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How do we achieve unitary perception of an object when it touches two parts of the sensory epithelium that are not contiguous? We investigated this problem with a simple psychophysical task, which we then used in an fMRI experiment. Two wooden rods were moved over two digits positioned to be spatially adjacent. The digits were either from one foot (or hand) or one digit was from either foot (or hand). When the rods were moving in phase, one object was reliably perceived. By contrast, when the rods were moving out of phase, two objects were reliably perceived. fMRI revealed four cortical areas where activity was higher when the moving rods were perceived as one object relative to when they were perceived as two separate objects. Areas in the right inferior parietal lobule, the left inferior temporal sulcus and the left middle frontal gyrus were activated for this contrast regardless of the anatomical configuration of the stimulated sensory epithelia. By contrast, the left intraparietal sulcus was activated specifically when integration across the midline was required, irrespective of whether the stimulation was applied to the hands or feet. These results reveal a network of brain areas involved in generating a unified percept of the presence of an object that comes into contact with different parts of the body surface.

This study quantitatively describes differences between participants with hemiparetic stroke and age-matched healthy participants in axial body segment and gait kinematics during a direction change task. Participants were required to change walking direction by 45 degrees, either to their left or right, at the midpoint of a 6-m path. Participants were visually cued either at the start of the walk (pre-planned) or one stride before they reached the turn point (reactive). The sequence and inter-segmental timing of axial orientation onset was preserved in participants with stroke. Analysis of a subgroup of stroke survivors indicated that participants with lesions affecting the basal ganglia (BG) took significantly longer time than control participants to initiate the reorientation synergy when making turns to their non-paretic side. We hypothesize that these differences are a product of asymmetrical activity of dopaminergic pathways in the brain resulting from compromised BG function.

Synchronizing our actions with external events is a task we perform without apparent effort. Its foundation relies on accurate temporal control that is widely accepted to take one of two different modes of implementation: explicit timing for discrete actions and implicit timing for smooth continuous movements. Here we assess synchronisation performance for different types of action and test the degree to which each action supports corrective updating following changes in the environment. Participants performed three different finger actions in time with an auditory pacing stimulus allowing us to assess synchronisation performance. Presenting a single perturbation to the otherwise regular metronome allowed us to examine corrections supported by movements varying in their mode of timing implementation. We find that discrete actions are less variable and support faster error correction. As such, discrete actions may be preferred when engaging in time-critical adaptive behaviour with people and objects in a dynamic environment.

We investigated the effect of correlation in background noise on the ability of participants to detect a kinaesthetic target on the index fingers of their hands. Participants had to judge whether the target (a smoothed ramp with quarter-sine onset and offset), was on the left or right finger (experiments 1 and 2). These targets were embedded in noise generated by pseudo-random up-down movements of both the left and right fingers. Positive correlation between the noise on the left and right fingers aided discrimination of the target signal relative to when the noise was uncorrelated. However, this benefit of positive correlation was subject to temporal limitations and diminished with increasing lag between the noise on the fingers (experiment 2). Tests of explicit detection of correlation in kinaesthetic noise (experiment 3) showed a similar pattern, with detection of correlation decreasing with the temporal lag between the noise on the fingers. The results suggest that kinaesthetic signal detection involves sensory integration across the fingers within a limited temporal window.

Typically, when a grasping response is made, the hand opens wider than the target object. We show that this "over-grasp" response is reduced when we reach to parts of our own face, relative to when we reach to other body parts or to neutral objects. This is not due to reaching to different parts of body space, as over-grasp responses are indifferent to whether or not other body parts or neutral objects are placed close to the face. It is also not due to differences in perceptual knowledge of the size of the target object. We conclude instead that the familiarity of face parts influences the grasping response directly. Subsequent experiments demonstrate that the movement representation determining any effect is not based on a torso-centred frame, and not abstracted from the specific hand used for grasping. We discuss the implications of the results for understanding and measuring motor representations for familiar actions.

Previous studies of paced repetitive movements with respect to an external beat have either emphasised (a) the form of movement trajectories or (b) timing errors made with respect to the external beat. The question of what kinds of movement trajectories assist timing accuracy has not previously been addressed. In an experiment involving synchronisation or syncopation with an external auditory metronome we show that the nervous system produces trajectories that are asymmetric with respect to time and velocity in the out and return phases of the repeating movement cycle. This asymmetry is task specific and is independent of motor implementation details (finger flexion vs. extension). Additionally, we found that timed trajectories are less smooth (higher mean squared jerk) than unpaced ones. The degree of asymmetry in the flexion and extension movement times is positively correlated with timing accuracy. Negative correlations were observed between synchronisation timing error and the movement time of the ensuing return phase, suggesting that late arrival of the finger is compensated by a shorter return phase and conversely for early arrival. We suggest that movement asymmetry in repetitive timing tasks helps satisfy requirements of precision and accuracy relative to a target event. © Springer-Verlag 2004.

Perception is linked to action via two routes: a direct route based on affordance information in the environment and an indirect route based on semantic knowledge about objects. The present study explored the factors modulating the recruitment of the two routes, in particular which factors affecting the selection of paired objects. In Experiment 1, we presented real objects among semantically related or unrelated distracters. Participants had to select two objects that can interact. The presence of distracters affected selection times, but not the semantic relations of the objects with the distracters. Furthermore, participants first selected the active object (e.g. teaspoon) with their right hand, followed by the passive object (e.g. mug), often with their left hand. In Experiment 2, we presented pictures of the same objects with no hand grip, congruent or incongruent hand grip. Participants had to decide whether the two objects can interact. Action decisions were faster when the presentation of the active object preceded the presentation of the passive object, and when the grip was congruent. Interestingly, participants were slower when the objects were semantically but not functionally related; this effect increased with congruently gripped objects. Our data showed that action decisions in the presence of strong affordance cues (real objects, pictures of congruently gripped objects) relied on sensory-motor representation, supporting the direct route from perception-to-action that bypasses semantic knowledge. However, in the case of weak affordance cues (pictures), semantic information interfered with action decisions, indicating that semantic knowledge impacts action decisions. The data support the dual-route account from perception-to-action.

In order to investigate the potential hand contribution to sideways balance, sideways pushes to the right, which subjects resisted using either the lower limbs ("hip only") or the lower limbs assisted by the right upper limb ("hand and hip"), were delivered to the pelvis. Analysis of force and electromyogram recordings from the legs, arm and hand in the hand-and-hip condition showed a close co-ordination of upper and lower limbs in terms of mean latencies and amplitudes. However, trial to trial fluctuations of forces generated by the hand and leg did not correlate, suggesting parallel pathways under central co-ordinative control.

This experiment investigated the prediction of load force (LF) in impulsive collisions inferred from an- ticipatory adjustments of grip force (GF) used to stabi- lise a hand-held object. Subjects used a precision grip to hold the object between thumb and index finger of their right hand and used the arm either: (1) to move the ob- ject to produce a collision by hitting the lower end of a pendulum, causing it to swing to one of three target an- gles, or (2) to hold the object still while receiving a colli- sion produced by the experimenter releasing the pendu- lum from one of three angles. Visual feedback of the pendulum's trajectory was available in the production task only. In all conditions, subjects increased GF in ad- vance of the collision. In receiving the collision without advance information, subjects set GF levels to the mid- range of the experienced forces. When subjects pos- sessed knowledge about the maximum angle of pendu- lum swing - either because they were going to produce it or because they were verbally informed - magnitude of the anticipatory-GF magnitude response was scaled to the predicted LF magnitude. Furthermore, GF was scaled to LF with a higher gain when producing compared to receiving the collision. This suggests that updating for- ward models through a semantic route is not as powerful as when the updating is achieved through the more direct route of dynamic exploration.

Previously, it has been shown that synchronising actions with periodic pacing stimuli are unaffected by ageing. However, synchronisation often requires combining evidence across multiple sources of timing information. We have previously shown the brain integrates multisensory cues to achieve a best estimate of the events in time and subsequently reduces variability in synchronised movements (Elliott et al. in Eur J Neurosci 31(10):1828–1835, 2010). Yet, it is unclear if sensory integration of temporal cues in older adults is degraded and whether this leads to reduced synchronisation performance. Here, we test for age-related changes when synchronising actions to multisensory temporal cues. We compared synchronisation performance between young (N = 15, aged 18–37 years) and older adults (N = 15, aged 63–80 years) using a finger-tapping task to auditory and tactile metronomes presented unimodally and bimodally. We added temporal jitter to the auditory metronome to determine whether participants would integrate auditory and tactile signals, with reduced weighting of the auditory metronome as its reliability decreased under bimodal conditions. We found that older adults matched the performance of young adults when synchronising to an isochronous auditory or tactile metronome. When the temporal regularity of the auditory metronome was reduced, older adults’ performance was degraded to a greater extent than the young adults in both unimodal and bimodal conditions. However, proportionally both groups showed similar improvements in synchronisation performance in bimodal conditions compared with the equivalent, auditory-only conditions. We conclude that while older adults become more variable in synchronising to less regular beats, they do not show any deficit in the integration of multisensory temporal cues, suggesting that using multisensory information may help mitigate any deficits in coordinating actions to complex timing cues.

An object held in precision grip creates predictable load forces on the hand during voluntary hand movement and these are associated with anticipatory modulation of grip force. Conflicting results have been obtained over whether predictable external load perturbations result in anticipatory grip force responses (e.g. Blakemore et al. in J Neurosci 18(18):7511-7518, 1998; Weeks et al. in Exp Brain Res 132:404-410, 2000). This paper investigated whether the discrepancies reflect differences in the methods used in estimating the time delay. Subjects held a manipulandum that delivered load force perturbations in the form of pulses of variable duration and interval or periodic 0.5 and 1 Hz square waves or sinusoids. The grip forces exerted by the subjects were measured. Two methods were used to assess the time delay of the grip force in relation to the load force: (1) cross-spectral analysis, (2) a single threshold method applied on time-locked averaged data. Despite a phase lag shown by the cross-spectral analysis, the threshold method revealed grip force increased 264.8+/-40.2 ms before the onset of the load force when 0.5 Hz square waves were used as the load force perturbation and 70.2+/-17.0 ms before the load force when 1 Hz square waves were used. Computer simulations indicated that the single threshold method gives a more sensitive estimate of the onset time than the cross-spectral analysis. We conclude that discrepancies in previous studies reflect differences in the methods used to assess the time-delay and that there is an anticipatory component in the grip force response to predictable external load perturbation.

Object symmetry is a visual attribute that may contribute to perceptual judgement and to action. We evaluated the effects of varying the physical symmetry of planar objects (presence versus absence) on both aspects. In Experiment 1, subjects estimated the magnitude of visually perceived symmetry of the objects. The results confirmed the influence of physical symmetry on perceived symmetry, and supported our binary categorisation of stimulus objects in terms of presence versus absence of physical symmetry. In Experiment 2, participants used a precision grip to grasp and stably lift the same planar objects varying in degree of symmetry. Choice of grasp points was unrestricted. Participants selected a grasp axis (between thumb and middle finger) that limited the perpendicular distance from CM (i.e., grasp-axis error) to just a few millimetres. Moreover, they took advantage of visual cues to object symmetry to better determine CM, thus reducing their grasp-axis error for symmetric (vs. asymmetric) objects by 31%. We interpret these findings in terms of user and object-geometry constraints on grasp-point selection.

We describe adjustments in grip force as a consequence of fluctuations in inertial load force during vertical movements of the upper limb in a patient with cerebellar degeneration. Normally grip force is adapted to load-force fluctuations, in particular to the maximum load force, which occurs early in upward movements and late in downward movements. Increased grip force during movement was observed in the patient, but the timing of maximum grip force was not different between upward and downward movements. This suggests impaired cerebellar prediction of the dynamic consequences of voluntary movement.

It has previously been shown that light contact with the finger tip on a fixed surface reduces centre of pressure (CoP) fluctuations in the frontal plane when standing in an unstable posture with the feet in line (tandem Romberg stance). Positive cross-correlations between horizontal finger forces and CoP fluctuations with finger forces exhibiting a phase lead suggest the hand provides sensory input for postural stability. The present study investigates whether this is the case for normal posture. We report reduced CoP fluctuations in the sagittal plane when light touch is permitted during normal bipedal stance. Moreover, we find positive crosscorrelations between finger tip forces and CoP fluctuations which are of similar magnitude and phase lag to those observed in tandem Romberg stance. This shows the utility of hand touch input for regulation of normal upright posture as well as inherently unstable postures such as tandem Romberg.

The effect of the predictability of perturbation to standing balance was evaluated in terms of the muscle activity and response dynamics of five subjects exposed to horizontal forces at the pelvis producing sideways or forward sway. Rapid (EMG onset latencies of 70–80 ms recorded from the left gluteus medius and gastrocnemius) and qualitatively different patterns of response were produced by forward pushes and pushes to either side. However, the EMG response to left push was constant in pattern and timing, whether the push direction was constant and, therefore, predictable over a block of trials or whether the left push trials were interleaved randomly with right push or forward push trials. Moreover, there were no systematic effects of perturbation direction uncertainty on the latency and rate of increase of ground reaction forces. We conclude that prior information does not speed postural responses that differ quantitatively according to the direction of perturbation to balance.

We have investigated the coordination of hand aperture with the spatial path of hand transport in prehensile movement by comparing straight prehensile movements with curved movements, in which subjects had to pass over a "via point" marked on the work surface before picking up an object in the target location. Spatial plots of hand aperture against hand transport showed that the preshaping of the hand to prepare an appropriate grasp was delayed in the curved movements relative to the straight movements, with most of the preshaping of the hand occurring after passing the via point, even when the via point occurred late in the course of the movement. The postponement of hand preshaping was apparently not due to subjects' segmenting the movement into two completely separate portions preceding and following the via point, since some degree of hand opening often occurred before the via point. We suggest that the delay in hand opening in curved movements involves a scheduling process, which uses information about hand transport to set an appropriate hand aperture.

We have investigated how the control of hand transport and of hand aperture are coordinated in prehensile movements by delivering mechanical perturbations to the hand transport component and looking for coordinated adjustments in hand aperture. An electric actuator attached to the subject's right arm randomly pulled the subject backwards, away from the target, or pushed them towards it, during a quarter of the experimental trials. A compensatory adjustment of hand aperture followed the immediate, mechanical effects of the perturbation of hand transport. The adjustment appeared to return the subject towards a stereotyped spatial relation between hand aperture and hand transport. These spatial patterns suggest how the two components may be coordinated during prehension. A simple model of this coordination, based on coupled position feedback systems, is presented.

In this paper, we examine grip forces and load forces during point-to-point arm movements with objects grasped with a precision grip. We demonstrate that grip force is finely modulated with load force. Variations in load force arise from inertial forces related to movement; grip force rises as the load force increases and falls as load force decreases. The same finding is observed in vertical and horizontal movements performed at various rates. In vertical movements, maximum grip force coincides in time with maximum load force. The maxima occur early in upward and later in downward movements. In horizontal movements, where peaks in load force are observed during both the acceleratory and deceleratory phases, grip force rises at the beginning of the movement and remains high until the end. The results suggest that when moving an object with the hand the programming of grip force is an integral part of the planning process.

In this paper we examine the coordination of grip force and load during brisk cyclic arm movements with a hand-held object under a range of conditions. We show that, regardless of the surface texture of the object or movement frequency, grip force is modulated in parallel with load. Thus, the tight coupling between grip force and load observed in short-duration tasks such as lifting or point-to-point movements is also seen in longer-duration cyclic movements. Moreover, the gain of the relation between grip force and load remains essentially constant over time. Across conditions, we find a dissociation between the gain of the relation between grip force and load and the grip force offset. With a more slippery surface texture both the gain and offset increase, whereas increases in frequency lead to an increase in the offset but a decrease in gain. This suggests that these two parameters are under independent high-level control. We also observe that when subjects were instructed to maintain a high-baseline grip force during the movement, grip force was still modulated with load even though an increase in grip was not necessary to prevent slip. This suggests that there is an obligatory coupling between grip force and load. This coupling might be subserved by low-level mechanisms not under high-level control.