Allocentric Reference Frame Research Articles

Rat head direction cell responses in zero-gravity parabolic flight.

Astronauts working in zero-gravity (0-G) often experience visual reorientation illusions (VRIs). For example, when floating upside down, they commonly misperceive the spacecraft floor as a ceiling and have a reversed sense of direction. Previous studies have identified a population of neurons in the rat's brain that discharge as a function of the rat's head direction (HD) in a gravitationally horizontal plane and is dependent on an intact vestibular system. Our goal was to characterize HD cell discharge under conditions of acute weightlessness. Seven HD cells in the anterior dorsal thalamus were monitored from rats aboard an aircraft in 0-G parabolic flight. Unrestrained rats locomoted in a clear plexiglas rectangular chamber that had wire mesh covering the floor, ceiling, and one wall. The chamber and surrounding visual environment were relatively up-down symmetrical. Each HD cell was recorded across forty 20-s episodes of 0-G. All HD cells maintained a significant direction-specific discharge when the rat was on the chamber floor during the 0-G and also during the hypergravity pull-out periods. Three of five cells also showed direction-specific responses on the wall in 1-G. In contrast, direction-specific discharge was usually not maintained when the rat locomoted on the vertical wall or ceiling in 0-G. The loss of direction-specific firing was accompanied by an overall increase in background firing. However, while the rat was on the ceiling, some cells showed occasional bursts of firing when the rat's head was oriented in directions that were flipped relative to the long axis of symmetry of the chamber compared with the cell's preferred firing direction on the floor. This finding is consistent with what might be expected if the rat had experienced a VRI. These responses indicate that rats maintain a normal allocentric frame of reference in 0-G and 1-G when on the floor, but may lose their sense of directional heading when placed on a wall or ceiling during acute exposures to 0-G.

Opposite visual field asymmetries for egocentric and allocentric spatial judgments.

The purpose of the present study was to investigate the relation between visual hemifields and spatial frames of reference, according to the idea that multiple representations of 3D space exist. Results from two experiments clearly show that an upper visual hemifield advantage only arises when allocentric spatial judgments are required in order to perform a location task, whereas a lower visual hemifield advantage arises when egocentric spatial judgments are required. Such a double dissociation was interpreted as due to the activity of two separate neural pathways operating specific transformations of visual input for different functional outputs: scene recognition, mostly relying on allocentric frames of reference and subserved by the ventral, occipito-temporal pathway of the visual system, and goal directed actions, mostly relying on egocentric systems and subserved by the dorsal, occipito-parietal pathway of the visual system.

Navigating in a virtual three-dimensional maze: how do egocentric and allocentric reference frames interact?

Spatial navigation in the presence of gravity restricts one's displacement to two-dimensional (2D) planes. Therefore, self-motion only includes translations and yaw rotations. In contrast, in weightlessness, one can translate and turn in any direction. In the first experiment, we compared the ability to memorize a virtual three-dimensional (3D) maze after passive exploration in three self-motion conditions, each using a different set of rotations for turning. Subjects indicated which pathway they traversed among four successive corridors presented from an outside perspective. Results showed that exploring in the terrestrial condition (including only yaw rotations, the viewer's virtual body remaining upright) allowed better recognition of the corridor than in the weightless condition (which included pitch and yaw rotations according to the turns), particularly for more complex 3D structures. The more frequently the viewer-defined (egocentric) and the global environment (allocentric) verticals were aligned during exploration, the more easily subjects could memorize the 3D maze, suggesting that simplifying the relationship between the egocentric and allocentric reference frames facilitates spatial updating. Nevertheless, with practice, performance in the weightless condition improved whereas in the natural terrestrial condition performance remained at its initial maximum, indicating that the cognitive processes involved were innate for this particular condition. The second experiment revealed that single rotations in the terrestrial condition must be performed around the body axis in order to obtain optimal spatial updating performance, and that the latter is independent of the conflict with gravity that might favor this condition when one is actually upright. This suggests that although humans can memorize 3D-structured environments their innate neurocognitive functions appear to be specialized for natural 2D navigation.

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 person’s 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.

Immediate spatial distortions of pointing movements induced by visual landmarks.

We tested the influence of two horizontally aligned visual landmarks on pointing movements to memorized targets, to investigate whether the visuomotor system can make use of an egocentric representation unaffected by visual context. The endpoints of pointing movements were systematically distorted toward the nearest visual landmark, indicating that spatial representations included both target and nontarget information. These distortions were not due to the presence of the landmarks during the movement but, rather, to their presence in the encoding phase. Qualitatively similar distortions were present even with the shortest possible retention phase, when the target was extinguished at movement onset. Finally, we found the same pattern of distortion when participants were forced to remember the target within an allocentric frame of reference. We argue that even early memory representations for pointing movements are influenced by visual information in the surrounding visual field.

Smooth anticipatory eye movements alter the memorized position of flashed targets.

Briefly flashed visual stimuli presented during smooth object- or self-motion are systematically mislocalized. This phenomenon is called the "flash-lag effect" (Nijhawan, 1994). All previous studies had one common characteristic, the subject's sense of motion. Here we asked whether motion perception is a necessary condition for the flash-lag effect to occur. In our first experiment, we briefly flashed a target during smooth anticipatory eye movements in darkness and subjects had to orient their gaze toward the perceived flash position. Subjects reported to have no sense of eye motion during anticipatory movements. In our second experiment, subjects had to adjust a cursor on the perceived position of the flash. As a result, we show that gaze orientation reflects the actual perceived flash position. Furthermore, a flash-lag effect is present despite the absence of motion perception. Moreover, the time course of gaze orientation shows that the flash-lag effect appeared immediately after the egocentric to allocentric reference frame transformation.

A step and a hop on the Müller-Lyer: illusion effects on lower-limb movements.

Past studies have shown that the effects of visual illusions on upper-limb movements can be modulated by the availability of current or recent visual information. The generality of this finding across body movements was examined by measuring the impact of the Müller-Lyer illusion on two types of lower limb actions. Subjects stood at one end of a Müller-Lyer figure, then either stepped or hopped to the other end. Visual feedback was manipulated through the use of visual closed-loop, open-loop, and open-loop three second delay conditions. For both stepping and hopping, effects of the illusion on movement accuracy were small in the closed-loop, moderate in the open-loop, and large in the three-second open-loop delay condition. These results were similar to those found in studies involving the Müller-Lyer illusion and upper-limb movements. The similar effects across different body movements suggests that a common visuomotor system subserves both upper and lower-limb movements, and that egocentric and allocentric reference frames make similar contributions for the two types of movements.

Aging Affects Pointing to Unseen Targets Encoded in an Allocentric Frame of Reference

The goal of the present study was to determine whether aging influences aiming performance to a remembered target encoded in an allocentric frame of reference. We presented four targets simultaneously. The targets were then withdrawn from the screen. Following a recall delay, only three of the targets were presented on the screen in the same configuration as before but at a different location from their first presentation. We asked younger ( M =21.9 years) and older participants ( M =73.8 years) to point first to the missing target. Because participants did not know where on the computer screen the targets would be presented at the end of the recall delay, they had no choice but to use an allocentric frame of reference to encode its location when the targets were first presented. The results indicate that older participants were significantly more variable when pointing to the remembered target than their younger counterparts, suggesting a decline of allocentric spatial memory with aging.

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.

Delay improves performance on a haptic spatial matching task.

Systematic deviations occur when blindfolded subjects set a test bar parallel to a reference bar in the horizontal plane using haptic information (Kappers and Koenderink 1999, Perception 28:781-795; Kappers 1999, Perception 28:1001-1012). These deviations are assumed to reflect the use of a combination of a biasing egocentric reference frame and an allocentric, more cognitive one (Kappers 2002, Acta Psychol 109:25-40). In two experiments, we have examined the effect of delay between the perception of a reference bar and the parallel setting of a test bar. In both experiments a 10-s delay improved performance. The improvement increased with a larger horizontal (left-right) distance between the bars. This improvement was interpreted as a shift from the egocentric towards the allocentric reference frame during the delay period.

Cognitive allocentric representations of visual space shape pointing errors

Subjects reached in three-dimensional space to a set of remembered targets whose position was varied randomly from trial to trial, but always fell along a "virtual" line (line condition). Targets were presented briefly, one-by-one and in an empty visual field. After a short delay, subjects were required to point to the remembered target location. Under these conditions, the target was presented in the complete absence of allocentric visual cues as to its position in space. However, because the subjects were informed prior to the experiment that all targets would fall on a straight line, they could conceivably imagine each point target as belonging to a single rigid object with a particular geometry and orientation in space, although this virtual object was never explicitly shown to the subjects. We compared the responses to repeated measurements of each target with those measured for targets presented in a directionally neutral configuration (sphere condition), and used the variable errors to infer the putative reference frames underlying the corresponding sensorimotor transformation. Performance in the different tasks was compared under two different lighting conditions (dim light or total darkness) and two memory delays (0.5 or 5 s). The pattern of variable errors differed significantly between the sphere condition and the line condition. In the former case, the errors were always accounted for by egocentric reference frames. By contrast the errors in the line condition revealed both egocentric and allocentric components, consistent with the hypothesis that target information can be defined concurrently in both egocentric and allocentric frames of reference, resulting in two independent coexisting representations.

Patterns of bimanual interference reveal movement encoding within a radial egocentric reference frame.

Constraints on interlimb coordination have been studied intensively in past years with a primary focus on temporal features. The present study addressed spatial constraints or the degree of directional interference as a function of different line combinations between the upper limbs as well as the modulation of this interference as a result of different board orientations within the performer's workspace. This paradigm was used to address a prominent theme in motor neuroscience, namely whether (bimanual) movements are encoded within an allocentric reference frame (pattern of interference invariant with respect to extrinsic space) or within an egocentric reference frame (pattern of interference invariant relative to the center of the performer's action space, i.e., intrinsic). The observed patterns of interference revealed that movements are primarily encoded within a radial egocentric reference frame in which the performer is the center of action space. The present psychophysical findings converge with primate single-cell recording studies in which the direction has been identified as a primary movement parameter that is encoded in various brain regions, thereby constituting a principal determinant of bilateral interference.

"Look where you're going!": gaze behaviour associated with maintaining and changing the direction of locomotion.

In order to fully understand how vision is used to guide locomotion it is necessary to know what people look at as they move through the environment. This study provides information, hitherto lacking, regarding gaze behaviour associated with both maintaining and changing the direction of locomotion: activities that are essential for efficient navigation through our cluttered environment. Participants' spatiotemporal gaze patterns were recorded whilst they performed a task requiring that they either maintained a straight walking trajectory or changed their direction of walking by 30 degrees or 60 degrees, left or right, at the midpoint of a 9-m path. Participants were either visually cued to turn when they stepped on a trigger mat placed one step before the mid-point of the walkway (cued trials) or given verbal instruction about the required route prior to the start of each trial (advance knowledge trials). Our clear finding was that for the large majority of the time participants' gaze was aligned with environmental features lying in their current plane of progression both prior to and following the onset of the transition stride during which the direction change was implemented. This gaze behaviour was observed both during cued trials (78% of total fixation time prior to the transition stride onset and 89% following the transition stride onset) and advance knowledge trials (67% prior to transition stride onset, 92% following transition stride onset). When not aligned with the plane of progression, gaze was normally fixated on environmental features related to either known or potential future routes. Prior to changing the direction of walking, individuals invariably made saccadic eye movements in order to align gaze with the end-point of the required travel path. This gaze realignment was invariably accompanied by head reorientation, which was initiated, on average, at the same time as the saccade. On average, participants fixated gaze on their goal (represented by the cue light at the travel path end-point) until after head realignment with the new path was achieved. Additionally, the head was consistently aligned with participants' current walking direction prior to and following the transition stride even on the minority of occasions when they were looking elsewhere. These findings challenge the ecological validity of existing theories of how visual information is used to determine heading direction and are consistent with the proposal that aligning the head with the desired travel direction through coordinated eye and head movements provides the CNS with an allocentric frame of reference that is used to control the movement of the body in space.

Position sensitivity in phasically discharging nucleus accumbens neurons of rats alternating between tasks requiring complementary types of spatial cues

To determine how hippocampal location-selective discharges might influence downstream structures for navigation, nucleus accumbens neurons were recorded in rats alternating between two tasks guided respectively by lit cues in the maze or by extramaze room cues. Of 144 phasically active neurons, 80 showed significant behavioral correlates including displacements, immobility prior to, or after reward delivery, as well as turning, similar to previous reports. Nine neurons were position-selective, 22 were sensitive to task and platform changes and 40 others were both.Although the accumbens neurons showed the same behavioral correlate in two or four functionally equivalent locations, these responses were stronger at some of these places, evidence for position sensitivity. To test whether position responses were selective for room versus platform cues, the experimental platform was rotated while the rat performed each of the two tasks. This revealed responses to changes in position relative to both platform and room cues, despite the fact that previous studies had shown that place responses of hippocampal neurons recorded in the same task are anchored to room cues only. After these manipulations and shifts between the two tasks, the responses varied among simultaneously recorded neurons, and even in single neurons in alternating visits to reward sites. Again this contrasts with the uniformity of place responses of hippocampal neurons recorded in this same task. Thus accumbens position responses may derive from hippocampal inputs, while responses to context changes are more likely to derive from other signals or intrinsic processing.Considering the accumbens as a limbic-motor interface, we conclude that position-modulated behavioral responses in the accumbens may be intermediate between the allocentric reference frame of position-selective discharges in the hippocampus and the egocentric coding required to organize movement control. The conflicting responses among simultaneously recorded neurons could reflect competition processes serving as substrates for action selection and learning.

What Causes Specificity of Practice in a Manual Aiming Movement: Vision Dominance or Transformation Errors?

The withdrawal of vision of the arm during a manual aiming task has been found to result in a large increase in aiming error, regardless of the amount of practice in normal vision before its withdrawal. In the present study, the authors investigated whether the increase in error reflects the domination of visual afferent information over the movement representation developed during practice to the detriment of other sources of afferent information or whether it reflects only transformation errors of the location of the target from an allocentric to an egocentric frame of reference. Participants (N = 40) performed aiming movements with their dominant or nondominant arm in a full-vision or target-only condition. The results of the present experiment supported both of those hypotheses. The data indicated that practice does not eliminate the need for visual information for optimizing movement accuracy and that learning is specific to the source or sources of afferent information more likely to ensure optimal accuracy during practice. In addition, the results indicated that movement planning in an allocentric frame of reference might require simultaneous vision of the arm and the target. Finally, practice in a target-only condition, with knowledge of results, was found to improve recoding of the target in an egocentric frame of reference.

Spatial learning in turtles

Turtles (Pseudemys scripta) were trained in place, cue and control open-field procedures. The turtles trained in both the place and the cue procedures were able to learn their respective tasks with accuracy. Subsequent probe tests revealed that the turtles trained in the place task relied on the information provided by the extramaze cues to locate the goal. However, for these animals, no single cue was essential for performance, as accurate navigation to the goal was still possible when subsets of extramaze cues were eliminated. Furthermore, the turtles trained in the place task were able to navigate accurately to the goal place from new start locations. These results suggest that the turtles trained in the place task used map-like, relational strategies, by encoding the simultaneous spatial relationships between the goal and the extramaze cues in an allocentric frame of reference. In contrast, the turtles trained in the cue procedure used guidance strategies, i.e. approaching the individual intramaze cue associated to the goal as it were a beacon and largely ignoring the extramaze cues. Thus, the results of this experiment suggest that turtles are able to employ spatial strategies that closely parallel those described in mammals and birds.

Shift of the beating field of vestibular nystagmus: an orientation strategy?

We investigated in humans whether the shift of the beating field, which is often observed during vestibular nystagmus, could be related to some strategy of orientation. Eye movements were measured with an infrared system during an experiment on self-motion perception in the dark. Subjects were asked to rotate, by means of a joystick, a mobile robot on which they were seated in order to reproduce a previously imposed passive rotation. We suggest that the shift of the ocular beating field is the manifestation of two different orientation strategies based on allocentric and egocentric reference frames, respectively. It is also proposed that subjects who preferably used the first strategy exhibited large shifts of the beating field, while the others who probably used egocentric memory did not exhibit any shift.

Representing Space in Language and Perception

Abstract: Space can be understood through perception and language, but are the processes that represent spatial information the same in both cases? This paper reviews psychological evidence for the functional equivalence of spatial representations based on perceptual and linguistic inputs. It is proposed that spatial information is processed by a specialised spatial representation system (SRS) that creates geometric representations of space. The SRS receives inputs from perceptual and linguistic systems and uses these basic inputs to construct mental spatial models of the observed or described environment. A mental spatial model is created by determining the coordinate locations of objects in the egocentric or allocentric frame of reference. The goal of the SRS is not to represent strictly what is perceived, but to model an environment that has an inherent three‐dimensional spatial structure.

Visual illusion and action

The role of allocentric cues on movement control was investigated in the present study. Pointing movements directed to the more distant vertex of closed and open configurations of the Müller-Lyer illusion, as well as to the vertex of control lines, were studied in four experimental conditions. In the first (full-vision condition) subjects saw both stimulus and their hand before and during movement, in the second (non-visual feedback condition) they saw the stimulus, but not their hand during movement. In the two remaining conditions (no-vision conditions) vision of the scene and the hand was precluded. Pointing was executed 0 sec (no vision 0 sec delay condition) or 5 sec (no-vision 5 sec delay condition) after the light was switched off. The Müller-Lyer illusion affected pointing kinematics with respect to the control lines. Subjects undershot and overshot the vertex location, respectively, of the closed and open configuration. Correspondingly, the entire kinematics were changed. The main result was, however, a gradually increasing effect of the perceptual illusion when pointing was executed from memory compared to the full-vision condition. These data are discussed according to the hypothesis that the system underlying visual perception in the allocentric frame of reference and that involved in motor action can functionally interact. The strength of this interaction depends upon the efficiency of the egocentric frame of reference by which motor actions are constructed.

Performance of goldfish trained in allocentric and egocentric maze procedures suggests the presence of a cognitive mapping system in fishes

Goldfish were trained to obtain food in a four-arm maze placed in a room with relevant spatial cues. Four experimental conditions were run: allocentric, egocentric, egocentric + allocentric, and control. Relative to controls, all groups were able to solve the different tasks with high accuracy after 1 week of training. Subsequent transfer tests revealed place and response strategies for allocentric and egocentric groups, respectively, and both types of strategies for the ego-allocentric group. Moreover, the allocentric group showed the capacity to choose the appropriate trajectory toward the goal, even from novel starting points, presumably by using the distal cues as a whole. The results suggest that, in addition to using egocentric strategies, goldfish are able to solve spatial tasks on the basis of allocentric frames of reference and to build complex spatial cognitive representations of their environment.