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The structure and shape of the urban environment influence our ability to find our way about in the city. Understanding how the physical properties of the environment affect spatial behavior and cognition is therefore a necessity. However, there are inherent difficulties in empirically studying complex and large-scale urban environments. These include the need to isolate the impact of specific urban features and to acquire data on the physical activity of individuals. In the present study, we attempted to overcome the above obstacles and examine the relation between urban environments and spatial cognition by testing the spatial behavior of rats. This idea originated from the resemblance in the operative brain functions and in the mechanisms and strategies employed by humans and other animals when acquiring spatial information and establishing an internal representation, as revealed in past studies. Accordingly, we tested rats in arenas that simulated a grid urban layout (e.g. Manhattan streets) and an irregular urban layout (e.g. Jerusalem streets). We found that in the grid layout, rat movement was more structured and extended over a greater area compared with their restricted movement in the irregular layout. These movement patterns recall those of humans in respective urban environments, illustrating that the structure and shape of the environment affect spatial behavior similarly in humans and rats. Overall, testing rats in environments that simulate facets of urban environments can provide new insights into human spatial cognition in urban environments.

BackgroundWe set out to solve two inherent problems in the study of animal spatial cognition (i) What is a “place”?; and (ii) whether behaviors that are not revealed as differing by one methodology could be revealed as different when analyzed using a different approach.MethodologyWe applied network analysis to scrutinize spatial behavior of rats tested in either a symmetrical or asymmetrical layout of 4, 8, or 12 objects placed along the perimeter of a round arena. We considered locations as the units of the network (nodes), and passes between locations as the links within the network.Principal FindingsWhile there were only minor activity differences between rats tested in the symmetrical or asymmetrical object layouts, network analysis revealed substantial differences. Viewing ‘location’ as a cluster of stopping coordinates, the key locations (large clusters of stopping coordinates) were at the objects in both layouts with 4 objects. However, in the asymmetrical layout with 4 objects, additional key locations were spaced by the rats between the objects, forming symmetry among the key locations. It was as if the rats had behaviorally imposed symmetry on the physically asymmetrical environment. Based on a previous finding that wayfinding is easier in symmetrical environments, we suggest that when the physical attributes of the environment were not symmetrical, the rats established a symmetric layout of key locations, thereby acquiring a more legible environment despite its complex physical structure.Conclusions and SignificanceThe present study adds a behavioral definition for “location”, a term that so far has been mostly discussed according to its physical attributes or neurobiological correlates (e.g. - place and grid neurons). Moreover, network analysis enabled the assessment of the importance of a location, even when that location did not display any distinctive physical properties.

This study was aimed at uncovering physical and geometric properties that make a particular landmark a target of exploration and navigation. Rats were tested in a square open-field arena with additional portable corners featuring the same properties as the arena corners. It was found that the routes of progression converged upon the added corners, whether located at the arena wall or the arena center. Route convergence upon the added corners involved numerous visits to these corners. However, time spent at the added corners was relatively short compared with the arena corners, including that from which rats were introduced into the arena. There was no differential effect of testing rats in light or dark, or with a low versus a high portable corner. It is suggested that the added corners were distinct against the background of the arena enclosure, whereas the four arena corners and walls were encoded by the rats as one geometric module. This distinctness, together with the greater accessibility of the added corners, made them salient landmarks and a target of exploration. Thus, the impact of a landmark extended beyond its specific self-geometry to include accessibility and distinctness, which are contextual properties. In addition to the contextual impact on locomotor behavior there was also a temporal effect, with security initially dominating the rats' behavior but then declining along with an increased attraction to salient landmarks. These spatiotemporal patterns characterized behavior in both lit and dark arenas, indicating that distal cues were secondary to local proximal cues in shaping routes.

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