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...l Right lingual gyrus (z = 3.28) (12, 86, 12) (BA 18/19) Right fusiform gyrus (z = 4.26) (28, 78, 8) (BA 37) Right parahippocampal gyrus (32, 38, 12) Left medial occipital–temporal Left lingual gyrus (24, 58, 12) (BA 18/19) Left fusiform gyrus (z = 4.46) (20, 46, 12) (BA 37) Left parahippocampal gyrus (32, 38, 8) Table shows regions activated for each spatial condition relative to the control task. For each region, the different anatomical areas involved are listed (Tzourio-Mazoyer, 2002; Duvernoy, 1991). For each anatomical label, coordinates in MNI space and Brodmann’s areas (Talairach and Tournoux, 1988) are given for voxels representing local maxima of activation or for representative voxels. In the former case, z-values are also provided. C o m m itteri et a l. 1 5 2 1 located inside the horizontal and descending segments of the intraparietal sulcus, ventrally becoming the intraoccipital sulcus. The activation involved the superior parietal lobules and both the superior and middle occipital gyri. A small portion of the right inferior parietal lobule was also activated during the landmarkcentered condition. On the medial cerebral surface, the activation encompassed the precuneus, a small reg...

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...l geometry. Finally, the parietal cortex (except for the left superior parietal lobule and precuneus, which have a preference for the landmark-centered condition) seems to play a general role in spatial distance estimation, together with the right PMv. METHODS Participants Fourteen neurologically normal volunteers (7 women and 7 men, mean age 26, range 23–33) gave their informed consent to participate in the experiment, whose procedures had received local ethical approval. Twelve subjects were right-handed and two were left-handed, as assessed by a modified version of the Edinburgh Inventory (Oldfield, 1971) (mean index = + 0.7; SD 0.5). The Virtual Environment A virtual reality software (3DStudio Max 4.2, Autodesk, Discreet) was used to create a three-dimensional realistic reconstruction of a complex environment, representing a square arena in front of the entrance of a palace. The arena is defined by the two short lateral wings and the long central wing of the palace. The remaining side is open and occupied by a lawn. The square has no Committeri et al. 1529 fixed furnishings, except for a fountain and four benches. The central frame in Figure 1 shows an aerial view of the environment. Stimuli ...

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... Maisog, Ungerleider, & Haxby, 1998). It is worth noting that the whole right superior frontal activation was shared by the viewer-centered and the landmarkcentered conditions, suggesting common computational mechanisms. The left ventral cortex, instead, showed a tendency to prefer the viewer-centered coding. Centered in the left IFG (BAs 44 and 45), it was already detected in previous experiments of body midline perception (Galati, Committeri, et al., 2001; Vallar et al., 1999), during active pointing (e.g., Lacquaniti et al., 1997), and also during passive observation of grasping movements (Rizzolatti, Fadiga, Gallese, & Fogassi, 1996), so that analogies have been drawn with monkey PMv (Rizzolatti, Fogassi, & Gallese, 2002). More specifically, area 44 is considered homologous to the rostral PMv (F5), whereas area 6 to the caudal PMv (F4). PMv is part of those circuits coding peripersonal (F4) and grasping-related space (F5), thus crucially involved in egocentric spatial representations (see Colby, 1998, for a review). Besides the motor role, the present data bring further support to a cognitive role for PMv (Rizzolatti et al., 2002). Landmark-centered Coding and the Ventromedial Occipital–Temporal Cortex One of the most in...

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...displacement. An automatic mechanism of spatial reorientation may well occur in all the spatial conditions, and even in the control task, but we argue that reorientation is much more implicated when information that are used to reorient ourselves, such as the environmental geometry (Wang & Spelke, 2002), have to be explicitly taken into consideration to perform the task. As proposed above, a comparison between the currently perceived environment (local view) and its stored spatial representation, would allow the disoriented subject to determine his position and heading within the environment (Gallistel, 1990). Landmark-centered Coding and the Parietal Cortex Significant differences between the spatial conditions have also been detected in dorsal regions, where the bilateral middle occipital gyrus and the left superior parietal cortex (superior parietal lobule and precuneus) appeared to be significantly more active during the landmark-centered than during the other two spatial conditions. Such a lateralization to the left comes from the fact that the landmark-centered condition induced a more symmetrical activity than the viewer- and object-centered conditions (which involved more the right hemisph...

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...luster. Localization and visualization of activations were achieved by using the in-house BrainShow software, which allows to superimpose statistical maps on brain slices and on folded, inflated, and flattened representations of the cortical surface. For visualization of group activations, we used the cortical surface of the single-subject MNI brain, reconstructed using the FreeSurfer software (surfer.nmr.mgh.harvard.edu). Anatomical labels were automatically assigned to activated areas by the BrainShow software, based on a macroscopical anatomical parcellation of the MNI singlesubject brain (Tzourio-Mazoyer et al., 2002), and then manually checked with reference to a standard atlas (Duvernoy, 1991). Acknowledgments We thank the UNAF/SHFJ/CEA and IFR44, Manuel Vidal and Nicholas Nebiolo for their help in building the 3-D environment, and Jean-Baptiste Poline for his methodological advice. This research was funded by Italian Ministry of Health and European Laboratory of Neurophysiology and Neuropsychology of Action (LENNA). Reprint requests should be sent to Giorgia Committeri, Laboratorio di Neuroimmagini, Fondazione Santa Lucia, via Ardeatina 306, 00179 Rome, Italy, or via e-mail: giorgia.committeri@ uniroma1...

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...., 2002). Landmark-centered Coding and the Ventromedial Occipital–Temporal Cortex One of the most interesting new findings of this study is the fact that the bilateral ventromedial occipito-temporal cortex, including the fusiform, lingual, and posterior parahippocampal gyri, was exclusively activated in the landmark-centered condition, where the geometrical structure of the environment had to be taken as a reference. Fusiform, lingual, and parahippocampal gyri have been all associated with the simple passive viewing of buildings/scenes (Haxby et al., 1999; Aguirre, Zarahn, & D’Esposito, 1998; Epstein & Kanwisher, 1998). In the lingual gyrus, this happens in a rostral region (‘‘lingual landmark area’’ [LLA]) which seems to be crucially involved in the recognition of stimuli with orienting 1526 Journal of Cognitive Neuroscience Volume 16, Number 9 value. In fact, a lesion here often causes ‘‘landmark agnosia’’ (see Aguirre & D’Esposito, 1999, for a review). On the other hand, a region within the posterior PH (‘‘parahippocampal place area’’ [PPA]), seems to be implicated more in the encoding of new place information into memory, than in recognition or navigation (Epstein, DeYoe, Press, Rosen, & Kanwisher, 2001...

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...onse function. Subject-specific contrast images were then entered at the second stage into one-sample t tests, testing the null hypothesis that the mean effect size was equal to zero in the whole population from which our participants were extracted. For each effect of interest, we obtained a statistical parametric map of the t statistic. Clusters of adjacent voxels surviving a threshold of p < .01 were formed and characterized in terms of Committeri et al. 1531 spatial extent. The significance of each cluster was estimated using distribution approximations from the theory of gaussian fields (Worsley, Marret, Neelin, Friston, & Evans, 1995; Worsley, Evans, et al., 1992), resulting in a corrected p value. Correction was applied over the whole brain, or in some cases, over a more limited search volume (see below). Activation clusters were retained as significant at p < .05 corrected. First, we inspected activation for each kind of spatial judgment with respect to the control task (Table 1). Then, we explored differential activation between the spatial conditions using two approaches (Table 2): (a) We performed pairwise voxel-level comparisons between the three spatial conditions, using p value correction over the set of voxels em...

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...sing and Analysis Image preprocessing and statistical analysis were performed using the SPM99 software platform (Wellcome Department of Cognitive Neurology, London, UK), implemented in MATLAB (The MathWorks, Natick, MA, USA). Head movements occurring between functional scans were calculated and a motion correction was applied when movement exceeded either 18 of rotation or 1 mm of translation. Functional MR images were then transformed into a standard space, using transformation parameters determined from the anatomical image through an automatic nonlinear stereotaxic normalization procedure (Friston et al., 1995). The template image was based on average data provided by the Montreal Neurological Institute (MNI brain: Mazziotta, Toga, Evans, Fox, & Lancaster, 1995) and conformed to a standard coordinate referencing system (Talairach & Tournoux, 1988). The final voxel size after normalization was an isotropic 4 mm. Functional images were then spatially smoothed using an isotropic gaussian kernel (8 mm full-width-at-halfmaximum). Group analysis was performed using a two-stage random-effect approach (Friston, Holmes, & Worsley, 1999; Holmes & Friston, 1998). At the first stage, the time series of function...

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...e incoming block appeared in the middle of the screen for 1.5 sec, followed by a 0.48 fixation cross for 1.5 sec. A series of six 2.5-sec trials followed. In each trial, a snapshot of the environment appeared for 1 sec, followed by the 1.5-sec fixation cross. Such a fast presentation rate was chosen in order to minimize eye movements. Imaging Parameters Echo-planar functional MR images (TR = 3 sec, TE = 60 msec, flip angle = 908, image matrix = 64 64, FOV = 240 240 mm, voxel size = 3.75 3.75 4 mm) were acquired in the axial plane using blood oxygenation level-dependent (BOLD) imaging (Kwong, Belliveau, & Chesler, 1992). Twenty-nine transverse slices were acquired without gaps. From the superior convexity, sampling included almost all the cerebral cortex, excluding only the ventral portion of the cerebellum. A three-dimensional high-resolution anatomical image was also acquired for each subject (124 sagittal slices, TR = 10 msec, TE = 2.2 msec, TI = 600 msec, flip angle = 108, image matrix = 256 192, FOV = 240 180 mm, voxel size = 0.94 0.94 1.2 mm). Image Processing and Analysis Image preprocessing and statistical analysis were performed using the SPM99 software platform (Wellcome Department of Cogn...

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... or large (Gouteux & Spelke, 2001; Learmonth, Newcombe, & Huttenlocher, 2001). Thus, the activation pattern of the object-centered condition should be more similar to that of the landmark-centered condition if the reference object was stable in the environment and did not change position from trial to trial. An intriguing result of the present study is the inferior occipital–temporal activation observed for the object-centered coding. It is located more anteriorly than the lateral occipital (LO) region, in which a certain amount of invariance for changes in object position has been described (Grill-Spector et al., 1999). Milner, Jonsrude, and Crane (1997) found a very similar activation during the recall phase of a task that required to memorize the location of objects within two-dimensional arrays. This was observed when the array was shifted with respect to the encoding phase, thus requiring the use of a spatial representation that coded the relationships between objects (as in our object-centered condition). Thus, object-centered coding seems to be implemented also in the ventrolateral object-related cortices, just as the landmark-centered coding is implemented in ventromedial building-specific regions. A...

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...ate referencing system (Talairach & Tournoux, 1988). The final voxel size after normalization was an isotropic 4 mm. Functional images were then spatially smoothed using an isotropic gaussian kernel (8 mm full-width-at-halfmaximum). Group analysis was performed using a two-stage random-effect approach (Friston, Holmes, & Worsley, 1999; Holmes & Friston, 1998). At the first stage, the time series of functional MR images obtained from each participant was analyzed separately. The effects of the experimental paradigm were estimated on a voxel-byvoxel basis, according to the general linear model (Friston, Worsley, Frackowiak, Mazziotta, & Evans, 1994; Worsley, Evans, Marret, & Neelin, 1992). The experimental blocks were modeled as box-car functions, convolved with a synthetic hemodynamic response function. Subject-specific contrast images were then entered at the second stage into one-sample t tests, testing the null hypothesis that the mean effect size was equal to zero in the whole population from which our participants were extracted. For each effect of interest, we obtained a statistical parametric map of the t statistic. Clusters of adjacent voxels surviving a threshold of p < .01 were formed and characterized in terms of Committeri ...

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...al finds himself in a particular location and with a particular direction. In a similar fashion, the monkey ‘‘spatial view cells’’ respond when the animal looks at a particular part of the environment (Georges-Francois, Rolls, & Robertson, 1999; Rolls & O’Mara, 1995). However, an alternate interpretation of the activation of the LLA/PPA cannot be excluded. Attention to buildings has been shown to modulate the activity of the PPA, even in conditions where the presented stimuli included other objects or faces (Avidan, Levy, Hendler, Zohary, & Malach, 2003; O’Craven, Downing, & Kanwisher, 1999; Tong, Nakayama, Vaughan, & Kanwisher, 1998). Thus, in the present study, attention to the building/scene, when it had to be used as reference, may have contributed to the observed activation, by modulating the activity of such specific areas, which do not respond to other kinds of objects (see, for example, Epstein & Kanwisher, 1998). Note however that the differential activation for the landmark-centered condition extended well beyond the PPA, for example, in the retrosplenial and parietal cortex (see below), where no specificity for buildings has been described. Thus, the attentional interpretation cannot hold for the whole pattern ...

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...how a clear dissociation in activated areas, with the ventromedial occipital–temporal and retrosplenial cortex firmly specialized for the landmark-centered judgements, and the lateral occipital–temporal cortex relatively specialized for the object-centered judgements. Thus, our data indicate that using either a movable object or a stable landmark (i.e., the environmental structure) as a spatial reference involves different brain regions. It is reasonable to believe that these results represent the neural correlates of a strong dissociation emerged at behavioral level. Recent evidence in rats (Cheng, 1986), monkeys (Gouteux, Thinus-Blanc, & Vauclair, 2001), and human infants and adults (Gouteux & Spelke, 2001; Wang & Spelke, 2000; Hermer & Spelke, 1994), shows that, during spatial reorientation tasks, the surrounding surface geometric layout is handled differently from nongeometric properties of the environment, such as the position or even the geometrical configuration of a set of (potentially movable) objects. Memory for object locations strongly relies on the surrounding surface layout, which solely survives disorientation and allows reorientation (Wang & Spelke, 2000, 2002). The detected ve...

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...1999). Several neuroimaging studies (Fink, Marshall, Shah, et al., 2000; Galati, Lobel, et al., 2000; Honda, Wise, Weeks, Deiber, & Hallett, 1998; Fink, Dolan, Halligan, Marshall, & Frith, 1997) have found activation in the posterior parietal cortex and the dorsal premotor cortex (PMd), mainly in the right hemisphere, and in early visual processing areas, during object-based spatial judgements. Finally, coding of topographical space is typically associated with the hippocampal formation. Cells with allocentric properties have been found in the hippocampal formation of both freely-moving rats (Taube, Muller, & Ranck, 1990; O’Keefe & Dostrovsky, 1971) and monkeys (Rolls & O’Mara, 1995; Ono, Nakamura, Nishijo, & Eifuku, 1993), as well as in the rat retrosplenial cortex (Chen, Lin, Green, Barnes, & McNaughton, 1994; McNaughton, Leonard, & Chen, 1989), but also in the monkey’s posterior parietal cortex (Snyder, Grieve, Brotchie, & Andersen, 1998). In humans, ventromedial lesions encroaching on the parahippocampal gyrus (Aguirre & D’Esposito, 1999; Habib & Sirigu, 1987) or the hippocampus proper (Spiers, Burgess, Hartley, Vargha-Khadem, & O’Keefe, 2001) lead to difficulties in storing and/or recalling identity and ...

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...ment of Cognitive Neurology, London, UK), implemented in MATLAB (The MathWorks, Natick, MA, USA). Head movements occurring between functional scans were calculated and a motion correction was applied when movement exceeded either 18 of rotation or 1 mm of translation. Functional MR images were then transformed into a standard space, using transformation parameters determined from the anatomical image through an automatic nonlinear stereotaxic normalization procedure (Friston et al., 1995). The template image was based on average data provided by the Montreal Neurological Institute (MNI brain: Mazziotta, Toga, Evans, Fox, & Lancaster, 1995) and conformed to a standard coordinate referencing system (Talairach & Tournoux, 1988). The final voxel size after normalization was an isotropic 4 mm. Functional images were then spatially smoothed using an isotropic gaussian kernel (8 mm full-width-at-halfmaximum). Group analysis was performed using a two-stage random-effect approach (Friston, Holmes, & Worsley, 1999; Holmes & Friston, 1998). At the first stage, the time series of functional MR images obtained from each participant was analyzed separately. The effects of the experimental paradigm were estimated on a voxel-byvoxel basis, ac...

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...els emerged as activated in at least one spatial condition relative to control. Also regions outside this limited volume are reported, when reaching a significant p value corrected over the whole brain; (b) We computed the mean signal increase for each spatial condition relative to the control task over each activated cluster and performed paired t tests on these data. Differences were retained as significant at p < .05. This kind of approach has less localizing power than the voxel-based approach, but offers more sensitivity when the spatial resolution of the signal is lower than voxel size (Friston, Holmes, Poline, Price, & Frith, 1996). Mean signal increases over a given cluster were computed in terms of the first eigenvariate of the BOLD response amplitudes estimated at the first analysis stage (i.e., in each subject) in all voxels of the cluster. Localization and visualization of activations were achieved by using the in-house BrainShow software, which allows to superimpose statistical maps on brain slices and on folded, inflated, and flattened representations of the cortical surface. For visualization of group activations, we used the cortical surface of the single-subject MNI brain, reconstructed using the FreeSurfer s...

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... was detected in the frontal lobe. It comprised the socalled deep frontal eye fields (FEFs) (Lobel et al., 2001) and the PMd, respectively coding oculomotor and reaching-related space in egocentric coordinates (e.g., Boussaoud & Bremmer, 1999; Goldberg & Bruce, 1990). The activation included the rostral portion of PMd, recently termed ‘‘pre-PMd,’’ as it is more closely related to cognitive processes than to motor processes (Picard & Strick, 2001). This region receives strong projections from the dorsolateral prefrontal cortex (DLPFC), which has been suggested to play a role in spatial memory (Wilson, Scalaidhe, & Goldman-Rakic, 1993). Interestingly, our activation extended rostrally inside the SFS, in the putative human homologue of DLPFC (Courtney, Petit, Maisog, Ungerleider, & Haxby, 1998). It is worth noting that the whole right superior frontal activation was shared by the viewer-centered and the landmarkcentered conditions, suggesting common computational mechanisms. The left ventral cortex, instead, showed a tendency to prefer the viewer-centered coding. Centered in the left IFG (BAs 44 and 45), it was already detected in previous experiments of body midline perception (Galati, Committeri, et al., 2001; Vallar et a...

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... et al., 2000; Mellet et al., 2000; Aguirre, Detre, et al., 1996). Such complex tasks entail spatial operations referred both to the viewer and to external references, which are difficult to disentangle. However, taking into account the subject’s navigational strategy, recent studies were able to show that different regions are involved when subjects use different spatial references: for example, the hippocampus is involved only when subjects rely on multiple landmarks available in the environment and their spatial relationships, and not when they use automatic stimulus–response associations (Hartley, Maguire, Spiers, & Burgess, 2003; Iaria, Petrides, Dagher, Pike, & Bohbot, 2003). In summary, there is evidence that distinct neural structures are involved depending on the spatial references employed during various spatial operations. The aim of the present study was to directly compare viewer-centered, object-centered, and landmark-centered spatial coding of the same realistic three-dimensional information, by using a simple perceptual task, which allows to explicitly manipulate the employed spatial reference. We built a 3-D virtual environment representing a square arena in a park, with some relevant landmarks such as a ...

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...ndmark-centered judgements, and the lateral occipital–temporal cortex relatively specialized for the object-centered judgements. Thus, our data indicate that using either a movable object or a stable landmark (i.e., the environmental structure) as a spatial reference involves different brain regions. It is reasonable to believe that these results represent the neural correlates of a strong dissociation emerged at behavioral level. Recent evidence in rats (Cheng, 1986), monkeys (Gouteux, Thinus-Blanc, & Vauclair, 2001), and human infants and adults (Gouteux & Spelke, 2001; Wang & Spelke, 2000; Hermer & Spelke, 1994), shows that, during spatial reorientation tasks, the surrounding surface geometric layout is handled differently from nongeometric properties of the environment, such as the position or even the geometrical configuration of a set of (potentially movable) objects. Memory for object locations strongly relies on the surrounding surface layout, which solely survives disorientation and allows reorientation (Wang & Spelke, 2000, 2002). The detected ventromedial occipital–temporal and retrosplenial activation during the landmark-centered condi1528 Journal of Cognitive Neuroscience Volume 16, Number ...

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...nt during the landmark-centered condition was the bilateral retrosplenial cortex. The retrosplenial cortex, because of its anatomical interposition between parietal and medial temporal regions and of its strong anatomical links with both areas, has been suggested as the transition zone between egocentric and allocentric/mnestic representations (e.g., Aguirre & D’Esposito, 1999). Together with visual areas of the ventral system, the superior temporal sulcus and the posterior parietal cortex, the retrosplenial cortex sends inputs to the parahippocampal gyrus and, through it, to the hippocampus (Suzuki & Amaral, 1994a, 1994b). Previous neuroimaging studies on navigation and orientation in large-scale space consistently detected a bilateral activity in the retrosplenial cortex (Hartley et al., 2003; Mellet et al., 2000; Maguire, Burgess, et al., 1998; Maguire, Frith, et al., 1998; Aguirre & D’Esposito, 1997; Ghaem et al., 1997; Maguire, Frackowiak, & Frith, 1997; Aguirre, Detre, et al., 1996). However, these studies contrasted very different conditions, such as, for example, name–place association and scrambled stimuli (Aguirre & D’Esposito, 1997), not allowing clear conclusions on the specific function of...

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... when the array was shifted with respect to the encoding phase, thus requiring the use of a spatial representation that coded the relationships between objects (as in our object-centered condition). Thus, object-centered coding seems to be implemented also in the ventrolateral object-related cortices, just as the landmark-centered coding is implemented in ventromedial building-specific regions. As pointed above, an attentional bias may also have contributed to the observed activation, by modulating the activity of regions which code objects different from buildings (e.g., Avidan et al., 2003; Wojciulik, Kanwisher, & Driver, 1998). Previous neuroimaging studies have investigated object-based spatial judgements, such as line bisection (Fink, Marshall, Shah, et al., 2000; Fink, Marshall, Weiss, et al., 2000; Galati et al., 2000; Weiss et al., 2000; Fink, Dolan, et al., 1997). With respect to them, the present object-centered condition activated similar posterior parietal regions, but failed in activating the PMd and early visual processing regions, particularly involved when dealing with far space (Weiss et al., 2000) or when the stimulus is more object-like (Fink, Marshall, Weiss, et al., 2000). Differently from the pr...

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... occipital cortex) were significantly more activated during the landmark-centered condition. The ventrolateral occipital–temporal cortex was particularly involved in object-centered coding. Results strongly demonstrate that viewer-centered (egocentric) coding is restricted to the dorsal stream and connected frontal regions, whereas a coding centered on external references requires both dorsal and ventral regions, depending on the reference being a movable object or a landmark. & INTRODUCTION Spatial locations may be defined either relative to the viewer or relative to some external reference (Howard, 1982). In the first case, spatial locations are coded egocentrically, with reference to relevant body parts, such as the eyes, head, trunk, and/or arm. This kind of coding is highly dynamic, needs continuous updating as one moves, and is particularly useful for the organization of movements towards objects in space. External references, instead, are used, for example, in object-based representations, which describe the spatial relationships among the component parts of an object or the relative position of multiple objects. Another common example comes from the study of spatial orientation and navi...

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...her support to a cognitive role for PMv (Rizzolatti et al., 2002). Landmark-centered Coding and the Ventromedial Occipital–Temporal Cortex One of the most interesting new findings of this study is the fact that the bilateral ventromedial occipito-temporal cortex, including the fusiform, lingual, and posterior parahippocampal gyri, was exclusively activated in the landmark-centered condition, where the geometrical structure of the environment had to be taken as a reference. Fusiform, lingual, and parahippocampal gyri have been all associated with the simple passive viewing of buildings/scenes (Haxby et al., 1999; Aguirre, Zarahn, & D’Esposito, 1998; Epstein & Kanwisher, 1998). In the lingual gyrus, this happens in a rostral region (‘‘lingual landmark area’’ [LLA]) which seems to be crucially involved in the recognition of stimuli with orienting 1526 Journal of Cognitive Neuroscience Volume 16, Number 9 value. In fact, a lesion here often causes ‘‘landmark agnosia’’ (see Aguirre & D’Esposito, 1999, for a review). On the other hand, a region within the posterior PH (‘‘parahippocampal place area’’ [PPA]), seems to be implicated more in the encoding of new place information into memory, than in recogniti...

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...ivation was shared by the viewer-centered and the landmarkcentered conditions, suggesting common computational mechanisms. The left ventral cortex, instead, showed a tendency to prefer the viewer-centered coding. Centered in the left IFG (BAs 44 and 45), it was already detected in previous experiments of body midline perception (Galati, Committeri, et al., 2001; Vallar et al., 1999), during active pointing (e.g., Lacquaniti et al., 1997), and also during passive observation of grasping movements (Rizzolatti, Fadiga, Gallese, & Fogassi, 1996), so that analogies have been drawn with monkey PMv (Rizzolatti, Fogassi, & Gallese, 2002). More specifically, area 44 is considered homologous to the rostral PMv (F5), whereas area 6 to the caudal PMv (F4). PMv is part of those circuits coding peripersonal (F4) and grasping-related space (F5), thus crucially involved in egocentric spatial representations (see Colby, 1998, for a review). Besides the motor role, the present data bring further support to a cognitive role for PMv (Rizzolatti et al., 2002). Landmark-centered Coding and the Ventromedial Occipital–Temporal Cortex One of the most interesting new findings of this study is the fact that the bilateral ventromedial occipito-...

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...iewer-centered and body-centered frames may be intrinsically different. A region significantly more active for the viewercentered coding than for the object-centered coding was detected in the frontal lobe. It comprised the socalled deep frontal eye fields (FEFs) (Lobel et al., 2001) and the PMd, respectively coding oculomotor and reaching-related space in egocentric coordinates (e.g., Boussaoud & Bremmer, 1999; Goldberg & Bruce, 1990). The activation included the rostral portion of PMd, recently termed ‘‘pre-PMd,’’ as it is more closely related to cognitive processes than to motor processes (Picard & Strick, 2001). This region receives strong projections from the dorsolateral prefrontal cortex (DLPFC), which has been suggested to play a role in spatial memory (Wilson, Scalaidhe, & Goldman-Rakic, 1993). Interestingly, our activation extended rostrally inside the SFS, in the putative human homologue of DLPFC (Courtney, Petit, Maisog, Ungerleider, & Haxby, 1998). It is worth noting that the whole right superior frontal activation was shared by the viewer-centered and the landmarkcentered conditions, suggesting common computational mechanisms. The left ventral cortex, instead, showed a tendency to prefer t...

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...ple objects. Another common example comes from the study of spatial orientation and navigation in humans and animals. Generally speaking, memory for places and way-finding are processes that are easier to understand in terms of coordinates external to the viewer, or allocentric (Berthoz, 1997). Indeed, most accounts of spatial orientation include some kind of enduring storage of environmental information, in the form of internal geocentric ‘‘cognitive maps’’ of topographical space (O’Keefe & Nadel, 1978), or of modules for the permanent storage of the geometrical structure of spatial layouts (Wang & Spelke, 2002). Egocentric frames are typically associated with the posterior parietal cortex. Neurons coding spatial position relative to body parts have been found in the monkey’s posterior parietal cortex and in connected regions of the premotor cortex (Cohen & Andersen, 2002; Colby, 1998). Patients with lesions to the posterior parietal cortex may either show inaccurate visuomotor coordination (optic ataxia: Perenin & Vighetto, 1988) or fail to explore the contralesional side of space (unilateral neglect: Vallar, 1998). In unilateral neglect, the affected sector of space is usually egocentrically define...

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...and mentally completed the visual scene. Furthermore, the left precuneus has been found more involved during mental navigation in an environment learned from a route perspective, than during spatial tasks on a map learned from a survey perspective (Mellet et al., 2000). To this regard, it is noteworthy that our subjects learned the environment from a route perspective and never saw the map. As for the middle occipital cortex, it was found to be activated in several previous neuroimaging studies of orientation in large-scale space (e.g., Maguire, Frith, et al., 1998). Moreover, a recent study (Hasson, Harel, Levy, & Malach, 2003) found a dorsal building-specific area located slightly beneath the present cluster of activity. The attentional modulation mechanism put forward in the previous paragraph may accounts for this activation. Landmark- versus Object-centered Coding It may be argued that, in the present perceptual paradigm, the presence of an observer and of incoming visual stimulation to be coded by this observer, may have introduced an automatic egocentric component in the other two conditions. Bearing in mind such theoretical argument, the present experiment has been designed for allowing direct comparisons be...

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...les for the permanent storage of the geometrical structure of spatial layouts (Wang & Spelke, 2002). Egocentric frames are typically associated with the posterior parietal cortex. Neurons coding spatial position relative to body parts have been found in the monkey’s posterior parietal cortex and in connected regions of the premotor cortex (Cohen & Andersen, 2002; Colby, 1998). Patients with lesions to the posterior parietal cortex may either show inaccurate visuomotor coordination (optic ataxia: Perenin & Vighetto, 1988) or fail to explore the contralesional side of space (unilateral neglect: Vallar, 1998). In unilateral neglect, the affected sector of space is usually egocentrically defined (see Bisiach, 1997; Vallar, Guariglia, & Rusconi, 1997, for related demonstrations). Both ataxic and neglect patients show specific deficits in perceiving the position of their body midsagittal plane (Pizzamiglio, Committeri, Galati, & Patria, 2000; Perenin, 1997), a fundamental egocentric spatial principle (Jeannerod & Biguer, 1989). A series of neuroimaging studies has provided direct evidence of the involvement of parietal–frontal circuits in the egocentric coding of space. A posterior parietal– frontal ...

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... or allocentric (Berthoz, 1997). Indeed, most accounts of spatial orientation include some kind of enduring storage of environmental information, in the form of internal geocentric ‘‘cognitive maps’’ of topographical space (O’Keefe & Nadel, 1978), or of modules for the permanent storage of the geometrical structure of spatial layouts (Wang & Spelke, 2002). Egocentric frames are typically associated with the posterior parietal cortex. Neurons coding spatial position relative to body parts have been found in the monkey’s posterior parietal cortex and in connected regions of the premotor cortex (Cohen & Andersen, 2002; Colby, 1998). Patients with lesions to the posterior parietal cortex may either show inaccurate visuomotor coordination (optic ataxia: Perenin & Vighetto, 1988) or fail to explore the contralesional side of space (unilateral neglect: Vallar, 1998). In unilateral neglect, the affected sector of space is usually egocentrically defined (see Bisiach, 1997; Vallar, Guariglia, & Rusconi, 1997, for related demonstrations). Both ataxic and neglect patients show specific deficits in perceiving the position of their body midsagittal plane (Pizzamiglio, Committeri, Galati, & Patria, 2000; Perenin, 1997...

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...f internal geocentric ‘‘cognitive maps’’ of topographical space (O’Keefe & Nadel, 1978), or of modules for the permanent storage of the geometrical structure of spatial layouts (Wang & Spelke, 2002). Egocentric frames are typically associated with the posterior parietal cortex. Neurons coding spatial position relative to body parts have been found in the monkey’s posterior parietal cortex and in connected regions of the premotor cortex (Cohen & Andersen, 2002; Colby, 1998). Patients with lesions to the posterior parietal cortex may either show inaccurate visuomotor coordination (optic ataxia: Perenin & Vighetto, 1988) or fail to explore the contralesional side of space (unilateral neglect: Vallar, 1998). In unilateral neglect, the affected sector of space is usually egocentrically defined (see Bisiach, 1997; Vallar, Guariglia, & Rusconi, 1997, for related demonstrations). Both ataxic and neglect patients show specific deficits in perceiving the position of their body midsagittal plane (Pizzamiglio, Committeri, Galati, & Patria, 2000; Perenin, 1997), a fundamental egocentric spatial principle (Jeannerod & Biguer, 1989). A series of neuroimaging studies has provided direct evidence of the involvement of pari...

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...n in new and/or familiar environments. Also patients with retrosplenial lesions are unable to orient themselves and to know which direction to take, as if they had lost their ‘‘heading’’ within the environment (Aguirre & D’Esposito, 1999). Neuroimaging studies using a variety of topographical tasks, such as landmark knowledge, orientation in largescale space and navigation, have detected activation in the posterior parahippocampal cortex (e.g., Gron, Wunderlich, Spitzer, Tomczak, & Riepe, 2000; Mellet et al., 2000; Maguire, Frith, Burgess, Donnett, & O’Keefe, 1998; Aguirre & D’Esposito, 1997; Ghaem et al., 1997; Maguire, Frackowiak, & Frith, 1997; Aguirre, Detre, Alsop, & D’Esposito, 1996), the hippocampus (Gron et al., 2000; Ghaem et al., 1997; Maguire, Frackowiak, & Frith, 1996, 1997), and the retrosplenial cortex (reviewed by Maguire, 2001), as well as the posterior parietal cortex (e.g., Gron et al., 2000; Mellet et al., 2000; Aguirre, Detre, et al., 1996). Such complex tasks entail spatial operations referred both to the viewer and to external references, which are difficult to disentangle. However, taking into account the subject’s navigational strategy, recent studies were able to show that d...

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...liar environments. Also patients with retrosplenial lesions are unable to orient themselves and to know which direction to take, as if they had lost their ‘‘heading’’ within the environment (Aguirre & D’Esposito, 1999). Neuroimaging studies using a variety of topographical tasks, such as landmark knowledge, orientation in largescale space and navigation, have detected activation in the posterior parahippocampal cortex (e.g., Gron, Wunderlich, Spitzer, Tomczak, & Riepe, 2000; Mellet et al., 2000; Maguire, Frith, Burgess, Donnett, & O’Keefe, 1998; Aguirre & D’Esposito, 1997; Ghaem et al., 1997; Maguire, Frackowiak, & Frith, 1997; Aguirre, Detre, Alsop, & D’Esposito, 1996), the hippocampus (Gron et al., 2000; Ghaem et al., 1997; Maguire, Frackowiak, & Frith, 1996, 1997), and the retrosplenial cortex (reviewed by Maguire, 2001), as well as the posterior parietal cortex (e.g., Gron et al., 2000; Mellet et al., 2000; Aguirre, Detre, et al., 1996). Such complex tasks entail spatial operations referred both to the viewer and to external references, which are difficult to disentangle. However, taking into account the subject’s navigational strategy, recent studies were able to show that different regions are involved when s...

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..., 2001) lead to difficulties in storing and/or recalling identity and spatial location of important landmarks, resulting in topographical disorientation in new and/or familiar environments. Also patients with retrosplenial lesions are unable to orient themselves and to know which direction to take, as if they had lost their ‘‘heading’’ within the environment (Aguirre & D’Esposito, 1999). Neuroimaging studies using a variety of topographical tasks, such as landmark knowledge, orientation in largescale space and navigation, have detected activation in the posterior parahippocampal cortex (e.g., Gron, Wunderlich, Spitzer, Tomczak, & Riepe, 2000; Mellet et al., 2000; Maguire, Frith, Burgess, Donnett, & O’Keefe, 1998; Aguirre & D’Esposito, 1997; Ghaem et al., 1997; Maguire, Frackowiak, & Frith, 1997; Aguirre, Detre, Alsop, & D’Esposito, 1996), the hippocampus (Gron et al., 2000; Ghaem et al., 1997; Maguire, Frackowiak, & Frith, 1996, 1997), and the retrosplenial cortex (reviewed by Maguire, 2001), as well as the posterior parietal cortex (e.g., Gron et al., 2000; Mellet et al., 2000; Aguirre, Detre, et al., 1996). Such complex tasks entail spatial operations referred both to the viewer and to external references, which are difficult t...

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...sphere, and in early visual processing areas, during object-based spatial judgements. Finally, coding of topographical space is typically associated with the hippocampal formation. Cells with allocentric properties have been found in the hippocampal formation of both freely-moving rats (Taube, Muller, & Ranck, 1990; O’Keefe & Dostrovsky, 1971) and monkeys (Rolls & O’Mara, 1995; Ono, Nakamura, Nishijo, & Eifuku, 1993), as well as in the rat retrosplenial cortex (Chen, Lin, Green, Barnes, & McNaughton, 1994; McNaughton, Leonard, & Chen, 1989), but also in the monkey’s posterior parietal cortex (Snyder, Grieve, Brotchie, & Andersen, 1998). In humans, ventromedial lesions encroaching on the parahippocampal gyrus (Aguirre & D’Esposito, 1999; Habib & Sirigu, 1987) or the hippocampus proper (Spiers, Burgess, Hartley, Vargha-Khadem, & O’Keefe, 2001) lead to difficulties in storing and/or recalling identity and spatial location of important landmarks, resulting in topographical disorientation in new and/or familiar environments. Also patients with retrosplenial lesions are unable to orient themselves and to know which direction to take, as if they had lost their ‘‘heading’’ within the environment (Aguirre & D’Esposito, 1999). Neuro...

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..., Detre, et al., 1996). Such complex tasks entail spatial operations referred both to the viewer and to external references, which are difficult to disentangle. However, taking into account the subject’s navigational strategy, recent studies were able to show that different regions are involved when subjects use different spatial references: for example, the hippocampus is involved only when subjects rely on multiple landmarks available in the environment and their spatial relationships, and not when they use automatic stimulus–response associations (Hartley, Maguire, Spiers, & Burgess, 2003; Iaria, Petrides, Dagher, Pike, & Bohbot, 2003). In summary, there is evidence that distinct neural structures are involved depending on the spatial references employed during various spatial operations. The aim of the present study was to directly compare viewer-centered, object-centered, and landmark-centered spatial coding of the same realistic three-dimensional information, by using a simple perceptual task, which allows to explicitly manipulate the employed spatial reference. We built a 3-D virtual environment representing a square arena in a park, with some relevant landmarks such as a three-winged palace and a fountain (see Figure ...

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...n in activated areas, with the ventromedial occipital–temporal and retrosplenial cortex firmly specialized for the landmark-centered judgements, and the lateral occipital–temporal cortex relatively specialized for the object-centered judgements. Thus, our data indicate that using either a movable object or a stable landmark (i.e., the environmental structure) as a spatial reference involves different brain regions. It is reasonable to believe that these results represent the neural correlates of a strong dissociation emerged at behavioral level. Recent evidence in rats (Cheng, 1986), monkeys (Gouteux, Thinus-Blanc, & Vauclair, 2001), and human infants and adults (Gouteux & Spelke, 2001; Wang & Spelke, 2000; Hermer & Spelke, 1994), shows that, during spatial reorientation tasks, the surrounding surface geometric layout is handled differently from nongeometric properties of the environment, such as the position or even the geometrical configuration of a set of (potentially movable) objects. Memory for object locations strongly relies on the surrounding surface layout, which solely survives disorientation and allows reorientation (Wang & Spelke, 2000, 2002). The detected ventromedial occipital–temporal and retrosplenial ac...

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...relies on the surrounding surface layout, which solely survives disorientation and allows reorientation (Wang & Spelke, 2000, 2002). The detected ventromedial occipital–temporal and retrosplenial activation during the landmark-centered condi1528 Journal of Cognitive Neuroscience Volume 16, Number 9 tion may represent the neural correlate of such encapsulated module for geometry (Wang & Spelke, 2002). Even if experimental data are still incongruent, it seems that nongeometric properties can be used in particular situations, as when they are particularly stable or large (Gouteux & Spelke, 2001; Learmonth, Newcombe, & Huttenlocher, 2001). Thus, the activation pattern of the object-centered condition should be more similar to that of the landmark-centered condition if the reference object was stable in the environment and did not change position from trial to trial. An intriguing result of the present study is the inferior occipital–temporal activation observed for the object-centered coding. It is located more anteriorly than the lateral occipital (LO) region, in which a certain amount of invariance for changes in object position has been described (Grill-Spector et al., 1999). Milner, Jonsrude, and Crane (1997) found a very...

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..., 1997). Indeed, most accounts of spatial orientation include some kind of enduring storage of environmental information, in the form of internal geocentric ‘‘cognitive maps’’ of topographical space (O’Keefe & Nadel, 1978), or of modules for the permanent storage of the geometrical structure of spatial layouts (Wang & Spelke, 2002). Egocentric frames are typically associated with the posterior parietal cortex. Neurons coding spatial position relative to body parts have been found in the monkey’s posterior parietal cortex and in connected regions of the premotor cortex (Cohen & Andersen, 2002; Colby, 1998). Patients with lesions to the posterior parietal cortex may either show inaccurate visuomotor coordination (optic ataxia: Perenin & Vighetto, 1988) or fail to explore the contralesional side of space (unilateral neglect: Vallar, 1998). In unilateral neglect, the affected sector of space is usually egocentrically defined (see Bisiach, 1997; Vallar, Guariglia, & Rusconi, 1997, for related demonstrations). Both ataxic and neglect patients show specific deficits in perceiving the position of their body midsagittal plane (Pizzamiglio, Committeri, Galati, & Patria, 2000; Perenin, 1997), a fundament...

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...splenial cortex firmly specialized for the landmark-centered judgements, and the lateral occipital–temporal cortex relatively specialized for the object-centered judgements. Thus, our data indicate that using either a movable object or a stable landmark (i.e., the environmental structure) as a spatial reference involves different brain regions. It is reasonable to believe that these results represent the neural correlates of a strong dissociation emerged at behavioral level. Recent evidence in rats (Cheng, 1986), monkeys (Gouteux, Thinus-Blanc, & Vauclair, 2001), and human infants and adults (Gouteux & Spelke, 2001; Wang & Spelke, 2000; Hermer & Spelke, 1994), shows that, during spatial reorientation tasks, the surrounding surface geometric layout is handled differently from nongeometric properties of the environment, such as the position or even the geometrical configuration of a set of (potentially movable) objects. Memory for object locations strongly relies on the surrounding surface layout, which solely survives disorientation and allows reorientation (Wang & Spelke, 2000, 2002). The detected ventromedial occipital–temporal and retrosplenial activation during the landmark-centered condi1528 Journal...

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...posterior parietal cortex and the dorsal premotor cortex (PMd), mainly in the right hemisphere, and in early visual processing areas, during object-based spatial judgements. Finally, coding of topographical space is typically associated with the hippocampal formation. Cells with allocentric properties have been found in the hippocampal formation of both freely-moving rats (Taube, Muller, & Ranck, 1990; O’Keefe & Dostrovsky, 1971) and monkeys (Rolls & O’Mara, 1995; Ono, Nakamura, Nishijo, & Eifuku, 1993), as well as in the rat retrosplenial cortex (Chen, Lin, Green, Barnes, & McNaughton, 1994; McNaughton, Leonard, & Chen, 1989), but also in the monkey’s posterior parietal cortex (Snyder, Grieve, Brotchie, & Andersen, 1998). In humans, ventromedial lesions encroaching on the parahippocampal gyrus (Aguirre & D’Esposito, 1999; Habib & Sirigu, 1987) or the hippocampus proper (Spiers, Burgess, Hartley, Vargha-Khadem, & O’Keefe, 2001) lead to difficulties in storing and/or recalling identity and spatial location of important landmarks, resulting in topographical disorientation in new and/or familiar environments. Also patients with retrosplenial lesions are unable to orient themselves and to know which direction to take,...

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... the viewer- and object-centered conditions (which involved more the right hemisphere). Together with the retrosplenial cortex, the posterior parietal cortex and the precuneus would participate in the perceptual processing of the visual scene necessary ‘‘to establish and maintain one’s bearings’’ (Hartley et al., 2003). In the monkey, for example, the majority of neurons in parietal area 7a have gain fields for head position that are referenced to the world (the experimental room) (Snyder et al., 1998). The precuneus seems also to have some relation with memory- and imagery-related processes (Fletcher et al., 1995). In our case, the spatial reference frame used in the landmark-centered condition was part of a stored representation, and was only partially visible. It is reasonable to think that our subjects recalled the stored information and mentally completed the visual scene. Furthermore, the left precuneus has been found more involved during mental navigation in an environment learned from a route perspective, than during spatial tasks on a map learned from a survey perspective (Mellet et al., 2000). To this regard, it is noteworthy that our subjects learned the environment from a route perspective a...

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..., & Frith, 1997) have found activation in the posterior parietal cortex and the dorsal premotor cortex (PMd), mainly in the right hemisphere, and in early visual processing areas, during object-based spatial judgements. Finally, coding of topographical space is typically associated with the hippocampal formation. Cells with allocentric properties have been found in the hippocampal formation of both freely-moving rats (Taube, Muller, & Ranck, 1990; O’Keefe & Dostrovsky, 1971) and monkeys (Rolls & O’Mara, 1995; Ono, Nakamura, Nishijo, & Eifuku, 1993), as well as in the rat retrosplenial cortex (Chen, Lin, Green, Barnes, & McNaughton, 1994; McNaughton, Leonard, & Chen, 1989), but also in the monkey’s posterior parietal cortex (Snyder, Grieve, Brotchie, & Andersen, 1998). In humans, ventromedial lesions encroaching on the parahippocampal gyrus (Aguirre & D’Esposito, 1999; Habib & Sirigu, 1987) or the hippocampus proper (Spiers, Burgess, Hartley, Vargha-Khadem, & O’Keefe, 2001) lead to difficulties in storing and/or recalling identity and spatial location of important landmarks, resulting in topographical disorientation in new and/or familiar environments. Also patients with retrosplenial lesions are unable to orient themselves a...

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...ad learned the environmental structure before the functional MR session. To perform the task, they had to recall the global environmental geometry from memory, because the central wing of the palace could only be recognized by means of its geometrical relationship with the two lateral wings. Therefore, the ventromedial activation could reflect the coding of the current spatial relationship between the viewer and the environmental geometry (see Epstein, Graham, & Downing, 2003), and its matching with the stored environmental representation. Such behavior recalls the ‘‘local-view place cells’’ (McNaughton, Chen, & Markus, 1991) in the rat hippocampal formation, which fire whenever the animal finds himself in a particular location and with a particular direction. In a similar fashion, the monkey ‘‘spatial view cells’’ respond when the animal looks at a particular part of the environment (Georges-Francois, Rolls, & Robertson, 1999; Rolls & O’Mara, 1995). However, an alternate interpretation of the activation of the LLA/PPA cannot be excluded. Attention to buildings has been shown to modulate the activity of the PPA, even in conditions where the presented stimuli included other objects or faces (Avidan, Levy, Hendler...

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...d with the hippocampal formation. Cells with allocentric properties have been found in the hippocampal formation of both freely-moving rats (Taube, Muller, & Ranck, 1990; O’Keefe & Dostrovsky, 1971) and monkeys (Rolls & O’Mara, 1995; Ono, Nakamura, Nishijo, & Eifuku, 1993), as well as in the rat retrosplenial cortex (Chen, Lin, Green, Barnes, & McNaughton, 1994; McNaughton, Leonard, & Chen, 1989), but also in the monkey’s posterior parietal cortex (Snyder, Grieve, Brotchie, & Andersen, 1998). In humans, ventromedial lesions encroaching on the parahippocampal gyrus (Aguirre & D’Esposito, 1999; Habib & Sirigu, 1987) or the hippocampus proper (Spiers, Burgess, Hartley, Vargha-Khadem, & O’Keefe, 2001) lead to difficulties in storing and/or recalling identity and spatial location of important landmarks, resulting in topographical disorientation in new and/or familiar environments. Also patients with retrosplenial lesions are unable to orient themselves and to know which direction to take, as if they had lost their ‘‘heading’’ within the environment (Aguirre & D’Esposito, 1999). Neuroimaging studies using a variety of topographical tasks, such as landmark knowledge, orientation in largescale space and naviga...

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...euroimaging studies using a variety of topographical tasks, such as landmark knowledge, orientation in largescale space and navigation, have detected activation in the posterior parahippocampal cortex (e.g., Gron, Wunderlich, Spitzer, Tomczak, & Riepe, 2000; Mellet et al., 2000; Maguire, Frith, Burgess, Donnett, & O’Keefe, 1998; Aguirre & D’Esposito, 1997; Ghaem et al., 1997; Maguire, Frackowiak, & Frith, 1997; Aguirre, Detre, Alsop, & D’Esposito, 1996), the hippocampus (Gron et al., 2000; Ghaem et al., 1997; Maguire, Frackowiak, & Frith, 1996, 1997), and the retrosplenial cortex (reviewed by Maguire, 2001), as well as the posterior parietal cortex (e.g., Gron et al., 2000; Mellet et al., 2000; Aguirre, Detre, et al., 1996). Such complex tasks entail spatial operations referred both to the viewer and to external references, which are difficult to disentangle. However, taking into account the subject’s navigational strategy, recent studies were able to show that different regions are involved when subjects use different spatial references: for example, the hippocampus is involved only when subjects rely on multiple landmarks available in the environment and their spatial relationships, and not wh...

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...ll as the virtual far space may have underestimated its preference for the near prehension space (Weiss et al., 2000), in both cases reducing the parietal involvement. Alternatively, viewer-centered and body-centered frames may be intrinsically different. A region significantly more active for the viewercentered coding than for the object-centered coding was detected in the frontal lobe. It comprised the socalled deep frontal eye fields (FEFs) (Lobel et al., 2001) and the PMd, respectively coding oculomotor and reaching-related space in egocentric coordinates (e.g., Boussaoud & Bremmer, 1999; Goldberg & Bruce, 1990). The activation included the rostral portion of PMd, recently termed ‘‘pre-PMd,’’ as it is more closely related to cognitive processes than to motor processes (Picard & Strick, 2001). This region receives strong projections from the dorsolateral prefrontal cortex (DLPFC), which has been suggested to play a role in spatial memory (Wilson, Scalaidhe, & Goldman-Rakic, 1993). Interestingly, our activation extended rostrally inside the SFS, in the putative human homologue of DLPFC (Courtney, Petit, Maisog, Ungerleider, & Haxby, 1998). It is worth noting that the whole right superior frontal activa...

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...e spatial conditions might suggest an eye movements effect. However, eye movements were controlled in the preliminary psychophysical study, thus they cannot have determined the present parietal activation. Even if little can be concluded about a negative result, the most likely interpretation for the parietal involvement is that it has a role in distance computation, that is, a common cognitive component across the three spatial conditions and absent in the control task. Distance estimation is in fact classically disturbed in subjects having lesions to the posterior dorsolateral cortex (e.g., Holmes, 1918; Balint, 1909). Conclusions Object localization in a complex environment requires the conjoint activity of several regions, depending on the spatial reference to be used: if viewer-centered, the parietal lobe and connected frontal premotor regions are recruited, whereas if centered on some external reference, both parietal–frontal and ventral regions play a role. Object-centered spatial judgements involve the ventrolateral occipital–temporal cortex, while landmarkcentered judgements require a strongly specialized circuit of ventromedial occipital–temporal and retrosplenial regions. In our op...

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...nt during the landmark-centered condition was the bilateral retrosplenial cortex. The retrosplenial cortex, because of its anatomical interposition between parietal and medial temporal regions and of its strong anatomical links with both areas, has been suggested as the transition zone between egocentric and allocentric/mnestic representations (e.g., Aguirre & D’Esposito, 1999). Together with visual areas of the ventral system, the superior temporal sulcus and the posterior parietal cortex, the retrosplenial cortex sends inputs to the parahippocampal gyrus and, through it, to the hippocampus (Suzuki & Amaral, 1994a, 1994b). Previous neuroimaging studies on navigation and orientation in large-scale space consistently detected a bilateral activity in the retrosplenial cortex (Hartley et al., 2003; Mellet et al., 2000; Maguire, Burgess, et al., 1998; Maguire, Frith, et al., 1998; Aguirre & D’Esposito, 1997; Ghaem et al., 1997; Maguire, Frackowiak, & Frith, 1997; Aguirre, Detre, et al., 1996). However, these studies contrasted very different conditions, such as, for example, name–place association and scrambled stimuli (Aguirre & D’Esposito, 1997), not allowing clear conclusions on the specific function of...

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...lling identity and spatial location of important landmarks, resulting in topographical disorientation in new and/or familiar environments. Also patients with retrosplenial lesions are unable to orient themselves and to know which direction to take, as if they had lost their ‘‘heading’’ within the environment (Aguirre & D’Esposito, 1999). Neuroimaging studies using a variety of topographical tasks, such as landmark knowledge, orientation in largescale space and navigation, have detected activation in the posterior parahippocampal cortex (e.g., Gron, Wunderlich, Spitzer, Tomczak, & Riepe, 2000; Mellet et al., 2000; Maguire, Frith, Burgess, Donnett, & O’Keefe, 1998; Aguirre & D’Esposito, 1997; Ghaem et al., 1997; Maguire, Frackowiak, & Frith, 1997; Aguirre, Detre, Alsop, & D’Esposito, 1996), the hippocampus (Gron et al., 2000; Ghaem et al., 1997; Maguire, Frackowiak, & Frith, 1996, 1997), and the retrosplenial cortex (reviewed by Maguire, 2001), as well as the posterior parietal cortex (e.g., Gron et al., 2000; Mellet et al., 2000; Aguirre, Detre, et al., 1996). Such complex tasks entail spatial operations referred both to the viewer and to external references, which are difficult to disentangle. Howeve...

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...tic nonlinear stereotaxic normalization procedure (Friston et al., 1995). The template image was based on average data provided by the Montreal Neurological Institute (MNI brain: Mazziotta, Toga, Evans, Fox, & Lancaster, 1995) and conformed to a standard coordinate referencing system (Talairach & Tournoux, 1988). The final voxel size after normalization was an isotropic 4 mm. Functional images were then spatially smoothed using an isotropic gaussian kernel (8 mm full-width-at-halfmaximum). Group analysis was performed using a two-stage random-effect approach (Friston, Holmes, & Worsley, 1999; Holmes & Friston, 1998). At the first stage, the time series of functional MR images obtained from each participant was analyzed separately. The effects of the experimental paradigm were estimated on a voxel-byvoxel basis, according to the general linear model (Friston, Worsley, Frackowiak, Mazziotta, & Evans, 1994; Worsley, Evans, Marret, & Neelin, 1992). The experimental blocks were modeled as box-car functions, convolved with a synthetic hemodynamic response function. Subject-specific contrast images were then entered at the second stage into one-sample t tests, testing the null hypothesis that the mean effect si...

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...of egocentric coding, as well as the virtual far space may have underestimated its preference for the near prehension space (Weiss et al., 2000), in both cases reducing the parietal involvement. Alternatively, viewer-centered and body-centered frames may be intrinsically different. A region significantly more active for the viewercentered coding than for the object-centered coding was detected in the frontal lobe. It comprised the socalled deep frontal eye fields (FEFs) (Lobel et al., 2001) and the PMd, respectively coding oculomotor and reaching-related space in egocentric coordinates (e.g., Boussaoud & Bremmer, 1999; Goldberg & Bruce, 1990). The activation included the rostral portion of PMd, recently termed ‘‘pre-PMd,’’ as it is more closely related to cognitive processes than to motor processes (Picard & Strick, 2001). This region receives strong projections from the dorsolateral prefrontal cortex (DLPFC), which has been suggested to play a role in spatial memory (Wilson, Scalaidhe, & Goldman-Rakic, 1993). Interestingly, our activation extended rostrally inside the SFS, in the putative human homologue of DLPFC (Courtney, Petit, Maisog, Ungerleider, & Haxby, 1998). It is worth noting that the whole righ...

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...Cognitive Neuroscience 16:9, pp. 1517–1535 localization of both visual and tactile stimuli (Galati, Committeri, Sanes, & Pizzamiglio, 2001). Object-based coding of space has been much less studied. Its existence has been demonstrated in the monkey’s frontal lobe (Olson & Gettner, 1995). In humans, unilateral neglect may concern the contralesional side of individual objects, independent of their egocentric position (‘‘object-based neglect’’: Driver, 1999). Several neuroimaging studies (Fink, Marshall, Shah, et al., 2000; Galati, Lobel, et al., 2000; Honda, Wise, Weeks, Deiber, & Hallett, 1998; Fink, Dolan, Halligan, Marshall, & Frith, 1997) have found activation in the posterior parietal cortex and the dorsal premotor cortex (PMd), mainly in the right hemisphere, and in early visual processing areas, during object-based spatial judgements. Finally, coding of topographical space is typically associated with the hippocampal formation. Cells with allocentric properties have been found in the hippocampal formation of both freely-moving rats (Taube, Muller, & Ranck, 1990; O’Keefe & Dostrovsky, 1971) and monkeys (Rolls & O’Mara, 1995; Ono, Nakamura, Nishijo, & Eifuku, 1993), as well as in the rat retrosplenial cortex (Chen, Lin, Gree...

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...them, the present object-centered condition activated similar posterior parietal regions, but failed in activating the PMd and early visual processing regions, particularly involved when dealing with far space (Weiss et al., 2000) or when the stimulus is more object-like (Fink, Marshall, Weiss, et al., 2000). Differently from the present experiment, those studies employed judgements relative to the spatial relationships among the component parts of a single object. This may suggest that ‘‘within-objects’’ and ‘‘between-objects’’ spatial coding is treated in different areas in the human brain (Humphreys & Riddoch, 1994). In our case, the important finding is the specificity of the object-centered ventrolateral activation with respect to a viewer-centered spatial coding. Distance Estimation and the Parietal Lobe Common parietal activity in the spatial conditions might suggest an eye movements effect. However, eye movements were controlled in the preliminary psychophysical study, thus they cannot have determined the present parietal activation. Even if little can be concluded about a negative result, the most likely interpretation for the parietal involvement is that it has a role in distance computation, that...

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...s performed on the same stimuli (Galati, Lobel, et al., 2000), and is found for body-centered 1SHFJ/CEA, Orsay, France, 2Fondazione Santa Lucia, Rome, Italy, 3University G. d’Annunzio, Chieti, Italy, 4CNRS—College de France, Paris, France, 5University of Rome La Sapienza, Italy D 2004 Massachusetts Institute of Technology Journal of Cognitive Neuroscience 16:9, pp. 1517–1535 localization of both visual and tactile stimuli (Galati, Committeri, Sanes, & Pizzamiglio, 2001). Object-based coding of space has been much less studied. Its existence has been demonstrated in the monkey’s frontal lobe (Olson & Gettner, 1995). In humans, unilateral neglect may concern the contralesional side of individual objects, independent of their egocentric position (‘‘object-based neglect’’: Driver, 1999). Several neuroimaging studies (Fink, Marshall, Shah, et al., 2000; Galati, Lobel, et al., 2000; Honda, Wise, Weeks, Deiber, & Hallett, 1998; Fink, Dolan, Halligan, Marshall, & Frith, 1997) have found activation in the posterior parietal cortex and the dorsal premotor cortex (PMd), mainly in the right hemisphere, and in early visual processing areas, during object-based spatial judgements. Finally, coding of topographical sp...

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... rostrally inside the SFS, in the putative human homologue of DLPFC (Courtney, Petit, Maisog, Ungerleider, & Haxby, 1998). It is worth noting that the whole right superior frontal activation was shared by the viewer-centered and the landmarkcentered conditions, suggesting common computational mechanisms. The left ventral cortex, instead, showed a tendency to prefer the viewer-centered coding. Centered in the left IFG (BAs 44 and 45), it was already detected in previous experiments of body midline perception (Galati, Committeri, et al., 2001; Vallar et al., 1999), during active pointing (e.g., Lacquaniti et al., 1997), and also during passive observation of grasping movements (Rizzolatti, Fadiga, Gallese, & Fogassi, 1996), so that analogies have been drawn with monkey PMv (Rizzolatti, Fogassi, & Gallese, 2002). More specifically, area 44 is considered homologous to the rostral PMv (F5), whereas area 6 to the caudal PMv (F4). PMv is part of those circuits coding peripersonal (F4) and grasping-related space (F5), thus crucially involved in egocentric spatial representations (see Colby, 1998, for a review). Besides the motor role, the present data bring further support to a cognitive role for PMv (Rizzolatti ...

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...continuous updating as one moves, and is particularly useful for the organization of movements towards objects in space. External references, instead, are used, for example, in object-based representations, which describe the spatial relationships among the component parts of an object or the relative position of multiple objects. Another common example comes from the study of spatial orientation and navigation in humans and animals. Generally speaking, memory for places and way-finding are processes that are easier to understand in terms of coordinates external to the viewer, or allocentric (Berthoz, 1997). Indeed, most accounts of spatial orientation include some kind of enduring storage of environmental information, in the form of internal geocentric ‘‘cognitive maps’’ of topographical space (O’Keefe & Nadel, 1978), or of modules for the permanent storage of the geometrical structure of spatial layouts (Wang & Spelke, 2002). Egocentric frames are typically associated with the posterior parietal cortex. Neurons coding spatial position relative to body parts have been found in the monkey’s posterior parietal cortex and in connected regions of the premotor cortex (Cohen & Andersen, 2002; Colby, ...

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... activated when stimuli are localized with respect to the body midsagittal plane (Vallar et al., 1999). This activation is much larger than when an object-based judgement is performed on the same stimuli (Galati, Lobel, et al., 2000), and is found for body-centered 1SHFJ/CEA, Orsay, France, 2Fondazione Santa Lucia, Rome, Italy, 3University G. d’Annunzio, Chieti, Italy, 4CNRS—College de France, Paris, France, 5University of Rome La Sapienza, Italy D 2004 Massachusetts Institute of Technology Journal of Cognitive Neuroscience 16:9, pp. 1517–1535 localization of both visual and tactile stimuli (Galati, Committeri, Sanes, & Pizzamiglio, 2001). Object-based coding of space has been much less studied. Its existence has been demonstrated in the monkey’s frontal lobe (Olson & Gettner, 1995). In humans, unilateral neglect may concern the contralesional side of individual objects, independent of their egocentric position (‘‘object-based neglect’’: Driver, 1999). Several neuroimaging studies (Fink, Marshall, Shah, et al., 2000; Galati, Lobel, et al., 2000; Honda, Wise, Weeks, Deiber, & Hallett, 1998; Fink, Dolan, Halligan, Marshall, & Frith, 1997) have found activation in the posterior parietal cortex and the dorsal premotor cortex (PMd...

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...cognized landmarks in order to find the way in familiar environments. Patients are unable to orient themselves and to know Committeri et al. 1527 which direction to take, as if they had lost their ‘‘heading’’ within the environment (Aguirre & D’Esposito, 1999). Accordingly, a small population of cells in the retrosplenial cortex of the rat fires only when the rat maintains a certain heading or orientation within the environment (Chen et al., 1994). This behavior was originally discovered in the so-called head-direction cells of the postsubiculum (a region of the dorsal hippocampal formation) (Zugaro, Arleo, Berthoz, & Wiener, 2003; Taube et al., 1990), which constitute the directional input to the hippocampal ‘‘place cells’’ (O’Keefe, Burgess, Donnett, Jeffery, & Maguire, 1999). In the present paradigm, the sudden change in the point of view which happens from trial to trial may be considered as analogous to disorienting subjects and having them to reorient on the exclusive basis of the visual scene, as no information are available about the occurred displacement. An automatic mechanism of spatial reorientation may well occur in all the spatial conditions, and even in the control task, but we argue that reorientation i...

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...anatomical image through an automatic nonlinear stereotaxic normalization procedure (Friston et al., 1995). The template image was based on average data provided by the Montreal Neurological Institute (MNI brain: Mazziotta, Toga, Evans, Fox, & Lancaster, 1995) and conformed to a standard coordinate referencing system (Talairach & Tournoux, 1988). The final voxel size after normalization was an isotropic 4 mm. Functional images were then spatially smoothed using an isotropic gaussian kernel (8 mm full-width-at-halfmaximum). Group analysis was performed using a two-stage random-effect approach (Friston, Holmes, & Worsley, 1999; Holmes & Friston, 1998). At the first stage, the time series of functional MR images obtained from each participant was analyzed separately. The effects of the experimental paradigm were estimated on a voxel-byvoxel basis, according to the general linear model (Friston, Worsley, Frackowiak, Mazziotta, & Evans, 1994; Worsley, Evans, Marret, & Neelin, 1992). The experimental blocks were modeled as box-car functions, convolved with a synthetic hemodynamic response function. Subject-specific contrast images were then entered at the second stage into one-sample t tests, testing the null hypothesi...

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...e typically associated with the posterior parietal cortex. Neurons coding spatial position relative to body parts have been found in the monkey’s posterior parietal cortex and in connected regions of the premotor cortex (Cohen & Andersen, 2002; Colby, 1998). Patients with lesions to the posterior parietal cortex may either show inaccurate visuomotor coordination (optic ataxia: Perenin & Vighetto, 1988) or fail to explore the contralesional side of space (unilateral neglect: Vallar, 1998). In unilateral neglect, the affected sector of space is usually egocentrically defined (see Bisiach, 1997; Vallar, Guariglia, & Rusconi, 1997, for related demonstrations). Both ataxic and neglect patients show specific deficits in perceiving the position of their body midsagittal plane (Pizzamiglio, Committeri, Galati, & Patria, 2000; Perenin, 1997), a fundamental egocentric spatial principle (Jeannerod & Biguer, 1989). A series of neuroimaging studies has provided direct evidence of the involvement of parietal–frontal circuits in the egocentric coding of space. A posterior parietal– frontal premotor network, bilateral but more extensive on the right, is activated when stimuli are localized with respect to the body midsagittal plan...

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...ndersen, 2002; Colby, 1998). Patients with lesions to the posterior parietal cortex may either show inaccurate visuomotor coordination (optic ataxia: Perenin & Vighetto, 1988) or fail to explore the contralesional side of space (unilateral neglect: Vallar, 1998). In unilateral neglect, the affected sector of space is usually egocentrically defined (see Bisiach, 1997; Vallar, Guariglia, & Rusconi, 1997, for related demonstrations). Both ataxic and neglect patients show specific deficits in perceiving the position of their body midsagittal plane (Pizzamiglio, Committeri, Galati, & Patria, 2000; Perenin, 1997), a fundamental egocentric spatial principle (Jeannerod & Biguer, 1989). A series of neuroimaging studies has provided direct evidence of the involvement of parietal–frontal circuits in the egocentric coding of space. A posterior parietal– frontal premotor network, bilateral but more extensive on the right, is activated when stimuli are localized with respect to the body midsagittal plane (Vallar et al., 1999). This activation is much larger than when an object-based judgement is performed on the same stimuli (Galati, Lobel, et al., 2000), and is found for body-centered 1SHFJ/CEA, Orsay, Franc...

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...ral coding. In a similar vein, the explicit perceptual judgement may have underestimated the primary action-oriented aspect of egocentric coding, as well as the virtual far space may have underestimated its preference for the near prehension space (Weiss et al., 2000), in both cases reducing the parietal involvement. Alternatively, viewer-centered and body-centered frames may be intrinsically different. A region significantly more active for the viewercentered coding than for the object-centered coding was detected in the frontal lobe. It comprised the socalled deep frontal eye fields (FEFs) (Lobel et al., 2001) and the PMd, respectively coding oculomotor and reaching-related space in egocentric coordinates (e.g., Boussaoud & Bremmer, 1999; Goldberg & Bruce, 1990). The activation included the rostral portion of PMd, recently termed ‘‘pre-PMd,’’ as it is more closely related to cognitive processes than to motor processes (Picard & Strick, 2001). This region receives strong projections from the dorsolateral prefrontal cortex (DLPFC), which has been suggested to play a role in spatial memory (Wilson, Scalaidhe, & Goldman-Rakic, 1993). Interestingly, our activation extended rostrally inside the SFS, in t...

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...more extensive in the right hemisphere, and the center of activity is quite superior and medial. Deficits in visuomotor coordination (e.g., ‘‘optic ataxia’’) can be observed in humans after dorsal lesions, and both the dorsal precuneus and the superior parietal lobule play a fundamental role in the genesis of such egocentric disorders (Perenin & Vighetto, 1988; Levine, Kaufman, & Mohr, 1978). Accordingly, optic ataxic patients have also a disturbed perception of their body midline (Perenin, 1997), like neglect patients with extensive lesions comprising posterior and superior parietal regions (Hasselback & Butter, 1997). Unexpectedly, none of the posterior parietal regions revealed a stronger involvement during the egocentric than during the other spatial coding. Galati, Lobel, et al. (2000) had instead detected a clear preference of the posterior parietal cortex for body- versus objectcentered tasks. The present viewer-centered condition, which refers to the viewer’s position as a whole rather than to a specific body part (such as the body midsagittal plane), may not have fully grasped the bodypart-centered nature of egocentric neural coding. In a similar vein, the explicit perceptual judgement may have und...

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...ntric frames are typically associated with the posterior parietal cortex. Neurons coding spatial position relative to body parts have been found in the monkey’s posterior parietal cortex and in connected regions of the premotor cortex (Cohen & Andersen, 2002; Colby, 1998). Patients with lesions to the posterior parietal cortex may either show inaccurate visuomotor coordination (optic ataxia: Perenin & Vighetto, 1988) or fail to explore the contralesional side of space (unilateral neglect: Vallar, 1998). In unilateral neglect, the affected sector of space is usually egocentrically defined (see Bisiach, 1997; Vallar, Guariglia, & Rusconi, 1997, for related demonstrations). Both ataxic and neglect patients show specific deficits in perceiving the position of their body midsagittal plane (Pizzamiglio, Committeri, Galati, & Patria, 2000; Perenin, 1997), a fundamental egocentric spatial principle (Jeannerod & Biguer, 1989). A series of neuroimaging studies has provided direct evidence of the involvement of parietal–frontal circuits in the egocentric coding of space. A posterior parietal– frontal premotor network, bilateral but more extensive on the right, is activated when stimuli are localized with ...

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...achusetts Institute of Technology Journal of Cognitive Neuroscience 16:9, pp. 1517–1535 localization of both visual and tactile stimuli (Galati, Committeri, Sanes, & Pizzamiglio, 2001). Object-based coding of space has been much less studied. Its existence has been demonstrated in the monkey’s frontal lobe (Olson & Gettner, 1995). In humans, unilateral neglect may concern the contralesional side of individual objects, independent of their egocentric position (‘‘object-based neglect’’: Driver, 1999). Several neuroimaging studies (Fink, Marshall, Shah, et al., 2000; Galati, Lobel, et al., 2000; Honda, Wise, Weeks, Deiber, & Hallett, 1998; Fink, Dolan, Halligan, Marshall, & Frith, 1997) have found activation in the posterior parietal cortex and the dorsal premotor cortex (PMd), mainly in the right hemisphere, and in early visual processing areas, during object-based spatial judgements. Finally, coding of topographical space is typically associated with the hippocampal formation. Cells with allocentric properties have been found in the hippocampal formation of both freely-moving rats (Taube, Muller, & Ranck, 1990; O’Keefe & Dostrovsky, 1971) and monkeys (Rolls & O’Mara, 1995; Ono, Nakamura, Nishijo, & Eifuku, 1993), as well as ...

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...hough we used real 3-D objects, rather than simple visual stimuli, and a distance judgment task, rather than a left–right position judgment, the results are very congruent. The detected parietal/frontal network is in fact bilateral but more extensive in the right hemisphere, and the center of activity is quite superior and medial. Deficits in visuomotor coordination (e.g., ‘‘optic ataxia’’) can be observed in humans after dorsal lesions, and both the dorsal precuneus and the superior parietal lobule play a fundamental role in the genesis of such egocentric disorders (Perenin & Vighetto, 1988; Levine, Kaufman, & Mohr, 1978). Accordingly, optic ataxic patients have also a disturbed perception of their body midline (Perenin, 1997), like neglect patients with extensive lesions comprising posterior and superior parietal regions (Hasselback & Butter, 1997). Unexpectedly, none of the posterior parietal regions revealed a stronger involvement during the egocentric than during the other spatial coding. Galati, Lobel, et al. (2000) had instead detected a clear preference of the posterior parietal cortex for body- versus objectcentered tasks. The present viewer-centered condition, which refers to the viewer’s position as...

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...nnected regions of the premotor cortex (Cohen & Andersen, 2002; Colby, 1998). Patients with lesions to the posterior parietal cortex may either show inaccurate visuomotor coordination (optic ataxia: Perenin & Vighetto, 1988) or fail to explore the contralesional side of space (unilateral neglect: Vallar, 1998). In unilateral neglect, the affected sector of space is usually egocentrically defined (see Bisiach, 1997; Vallar, Guariglia, & Rusconi, 1997, for related demonstrations). Both ataxic and neglect patients show specific deficits in perceiving the position of their body midsagittal plane (Pizzamiglio, Committeri, Galati, & Patria, 2000; Perenin, 1997), a fundamental egocentric spatial principle (Jeannerod & Biguer, 1989). A series of neuroimaging studies has provided direct evidence of the involvement of parietal–frontal circuits in the egocentric coding of space. A posterior parietal– frontal premotor network, bilateral but more extensive on the right, is activated when stimuli are localized with respect to the body midsagittal plane (Vallar et al., 1999). This activation is much larger than when an object-based judgement is performed on the same stimuli (Galati, Lobel, et al., 2000), and is found for body-centered 1SHFJ/C...

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... Chen, & Markus, 1991) in the rat hippocampal formation, which fire whenever the animal finds himself in a particular location and with a particular direction. In a similar fashion, the monkey ‘‘spatial view cells’’ respond when the animal looks at a particular part of the environment (Georges-Francois, Rolls, & Robertson, 1999; Rolls & O’Mara, 1995). However, an alternate interpretation of the activation of the LLA/PPA cannot be excluded. Attention to buildings has been shown to modulate the activity of the PPA, even in conditions where the presented stimuli included other objects or faces (Avidan, Levy, Hendler, Zohary, & Malach, 2003; O’Craven, Downing, & Kanwisher, 1999; Tong, Nakayama, Vaughan, & Kanwisher, 1998). Thus, in the present study, attention to the building/scene, when it had to be used as reference, may have contributed to the observed activation, by modulating the activity of such specific areas, which do not respond to other kinds of objects (see, for example, Epstein & Kanwisher, 1998). Note however that the differential activation for the landmark-centered condition extended well beyond the PPA, for example, in the retrosplenial and parietal cortex (see below), where no specificity for buildings has been ...

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...sterior parietal cortex may either show inaccurate visuomotor coordination (optic ataxia: Perenin & Vighetto, 1988) or fail to explore the contralesional side of space (unilateral neglect: Vallar, 1998). In unilateral neglect, the affected sector of space is usually egocentrically defined (see Bisiach, 1997; Vallar, Guariglia, & Rusconi, 1997, for related demonstrations). Both ataxic and neglect patients show specific deficits in perceiving the position of their body midsagittal plane (Pizzamiglio, Committeri, Galati, & Patria, 2000; Perenin, 1997), a fundamental egocentric spatial principle (Jeannerod & Biguer, 1989). A series of neuroimaging studies has provided direct evidence of the involvement of parietal–frontal circuits in the egocentric coding of space. A posterior parietal– frontal premotor network, bilateral but more extensive on the right, is activated when stimuli are localized with respect to the body midsagittal plane (Vallar et al., 1999). This activation is much larger than when an object-based judgement is performed on the same stimuli (Galati, Lobel, et al., 2000), and is found for body-centered 1SHFJ/CEA, Orsay, France, 2Fondazione Santa Lucia, Rome, Italy, 3University G. d’Annunzio, Chi...

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...In the lingual gyrus, this happens in a rostral region (‘‘lingual landmark area’’ [LLA]) which seems to be crucially involved in the recognition of stimuli with orienting 1526 Journal of Cognitive Neuroscience Volume 16, Number 9 value. In fact, a lesion here often causes ‘‘landmark agnosia’’ (see Aguirre & D’Esposito, 1999, for a review). On the other hand, a region within the posterior PH (‘‘parahippocampal place area’’ [PPA]), seems to be implicated more in the encoding of new place information into memory, than in recognition or navigation (Epstein, DeYoe, Press, Rosen, & Kanwisher, 2001; Epstein, Harris, Stanley, & Kanwisher, 1999). Neuroimaging studies repeatedly found a bilateral (Mellet et al., 2000; Aguirre, Detre, et al., 1996; Maguire, Frackowiak, & Frith, 1996) or unilateral right (Maguire, Burgess, et al., 1998; Aguirre & D’Esposito, 1997) PH activation during topographical learning, but also during the recall of recently learned environmental information (Aguirre & D’Esposito, 1997; Ghaem et al., 1997), and when the environment (O’Craven & Kanwisher, 2000) or the landmarks (Ghaem et al., 1997) were only mentally imagined. Our activation, which encompassed both areas, cannot be due to the simple passive viewing...