thus, are not specific to any particular environment or location.”

In the paper

“The integration of multiple gists, in turn, could give rise to spatial schemas [G] that are independent of a specific environment and the HPC.”

In the paper

“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 .”

What the sources say

• Not checked yetcites [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
• Not checked yetcites [9] O’Keefe, J. & Nadel, L. The Hippocampus as a Cognitive Map (Clarendon Press, 1978).

In the paper

“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 .”

What the sources say

• Not checked yetcites [65] Rolls, E. T. On pattern separation in the primate, including human, hippocampus. Trends Cogn. Sci. 25, 920–922 (2021). source
• Not checked yetcites [66] Yassa, M. A. & Stark, C. E. Pattern separation in the hippocampus. Trends Neurosci. 34, 515–525 (2011). source
• Not checked yetcites [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

In the paper

“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.”

In the paper

“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.”

In the paper

“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.”

In the paper

“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.”

In the paper

“A schema, by contrast, might facilitate learning of a novel environment that resembles a familiar one.”

In the paper

“The difference between spatial gists and spatial schemas should be revealed at a neuroanatomical level.”

In the paper

“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.”

In the paper

“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.”

What the sources say

• Not checked yetcites [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

In the paper

“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.”

What the sources say

• Not checked yetcites [27] Spiers, H. J. & Maguire, E. A. The neuroscience of remote spatial memory: a tale of two cities. Neuroscience 149, 7–27 (2007). source
• Not checked yetcites [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 yetcites [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

In the paper

“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 .”

What the sources say

• Not checked yetcites [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

In the paper

“This was accompanied by a more rapid shift of activity from HPC to vlPFC, reflecting reduced forgetting and accelerated systems consolidation.”

In the paper

“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.”

What the sources say

• Not checked yetcites [72] Sommer, T. The emergence of knowledge and how it supports the memory for novel related information. Cereb. Cortex 27, 1906–1921 (2017).

In the paper

“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 .”

What the sources say

• Not checked yetcites [27] Spiers, H. J. & Maguire, E. A. The neuroscience of remote spatial memory: a tale of two cities. Neuroscience 149, 7–27 (2007). source
• Not checked yetcites [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 yetcites [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

In the paper

“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.”

What the sources say

• Not checked yetcites [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

In the paper

“A spatial schema reflects the integration of spatial gists across multiple environments that have shared features, identified via statistical abstraction.”

In the paper

“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 .”

What the sources say

• Not checked yetcites [89] Lewis, P. A. & Durrant, S. J. Overlapping memory replay during sleep builds cognitive schemata. Trends Cogn. Sci. 15, 343–351 (2011). source
• Not checked yetcites [101] Lee, A. K. & Wilson, M. A. Memory of sequential experience in the hippocampus during slow wave sleep. Neuron 36, 1183–1194 (2002). source
• Not checked yetcites [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

In the paper

“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 .”

What the sources say

• Not checked yetcites [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
• Not checked yetcites [105] Quiroga, R. Q. Neural representations across species. Science 363, 1388–1389 (2019). source
• Not checked yetcites [106] Quiroga, R. Concept cells: the building blocks of declarative memory functions. Nat. Rev. Neurosci. 13, 587–597 (2012). source

In the paper

“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 .”

What the sources say

• Not checked yetcites [89] Lewis, P. A. & Durrant, S. J. Overlapping memory replay during sleep builds cognitive schemata. Trends Cogn. Sci. 15, 343–351 (2011). source
• Not checked yetcites [97] Stickgold, R. & Walker, M. P. Sleep-dependent memory triage: evolving generalization through selective processing. Nat. Neurosci. 16, 139–145 (2013). source

In the paper

“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 .”

What the sources say

• Not checked yetcites [92] Winocur, G. & Moscovitch, M. Memory transformation and systems consolidation. J. Int. Neuropsychol. Soc. 17, 766–780 (2011). source
• Not checked yetcites [93] Sekeres, M. J., Winocur, G. & Moscovitch, M. The hippocampus and related neocortical structures in memory transformation. Neurosci. Lett. 680, 39–53 (2018). source

In the paper

“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.”

What the sources say

• Not checked yetcites [101] Lee, A. K. & Wilson, M. A. Memory of sequential experience in the hippocampus during slow wave sleep. Neuron 36, 1183–1194 (2002). source
• Not checked yetcites [102] Wilson, M. A. & McNaughton, B. L. Reactivation of hippocampal ensemble memories during sleep. Science 265, 676–679 (1994). source
• Not checked yetcites [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

In the paper

“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 .”

What the sources say

• Not checked yetcites [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
• Not checked yetcites [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
• Not checked yetcites [114] Latuske, P., Kornienko, O., Kohler, L. & Allen, K. Hippocampal remapping and its entorhinal origin. Front. Behav. Neurosci. 11, 253 (2017). source
• Not checked yetcites [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

In the paper

“In this way, the HPC, guided by schemas, could extract the particular gist of the environment without attention to specific details.”

In the paper

“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 .”

What the sources say

• Not checked yetcites [116] Sanders, H., Wilson, M. A. & Gershman, S. J. Hippocampal remapping as hidden state inference. eLife 9, e51140 (2020). source

In the paper

“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 .”

What the sources say

• Not checked yetcites [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
• Not checked yetcites [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
• Not checked yetcites [105] Quiroga, R. Q. Neural representations across species. Science 363, 1388–1389 (2019). source

In the paper

“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 .”

What the sources say

• Not checked yetcites [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

In the paper

“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.”

In the paper

“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 .”

What the sources say

• Not checked yetcites [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

In the paper

“However, schemas themselves may be better represented by cell assemblies outside of the HPC than by single schema cells, at least in humans.”

In the paper

“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.”

In the paper

“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 .”

What the sources say

• Not checked yetcites [136] Tversky, B. Distortions in cognitive maps. Geoforum 23, 131–138 (1992). source
• Not checked yetcites [137] Poucet, B. Spatial cognitive maps in animals: new hypotheses on their structure and neural mechanisms. Psychol. Rev. 100, 163–182 (1993). source
• Not checked yetcites [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

In the paper

“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 .”

What the sources say

• Not checked yetcites [25] Frankland, P. W. & Bontempi, B. The organization of recent and remote memories. Nat. Rev. Neurosci. 6, 119–130 (2005). source
• Not checked yetcites [27] Spiers, H. J. & Maguire, E. A. The neuroscience of remote spatial memory: a tale of two cities. Neuroscience 149, 7–27 (2007). source
• Not checked yetcites [127] Rosenbaum, R. S. et al. fMRI studies of remote spatial memory in an amnesic person. Brain Cogn. 54, 170–172 (2004).

In the paper

“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] .”

What the sources say

• Not checked yetcites [27] Spiers, H. J. & Maguire, E. A. The neuroscience of remote spatial memory: a tale of two cities. Neuroscience 149, 7–27 (2007). source
• Not checked yetcites [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
• Not checked yetcites [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

In the paper

“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 .”

What the sources say

• Not checked yetcites [11] Moser, E. I., Moser, M.-B. & McNaughton, B. L. Spatial representation in the hippocampal formation: a history. Nat. Neurosci. 20, 1448 (2017). source

In the paper

“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 .”

What the sources say

• Not checked yetcites [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

In the paper

“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 ).”

What the sources say

• Not checked yetcites [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
• Not checked yetcites [146] Bicanski, A. & Burgess, N. A neural-level model of spatial memory and imagery. eLife 7, e33752 (2018). source

In the paper

“Rodent HPC cells show repeated firing in multicompartment environments, with increased clustering of activity when the animal is near open doorways 15,16 .”

What the sources say

• Not checked yetcites [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
• Not checked yetcites [16] Derdikman, D. et al. Fragmentation of grid cell maps in a multicompartment environment. Nat. Neurosci. 12, 1325–1332 (2009). source

In the paper

“In either case, regionalization using physical barriers or metric distance travelled are both determined by the global properties of the environment.”

In the paper

“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 .”

What the sources say

• Not checked yetcites [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

In the paper

“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 .”

What the sources say

• Not checked yetcites [146] Bicanski, A. & Burgess, N. A neural-level model of spatial memory and imagery. eLife 7, e33752 (2018). source
• Not checked yetcites [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
• Not checked yetcites [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

In the paper

“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 .”

What the sources say

• Not checked yetcites [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

In the paper

“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 .”

What the sources say

• Not checked yetcites [157] Behrens, T. E. J. et al. What is a cognitive map? Organizing knowledge for flexible behavior. Neuron 100, 490–509 (2018). source

In the paper

“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 .”

What the sources say

• Not checked yetcites [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
• Not checked yetcites [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

In the paper

“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 .”

What the sources say

• Not checked yetcites [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
• Not checked yetcites [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
• Not checked yetcites [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

In the paper

“We suggest that schema instantiation involves pattern recognition and the regionalization of previously encountered spaces to extract a common geometric structure.”

In the paper

“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 .”

What the sources say

• Not checked yetcites [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

In the paper

“One approach to understanding how the built environment is organized and related to human behaviour is through the field of space syntax 166 .”

What the sources say

• Not checked yetcites [166] Hillier, B. & Julienne, H. The Social Logic of Space (Cambridge University Press, 1989).

In the paper

“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 .”

What the sources say

• Not checked yetcites [158] Javadi, A. H. et al. Hippocampal and prefrontal processing of network topology to simulate the future. Nat. Commun. 8, 14652 (2017). source

In the paper

“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.”

What the sources say

• Not checked yetcites [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
• Not checked yetcites [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 yetcites [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).

In the paper

“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 .”

What the sources say

• Not checked yetcites [166] Hillier, B. & Julienne, H. The Social Logic of Space (Cambridge University Press, 1989).
• Not checked yetcites [167] Pallasmaa, J. The Eyes of the Skin: Architecture and the Senses (John Wiley & Sons, 2012).

In the paper

“'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] .”

What the sources say

• Not checked yetcites [162] Koolhaas, R. The Generic City (Sikkens Foundation, 1995).
• Not checked yetcites [163] Kristan, W. B. & Katz, P. Form and function in systems neuroscience. Curr. Biol. 16, R828–R831 (2006). source
• Not checked yetcites [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

In the paper

“Buildings also establish social importance and 'meaning' through embellishments and modifications to their surface properties and structural features 165 .”

What the sources say

• Not checked yetcites [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

In the paper

“Space syntax aims to identify the series of construction steps or local rules that result in a global spatial pattern.”

In the paper

“For example, a building regulates the flow of air and individuals across its boundaries, similar to a living system 165 .”

What the sources say

• Not checked yetcites [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

In the paper

“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).”

What the sources say

• Not checked yetcites [161] Summerfield, C., Luyckx, F. & Sheahan, H. Structure learning and the parietal cortex. Prog. Neurobiol. 184, 101717 (2020). source

In the paper

“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.”

In the paper

“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.”

In the paper

“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 .”

What the sources say

• Not checked yetcites [33] Lynch, K. The Image of the City. Vol. 11 (MIT Press, 1960).

In the paper

“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.”

In the paper

“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.”

What the sources say

• Not checked yetcites [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 yetcites [138] Waller, D. E. & Nadel, L. E. Handbook of Spatial Cognition (American Psychological Association, 2013). source

In the paper

“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 .”

What the sources say

• Not checked yetcites [166] Hillier, B. & Julienne, H. The Social Logic of Space (Cambridge University Press, 1989).
• Not checked yetcites [170] Levi-Strauss, C. Structural Anthropology Translated from the French by Jacobson, C. & Grundfest Schoepf, B. (Basic Books, 1963).

In the paper

“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 .”

What the sources say

• Not checked yetcites [176] Bettencourt, L. & West, G. A unified theory of urban living. Nature 467, 912 (2010). source

In the paper

“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 .”

What the sources say

• Not checked yetcites [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 yetcites [129] Batty, M. Cities and Complexity: Understanding Cities with Cellular Automata, Agent-Based Models, and Fractals (The MIT Press, 2007).

In the paper

“First, a city street network is transformed into a network of lines using its topological properties 34,[173][174][175] .”

What the sources say

• Not checked yetcites [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 yetcites [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
• Not checked yetcites [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

In the paper

“Space syntax has been used to study prototypical spatial patterns on a global scale (for example, across different street grids) 166,172 .”

What the sources say

• Not checked yetcites [166] Hillier, B. & Julienne, H. The Social Logic of Space (Cambridge University Press, 1989).
• Not checked yetcites [172] Peponis, J., Zimring, C. & Choi, Y. K. Finding the building in wayfinding. Environ. Behav. 22, 555–590 (1990). source

In the paper

“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).”

In the paper

“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].”

In the paper

“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.”

What the sources say

• Not checked yetcites [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

In the paper

“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 .”

What the sources say

• Not checked yetcites [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 yetcites [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 yetcites [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

In the paper

“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).”

In the paper

“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 .”

What the sources say

• Not checked yetcites [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

In the paper

“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.”

In the paper

“A spatial schema is a consolidated distribution of spatial gists for a given prototype (panels a,d).”

In the paper

“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.”

What the sources say

• Not checked yetcites [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

In the paper

“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.”

In the paper

“Environment geometry input is received by the entorhinal cortex (ERC).”

In the paper

“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.”

In the paper

“HPC-lesioned rats were severely impaired in room change and village rotation conditions, but performed well in the cue distortion condition.”

What the sources say

• Not checked yetcites [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

In the paper

“Schemas are dynamic knowledge structures formed by extracting generalizable common features of particular detailed episodic events.”

In the paper

“A network of posterior regions, including parahippocampal cortex (PHC) and posterior parietal cortex (PPC), is involved in representing perceptual features and spatial relations.”

In the paper

“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.”

In the paper

“The medial prefrontal cortex (mPFC) facilitates schema development by linking spatial features with goal-relevant actions and behaviours.”

In the paper

“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.”

In the paper

“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.”

What the sources say

• Not checked yetcites [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

In the paper

“Although the overall spatial configuration of K.C.'s sketch map was intact, he included fewer streets and landmarks than did control participants.”

What the sources say

• Not checked yetcites [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

In the paper

“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 .”

What the sources say

• Not checked yetcites [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 yetcites [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