Differential developmental trajectories for egocentric, environmental and intrinsic frames of reference in spatial memory

We studied the development of spatial frames of reference in children aged 3-6 years, who retrieved hidden toys from an array of identical containers bordered by landmarks under four conditions. By moving the child and/or the array between presentation and test, we varied the consistency of the hidden toy with (i) the body, and (ii) the testing room. The toy's position always remained consistent with (iii) the array and bordering landmarks. We found separate, additive performance advantages for consistency with body and room. These effects were already present at 3 years. A striking finding was that the room effect, which implies allocentric representations of the room and/or egocentric representations updated by self-motion, was much stronger in the youngest children than the body effect, which implies purely egocentric representations. Children as young as 3 years therefore had, and greatly favoured, spatial representations that were not purely egocentric. Viewpoint-independent recall based only on the array and bordering landmarks emerged at 5 years. There was no evidence that this later-developing ability, which implies object-referenced (intrinsic) representations, depended on verbal encodings. These findings indicate that core components of adult spatial competence, including parallel egocentric and nonegocentric representations of space, are present as early as 3 years. These are supplemented by later-developing object-referenced representations.     

Reference directions and reference objects in spatial memory of a briefly viewed layout

Two experiments investigated participants' spatial memory of a briefly viewed layout. Participants saw an array of five objects on a table and, after a short delay, indicated whether the target object indicated by the experimenter had been moved. Experiment 1 showed that change detection was more accurate when non-target objects were stationary than when non-target objects were moved. This context effect was observed when participants were tested both at the original learning perspective and at a novel perspective. In Experiment 2, the arrays of five objects were presented on a rectangular table and two of the non-target objects were aligned with the longer axis of the table. Change detection was more accurate when the target object was presented with the two objects that were aligned with the longer axis of the table during learning than when the target object was presented with the two objects that were not aligned with the longer axis of the table during learning. These results indicated that the spatial memory of a briefly viewed layout has interobject spatial relations represented and utilizes an allocentric reference direction.     

Landmark Stability: Further Studies Pointing to a Role in Spatial Learning

Two experiments were conducted to investigate the possible role of landmark stability in spatial learning. Rats were trained to search in a large arena for food hidden at a consistent distance and direction from either a single radially symmetric landmark or an array of two landmarks. We varied the relative degree to which the landmark array and/or the cues of the training context predicted the location of food, without varying the conditional probability of food being available given either cue. Experiment 1 used vestibular disorientation to ensure control of search location by experimenter-controlled cues. The results showed that making either a single landmark or a cluster of two adjacent landmarks the sole spatial predictor of reward location reduced the accuracy of search compared to a condition where both the landmark array and context cues were reliable spatial predictors. Varying global landmark stability had no effect when training was conducted using an array of two landmarks located some distance from each other. Context cues, when tested alone, triggered very little searching in appropriate locations, and the absolute magnitude of control over search was insufficient to account for the superiority of stable landmarks. The better learning with a stable landmark, and the dependence of this effect on the geometrical arrangement of landmarks, points to the conditions of spatial learning involving additional principles to those of simple associative conditioning. Experiment 2 examined landmark stability using a single landmark and fixed directional cues in the absence of vestibular disorientation. This also revealed a relative advantage of landmark stability, but animals with a landmark that moved from trial to trial did show some evidence of learning. Context cues when tested alone had minimal influence. Parametric manipulation of landmark stability offers a novel way of influencing spatial learning and thus understanding better the process through which egocentric representations of perceived space are transformed into allocentric representations of the real world. The purpose of this paper is to describe two experiments concerned with identifying the psychological processes of allocentric spatial learning. The results point to the idea that landmark stability is an important factor in spatial learning. Specifically, they reveal that whether or not a landmark will be used for the purpose of representing the location of another object (including hidden objects) is influenced by whether it is perceived as geometrically stable with respect to at least one other landmark and/or certain geometric features of the environment. This phenomenon is relevant to the application of associative learning principles to the spatial domain.     

Object localisation and frames of reference

In this paper, we explore which spatial frames of reference, egocentric or allocentric, are used to locate objects either in relation to ourselves (i.e. subject-to-object localisation) or to other objects (i.e. object-to-object localisation). In particular, we wanted to know whether the same or different frames of reference are used in these two different kinds of localisation after learning the environment in an egocentric way. Egocentric frames of reference are determined by the position of the person in relation to the spatial layout, whereas allocentric frames of reference are centred on the environment or on objects, independent of a persons position. We hypothesised that subject-to-object localisation is based on egocentric spatial representations, whereas object-to-object localisation is based on allocentric spatial representations. Participants were asked to study eight common objects, placed in a circle. Next, half of the participants had to point to an object in relation to their imagined position (egocentric condition) and the other half to an object in relation to another object (allocentric condition). The overall results show no difference between subject-to-object and object-to-object localisation. In both cases, access to positions corresponding to the front/back body axis was facilitated, in terms of both latency and error. Moreover, participants were able to retrieve objects positions better from the perspective from which they had learned the spatial array than from new perspectives. These results support the conclusion that egocentric coordinates, which are selected on the basis of our body-centred experience of the environment, define spatial representations underlying both subject-to-object and object-to-object localisation.     

Egocentric metric representations in peripersonal space: A bridge between motor resources and spatial memory

Research on visuospatial memory has shown that egocentric (subject-to-object) and allocentric (object-to-object) reference frames are connected to categorical (non-metric) and coordinate (metric) spatial relations, and that motor resources are recruited especially when processing spatial information in peripersonal (within arm reaching) than extrapersonal (outside arm reaching) space. In order to perform our daily-life activities, these spatial components cooperate along a continuum from recognition-related (e.g., recognizing stimuli) to action-related (e.g., reaching stimuli) purposes. Therefore, it is possible that some types of spatial representations rely more on action/motor processes than others. Here, we explored the role of motor resources in the combinations of these visuospatial memory components. A motor interference paradigm was adopted in which participants had their arms bent behind their back or free during a spatial memory task. This task consisted in memorizing triads of objects and then verbally judging what was the object: (1) closest to/farthest from the participant (egocentric coordinate); (2) to the right/left of the participant (egocentric categorical); (3) closest to/farthest from a target object (allocentric coordinate); and (4) on the right/left of a target object (allocentric categorical). The triads appeared in participants' peripersonal (Experiment 1) or extrapersonal (Experiment 2) space. The results of Experiment 1 showed that motor interference selectively damaged egocentric-coordinate judgements but not the other spatial combinations. The results of Experiment 2 showed that the interference effect disappeared when the objects were in the extrapersonal space. A third follow-up study using a within-subject design confirmed the overall pattern of results. Our findings provide evidence that motor resources play an important role in the combination of coordinate spatial relations and egocentric representations in peripersonal space.     

Theories of spatial representations and reference frames: What can configuration errors tell us?

The issue of reference frame is central to theories of spatial representations. Various classifications have been made for different types of reference frames, along with prototypical research paradigms to distinguish between them. This article focuses on the configuration error paradigm proposed by Wang and Spelke (Cognition 77:215-250, 2000) that has been used to examine the nature of the spatial representations underlying object localization during self-movement. Three basic models of spatial memory and spatial updating are discussed, as well as the assumptions behind the configuration error paradigm, to distinguish between static representations, such as the traditional allocentric cognitive map and the egocentric snapshots, and dynamic representations, such as the egocentric updating system. Recent experimental findings are reexamined and shown to be consistent with multiple models, among which the egocentric-updating-and-reload model with an enduring egocentric component provides the simplest interpretations.     

Remembering object position in the absence of vision: egocentric, allocentric, and egocentric decentred frames of reference

In three experiments we examined whether memory for object locations in the peri-personal space in the absence of vision is affected by the correspondence between encoding and test either of the body position or of the reference point. In particular, the study focuses on the distinction between different spatial representations, by using a paradigm in which participants are asked to relocate objects explored haptically. Three frames of reference were systematically compared. In experiment 1, participants relocated the objects either from the same position of learning by taking as reference their own body (centred egocentric condition) or from a 90 degrees decentred position (allocentric condition). Performance was measured in terms of linear distance errors and angular distance errors. Results revealed that the allocentric condition was more difficult than the centred egocentric condition. In experiment 2, participants performed either the centred egocentric condition or a decentred egocentric condition, in which the body position during the test was the same as at encoding (egocentric) but the frame of reference was based on a point decentred by 90 degrees. The decentred egocentric condition was found to be more difficult than the centred egocentric condition. Finally, in experiment 3, participants performed in the decentred egocentric condition or the allocentric condition. Here, the allocentric condition was found to be more difficult than the decentred egocentric condition. Taken together, the results suggest that also in the peripersonal space and in the absence of vision different frames of reference can be distinguished. In particular, the decentred egocentric condition involves a frame of reference which seems to be neither allocentric nor totally egocentric.     

Egocentric-updating during navigation facilitates episodic memory retrieval

Influential models suggest that spatial processing is essential for episodic memory [O’Keefe, J., & Nadel, L. (1978). The hippocampus as a cognitive map. London: Oxford University Press]. However, although several types of spatial relations exist, such as allocentric (i.e. object-to-object relations), egocentric (i.e. static object-to-self relations) or egocentric updated on navigation information (i.e. self-to-environment relations in a dynamic way), usually only allocentric representations are described as potentially subserving episodic memory [Nadel, L., & Moscovitch, M. (1998). Hippocampal contributions to cortical plasticity. Neuropharmacology, 37(4-5), 431-439]. This study proposes to confront the allocentric representation hypothesis with an egocentric updated with self-motion representation hypothesis. In the present study, we explored retrieval performance in relation to these two types of spatial processing levels during learning. Episodic remembering has been assessed through Remember responses in a recall and in a recognition task, combined with a “Remember-Know-Guess” paradigm [Gardiner, J. M. (2001). Episodic memory and autonoetic consciousness: A first-person approach. Philosophical Transactions of the Royal Society B: Biological Sciences, 356(1413), 1351-1361] to assess the autonoetic level of responses. Our results show that retrieval performance was significantly higher when encoding was performed in the egocentric-updated condition. Although egocentric updated with self-motion and allocentric representations are not mutually exclusive, these results suggest that egocentric updating processing facilitates remember responses more than allocentric processing. The results are discussed according to Burgess and colleagues’ model of episodic memory [Burgess, N., Becker, S., King, J. A., & O’Keefe, J. (2001). Memory for events and their spatial context: models and experiments. Philosophical Transactions of the Royal Society of London. Series B: Biological Sciences, 356(1413), 1493-1503].     

Updating egocentric representations in human navigation

Seven experiments tested whether human navigation depends on enduring representations, or on momentary egocentric representations that are updated as one moves. Human subjects pointed to unseen targets, either while remaining oriented or after they had been disoriented by self-rotation. Disorientation reduced not only the absolute accuracy of pointing to all objects ('heading error') but also the relative accuracy of pointing to different objects ('configuration error'). A single light providing a directional cue reduced both heading and configuration errors if it was present throughout the experiment. If the light was present during learning and test but absent during the disorientation procedure, however, subjects showed low heading errors (indicating that they reoriented by the light) but high configuration errors (indicating that they failed to retrieve an accurate cognitive map of their surroundings). These findings provide evidence that object locations are represented egocentrically. Nevertheless, disorientation had little effect on the coherence of pointing to different room corners, suggesting both (a) that the disorientation effect on representations of object locations is not due to the experimental paradigm and (b) that room geometry is captured by an enduring representation. These findings cast doubt on the view that accurate navigation depends primarily on an enduring, observer-free cognitive map, for humans construct such a representation of extended surfaces but not of objects. Like insects, humans represent the egocentric distances and directions of objects and continuously update these representations as they move. The principal evolutionary advance in animal navigation may concern the number of unseen targets whose egocentric directions and distances can be represented and updated simultaneously, rather than a qualitative shift in navigation toward reliance on an allocentric map.     

Voluntary head movement and allocentric perception of space

Although visual input is egocentric, at least some visual perceptions and representations are allocentric, that is, independent of the observer's vantage point or motion. Three experiments investigated the visual perception of three-dimensional object motion during voluntary and involuntary motion in human subjects. The results show that the motor command contributes to the objective perception of space: Observers are more likely to apply, consciously and unconsciously, spatial criteria relative to an allocentric frame of reference when they are executing voluntary head movements than while they are undergoing similar involuntary displacements (which lead to a more egocentric bias). Furthermore, details of the motor command are crucial to spatial vision, as allocentric bias decreases or disappears when self-motion and motor command do not match.     

Spatial memory: how egocentric and allocentric combine

Recent experiments indicate the need for revision of a model of spatial memory consisting of viewpoint-specific representations, egocentric spatial updating and a geometric module for reorientation. Instead, it appears that both egocentric and allocentric representations exist in parallel, and combine to support behavior according to the task. Current research indicates complementary roles for these representations, with increasing dependence on allocentric representations with the amount of movement between presentation and retrieval, the number of objects remembered, and the size, familiarity and intrinsic structure of the environment. Identifying the neuronal mechanisms and functional roles of each type of representation, and of their interactions, promises to provide a framework for investigation of the organization of human memory more generally.     

Spatial updating relies on an egocentric representation of space: Effects of the number of objects

Models of spatial updating attempt to explain how representations of spatial relationships between the actor and objects in the environment change as the actor moves. In allocentric models, object locations are encoded in an external reference frame, and only the actor’s position and orientation in that reference frame need to be updated. Thus, spatial updating should be independent of the number of objects in the environment (set size). In egocentric updating models, object locations are encoded relative to the actor, so the location of each object relative to the actor must be updated as the actor moves. Thus, spatial updating efficiency should depend on set size. We examined which model better accounts for human spatial updating by having people reconstruct the locations of varying numbers of virtual objects either from the original study position or from a changed viewing position. In consistency with the egocentric updating model, object localization following a viewpoint change was affected by the number of objects in the environment.     

Quantifying the interactions between allo- and egocentric representations of space

Under many circumstances, humans do not judge the location of objects in space where they really are. For instance, when a background is added to a target object, the judged position of a target with respect to oneself (egocentric position) is shifted in the opposite direction as the placement of such a background with respect to the body midline. It is an ongoing debate whether such effects are due to a uni- or bi-directional interaction between allo- and egocentric spatial representations in the brain, or reflect a response strategy, known as the perceived midline shift. In this study, the effects of allocentric stimulus coordinates on perceived egocentric position were examined more precisely and in a quantitative manner. Furthermore, it was investigated whether the judged allocentric position (with respect to a background) is also influenced by the egocentric position in space of that object. Allo- and egocentric coordinates were varied independently. Also, the effect of background luminance on the observed interactions between spatial coordinates was determined. Since background luminance had an effect on the size of the interaction between allocentric stimulus coordinates and egocentric judgments, and no reverse interaction was found, it seems that interactions between ego- and allocentric reference frames is most likely only unidirectional, with the latter affecting the former. This interaction effect was described in a quantitative manner.     

听障和听力正常人群空间主导性和空间参照框架的交互作用

通过要求被试分别在近处空间和远处空间完成空间参照框架的判断任务, 考察了听障和听力正常人群空间主导性和空间参照框架的交互作用。结果表明：(1)相对于听力正常人群, 听障人群完成自我参照框架判断任务的反应时更长, 而在完成环境参照框架判断任务无显著差异; (2)听障人群和听力正常人群空间主导性和空间参照框架交互作用呈现出相反模式。研究表明, 听障人群在听力功能受损后, 其空间主导性和空间参照框架的交互作用也产生了变化。     

Landmark Discrimination Learning in the Dog

Allocentric spatial memory was studied in dogs of varying ages and sources using a landmark discrimination task. The primary goal of this study was to develop a protocol to test landmark discrimination learning in the dog. Using a modified version of a landmark test developed for use in monkeys, we successfully trained dogs to make a spatial discrimination on the basis of the position of a visual landmark relative to two identical discriminanda. Task performance decreased, however, as the distance between the landmark and the “discriminandum” was increased. A subgroup of these dogs was also tested on a delayed nonmatching to position spatial memory task (DNMP), which relies on egocentric spatial cues. These findings suggest that dogs can acquire both allocentric and egocentric spatial tasks. These data provide a useful tool for evaluating the ability of canines to use allocentric cues in spatial learning.     

Spatial memory during progressive disorientation

Human spatial representations of object locations in a room-sized environment were probed for evidence that the object locations were encoded relative not just to the observer (egocentrically) but also to each other (allocentrically). Participants learned the locations of 4 objects and then were blindfolded and either (a) underwent a succession of 70 degrees and 200 degrees whole-body rotations or (b) were fully disoriented and then underwent a similar sequence of 70 degrees and 200 degrees rotations. After each rotation, participants pointed to the objects without vision. Analyses of the pointing errors suggest that as participants lost orientation, represented object directions generally "drifted" off of their true directions as an ensemble, not in random, unrelated directions. This is interpreted as evidence that object-to-object (allocentric) relationships play a large part in the human spatial updating system. However, there was also some evidence that represented object directions occasionally drifted off of their true directions independently of one another, suggesting a lack of allocentric influence. Implications regarding the interplay of egocentric and allocentric information are considered.     

Remembering the past and imagining the future: a neural model of spatial memory and imagery

The authors model the neural mechanisms underlying spatial cognition, integrating neuronal systems and behavioral data, and address the relationships between long-term memory, short-term memory, and imagery, and between egocentric and allocentric and visual and ideothetic representations. Long-term spatial memory is modeled as attractor dynamics within medial-temporal allocentric representations, and short-term memory is modeled as egocentric parietal representations driven by perception, retrieval, and imagery and modulated by directed attention. Both encoding and retrieval/imagery require translation between egocentric and allocentric representations, which are mediated by posterior parietal and retrosplenial areas and the use of head direction representations in Papez's circuit. Thus, the hippocampus effectively indexes information by real or imagined location, whereas Papez's circuit translates to imagery or from perception according to the direction of view. Modulation of this translation by motor efference allows spatial updating of representations, whereas prefrontal simulated motor efference allows mental exploration. The alternating temporal-parietal flows of information are organized by the theta rhythm. Simulations demonstrate the retrieval and updating of familiar spatial scenes, hemispatial neglect in memory, and the effects on hippocampal place cell firing of lesioned head direction representations and of conflicting visual and ideothetic inputs.     

Feature bindings endure without attention: Evidence from an explicit recall task

Are integrated objects the unit of capacity of visual working memory, or is continued attention needed to maintain bindings between independently stored features? In a delayed recall task, participants reported the color and shape of a probed item from a memory array. During the delay, attention was manipulated with an exogenous cue. Recall was elevated at validly cued positions, indicating that the cue affected item memory. On invalid trials, participants most frequently recalled either both features (perfect object memory) or neither of the two features (no object memory); the frequency with which only one feature was recalled was significantly lower than predicted by feature independence as determined in a single-feature recall task. These data do not support the view that features are remembered independently when attention is withdrawn. Instead, integrated objects are stored in visual working memory without need for continued attention.     

Caudate nucleus and memory for egocentric localization

A large body of evidence suggests that the caudate nucleus (CN) plays a critical role in the processing of spatial localization information. Furthermore, evidence has begun to accumulate that the CN is involved in the processing of a very specific class of spatial cues, namely, egocentric cues (localization with reference to the organism). This is in contrast to allocentric localization, where an organism localizes on the basis of cues external to the organism. One would then predict that lesions to the CN should disrupt performance on any tasks that depend chiefly on egocentric spatial cues, while leaving performance on allocentric tasks intact. To test this prediction, two groups of rats were trained on two different egocentric memory tasks and two different allocentric memory tasks. Specifically, one group of rats was trained on an adjacent-arm (egocentric) and an 8-arm radial maze task (allocentric). A second group of rats was trained on a right-left discrimination (egocentric) and a place-learning task (allocentric). After training, both groups received bilateral lesions of the CN. Results showed that CN-lesioned animals were profoundly impaired on retention of the egocentric tasks. In sharp contrast to this, the same animals were not or were only transiently impaired or transiently affected on allocentric tasks. Sham-operated controls were either unimpaired or transiently affected on all tasks. These findings further support the idea that the CN plays a critical modulatory role in the processing of egocentric spatial and not allocentric spatial cues.     

序列本体感觉学习获得稳定的自我中心空间表征*

通过同步视觉或者序列本体感觉,被试学习不规则场景。学习完毕之后,在面对学习方向、自主旋转240°、和持续旋转直至迷向3种运动条件下,被试随机指出各个物体的位置。迷向导致同步视觉学习组指向的内部一致性显著变差,而序列本体感觉学习组不受迷向影响。离线的相对位置判断任务表明两组被试的环境中心空间表征没有差异。这证明通过序列本体感觉学习被试也可以形成稳定的自我中心空间表征,支持了空间快照理论的扩展和空间认知的功能等价假说。