The influence of perceptual and motor factors on bimanual coordination in a polyrhythmic tapping task

Summary In this study the role of perceptual and motor factors on the motor organization (integrated versus parallel) adopted by musically skilled and unskilled subjects in a polyrhythmic tapping task was investigated. Subjects tapped a 3:2 polyrhythm to match the timing of two isochronous tone trains, one tone train for each hand. Perceptual factors were examined by the manipulation of the frequency difference between the tone trains to produce either an integrated or a streamed percept. Motor factors were examined by comparison of performance on two versions of the 3:2 polyrhythm. In one (simultaneous) version, each cycle of the polyrhythm began with a simultaneous left- and right-hand tap. In the other (shifted) version a 100-ms interval was introduced between the initial left and right taps in each cycle. Examination of the pattern of variances and covariances among intertap intervals suggested that most of the subjects in this study adopted an integrated motor organization that involved interleaving the timing of the two hands. Further analysis revealed that a serial chained model described the pattern of covariances best for the simultaneous pattern, whereas a hierarchical organization described the pattern of covariances for the shifted pattern best. The finding that performance was more accurate with integrated tones than with streamed tones provides some support for a perceptual-motor facilitation hypothesis.     

Expanding the tactual field of view

Can tactual information be acquired simultaneously by several different fingers? Blind and sighted Ss were asked to scan vertical displays of braille (consisting of either one or two dots) with the index and middle fingers on each hand-using one, two, or four fingers at the same time. Stimuli were recognized most rapidly when the displays were scanned by two fingers on different hands and least rapidly when two fingers on the same hand were used; performance was similar with one finger and with four fingers. The results indicated some parallel perceptual processing of the inputs to the two hands and mutual interference in processing inputs from fingers on the same hand.     

Response preparation with static versus moving hands

This research tested the response inhibition account of the hand-advantage found in the finger precuing task. According to this account, the advantage of preparing two fingers on one hand (represented in one hemisphere) as opposed to preparing two fingers on two hands (represented in two hemispheres) is due, in part, to a response inhibition process that operates more efficiently within than between hemispheres. In this view, supplying extra activation to both hemispheres by moving the hands should decrease the within-hemisphere inhibition advantage. Twelve participants performed the finger precuing task with static and moving hands. As predicted by the response inhibition account, the hand-advantage, present with the hands at rest, decreased with the hands moving.     

Response preparation with static versus moving hands

This research tested the response inhibition account of the hand-advantage found in the finger pre-cuing task. According to this account, the advantage of preparing two fingers on one hand (represented in one hemisphere) as opposed to preparing two fingers on two hands (represented in two hemispheres) is due, in part, to a response inhibition process that operates more efficiently within than between hemispheres. In this view, supplying extra activation to both hemispheres by moving the hands should decrease the within-hemisphere inhibition advantage. Twelve participants performed the finger pre-cuing task with static and moving hands. As predicted by the response inhibition account, the hand-advantage, present with the hands at rest, decreased with the hands moving.     

Difference of motor overflow depending on the impaired or unimpaired hand in stroke patients

The aim of this study was to investigate the patterns of contralateral motor overflow (i.e. mirror movement) between the homologous body parts on the right and left side, in stroke patients during single-finger and multi-finger maximum force production tasks. Forty subjects, including stroke (n = 20) and normal subjects (n = 20), participated in this study. The stroke subjects maximally pressed force sensors with their fingers in a flexed position using a single (index, middle, ring, or little) or all fingers (all 4 fingers) using the impaired (IH) or unimpaired (UIH) hand, while the non-patient subjects used their right hands for the same tasks. The maximal voluntary forces in the ipsilateral and unintended pressing forces of each contralateral finger were recorded during the tasks. The magnitude of motor overflow to the contralateral side was calculated using the index of contralateral independence (CI). During the single finger tasks, the finger CI was significantly decreased in the UIH (91%) compared with that in the IH (99%) or normal hands (99%). Likewise, the multiple finger tasks showed that the CI was significantly lower in the UIH (84%) compared with that in the IH (96%) or normal hands (99%). However, the maximal forces were significantly lower in the IH relative to those in the UIH and normal hands. These data demonstrate that stroke patients have greater motor overflow from the UIH to the IH.     

Response-repetition effects depend on motor set: Evidence for anatomical coding in response selection

Successful motor performance requires a process of response selection that chooses the correct response out of a set of possible ones. Most theories of response selection assume that this selection process operates on spatial codes, which define the location of stimuli and responses in environmental coordinates, with little or no role for the anatomical codes of the effectors involved. In this study, we tested this assumption by investigating response-repetition effects in a response-cuing paradigm using two motor sets (fingers on one hand vs. fingers on two hands). Reaction time results demonstrated a robust response-repetition benefit that was greater for the one-hand set than for the two-hands set. Furthermore, with the one-hand set the repetition benefit was independent of cue type and cue-stimulus interval on the previous trial, whereas with the two-hands set it was strongly modulated by these two factors. These differential response-repetition effects for one- and two-hands motor sets demonstrate the important role of the neuro-anatomical hand distinction in response selection, thereby supporting multiple coding notions.     

Visual and tactile action effects determine bimanual coordination performance

Effect-based models of motor control assign a crucial role to anticipated perceptual feedback in action planning. Two experiments were conducted to test the validity of this proposal for discrete bimanual key press responses. The results revealed that the normally observed performance advantage for the preparation of two responses with homologous rather than non-homologous fingers becomes inverted when homologous fingers produce non-identical visual effects, and non-homologous fingers produce identical visual effects. In the second experiment the finger homology effect was strongly reduced when homologous fingers produced non-identical tactile feedback. The results show that representations of to-be-produced visual and tactile action effects both contribute to action planning, though possibly to a varying degree. Implications of these results for effect-based models of motor control are considered.     

Do perception and motor production share common timing mechanisms: A correlational analysis

The two experiments of this study exploited individual variation in timing ability to ask whether the production of time intervals by different motor effectors and the judgement of perceptually based time intervals all share common timing mechanisms. In one task subjects produced a series of taps, attempting to maintain constant intervals between them. Individual differences in variability of the produced intervals correlated across the effectors of finger and foot. That is, people that were ‘good timers’ with one effector tended to be ‘good timers’ with another. Besides timing motor production, the subjects also judged durations of brief perceptual events. The acuity of perceptual judgements correlate substantially with regularity of motor production. Further results involving maximum speed of motor production suggested that variability of motor timing comes from two sources, one source in common with perception, and hence called clock variability, and the other source in common with motor speed, and hence called motor implementation variability. The second experiment showed that people high in skill on the piano were better at both types of timing on the average than control subjects with no expertise.     

On marching to two different drummers: perceptual aspects of the difficulties

People have remarkable difficulty generating two responses that must follow different temporal sequences, unless the temporal patterns are simply related (e.g., periods in 2:1, 3:1 relation). For example, it is hard to tap to two conflicting rhythms presented concurrently (i.e., a polyrhythm) using the right and left hands (Klapp, 1979), or to tap while articulating a conflicting speech utterance (Klapp, 1981). The present experiments indicate that difficulties in processing conflicting rhythms occur even when people must (a) merely monitor the stimuli and indicate the termination of one rhythmic sequence or (b) tap with a single hand. Responding to polyrhythms is thus difficult even without multiple limb coordination. Furthermore, the difficulty of two-handed tapping to polyrhythms that involve two different tones was found to decrease as the pitch difference between the tones was decreased. This result indicates that the difficulty of rhythmic coordination can be perceptually manipulated in a striking fashion. Polyrhythmic performance thus provides an excellent opportunity for examining possible interactions of perceptual and motor organizations.     

Hierarchical organisation of neuro-anatomical constraints in interlimb coordination

Based on the observation that bimanual finger tapping movements tend toward mirror symmetry with respect to the body midline, despite the synchronous activation of non-homologous muscles, F. Mechsner, D. Kerzel, G. Knoblich, and W. Prinz (2001) [Perceptual basis of bimanual coordination. Nature, 414, 69-73] suggested that the basis of rhythmic coordination is purely spatial/perceptual in nature, and independent of the neuro-anatomical constraints of the motor system. To investigate this issue further, we employed a four finger tapping task similar to that used by F. Mechsner and G. Knoblich (2004) [Do muscle matter in bimanual coordination? Journal of Experimental Psychology: Human Perception and Performance, 30, 490-503] in which six male participants were required to alternately tap combinations of adjacent pairs of index (I), middle (M) and ring (R) fingers of each hand in time with an auditory metronome. The metronome pace increased continuously from 1 Hz to 3 Hz over the course of a 30-s trial. Each participant performed three blocks of trials in which finger combination for each hand (IM or MR) and mode of coordination (mirror or parallel) were presented in random order. Within each block, the right hand was placed in one of three orientations; prone, neutral and supine. The order of blocks was counterbalanced across the six participants. The left hand maintained a prone position throughout the experiment. On the basis of discrete relative phase analyses between synchronised taps, the time at which the initial mode of coordination was lost was determined for each trial. When the right hand was prone, transitions occurred only from parallel symmetry to mirror symmetry, regardless of finger combination. In contrast, when the right hand was supine, transitions occurred only from mirror symmetry to parallel but no transitions were observed in the opposite direction. In the right hand neutral condition, mirror and parallel symmetry are insufficient to describe the modes of coordination since the hands are oriented orthogonally. When defined anatomically, however, the results in each of the three right hand orientations are consistent. That is, synchronisation of finger tapping is determined by a hierarchy of control of individual fingers based on their intrinsic neuro-mechanical properties rather than on the basis of their spatial orientation.     

Tactile localization

Clinical studies suggested that errors of tactile localization made by leucotomized subjects were due to a failure of analysis of a conceptual type.

The present experiment examines the normal mechanisms of localizing stimuli to different sites on the hands by measuring the time taken to make a correct response as well as by an analysis of the errors made under conditions of stress (the reaction-time situation). It is demonstrated that healthy subjects of above-average intelligence have appreciable difficulty in determining which of their fingers has been touched. The degree of this difficulty depends on which finger has been stimulated. The results strongly support the hypothesis that normal tactile localization involves the analysis of primary sensory data in terms of a few simple parameters of orientation. It is suggested that some of the errors which occur are due to contamination between different systems of conceptual analysis.     

A priming method for investigating the selection of motor responses

We describe a priming method for investigating the mechanisms underlying the selection of motor responses. The empirical question addressed with the method is how the choice reaction time for a response depends on its relationship to a response that the subject was primed to perform. We explore the method in a study of manual response selection where we investigate the effects of requiring that two possible responses use the same finger or hand. A requirement of the method — that subjects get ready to perform primed responses only — is not met in some conditions of the experiment. When the two possible responses are made with different hands, it appears that multiple response preparation occurs prior to detection of the reaction signal, whereas when the two possible responses are made with different fingers of the same hand it appears that advance preparation is limited to a single response. This finding implies that subjects engage in different kinds of response preparation depending on the relationship between the alternative possible responses. We discuss the implications of this hypothesis for the priming method introduced here as well as for theories of response selection generally.     

Characteristics of finger force production during one- and two-hand tasks

Relations among finger forces were studied during one-hand and two-hand isometric maximal force production tasks in right- and left-handers. We particularly focused on the phenomena of force deficit during one-hand multi-finger tasks and of bilateral force deficit during two-hand tasks. Ten healthy subjects (five of them left-handed) performed maximal voluntary force production tasks with different finger combinations involving fingers of one of the hands or of both hands together. In one-hand tasks, finger enslaving (forces produced by fingers that were not instructed to produce force) was larger in the dominant hand, while force deficit (drop in individual finger peak force during multi-finger tasks) showed no differences between the hands. An additional drop in finger forces was seen in two-hand tests (bilateral deficit). The magnitude of the bilateral deficit for a hand was larger for tasks involving fewer fingers within the hand and more fingers in the other hand, with a ceiling effect. Smaller bilateral deficit was seen in tasks involving symmetrical finger combinations. In two-hand tasks that could potentially lead to the generation of large total moments in the frontal plane, the hand that was expected to generate larger moments showed larger bilateral deficit, so that the magnitude of the total moment was reduced. These observations suggest that force deficit within a hand and bilateral deficit have different origins but their effects are combined at a certain level of the multi-finger control hierarchy. Bilateral deficit may display task dependence reflecting, in particular, the principle of minimization of secondary moments. A double-representation, mirror-image hypothesis is suggested to provide a neurophysiological basis for the observed patterns of bilateral deficit.     

Stimulus-dependent modulation of perceptual and motor learning in a serial reaction time task

In two experiments, we investigated the impact of spatial attributes on the representation acquired during a serial reaction time task. Two sequences were used, in which structural regularities occurred either in the horizontal or in the vertical locations of successive stimuli. After training with the dominant hand, participants were required to respond with the non-dominant hand to either the original sequence or to a mirror-ordered version of the original sequence that required finger movements homologous to those used during training. We observed that a difference in reaction times between the two transfer conditions was smaller in the vertical sequence than in the horizontal sequence. This pattern of results was independent of whether three fingers (Experiment 1) were used or only one finger (Experiment 2) was used for responding. This result suggests that perceptual and motor learning mechanisms may be weighted differently depending on the context in which the stimulus is presented.     

The influence of response grouping on free-choice decision making in a response selection task

Previous research has demonstrated an advantage for the preparation of fingers on one hand over the preparation of fingers on two hands, and for the preparation of homologous fingers over that of non-homologous fingers. In the present study, we extended the precuing effects observed with finger responses to response selection under free-choice conditions. Participants were required to choose from a range of possible responses following the presentation of a precue that indicated which response to prepare (go-to precue) or prevent (no-go-to precue). In Experiment 1 the choice was between homologous and non-homologous finger responses on the hand opposite to the precue while in Experiment 2 the choice was between finger responses on the same or different hand to the precue. In the go-to precue condition, the frequency of homologous finger choices was more frequent than non-homologous finger responses. Similarly, participants chose finger responses on the same hand as the precue regardless of whether they were instructed to prepare or prevent the precued response. The hand effect bias was stronger than the finger effect bias. These findings are consistent with the Grouping Model (Adam, Hommel, & Umilta, 2003).     

Manipulating response set in a choice reaction task: evidence for anatomical coding in response selection

Accurate motor performance requires a process of response selection that chooses the correct response out of a set of possible ones. Most theories of response selection assume that this selection process operates on spatial codes, which define the location of stimuli and responses in environmental coordinates, with little or no role for the anatomical codes of the effectors involved. In this study, I systematically manipulated the anatomical identities of the response fingers in a four-choice reaction task, while keeping the stimulus and response locations constant. Results showed that homologous fingers on different hands modulated the reaction time profiles and error patterns in a mirror-symmetrical way. This finding indicates that anatomical factors may play a more substantial role in response selection than often assumed.     

The relationship of target and response complexity in coincident timing performance

This paper reports two experiments concerned with the interaction of response and target factors in a coincident timing situation. Coincident timing involves executing a response to intercept a moving target. The target response relationship was manipulated in the first experiment by providing some targets with no structure (linear path) and others containing 1 and 2 bounce points. In the second experiment, responses required 0, 1, or 2 reversals in direction. The results provide tentative support for the notion that structure in the target path could be linked to the key temporal response dynamics for improved performance consistency, and that subjects may alter their response structure to incorporate this linkage. This can be further extended to the advantage of simply moving while perceiving movement. This paper stresses the importance of examining the perceptual and motor requirements of the coincident timing skill in unison.     

Ideomotor effects of pitch on continuation tapping

The ideomotor principle predicts that perception will modulate action where overlap exists between perceptual and motor representations of action. This effect is demonstrated with auditory stimuli. Previous perceptual evidence suggests that pitch contour and pitch distance in tone sequences may elicit tonal motion effects consistent with listeners' implicit awareness of the lawful dynamics of locomotive bodies. To examine modulating effects of perception on action, participants in a continuation tapping task produced a steady tempo. Auditory tones were triggered by each tap. Pitch contour randomly and persistently varied within trials. Pitch distance between successive tones varied between trials. Although participants were instructed to ignore them, tones systematically affected finger dynamics and timing. Where pitch contour implied positive acceleration, the following tap and the intertap interval (ITI) that it completed were faster. Where pitch contour implied negative acceleration, the following tap and the ITI that it completed were slower. Tempo was faster with greater pitch distance. Musical training did not predict the magnitude of these effects. There were no generalized effects on timing variability. Pitch contour findings demonstrate how tonal motion may elicit the spontaneous production of accents found in expressive music performance.     

The Relationship of Target and Response Complexity in Coincident Timing Performance

This paper reports two experiments concerned with the interaction of response and target factors in a coincident timing situation. Coincident timing involves executing a response to intercept a moving target. The target response relationship was manipulated in the first experiment by providing some targets with no structure (linear path) and others containing 1 and 2 bounce points. In the second experiment, responses required 0, 1, or 2 reversals in direction. The results provide tentative support for the notion that structure in the target path could be linked to the key temporal response dynamics for improved performance consistency, and that subjects may alter their response structure to incorporate this linkage. This can be further extended to the advantage of simply moving while perceiving movement. This paper stresses the importance of examining the perceptual and motor requirements of the coincident timing skill in unison.     

Forgetting motor programmes: Retrieval dynamics in procedural memory

When motor sequences are stored in memory in a categorised manner, selective retrieval of some sequences can induce forgetting of the non-retrieved sequences. We show that such retrieval-induced forgetting (RIF) occurs not only in cued recall but also in a test assessing memory indirectly by providing novel test cues without involving recall of items. Participants learned several sequential finger movements (SFMs), each consisting of the movement of two fingers of either the left or the right hand. Subsequently, they performed retrieval practice on half of the sequences of one hand. A final task then required participants to enter letter dyads. A subset of these dyads corresponded to the previously learned sequences. RIF was present in the response times during the entering of the dyads. The finding of RIF in the slowed-down execution of motor programmes overlapping with initially trained motor sequences suggests that inhibition resolved interference between procedural representations of the acquired motor sequences of one hand during retrieval practice.