Spatial priming in visual search: memory for body-centred information

Spatial priming allows memory for target locations to be evaluated, whereby when a target appears in the same location across trials, participants become more efficient at locating it and consequently their search times decrease. Previously, we reported priming effects when the location of a target was repeated with respect to the participant’s body but not when it was repeated relative to their eye position; therefore, suggesting that body-centred information is available after a delay of at least a few seconds (Ball et al. in Exp Brain Res 204:585–594, 2010). However, we were unable to rule out the possibility that stable allocentric cues within the room may have contributed to the priming effects that we observed. In this current study, we introduced a condition where despite participants moving to a new location between trials, their position relative to the target was maintained. This movement disrupted any potential room-based cues about the target location. While we replicated our previous finding of priming when the location of the target was repeated relative to the viewer when no movement was required, we also found robust priming effects when participants moved to a new location in between trials. Thus, we provide clear evidence that in our spatial priming task, the location of the target was being coded in a body-centred reference frame and that this information is available after a delay.     

Frames of reference and categorical and coordinate spatial relations: a hierarchical organisation

This research is about the role of categorical and coordinate spatial relations and allocentric and egocentric frames of reference in processing spatial information. To this end, we asked whether spatial information is firstly encoded with respect to a frame of reference or with respect to categorical/coordinate spatial relations. Participants had to judge whether two vertical bars appeared on the same side (categorical) or at the same distance (coordinate) with respect to the centre of a horizontal bar (allocentric) or with respect to their body midline (egocentric). The key manipulation was the timing of the instructions: one instruction (reference frame or spatial relation) was given before stimulus presentation, the other one after. If spatial processing requires egocentric/allocentric encoding before coordinate/categorical encoding, then spatial judgements should be facilitated when the frame of reference is specified in advance. In contrast, if categorical and coordinate dimensions are primary, then a facilitation should appear when the spatial relation is specified in advance. Results showed that participants were more accurate and faster when the reference frame rather than the type of spatial relation was provided before stimulus presentation. Furthermore, a selective facilitation was found for coordinate and categorical judgements after egocentric and allocentric cues, respectively. These results suggest a hierarchical structure of spatial information processing where reference frames play a primary role and selectively interact with subsequent processing of spatial relations.     

Goal-directed reaching: movement strategies influence the weighting of allocentric and egocentric visual cues

The location of an object in peripersonal space can be represented with respect to our body (i.e., egocentric frame of reference) or relative to contextual features and other objects (i.e., allocentric frame of reference). In the current study, we sought to determine whether the frame, or frames, of visual reference supporting motor output is influenced by reach trajectories structured to maximize visual feedback utilization (i.e., controlled online) or structured largely in advance of movement onset via central planning mechanisms (i.e., controlled offline). Reaches were directed to a target embedded in a pictorial illusion (the induced Roelofs effect: IRE) and advanced knowledge of visual feedback was manipulated to influence the nature of reaching control as reported by Zelaznik et al. (J Mot Behav 15:217–236, 1983). When vision could not be predicted in advance of movement onset, trajectories showed primary evidence of an offline mode of control (even when vision was provided) and endpoints demonstrated amplified sensitivity to the illusory (i.e., allocentric) features of the IRE. In contrast, reaches performed with reliable visual feedback evidenced a primarily online mode of control and showed increased visuomotor resistance to the IRE. These findings suggest that the manner a reaching response is structured differentially influences the weighting of allocentric and egocentric visual information. More specifically, when visual feedback is unavailable or unpredictable, the weighting of allocentric visual information for the advanced planning of a reach trajectory is increased.

Frame of reference for visual perception in young infants during change of body position

The visual and vestibular systems begin functioning early in life. However, it is unclear whether young infants perceive the dynamic world based on the retinal coordinate (egocentric reference frame) or the environmental coordinate (allocentric reference frame) when they encounter incongruence between frames of reference due to changes in body position. In this study, we performed the habituation–dishabituation procedure to assess novelty detection in a visual display, and a change in body position was included between the habituation and dishabituation phases in order to test whether infants dishabituate to the change in stimulus on the retinal or environmental coordinate. Twenty infants aged 3–4 months were placed in the right-side-down position (RSDp) and habituated to an animated human-like character that walked horizontally in the environmental frame of reference. Subsequently, their body position was changed in the roll plane. Ten infants were repositioned to the upright position (UPp) and the rest, to the RSDp after rotation. In the test phase, the displays that were spatially identical to those shown in the habituation phase and 90° rotated displays were alternately presented, and visual preference was examined. The results revealed that infants looked longer at changes in the display on the retinal coordinate than at changes in the display on the environmental coordinate. This suggests that changes in body position from lying to upright produced incongruence of the egocentric and allocentric reference frames for perception of dynamic visual displays and that infants may rely more on the egocentric reference frame.     

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. Electronic Publication     

Reference systems for coding spatial information in normal subjects and a deafferented patient

To produce accurate goal-directed arm movements, subjects must determine the precise location of target object. Position of extracorporeal objects can be determined using: (a) an egocentric frame of reference, in which the target is localized in relation to the position of the body; and/or (b) an allocentric system, in which target position is determined in relation to stable visual landmarks surrounding the target (Bridgeman 1989; Paillard 1991). The present experiment was based on the premise that (a) the presence of a structured visual environment enables the use of an allocentric frame of reference, and (b) the sole presence of a visual target within a homogeneous background forces the registration of the target location by an egocentric system. Normal subjects and a deafferented patient (i.e., with an impaired egocentric system) pointed to visual targets presented in both visual environments to evaluate the efficiency of the two reference systems. For normals, the visual environment conditions did not affect pointing accuracy. However, kinematic parameters were affected by the presence or absence of a structured visual surrounding. For the deafferented patient, the presence of a structured visual environment permitted a decrease in spatial errors when compared with the unstructured surrounding condition (for movements with or without visual feedback of the trajectory). Overall, results support the existence of an egocentric and an allocentric reference system capable of organizing extracorporeal space during arm movements directed toward visual targets.     

Temporal dynamics of object location processing in allocentric reference frame

The spatial location of objects is processed in egocentric and allocentric reference frames, the early temporal dynamics of which have remained relatively unexplored. Previous experiments focused on ERP components related only to egocentric navigation. Thus, we designed a virtual reality experiment to see whether allocentric reference frame‐related ERP modulations can also be registered. Participants collected reward objects at the end of the west and east alleys of a cross maze, and their ERPs to the feedback objects were measured. Participants made turn choices from either the south or the north alley randomly in each trial. In this way, we were able to discern place and response coding of object location. Behavioral results indicated a strong preference for using the allocentric reference frame and a preference for choosing the rewarded place in the next trial, suggesting that participants developed probabilistic expectations between places and rewards. We also found that the amplitude of the P1 was sensitive to the allocentric place of the reward object, independent of its value. We did not find evidence for egocentric response learning. These results show that early ERPs are sensitive to the location of objects during navigation in an allocentric reference frame.     

Frames of reference during implicit and explicit learning

There is a significant overlap between the processes and neural substrates of spatial cognition and those subserving memory and learning. However, for procedural learning, which often is spatial in nature, we do not know how different forms of spatial knowledge, such as egocentric and allocentric frames of reference, are utilized nor whether these frames are differentially engaged during implicit and explicit processes. To address this issue, we trained human subjects on a movement sequence presented on a bi-dimensional (2D) geometric frame. We then systematically manipulated the geometric frame (allocentric) or the sequence of movements (egocentric) or both, and retested the subjects on their ability to transfer the sequence knowledge they had acquired in training and also determined whether the subjects had learned the sequence implicitly or explicitly. None of the subjects (implicit or explicit) showed evidence of transfer when both frames of reference were changed which suggests that spatial information is essential. Both implicit and explicit subjects transferred when the egocentric frame was maintained indicating that this representation is common to both processes. Finally, explicit subjects were also able to benefit from the allocentric frame in transfer, which suggests that explicit procedural knowledge may have two tiers comprising egocentric and allocentric representations.     

Judging hand laterality from my or your point of view: Interactions between motor imagery and visual perspective

Motor imagery tasks (hand laterality judgment) are usually performed with respect to a self-body (egocentric) representation, but manipulations of stimulus features (hand orientation) can induce a shift to other's body (allocentric) reference frame. Visual perspective taking tasks are also performed in self-body perspective but a shift to an allocentric frame can be triggered by manipulations of context features (e.g., another person present in the to-be-judged scene). Combining hand laterality task and visual perspective taking, we demonstrated that both stimulus and context features can modulate motor imagery performance. In Experiment 1, participants judged laterality of a hand embedded in a human or non-human silhouette. Results showed that observing a human silhouette interfered with judgments on “egocentric hand stimuli” (right hand, fingers up). In Experiment 2, participants were explicitly required to judge laterality of a hand embedded in a human silhouette from their own (egocentric group) or from the silhouette's perspective (allocentric group). Consistent with previous results, the egocentric group was significantly faster than the allocentric group in judging fingers-up right hand stimuli. These findings showed that concurrent activation of egocentric and allocentric frames during mental transformation of body parts impairs participants’ performance due to a conflict between motor and visual mechanisms.     

Alpha modulation in parietal and retrosplenial cortex correlates with navigation performance

The present study investigated the brain dynamics accompanying spatial navigation based on distinct reference frames. Participants preferentially using an allocentric or an egocentric reference frame navigated through virtual tunnels and reported their homing direction at the end of each trial based on their spatial representation of the passage. Task-related electroencephalographic (EEG) dynamics were analyzed based on independent component analysis (ICA) and subsequent clustering of independent components. Parietal alpha desynchronization during encoding of spatial information predicted homing performance for participants using an egocentric reference frame. In contrast, retrosplenial and occipital alpha desynchronization during retrieval covaried with homing performance of participants using an allocentric reference frame. These results support the assumption of distinct neural networks underlying the computation of distinct reference frames and reveal a direct relationship of alpha modulation in parietal and retrosplenial areas with encoding and retrieval of spatial information for homing behavior.     

The neural basis of egocentric and allocentric coding of space in humans: a functional magnetic resonance study

The spatial location of an object can be represented in the brain with respect to different classes of reference frames, either relative to or independent of the subject's position. We used functional magnetic resonance imaging to identify regions of the healthy human brain subserving mainly egocentric or allocentric (object-based) coordinates by asking subjects to judge the location of a visual stimulus with respect to either their body or an object. A color-judgement task, matched for stimuli, difficulty, motor and oculomotor responses, was used as a control. We identified a bilateral, though mainly right-hemisphere based, fronto-parietal network involved in egocentric processing. A subset of these regions, including a much less extensive unilateral, right fronto-parietal network, was found to be active during object-based processing. The right-hemisphere lateralization and the partial superposition of the egocentric and the object-based networks is discussed in the light of neuropsychological findings in brain-damaged patients with unilateral spatial neglect and of neurophysiological studies in the monkey.     

Numbers are represented in egocentric space: Effects of numerical cues and spatial reference frames on hand laterality judgements

Convergent findings demonstrate that numbers can be represented according to a spatially oriented mental number line. However, it is not established whether a default organization of the mental number line exists (i.e., a left-to-right orientation) or whether its spatial arrangement is only the epiphenomenon of specific task requirements. To address this issue we performed two experiments in which subjects were required to judge laterality of hand stimuli preceded by small, medium or large numerical cues; hand stimuli were compatible with egocentric or allocentric perspectives. We found evidence of a left-to-right number–hand association in processing stimuli compatible with an egocentric perspective, whereas the reverse mapping was found with hands compatible with an allocentric perspective. These findings demonstrate that the basic left-to-right arrangement of the mental number line is defined with respect to the body-centred egocentric reference frame.     

Supra-spinal circuits shape inhibitory postural adjustments anticipating voluntary index-finger flexion

We investigated the influence of gaze elevation on judging the possibility of passing under high obstacles during pitch body tilts, while stationary, in absence of allocentric cues. Specifically, we aimed at studying the influence of egocentric references upon geocentric judgements. Seated subjects, orientated at various body orientations, were asked to perceptually estimate the possibility of passing under a projected horizontal line while keeping their gaze on a fixation target and imagining a horizontal body displacement. The results showed a global overestimation of the possibility of passing under the line, and confirmed the influence of body orientation reported by Bringoux et al. (Exp Brain Res 185(4):673–680, 2008). More strikingly, a linear influence of gaze elevation was found on perceptual estimates. Precisely, downward eye elevation yielded increased overestimations, and conversely upward gaze elevation yielded decreased overestimations. Furthermore, body and gaze orientation effects were independent and combined additively to yield a global egocentric influence with a weight of 45 and 54%, respectively. Overall, our data suggest that multiple egocentric references can jointly affect the estimated possibility of passing under high obstacles. These results are discussed in terms of “interpenetrability” between geocentric and egocentric reference frames and clearly demonstrate that gaze elevation is involved, as body orientation, in geocentric spatial localization.     

Visuospatial memory computations during whole-body rotations in roll

We used a memory-saccade task to test whether the location of a target, briefly presented before a whole-body rotation in roll, is stored in egocentric or in allocentric coordinates. To make this distinction, we exploited the fact that subjects, when tilted sideways in darkness, make systematic errors when indicating the direction of gravity (an allocentric task) even though they have a veridical percept of their self-orientation in space. We hypothesized that if spatial memory is coded allocentrically, these distortions affect the coding of remembered targets and their readout after a body rotation. Alternatively, if coding is egocentric, updating for body rotation becomes essential and errors in performance should be related to the amount of intervening rotation. Subjects (n = 6) were tested making saccades to remembered world-fixed targets after passive body tilts. Initial and final tilt angle ranged between -120 degrees CCW and 120 degrees CW. The results showed that subjects made large systematic directional errors in their saccades (up to 90 degrees ). These errors did not occur in the absence of intervening body rotation, ruling out a memory degradation effect. Regression analysis showed that the errors were closely related to the amount of subjective allocentric distortion at both the initial and final tilt angle, rather than to the amount of intervening rotation. We conclude that the brain uses an allocentric reference frame, possibly gravity-based, to code visuospatial memories during whole-body tilts. This supports the notion that the brain can define information in multiple frames of reference, depending on sensory inputs and task demands.     

Egocentric and allocentric localization during induced motion

This research examined motor measures of the apparent egocentric location and perceptual measures of the apparent allocentric location of a target that was being seen to undergo induced motion (IM). In Experiments 1 and 3, subjects fixated a stationary dot (IM target) while a rectangular surround stimulus (inducing stimulus) oscillated horizontally. The inducing stimulus motion caused the IM target to appear to move in the opposite direction. In Experiment 1, two dots (flashed targets) were flashed above and below the IM target when the surround had reached its leftmost or rightmost displacement from the subject’s midline. Subjects pointed open-loop at either the apparent egocentric location of the IM target or at the bottom of the two flashed targets. On separate trials, subjects made judgments of the Vernier alignment of the IM target with the flashed targets at the endpoints of the surround’s oscillation. The pointing responses were displaced in the direction of the previously seen IM for the IM target and to a lesser degree for the bottom flashed target. However, the allocentric Vernier judgments demonstrated no perceptual displacement of the IM target relative to the flashed targets. Thus, IM results in a dissociation of egocentric location measures from allocentric location measures. In Experiment 2, pointing and Vernier measures were obtained with stationary horizontally displaced surrounds and there was no dissociation of egocentric location measures from allocentric location measures. These results indicate that the Roelofs effect did not produce the pattern of results in Experiment 1. In Experiment 3, pointing and Vernier measures were obtained when the surround was at the midpoint of an oscillation. In this case, egocentric pointing responses were displaced in the direction of surround motion (opposite IM) for the IM target and to a greater degree for the bottom flashed target. However, there was no apparent displacement of the IM target relative to the flashed targets in the allocentric Vernier judgments. Therefore, in Experiment 3 egocentric location measures were again dissociated from allocentric location measures. The results of this experiment also demonstrate that IM does not generate an allocentric displacement illusion analogous to the “flash-lag” effect.

Hand orientation is insufficiently compensated for in haptic spatial perception

What humans haptically perceive as parallel is often far from physically parallel. These deviations from parallelity are highly significant and very systematic. There exists accumulating evidence, both psychophysical and neurophysiological, that what is haptically parallel is decided in a frame of reference intermediate to an allocentric and an egocentric one. The central question here concerns the nature of the egocentric frame of reference. In the literature, various kinds of egocentric reference frames are mentioned for haptic spatial tasks, such as hand-centered, arm-centered, and body-centered frames of reference. Thus far, it has not been possible to distinguish between body-centered, arm-centered, and hand-centered reference frames in our experiments, as hand and arm orientation always covaried with distance from the body-midline. In the current set of experiments the influence of body-centered and hand-centered reference frames could be dissociated. Subjects were asked to make a test bar haptically parallel to a reference bar in five different conditions, in which their hands were oriented straight ahead, rotated to the left, rotated to the right, rotated outward or rotated inward. If the reference frame is body-centered, the deviations should be independent of condition. If, on the other hand, the reference frame is hand-centered, the deviations should vary with condition. The results show that deviation size varies strongly with condition, exactly in the way predicted by the influence of a hand-centered egocentric frame of reference. Interestingly, this implies that subjects do not sufficiently take into account the orientation of their hands.     

Pointing at targets by children with congenital and transient blindness

The study investigated pointing at memorized targets in reachable space in congenitally blind (CB) and blindfolded sighted (BS) children (6, 8, 10 and 12 years; ten children in each group). The target locations were presented on a sagittal plane by passive positioning of the left index finger. A go signal for matching the target location with the right index finger was provided 0 or 4 s after demonstration. An age effect was found only for absolute distance errors and the surface area of pointing was smaller for the CB children. Results indicate that early visual experience and age are not predictive factors for pointing in children. The delay was an important factor at all ages and for both groups, indicating distinct spatial representations such as egocentric and allocentric frames of reference, for immediate and delayed pointing, respectively. Therefore, the CB like the BS children are able to use both ego- and allocentric frames of reference.     

Differential effects of non-informative vision and visual interference on haptic spatial processing

The primary purpose of this study was to examine the effects of non-informative vision and visual interference upon haptic spatial processing, which supposedly derives from an interaction between an allocentric and egocentric reference frame. To this end, a haptic parallelity task served as baseline to determine the participant-dependent biasing influence of the egocentric reference frame. As expected, large systematic participant-dependent deviations from veridicality were observed. In the second experiment we probed the effect of non-informative vision on the egocentric bias. Moreover, orienting mechanisms (gazing directions) were studied with respect to the presentation of haptic information in a specific hemispace. Non-informative vision proved to have a beneficial effect on haptic spatial processing. No effect of gazing direction or hemispace was observed. In the third experiment we investigated the effect of simultaneously presented interfering visual information on the haptic bias. Interfering visual information parametrically influenced haptic performance. The interplay of reference frames that subserves haptic spatial processing was found to be related to both the effects of non-informative vision and visual interference. These results suggest that spatial representations are influenced by direct cross-modal interactions; inter-participant differences in the haptic modality resulted in differential effects of the visual modality.     

Whose hand is this? Handedness and visual perspective modulate self/other discrimination

We required healthy subjects to recognize visually presented one’s own or others’ hands in egocentric or allocentric perspective. Both right- and left-handers were faster in recognizing dominant hands in egocentric perspective and others’ non-dominant hand in allocentric perspective. These findings demonstrated that body-specific information contributes to sense of ownership, and that the “peri-dominant-hand space” is the preferred reference frame to distinguish self from not-self body parts.     

Allocentric versus egocentric spatial memory after unilateral temporal lobectomy in humans

Thirty patients who had undergone either a right or left unilateral temporal lobectomy (14 RTL; 16 LTL) and 16 control participants were tested on a computerized human analogue of the Morris Water Maze. The procedure was designed to compare allocentric and egocentric spatial memory. In the allocentric condition, participants searched for a target location on the screen, guided by object cues. Between trials, participants had to walk around the screen, which disrupted egocentric memory representation. In the egocentric condition, participants remained in the same position, but the object cues were shifted between searches to prevent them from using allocentric memory. Only the RTL group was impaired on the allocentric condition, and neither the LTL nor RTL group was impaired on additional tests of spatial working memory or spatial manipulation. The results support the notion that the right anterior temporal lobe stores long-term allocentric spatial memories.