The identification of coordination constraints across planes of motion

Two dominant coordination constraints have been identified during isofrequency conditions in previous work: the egocentric constraint, i.e., simultaneous activation of homologous muscle groups, and the allocentric constraint, i.e., moving the segments in the same direction in extrinsic space. To verify their generalization, bimanual drawing movements were performed in different planes of motion (transverse, frontal, sagittal, frontal-transverse) according to the in-phase and anti-phase mode along the X- and Y-axes. Convergent findings were obtained across the transverse, frontal, and frontal-transverse planes. The in-phase mode along both axes was performed most accurately/consistently, whereas the anti-phase mode resulted in a deterioration of the coordination pattern and this effect was most pronounced when the latter mode was introduced with respect to both dimensions. For sagittal plane motions, the in-phase mode was again superior but the second most optimal configuration was the anti-phase mode along both axes. This finding was hypothesized to result from the familiarity with the pattern since it resembles cycling behavior. It illustrates how cognitive mapping is superimposed onto the dynamics of interlimb coordination. Overall, these results support the presence of both the egocentric and allocentric constraint during bimanual movement production. Received: 21 August 1998 / Accepted: 12 February 1999     

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.     

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.     

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.     

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.

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.     

Interactions between interlimb and intralimb coordination during the performance of bimanual multijoint movements

The simultaneous performance of movements involving different effectors gives rise to neural and biomechanical interactions between and within limbs. The present study addressed the role of interlimb and intralimb constraints during the control of bimanual multijoint movements. Thirteen participants performed eight tasks involving the bilateral elbows and wrists under different coordination conditions. With respect to interlimb coordination, coordination patterns referred to the in-phase and anti-phase coordination modes, involving the simultaneous timing of homologous versus non-homologous muscles, respectively. With respect to inter-segmental (intralimb) coordination, the isodirectional mode referred to simultaneous flexions and extensions in the ipsilateral wrist and elbow joints, whereas the non-isodirectional mode involved simultaneous flexion in one joint together with extension in the other joint, or vice versa. The analysis of the data focused upon measures of relative phasing between proximal and distal joints within a limb as well as between the homologous joints of both limbs. With respect to interlimb coordination, findings revealed that adoption of the in-phase mode resulted in a higher quality of interlimb coordination than the anti-phase mode. However, the mode adopted in the distal joints had a larger impact on the quality of interlimb coordination than the mode adopted in the proximal joints. More specifically, in-phase coordination of the distal joints had a positive, and anti-phase coordination a negative, influence on the global coordinative behavior of the system. Minor effects of intralimb coordination modes on interlimb coordination were observed. With respect to intralimb coordination between the ipsilateral elbow and wrist, the isodirectional mode was performed with higher stability than the non-isodirectional mode. The mode of interlimb coordination also affected the quality of intralimb coordination, such that generating anti-phase coordination patterns in the distal joints had a negative influence on the accuracy and stability of intralimb coordination. Taken together, the present findings suggest a hierarchical structure whereby interlimb coordination constraints have a stronger impact on the global coordinative behavior of the system than intralimb coordination constraints. Moreover, the global coordinative state of the system is more affected by the coordination between the distal than between the proximal joints. Overall, the findings suggest that the mirror-image symmetry constraint has a powerful influence on bimanual multijoint coordination.     

Place recognition responses of neurons in monkey hippocampus

Neuronal activity in the monkey hippocampus was recorded while the monkey sat in a rotatable cab which it could cause to move from one location to another by pressing bars, and while it was presented various visual stimulation from several horizontal directions (directional stimulation). Of 174 hippocampal neurons recorded, 20 were selective to direction of the stimulus without place relation. Responses of these neurons could be described in egocentric coordinates for some and allocentric coordinates for others. Seventy-seven neurons had place related activity (place related neurons). Of these place related neurons, 21 were also directionally selective with responses described in egocentric or allocentric coordinates or both. The results show close relations between the coding of environmental space cues in egocentric and allocentric coordinates, and place related activity in the primate hippocampus.     

Reaction times for allocentric movements are 35 ms slower than reaction times for target-directed movements

Many movements that people perform every day are directed at visual targets, e.g., when we press an elevator button. However, many other movements are not target-directed, but are based on allocentric (object-centered) visual information. Examples of allocentric movements are gesture imitation, drawing or copying. Here, show a reaction time difference between these two types of movements in four separate experiments. In Exp. 1, subjects moved their eyes freely and used direct hand movements. In Exp. 2, subjects moved their eyes freely and their movements were tool-mediated (computer mouse). In Exp. 3, subjects fixated a central target and the visual field in which visual information was presented was manipulated. Experiment 4 was identical to Exp. 3 except for the fact that visual information about targets disappeared before movement onset. In all four experiments, reaction times in the allocentric task were approximately 35 ms slower than they were in the target-directed task. We suggest that this difference in reaction time between the two tasks reflects the fact that allocentric, but not target-directed, movements recruit the ventral stream, in particular lateral occipital cortex, which increases processing time. We also observed an advantage for movements made in the lower visual field as measured by movement variability, whether or not those movements were allocentric or target-directed. This latter result, we argue, reflects the role of the dorsal visual stream in the online control of movements in both kinds of tasks.     

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.     

Learned association of allocentric and egocentric information in the hippocampus

Single-cell recording was conducted in the hippocampus of rats that performed a spontaneous alternation task in a modified T-maze. In the central arm of the maze, 4 out of 45 cells (8%) were found that fired selectively depending on which turn the animals would take. This result is in disagreement with a previous study in which two-thirds of cells (22 out of 33) showed a clear bias for direction of turns. The interpretation was that the cells coded information of episodic memory. Our results do not support this hypothesis. Interestingly, over the course of training, an increasing number of cells were found that fired in correlation with the rats movements. It is proposed that these cells associate egocentric motor information with allocentric spatial information rather than encode episodic memory.     

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     

Allocentrically implied target locations are updated in an eye-centred reference frame

When reaching to remembered target locations following an intervening eye movement a systematic pattern of error is found indicating eye-centred updating of visuospatial memory. Here we investigated if implicit targets, defined only by allocentric visual cues, are also updated in an eye-centred reference frame as explicit targets are. Participants viewed vertical bars separated by varying distances, and horizontal lines of equivalently varying lengths, implying a “target” location at the midpoint of the stimulus. After determining the implied “target” location from only the allocentric stimuli provided, participants saccaded to an eccentric location, and reached to the remembered “target” location. Irrespective of the type of stimulus reaching errors to these implicit targets are gaze-dependent, and do not differ from those found when reaching to remembered explicit targets. Implicit target locations are coded and updated as a function of relative gaze direction with respect to those implied locations just as explicit targets are, even though no target is specifically represented.     

Altered navigational strategy use and visuospatial deficits in hAPP transgenic mice

Navigation deficits are prominent in Alzheimer's disease (AD) patients and transgenic mice expressing familial AD-mutant hAPP and A beta peptides. To determine the impact of strategy use on these deficits, we assessed hAPP and nontransgenic mice in a cross maze that can be solved by allocentric (world-based) or egocentric (self-based) strategies. Most nontransgenic mice used allocentric strategies, whereas half of hAPP mice were egocentric. At 3 months, all mice learned the cross maze rapidly; at 6 months, only allocentric hAPP mice were impaired. At 3 and 6 months, hAPP mice had reduced hippocampal Fos expression, which correlated with cross maze learning in older mice. Striatal pCREB expression was unaltered in hAPP mice, suggesting striatal sparing. We conclude that egocentric strategy use may be an earlier indicator of hAPP/A beta-induced hippocampal impairment than spatial learning deficits. Persistent use of allocentric strategies when egocentric strategies are available is maladaptive when there is hippocampal damage. Interventions promoting flexibility in selecting learning strategies might help circumvent otherwise debilitating navigational deficits caused by AD-related hippocampal dysfunction.     

The order of gaze shifts affects spatial and temporal aspects of discrete bimanual pointing movements

We investigated whether the order of gaze shifts affected spatial and temporal aspects of discrete bimanual pointing movements. Ten male participants concurrently executed bimanual pointing movements as quickly and accurately as possible to left and right lateral targets presented with the same and different amplitudes. They were asked to gaze initially at the left target and subsequently at the right target, or vice versa. Each hand showed less variable error and a faster reaction when the initial gaze shifted to the corresponding target than when the subsequent gaze shifted to it. For the same-amplitude targets, constant error (CE) was not influenced by the gaze order conditions. However, for the different-amplitude targets, CE for the short-amplitude target became larger when they initially gazed at the long-amplitude target than when they initially gazed at the short-amplitude target. The larger overshoot of the hand for the short-amplitude target occurred when the participants could not afford to foveate the target. Our results suggest that the order of gaze shifts determines whether asymmetric amplitude assimilation between the two hands occurs or not. Fast, consistent, and accurate bimanual pointing movements might be attributable to updating gaze-centered representations of target positions.     

The coordination patterns observed when two hands reach-to-grasp separate objects

What determines coordination patterns when both hands reach to grasp separate objects at the same time? It is known that synchronous timing is preferred as the most stable mode of bimanual coordination. Nonetheless, normal unimanual prehension behaviour predicts asynchrony when the two hands reach towards unequal targets, with synchrony restricted to targets equal in size and distance. Additionally, sufficiently separated targets require sequential looking. Does synchrony occur in all cases because it is preferred in bimanual coordination or does asynchrony occur because of unimanual task constraints and the need for sequential looking? We investigated coordinative timing when participants (n = 8) moved their right (preferred) hand to the same object at a fixed distance but the left hand to objects of different width (3, 5, and 7 cm) and grip surface size (1, 2, and 3 cm) placed at different distances (20, 30, and 40 cm) over 270 randomised trials. The hand movements consisted of two components: (1) an initial component (IC) during which the hand reached towards the target while forming an appropriate grip aperture, stopping at (but not touching) the object; (2) a completion component (CC) during which the finger and thumb closed on the target. The two limbs started the IC together but did not interact until the deceleration phase when evidence of synchronisation began to appear. Nonetheless, asynchronous timing was present at the end of the IC and preserved through the CC even with equidistant targets. Thus, there was synchrony but requirements for visual information ultimately yielded asynchronous coordinative timing.     

Head movements destabilize cyclical in-phase but not anti-phase homologous limb coordination in humans

The present study addressed the role of head movements in the coordination of the homologous upper or lower limbs in supine normal subjects. Consistent with previous research, in-phase mirror symmetrical movements were performed more accurately and consistently than anti-phase movements. However, inclusion of head movements destabilized in-phase but not anti-phase homologous limb coordination, in contrast to previous work demonstrating a higher vulnerability of anti-phase than in-phase coordination to various experimental perturbations. It was observed that the head moved in the same direction as the limbs during anti- but not during in-phase coordination. Furthermore, the interlimb patterns also affected the head rotations that were lower in spatiotemporal consistency and less consistently coupled with the limbs during in-phase than during anti-phase coordination. These findings provide new insights into the coalition of egocentric and allocentric constraints during interlimb coordination.     

Prefrontal serotonin depletion impairs egocentric, but not allocentric working memory in rats

Working memory is a cognitive ability chiefly organized by the prefrontal cortex. Working memory tests may be resolved based on allocentric or egocentric spatial strategies. Serotonergic neurotransmission is closely involved in working memory, but its role in spatial strategies for working memory performance is unknown. To address this issue, prefrontal serotonin depletion was induced to adult male rats, and three days after the behavioral expression of both allocentric and egocentric strategies were evaluated in the "Y" maze and in a crossed-arm maze, respectively. Serotonin depletion caused no effects on allocentric-related behavioral performance, but lesioned rats performed deficiently when the egocentric working memory was evaluated. These results suggest that serotonin may be more closely related with the organization of working memory that uses own movement-guided responses than with that involving the use of external visuospatial signals. Further neurochemical studies are needed to elucidate possible interactions between serotonergic activity and other neurotransmitter systems in the organization of working memory-related allocentric and egocentric strategies.     

Coordination and concurrency in bimanual rotation tasks when moving away from and toward the body

In the present series of experiments we investigated how object transport and rotate movements are performed when they are directed away from (Experiment 1) and toward (Experiment 2) the body under both unimanual and bimanual conditions. Our results indicated that unimanual conditions are faster and more efficiently produced than bimanual movements in far peripersonal space, suggesting that there is a cost to performing bimanual movements. However, in near peripersonal space, bimanual same movements were performed in a manner similar to unimanual movements, indicating that there is no significant cost associated with similar bimanual movements that are performed using the lower visual field and in near peripersonal space. Both experiments also indicate that the two hands are tightly synchronized when the two movements being performed require the same rotation. However, when performing bimanual movements where the rotation being performed by the two hands is different, this synchronization is weaker. Finally, the combined results from the two experiments indicated that movements made toward the body are not performed in a similar manner to movements that are made away from the body. Specifically, it is clear from the current studies that movements toward the body are performed faster and possibly that the hands are less synchronized for bimanual movements requiring different rotations by the two hands.

Unimanual and bimanual voluntary movement in Huntington's disease

Unimanual and bimanual cyclical forearm movements were studied in 15 Huntington's disease (HD) patients and 15 healthy, gender- and age-matched controls. Whereas the unimanual task was only performed at maximal speed, the bimanual movements were performed according to the in-phase and anti-phase mode at different cycling frequencies. The HD patients also performed the tasks after 12 months of follow-up. Findings revealed that maximal cycling frequency during unimanual movement was significantly lower in HD patients as compared with controls. In addition, measures of relative phasing established that bimanual cyclical movements were performed with lower accuracy and higher variability in HD patients. The differential variability between both groups was magnified by increasing the cycling frequency and coordinative complexity whereas only coordinative complexity differentially affected the accuracy of relative phasing. The obtained performance measures were found to be significantly correlated with disease duration (unimanual) and with the score on the total motor scale, the Mini-Mental State Examination and the Stroop Interference Test (uni- and bimanual). After 12 months, maximal cycling frequency of unimanual elbow flexion–extension was significantly decreased in HD patients whereas the quality of the in-phase and anti-phase movement patterns remained stable. Electronic Publication