'Side-effects': intrinsic and task-induced asymmetry in bimanual rhythmic coordination

Multifrequency coordination studies have shown the importance of hand-role in addition to hand-preference in bimanual rhythmic coordination. In these studies, hand-role has been defined by the task of the individual hands (moving fast or slow). In the present study, the hands were coordinated at the same frequency and hand-role was defined by the asymmetry of the coordination pattern. Eleven consistent left-handers and 13 consistent right-handers tapped three patterns (anti-phase, left-gallop, right-gallop) in four visual feedback conditions (no feedback, left-hand feedback, right-hand feedback, full feedback). The analysis focused on phase shifts, phase variability, intertap interval variability, and correlations between intertap intervals. The manipulation of visual feedback had only minor effects. In the anti-phase pattern, a symmetric coupling mechanism was found. The results support the idea that coordination in the gallop pattern is governed by a hierarchical control mechanism. In contrast to the multifrequency studies, however, successful control in the gallop is not dependent on a hand arrangement that accommodates the preferred hand as the leading hand. An adjustment to the model of Summers et al. (1993, J Exp Psychol Hum Percept Perform 19:416–428) is presented for the case of the gallop pattern.     

Perception of relative throw-ability

Bingham et al. (J Exp Psychol Hum Percept Perform 15(3):507–528, 1989) showed that skilled throwers can perceive optimal objects for throwing to a maximum distance. Zhu and Bingham (J Exp Psychol Hum Percept Perform 34(4):929, 2008, 36(4):862–875, 2010) replicated this finding and then showed that felt heaviness is used to perceive this affordance (see also Zhu and Bingham in Evol Hum Behav 32(4):288–293, 2011; Zhu et al. in Exp Brain Res 224(2):221–231, 2013). Throwers pick the best weight for spherical projectiles in each graspable size. Bingham et al. (J Exp Psychol Hum Percept Perform 15(3):507–528, 1989) speculated that relative throw-ability might be perceptible. This would mean that the ordering of distances achieved by maximum effort throws of different objects could be judged. This affordance property is not the same as optimal throw-ability, because it requires all projectiles to be evaluated relative to one another with respect to ordinally scaled distances, not just a discrete optimum. We now used a magnitude estimation task to test this hypothesis, comparing the resultant ordering with that exhibited by distances of throws in previous studies. The findings show that participants were able to perform the perceptual task. However, discrimination among objects of different weight within a size was better than between sizes. The implications of these results for understanding of the information used to perform this task are discussed.     

Bimanual Fitts’ tasks: Kelso,Southard, and Goodman, 1979 revisited

The experiment was designed to replicate and extend to an integrated feedback condition the pattern of movement time results found by Kelso et al. (J Exp Psychol Hum Percept Perform 5:229–238, 1979a, Science 204:1029–1031, 1979b) where the simultaneous movement of one hand to a low ID target and the other to a higher ID target indicated “a tight coordinate coupling between the hands” (p. 229). In the present experiment, a control group was provided feedback that depicted the independent movement of the two limbs under low and higher indexes of difficulty (ID). A Lissajous group was provided integrated feedback in the form of a Lissajous plot. The results indicated a pattern of results for the control and Lissajous groups similar to that found by Kelso et al. for one and two-limb movements to the same difficulty targets. The control group also replicated the finding for two-limb movements to mixed ID tasks. However, the Lissajous group simultaneously produced disparate movement in the mixed target conditions. The results are consistent with recent findings indicating that when provided salient integrated feedback participants can effectively produce disparate movements of the two limbs.     

Revisited: the inertia tensor as a proprioceptive invariant in humans.

A multitude of tasks that we perform on a daily basis require precise information about the orientation of our limbs with respect to the environment and the objects located within it. Recent studies have suggested that the inertia tensor, a physical property whose values are time- and co-ordinate-independent, may be an important informational invariant used by the proprioceptive system to control the movements of our limbs (Pagano et al., Ecol. Psychol. 8 (1996) 43; Pagano and Turvey, Percept. Psychophys. 52 (1992) 617; Pagano and Turvey, J. Exp. Psychol. Hum. Percept. Perform. 21 (1995) 1070). We tested this hypothesis by recording the angular errors made by subjects when pointing to virtual targets in the dark. Close examination of the pointing errors made did not show any significant effects of the inertia tensor modifications on pointing accuracy. The kinematics of the pointing movements did not indicate that any on-line adjustments were being made to compensate for the inertia tensor changes. The implications of these findings with respect to the functioning of the proprioceptive system are discussed.     

Spatial constraints in bimanual coordination: influences of effector orientation

Two experiments are reported that examined the influence of spatial orientation of the upper limbs in bimanual coordination. In both experiments, the upper limbs were oriented in either parallel, orthogonal, or obtuse spatial configurations and participants were asked to move their limbs continuously in temporal (1:1) synchrony, prepared in either in-phase or anti-phase modes of coordination. Bimanual coordination trials in Experiment 1 were paced by a metronome at one of four frequencies (1.0, 1.5, 2.0 or 2.5 Hz). Measures of relative phase accuracy and stability both revealed that, as metronome frequency increased, in-phase coordination dominated for the parallel spatial orientation, anti-phase coordination dominated for the orthogonal spatial orientation, and neither pattern dominated for the obtuse spatial orientations. In Experiment 2, an intentional switch method replicated and extended these influences of spatial orientation. The time to voluntarily switch from an anti-phase pattern to an in-phase pattern was faster than an in-phase to anti-phase switch (confirming support for the dominance of the in-phase pattern), but this was true only for the parallel spatial orientation. The reverse was true for the orthogonal spatial orientation (i.e., faster from in-phase to anti-phase), and no difference in switch times was observed for an obtuse spatial orientation. These findings support and extend previous research regarding the influence of spatial orientation in bimanual coordination and may be attributed to the role of, and potential interactions between, egocentric, allocentric, and mechanical constraints during action.     

Limitations of feedforward control in multiple-phase steering movements

When attempting to perform bi-phasic steering movements (such as a lane change) in the absence of visual and inertial feedback, drivers produce a systematic heading error in the direction of the lane change (Wallis et al., Curr Biol 12(4):295–299, 2002; J Exp Psychol Hum Percept Perform 33(55):1127–1144, 2007). Theories of steering control which employ exclusively open-loop control mechanisms cannot accommodate this finding. In this article we show that a similar steering error occurs with obstacle avoidance, and offer compelling evidence that it stems from a seemingly general failure of human operators to correctly internalise the dynamics of the steering wheel. With respect to lateral position, the steering wheel is an acceleration control device, but we present data indicating that drivers treat it as a rate control device. Previous findings from Wallis et al. can be explained the same way. Since an open-loop control mechanism will never succeed when the dynamics of the controller are internalised improperly, we go on to conclude that regular, appropriately timed sensory feedback—predominantly from vision—is necessary for regulating heading, even during well-practiced, everyday manoeuvres such as lane changing and obstacle avoidance.     

Locomotor behaviour of children while navigating through apertures

During everyday locomotion, we encounter a range of obstacles requiring specific motor responses; a narrow aperture which forces us to rotate our shoulders in order to pass through is one example. In adults, the decision to rotate their shoulders is body scaled (Warren and Whang in J Exp Psychol Hum Percept Perform 13:371–383, 1987), and the movement through is temporally and spatially tailored to the aperture size (Higuchi et al. in Exp Brain Res 175:50–59, 2006; Wilmut and Barnett in Hum Mov Sci 29:289–298, 2010). The aim of the current study was to determine how 8-to 10-year-old children make action judgements and movement adaptations while passing through a series of five aperture sizes which were scaled to body size (0.9, 1.1, 1.3, 1.5 and 1.7 times shoulder width). Spatial and temporal characteristics of movement speed and shoulder rotation were collected over the initial approach phase and while crossing the doorway threshold. In terms of making action judgements, results suggest that the decision to rotate the shoulders is not scaled in the same way as adults, with children showing a critical ratio of 1.61. Shoulder angle at the door could be predicted, for larger aperture ratios, by both shoulder angle variability and lateral trunk variability. This finding supports the dynamical scaling model (Snapp-Childs and Bingham in Exp Brain Res 198:527–533, 2009). In terms of movement adaptations, we have shown that children, like adults, spatially and temporally tailor their movements to aperture size.     

The stability of rhythmic movement coordination depends on relative speed: the Bingham model supported

Following many studies showing that the coupling in bimanual coordination can be perceptual, Bingham (Ecol Psychol in 16:45–53, 2001; 2004a, b) proposed a dynamical model of such movements. The model contains three key hypotheses: (1) Being able to produce stable coordinative movements is a function of the ability to perceive relative phase, (2) the information to perceive relative phase is relative direction of motion, and (3) the ability to resolve this information is conditioned by relative speed. The first two hypotheses have been well supported (Wilson and Bingham in Percept Psychophys 70:465–476, 2008; Wilson et al. in J Exp Psychol Hum 36:1508–1514, 2010a), but the third was not supported when tested by de Rugy et al. (Exp Brain Res 184:269–273, 2008) using a visual coordination task that required simultaneous control of both the amplitude and relative phase of movement. The purposes of the current study were to replicate this task with additional measures and to modify the original model to apply it to the new task. To do this, we conducted two experiments. First, we tested the ability to produce 180° visual coordination at different frequencies to determine frequencies suitable for testing in the de Rugy et al. task. Second, we tested the de Rugy et al. task but included additional measures that yielded results different from those reported by de Rugy et al. These results were used to elaborate the original model. First, one of the phase-driven oscillators was replaced with a harmonic oscillator, so the resulting coupling was unidirectional. This change resulted in the model producing less stable 180° coordination behavior beyond 1.5 Hz consistent with the results obtained in Experiment 1. Next, amplitude control and phase correction elements were added to the model. With these changes, the model reproduced behaviors observed in Experiment 2. The central finding was that the stability of rhythmic movement coordination does depend on relative speed and, thus, all three of the hypotheses contained in the original Bingham model are supported.     

Two mechanisms underlying inhibition of return

Inhibition of return (IOR) refers to slower reaction times to targets presented at previously stimulated or inspected locations. Taylor and Klein (J Exp Psychol Hum Percept Perform 26(5):1639–1656, 2000) showed that IOR can affect either attentional/perceptual or motor processes, depending on whether the oculomotor system is in a quiescent or in an activated state, respectively. If the motoric flavour of IOR is truly non-perceptual and non-attentional, no IOR should be observed when the responses to targets are not based on spatial information. In the present experiments, we demonstrated that when the eyes moved to the peripheral cue and back to centre before the target appeared (to generate the motoric flavour), IOR was observed in detection tasks, for which the spatial location is an integral feature of the onset that is reported, but not in colour discrimination tasks, for which the outcome of a non-spatial perceptual discrimination is reported. When eye movements were prevented, both tasks showed robust IOR. We, therefore, conclude that the motoric flavour of IOR, elicited by oculomotor activation, does not affect attention or perceptual processing.     

Postural coordination modes and transition: dynamical explanations

While research to date has been successful in quantifying postural behaviour, this paper examines the causes of transition between postural coordination mode using dynamical variables and, by inference, efficient control strategies underlying postural behaviour. To this end, six subjects in bipedal stance were instructed to maintain a constant distance between their head and a visual target that oscillated along the line of sight. Within sessions, participants were exposed to gradual changes in increasing target motion frequency. Kinematic results showed a sudden transition between in-phase and anti-phase postural coordination modes in visual target tracking. The dynamical analysis pointed out that (1) the center of pressure (CoP) position parameter is a crucial parameter in the determination of the adopted coordination mode, (2) the change occurred in response to limits bordered by the system: the interaction between equilibrium constraints (A/P displacements of CoP), physiological limits (net joint moments) support the emergence of different postural behaviours and, (3) finally, the anti-phase mode presents a better distribution of muscular moment between hip and ankle joints and is more effective to achieve high frequency oscillations with limited CoP displacements.     

Visual–spatial and anatomical constraints interact in a bimanual coordination task with transformed visual feedback

There is a debate in the literature about the influence of spatial and anatomical constraints on bimanual coordination dynamics. In the present experiment, participants swung hand-held pendulums about the wrist while attending to visual feedback about relative phase (superimposed phase plots of each pendulum) that was displayed on a screen. Participants were instructed to maintain in-phase or anti-phase coordination in the visual display. Visual–spatial and anatomical constraints were dissociated by introducing a phase shift in the visual display so that visual feedback differed from the movements being performed by the participants in 15° increments from −180° to +180°. Analysis of mean relative phase and its variability suggested that visual–spatial and anatomical constraints interact in bimanual coordination dynamics.     

Short duty cycle destabilizes a half-center oscillator, but gap junctions can restabilize the anti-phase pattern

Mutually inhibitory pacemaker neurons with duty cycle close to 50% operate as a half-center oscillator (anti-phase coordination, i.e., 180 degrees out of phase), even in the presence of weak to modest gap junctional coupling. For electrical coupling strength above a critical value synchronization occurs. But, as shown here with modeling studies, the effects of electrical coupling depend critically on a cell's duty cycle. Instead of oscillating either in-phase or anti-phase, model cells with short duty cycle express additional rhythmic patterns, and different transitions between them, depending on electrical coupling strength. For weak or no electrical coupling, cells do not oscillate in anti-phase but instead exhibit almost in-phase activity. Strengthening this weak coupling leads to stable anti-phase activity. With yet stronger electrical coupling stable inphase (synchrony) emerges but it coexists with the anti-phase pattern. Thus the network shows bistability for an intermediate range of coupling strength. For sufficiently strong electrical coupling synchrony is the network's only attracting rhythmic state. Our results, numerical and analytical (phase plane analysis), are based on a minimal but biophysically motivated pacemaker model for the slowly oscillating envelope of bursting neurons. However, illustrations for an Hodgkin-Huxley model suggest that some of our results for short duty cycle may extend to patterning of repetitive spikes. In particular, electrical coupling of intermediate strength may promote anti-phase activity and provide bistability of anti-phase and in-phase spiking.     

Effects of plantar flexor muscle fatigue induced by electromyostimulation on postural coordination

The aim of the present study was to investigate the influence of a modification of an intrinsic capacity (plantar flexor strength) on the implementation of in-phase and anti-phase mode of coordination. Analysis of hip and ankle relative phases during fore-aft tracking task was done before and after an electromyostimulation fatigue protocol on the soleus muscles. Results showed participants used exclusively in-phase and anti-phase modes of coordination, with a sudden switch from one to the other with target frequency increase. Regarding tracking tasks, fatigue induces a decrease of performance for lower frequencies, and a significant decrease of switch frequency (−0.08 Hz) for each subject. In conclusion, changes in mode of coordination implementation suggest that the in-phase mode implementation is highly linked to the strength production capacity at the ankle joint.     

Evidence for an attentional component in saccadic inhibition of return

After presentation of a peripheral cue, facilitation at the cued location is followed by inhibition of return (IOR). It has been recently proposed that IOR may originate at different processing stages for manual and ocular responses, with manual IOR resulting from inhibited attentional orienting, and ocular IOR resulting form inhibited motor preparation. Contrary to this interpretation, we found an effect of target contrast on saccadic IOR. The effect of contrast decreased with increasing reaction times (RTs) for saccades, but not for manual key-press responses. This may have masked the effect of contrast on IOR with saccades in previous studies (Hunt and Kingstone in J Exp Psychol Hum Percept Perform 29:1068–1074, 2003) because only mean RTs were considered. We also found that background luminance strongly influenced the effects of gap and target contrast on IOR.

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     

The behavioural and electrophysiological effects of visual task difficulty and bimanual coordination mode during dual-task performance

The difficulty of a visual three stimulus and a bimanual coordination task was manipulated by varying discrimination difficulty (easy, hard) and coordination mode (in-phase, anti-phase) respectively. Electroencephalographic activity was recorded from 32 sites whilst participants (n = 16) completed four dual-task conditions in counterbalanced order. Longer reaction time and lower accuracy were found for the hard relative to the easy visual task and, for the hard visual task, accuracy was lower under anti-phase relative to in-phase conditions. Amplitude and latency of event-related potential components P3a and P3b were recorded and measured. There was a reduction in P3b amplitude and increase in P3a amplitude for the hard visual task overall and a further reduction in frontal P3b amplitude under the more demanding anti-phase condition. For the easy visual task, however, P3b and P3a amplitude were greater under the anti-phase relative to in-phase coordination condition at left hemisphere frontal sites. These findings suggest that the attentional cost of stabilising anti-phase bimanual coordination is largely associated with top-down automatic processes subserved by the frontal attentional network.     

Using scanning trials to assess intrinsic coordination dynamics

Bimanual 1:1 coordination patterns other than in-phase (0°) and anti-phase (180°) have proven difficult to perform even with extended practice. The difficulty has been attributed to phase attraction that draws the coordination between the limbs towards the bimanual patterns of in-phase and anti-phase and variability associated with the activation of non-homologous muscles via crossed and uncrossed cortical pathways. We found participants could very effectively produce a large range of supposedly unstable coordination patterns (between 0° and 180° in 30° increments) after only 3 min of practice when integrated feedback (Lissajous plots) was provided and other perceptual and attentional distractions were minimized. These findings clearly indicate that the perception-action system is fully capable of producing a wide range of bimanual coordination patterns and that the reason for the failure to produce these patterns in previous experiments reside in the perceptual information and attentional requirements typically found in experimental testing environments.     

Stimulus-driven selection of routes to imitation

Several models have proposed that an action can be imitated via one of two routes: a direct visuospatial route, which can in principle mediate imitation of both meaningful (MF) and meaningless (ML) actions, and an indirect semantic route, which can be used only for MF actions. The present study investigated whether selection between the direct and indirect routes is strategic or stimulus driven. Tessari and Rumiati (J Exp Psychol Hum Percept Perform 30:1107–1116, 2004) have previously shown, using accuracy measures, that imitation of MF actions is superior to imitation of ML actions when the two action types are presented in separate blocks, and that the advantage of MF over ML items is smaller or absent when they are presented in mixed blocks. We first replicated this finding using an automated reaction time (RT), as well as accuracy, measure. We then examined imitation of MF and ML actions in the mixed condition as a function of the action type presented in the previous trial and in relation to the number of previous test trials. These analyses showed that (1) for both action types, performance was worse immediately after ML than MF trials, and (2) even at the beginning of the mixed condition, responding to MF actions was no better than responding to ML items. These results suggest that the properties of the action stimulus play a substantial role in determining whether imitation is mediated by the direct or the indirect route, and that effects of block composition on imitation need not be generated through strategic switching between routes.     

Is perception of upper body orientation based on the inertia tensor? Normogravity versus microgravity conditions

During lateral leg raising, a synergistic inclination of the supporting leg and trunk in the opposite direction to the leg movement is performed in order to preserve equilibrium. As first hypothesized by Pagano and Turvey (J Exp Psychol Hum Percept Perform, 1995, 21:1070–1087), the perception of limb orientation could be based on the orientation of the limbs inertia tensor. The purpose of this study was thus to explore whether the final upper body orientation (trunk inclination relative to vertical) depends on changes in the trunk inertia tensor. We imposed a loading condition, with total mass of 4 kg added to the subjects trunk in either a symmetrical or asymmetrical configuration. This changed the orientation of the trunk inertia tensor while keeping the total trunk mass constant. In order to separate any effects of the inertia tensor from the effects of gravitational torque, the experiment was carried out in normo- and microgravity. The results indicated that in normogravity the same final upper body orientation was maintained irrespective of the loading condition. In microgravity, regardless of loading conditions the same (but different from the normogravity) orientation of the upper body was achieved through different joint organizations: two joints (the hip and ankle joints of the supporting leg) in the asymmetrical loading condition, and one (hip) in the symmetrical loading condition. In order to determine whether the different orientations of the inertia tensor were perceived during the movement, the interjoint coordination was quantified by performing a principal components analysis (PCA) on the supporting and moving hips and on the supporting ankle joints. It was expected that different loading conditions would modify the principal component of the PCA. In normogravity, asymmetrical loading decreased the coupling between joints, while in microgravity a strong coupling was preserved whatever the loading condition. It was concluded that the trunk inertia tensor did not play a role during the lateral leg raising task because in spite of the absence of gravitational torque the final upper body orientation and the interjoint coupling were not influenced.     

Young and older adults use body-scaled information during a non-confined aperture crossing task

Young and older adults demonstrate differences in action when passing through confined spaces (Warren and Whang in J Exp Psychol Hum Percept Perform 13:371–383, 1987; Hackney and Cinelli in Gait Posture 33:733–736, 2011). However, it is unknown whether or not these differences in actions exist during non-confined multiple obstacle avoidance tasks. The current study aimed to determine: (1) the differences in actions between young and older adults when given a choice in path selection and (2) establish the variables that may account for these differences in action. Older adults (N = 12) and young adults (N = 12) walked along a 10-m path towards a goal and avoided two vertical poles placed halfway down the path on either side of the midline (ranging between 0.6 and 1.8× shoulder width). Results revealed that in non-confined space, both age groups use body-scaled information to determine the passability of apertures and maintain similar Critical Points to those reported in confined aperture crossing (1.4 for young adults and 1.6 for older adults). Variability of the medial–lateral centre of mass movement (i.e. how much the trunk moved side to side) between the groups most likely accounted for the larger aperture sizes (i.e. Critical Points) required by the older adults to pass through the apertures. Therefore, it appears that body-scaled information may include an individual’s knowledge of both actual body size and body sway magnitude.