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Laterally focused attention modulates asymmetric coupling in rhythmic interlimb coordination.

Peters (J Motor Behav 21:151-155, 1989; Interlimb coordination: neural, dynamical and cognitive constraints, Academic, Orlando, pp 595-615, 1994) suggested that expressions of handedness in bimanual coordination may be reflections of an inherent attentional bias. Indeed, previous results indicated that focusing attention on one of the limbs affected the relative phasing between the limbs in a manner comparable to the effects of hand dominance. The present study extended the comparison between the effects of attentional focus and handedness by testing their impact on the interactions between the limbs. Both left-handed and right-handed participants performed rhythmic bimanual coordination tasks (in-phase and antiphase coordination), while directing attention to either limb. Using brief mechanical perturbations, the degree to which the limbs were influenced by each other was determined. The results revealed that the non-dominant limb was more strongly affected by the dominant limb than vice versa and that, in line with Peters' proposition, this handedness-related asymmetry in coupling strength was reduced when attention was focused on the non-dominant limb, thereby highlighting the potential relation between inherent (handedness-related) asymmetries and voluntary attentional asymmetries. In contrast to previous findings, the (commonly observed) phase lead of the dominant limb was attenuated (rather than accrued) when attention was focused on this limb. This unexpected result was explained in terms of the observed attention-related difference in amplitude between the limbs.

Adult↗

A theoretical note on models of interlimb coordination.

This article clarifies some points about models of interlimb coordination. The authors discuss in detail the predictions and extensions of the H. Haken, J.A.S. Kelso, and H. Bunz (1985) model and compare these to a truncated version used by R.C. Schmidt and colleagues. It is demonstrated that procedures commonly used to fit experimental data to model parameters require the fulfillment of certain assumptions. Finally, the authors show how recent experimental findings can be modeled straightforwardly using a system of coupled nonlinear oscillators.

Extremities↗

Relative phase destabilization during interlimb coordination: the disruptive role of kinesthetic afferences induced by passive movement.

The disruption of three patterns of two-limb coordination, involving cyclical flexion-extension movements performed in the same or in different directions, was investigated through application of passive movement to a third limb by the experimenter. The three patterns referred to the homologous, homolateral, and heterolateral (diagonal) limb combinations which were performed in the sagittal plane. The passive movement involved a spatiotemporal trajectory that differed from the movements controlled actively. Even though subjects were instructed to completely ignore the passive limb movement, the findings of experiment 1 demonstrated a moderate to severe destabilization of the two-limb patterns, as revealed by analyses of power spectra, relative phase, cycle duration, and amplitude. This disruption was more pronounced in the homolateral and heterolateral than in the homologous effector combinations, suggesting stronger coupling between homologous than nonhomologous limb pairs. Moreover, passive mobilization affected antiphase (nonisodirectional) movements more than inphase (isodirectional) movements, pointing to the differential stability of these patterns. Experiment 2 focused on homolateral coordination and demonstrated that withdrawal of visual information did not alter the effects induced by passive movement. It was therefore hypothesized that the generation of extra kinesthetic afferences through passive limb motion was primarily responsible for the detriment in interlimb coordination, possibly conflicting with the sensory information accompanying active movement production. In addition, it was demonstrated that the active limbs were more affected by their homologous passive counterpart than by their nonhomologous counterpart, favoring the notion of "specific" interference. The findings are discussed in view of the potential role of kinesthetic afferences in human interlimb coordination, more specifically the preservance of relative phasing through a kinesthetic feedback loop.

Adolescent↗

Example of 2:1 interlimb coordination during fictive rostral scratching in a spinal turtle.

The usual interlimb coordination pattern during fictive rostral scratching in turtles is 1:1 coordination. We describe an example in a turtle of 2:1 coordination during fictive rostral scratching in which there were two cycles of ipsilateral hip flexor activity during each cycle of contralateral hip extensor activity. During 2:1 coordination, there were fluctuations in the ipsilateral hip flexor cycle period such that a larger ipsilateral hip flexor normalized period, which was associated with the onset of a contralateral hip extensor burst, alternated with a smaller ipsilateral hip flexor normalized period, which was associated with the absence of the onset of a contralateral hip extensor burst. These observations support the concept that contralateral circuitry modulates the timing of ipsilateral motor rhythms and therefore contributes to the production of the ipsilateral motor pattern for rostral scratching.

Animals↗

Patterns of human interlimb coordination emerge from the properties of non-linear, limit cycle oscillatory processes: theory and data.

The present article represents an initial attempt to offer a principled solution to a fundamental problem of movement identified by Bernstein (1967), namely, how the degrees of freedom of the motor system are regulated. Conventional views of movement control focus on motor programs or closed-loop devices and have little or nothing to say on this matter. As an appropriate conceptual framework we offer Iberall and his colleagues' physical theory of homeokinetics first elaborated for movement by Kugler, Kelso, and Turvey (1980). Homeo kinetic theory characterizes biological systems as ensembles of non-linear, limit cycle oscillatory processes couple and mutually entrained at all the levels of organization. Patterns of interlimb coordination may be predicted from the properties of non-linear, limit cycle oscillators. In a set of experiments and formal demonstrations we show that cyclical, two-handed movements maintain fixed amplitude and frequency ( a stable limit cycle organization) under the following conditions: (a) when brief and constantly applied load perturbations are imposed on one hand or the other, (b) regardless of the presence or absence of fixed mechanical constraints, and (c) in the face of a range of external driving frequencies from a visual source. In addition, we observe a tight phasic relationship between the hands before and after perturbations (quantified by cross-correlation techniques), a tendency of one limb to entrain the other (mutual entrainment) and that limbs cycling at different frequencies reveal non-arbitrary, sub-harmonic relationships (small integer, subharmonic entrainment). In short, all the above patterns of interlimb coordination fall out of a non-linear oscillatory design. Discussion focuses on the compatibility of these results with past and present neurobiological work, and the theoretical insights into problems of movement offered by homeokinetic physics. Among these are, we think, the beginnings of a principled solution to the degrees of freedom problem, and the tentative claim that coordination and control are emergent consequences of dynamical interaction among non-linear, limit cycle oscillatory processes.

Journal Article↗

Effect of hindlimb unloading on interlimb coordination during treadmill locomotion in the rat.

Effects of hindlimb unloading on interlimb coordination were examined in adult rats walking on a treadmill at moderate speed. In the first group of animals, the electromyographic activity (EMG) of soleus muscle of both hindlimbs was recorded after 7 and 14 days of unloading. In the second group, the EMG was recorded daily until the 14th day of unloading. The general organization of locomotion was preserved in the two groups whatever the duration of the unloading. The step cycles of the two hindlimbs were always strictly alternating. However, the locomotor pattern was very irregular. A lateral instability was observed. It was accompanied by an abduction of the hindlimbs, and frequent hyperextensions of the ankle when walking. The EMG analysis showed an increase in step cycle duration and in coactivation duration of the soleus muscles (i.e. in the double stance duration). In the rats recorded daily, mean EMG was dramatically reduced the 1st day of unloading, suggesting a decrease in the neural drive. Taken together, these data indicate that 14 days of hindlimb unloading can alter the neuromuscular pattern during locomotion. It is proposed that these changes are related to changes in the peripheral sensory information.

Animals↗

Interlimb coordination in 20-day-old rat fetuses.

Evidence for short sequences of interlimb coordination was found in 20-day-old rat fetuses. Frame-by-frame analysis of videotape records showed phase relationships indicating a pattern of alternation in sequences involving forelimb-forelimb and hindlimb-hindlimb coordination. Forelimb-hindlimb coordination was not observed.

Animals↗

A developmental study of the interlimb coordination in running and galloping.

Using a dynamical systems perspective on motor behavior, it was predicted that interlimb coordination of running and galloping would behave like coupled, nonlinear, limit-cycle oscillators, which show the properties of phase locking, entrainment, and structural stability. Female subjects ranging in age from 2.5 years to adult were filmed while running and galloping with and without a weight perturbation. Analysis of both temporal- and amplitude-phasing measures revealed that both gaits demonstrated oscillatory properties. Differences between gaits and across age were primarily a matter of degree. In general, children 4 years of age and below had slightly less table phasing patterns, and all age groups showed slightly less ability in the gallop, particularly with amplitude phasing.

Journal Article↗

Coupling dynamics in interlimb coordination.

In 1:1 frequency locking, the interlimb phase difference phi is an order parameter quantifying the spatial-temporal organization of 2 rhythmic subsystems. Dynamical modeling and experimental analyses indicate that an intentional parameter phi psi (intended coordination mode, phi = 0 degrees or phi = 180 degrees) and 2 control parameters omega c (coupled frequency) and delta omega (difference between uncoupled eigen-frequencies) affect phi. An experiment was conducted on 1:1 frequency locking in which phi psi, omega c, and delta omega were manipulated using a paradigm in which a person swings hand-held pendulums. As delta omega deviated from 0, the observed phi deviated from the phi psi, indicating a displacement in the phi attractor point. The displacements were exaggerated by increasing omega c. The displacements were coordinated with a decrease in the stability of phi and with higher harmonics in power spectrum of phi. Implications of the results for modeling interlimb coordination are discussed.

Adult↗

The development of interlimb coordination during bimanual finger tapping.

Normal subjects aged 7-25 years were asked to tap the index fingers of both hands: a) in four different patterns of interlimb coordination; b) at two different response frequencies; and c) both before and after the entraining metronome was turned off. The outcome variables of primary interest were the within-subject variability of interresponse intervals (IRI) as an index of timing precision; and deviations from prescribed response frequency, as an index of temporal tracking accuracy. Stability of timing precision and accuracy of temporal tracking increased significantly from 7 to 9 and from 9 to 11 years, with only minor advances thereafter. There were significant right-left performance asymmetries in all bimanual tasks; variability of IRI and deviations from prescribed rate were greater at the faster of the two response frequencies tested; and stability of IRI and accuracy of temporal tracking were greater with than without the metronome. Stability of IRI and accuracy of temporal tracking were strongly correlated in some bimanual tasks. The findings are discussed in terms of the two major theoretical perspectives on human brain-behavior relationships that have specifically addressed the issue of bimanual coordination.

Adolescent↗

Propriospinal circuitry underlying interlimb coordination in mammalian quadrupedal locomotion.

Soon after birth, freely moving quadrupeds can express locomotor activity with coordinated forelimb and hindlimb movements. To investigate the neural mechanisms underlying this coordination, we used an isolated spinal cord preparation from neonatal rats. Under bath-applied 5-HT, N-methyl-d,l-aspartate (NMA), and dopamine (DA), the isolated cord generates fictive locomotion in which homolateral cervicolumbar extensor motor bursts occur in phase opposition, as does bursting in homologous (left-right) extensor motoneurons. This coordination corresponded to a walking gait monitored with EMG recordings in the freely behaving animal. Functional decoupling of the cervical and lumbar generators in vitro by sucrose blockade at the thoracic cord level revealed independent rhythmogenic capabilities with similar cycle frequencies in the two locomotor regions. When the cord was partitioned at different thoracic levels and 5-HT/NMA/DA was applied to the more caudal compartment, the ability of the lumbar generators to drive their cervical counterparts increased with the proportion of chemically exposed thoracic segments. Blockade of synaptic inhibition at the lumbar level caused synchronous bilateral lumbar rhythmicity that, surprisingly, also was able to impose bilaterally synchronous bursting at the unblocked cervical level. Furthermore, after a midsagittal section from spinal segments C1 to T7, and during additional blockade of cervical synaptic inhibition, the cord exposed to 5-HT/NMA/DA continued to produce a coordinated fictive walking pattern similar to that observed in control. Thus, in the newborn rat, a caudorostral propriospinal excitability gradient appears to mediate interlimb coordination, which relies more on asymmetric axial connectivity (both excitatory and inhibitory) between the lumbar and cervical generators than on differences in their inherent rhythmogenic capacities.

Animals↗

Models of interlimb coordination--equilibria, local analyses, and spectral patterning: comment on Fuchs and Kelso (1994).

This article is a comment on A. Fuchs and J. A. S. Kelso's (1994) theoretical note on models of interlimb coordination. The generality of the order parameter equation derived by H. Haken, J. A. S. Kelso, and H. Bunz (HKB; 1985) for correlated limb movements is underscored through the correct and nonintuitive predictions it makes about steady state behavior. Local and global dynamical models are contrasted, and experimental situations in which local models are pragmatic alternatives to global models, like the HKB equation, are described. Questions are raised about the basis for interpreting detuning in HKB as a simple frequency difference and about the ability of the system of coupled nonlinear oscillators proposed by A. Fuchs and J. A. S. Kelso (1994) to model fully the experimental findings on the spectrum of relative phase.

Extremities↗

Central generation of grooming motor patterns and interlimb coordination in locusts.

Coordinated bursts of leg motoneuron activity were evoked in locusts with deefferented legs by tactile stimulation of sites that evoke grooming behavior. This suggests that insect thoracic ganglia contain central pattern generators for directed leg movements. Motoneuron recordings were made from metathoracic and mesothoracic nerves, after eliminating all leg motor innervation, as well as all input from the brain, subesophageal ganglion, and prothoracic ganglion. Strong, brief trochanteral levator motoneuron bursts occurred, together with silence of the slow and fast trochanteral depressor motoneurons and activation of the common inhibitor motoneuron. The metathoracic slow tibial extensor motoneuron was active in a pattern distinct from its activity during walking or during rhythms evoked by the muscarinic agonist pilocarpine. Preparations in which the metathoracic ganglion was isolated from all other ganglia could still produce fictive motor patterns in response to tactile stimulation of metathoracic locations. Bursts of trochanteral levator and depressor motoneurons were clearly coordinated between the left and right metathoracic hemiganglia and also between the mesothoracic and the ipsilateral metathoracic ganglia. These data provide clear evidence for centrally generated interlimb coordination in an insect.

Animals↗

Interlimb coordination in patients with Parkinson's disease: motor learning deficits and the importance of augmented information feedback.

The basal ganglia have traditionally been associated with motor control functions and this view has prevailed since the late nineteenth century. Recent experimental studies suggest that this neuroanatomical system is also critically involved in motor learning. In the present study, motor learning/transfer capabilities were compared between patients with Parkinson's disease and a group of normal elderly people. Subjects practiced a bimanual coordination task that required continuous flexion-extension movements in the transverse plane with a 90 degrees phase offset between the forearms. During acquisition, augmented visual feedback of the relative motions was provided in real time. The findings revealed improvements in the bimanual coordination pattern across practice in both groups when the augmented concurrent feedback was present. However, when transferred to performance conditions in which the augmented information was withheld, performance deteriorated (relative to the augmented condition) and this effect was more prevalent in the Parkinson patients. More specifically, no improvement in interlimb coordination was observed under nonaugmented feedback conditions across practice. Instead, a drift toward the preferred in-phase and antiphase coordination patterns was evident. The present findings suggest that Parkinson patients can improve their performance on a new motor task, but they remain strongly dependent on augmented visual information to guide these newly acquired movements. The apparent adoption of a closed-loop control mode is accompanied with decreases in movement speed in order to use the feedback to ensure accuracy. When the augmented feedback is withheld and the movement pattern is to be controlled by means of intrinsic information feedback sources, performance is severely hampered. The findings are hypothesized to indicate that learning/transfer is affected in Parkinson patients who apparently prefer some constancy in the environmental contingencies under which practice takes place. The present findings are consistent with the notion that the basal ganglia form a critical neuroanatomical substrate for motor learning.

Aged↗

The effect of postural stability and spatial orientation of the upper limbs on interlimb coordination.

It has recently been reported that the spatial orientation of two moving limbs has a determining influence on the relative accuracy and stability of coordination patterns. The purpose of the present experiments was to test perceptual and neuromuscular explanations of these spatial orientation effects. Experiment 1 was an initial test of the hypotheses and an extension of a previous study [Lee et al. (2002) Exp Brain Res 146:205-212] that required participants to coordinate inphase and antiphase movement patterns in four spatial orientations: two symmetric orientations (90 degrees and 180 degrees separation between the limbs) and two asymmetric orientations (90 degrees and 135 degrees separation between the limbs). Results of Experiment 1 suggest that the symmetry of movement may be a key factor influencing spatial orientation effects observed during interlimb coordination. In Experiment 2, participants again performed inphase and antiphase movement patterns in symmetric and asymmetric spatial orientations. However, one-half of the participants in Experiment 2 were provided with mechanical constraints during the performance of the desired coordination patterns. The mechanical constraints provided postural support but did not influence the visual experience. Results showed that the addition of the postural support improved performance. These findings suggest that neuromuscular, and perhaps biomechanical, constraints contribute more to the influence of spatial orientation than visual-perceptual constraints.

Adult↗

Interlimb coordination deficits in patients with Parkinson's disease during the production of two-joint oscillations in the sagittal plane.

Two-limb coordination patterns involving cyclical flexion-extension movements, performed in the same or in different directions, were studied in patients with Parkinson's disease and a group of elderly subjects. The three patterns referred to the homologous (both arms or legs), homolateral (right or left arm and leg), and heterolateral (right arm and left leg or vice versa) limb segment combinations that were performed in the sagittal plane from a seated position. Findings revealed that interlimb coordination deficits were evident in patients with Parkinson's disease. Moreover, mean cycle duration and its variability were increased, particularly during the production of nonhomologous limb movements in different directions. These temporal findings suggest that movement slowness was not a primary consequence of an intrinsic inability to move the limb segments at the required speed but rather reflected an intentional strategy to cope with the complexity of the coordination pattern. Finally, movement amplitude was substantially smaller and more variable in patients with Parkinson's disease, suggestive of hypometria during the production of these cyclical tasks.

Aged↗

Dynamics of effortful touch and interlimb coordination.

In biological movement systems, the level of muscular-articular links is responsible for assembling highly reliable, stable, and reproducible coordination patterns involving very many joints and muscles. Research shows that the important perceptual capabilities of this level arise from the bulk sensitivity of muscles and tendons, so-called effortful or dynamic touch, to the quantities of rotational dynamics that remain invariant (such as the inertia tensor) over variations in rotational forces and motions. The power laws characterizing this sensitivity point to underlying fractal (self-similar) processes. Other research shows that the hallmark ability of this level to produce repetitive interlimb coordinations can be addressed through a dynamics of coordination in which equations express the time-evolution of collective neuromuscular states. This research also suggests that the assembled rhythms exploit the unique blend of stability and variability characteristic of low-dimensional chaotic motion on strange attractors. In overview, research into the capabilities of the level of muscular-articular links highlights the importance of applying classical and modern (nonlinear) dynamics to understanding the assembly and perceptual control of biological movements.

Extremities↗

Average phase difference theory and 1:1 phase entrainment in interlimb coordination.

The dynamics of coupled biological oscillators can be modeled by averaging the effects of coupling over each oscillatory cycle so that the coupling depends on the phase difference phi between the two oscillators and not on their specific states. Average phase difference theory claims that mode locking phenomena can be predicted by the average effects of the coupling influences. As a starting point for both empirical and theoretical investigations, Rand et al. (1988) have proposed d phi/dt = delta omega--K sin phi, with phase-locked solutions phi = arcsin(delta omega/K), where delta omega is the difference between the uncoupled frequencies and K is the coupling strength. Phase-locking was evaluated in three experiments using an interlimb coordination paradigm in which a person oscillates hand-held pendulums. Delta omega was controlled through length differences in the left and right pendulums. The coupled frequency omega c was varied by a metronome, and scaled to the eigenfrequency omega v of the coupled system; K was assumed to vary inversely with omega c. The results indicate that: (1) delta omega and K contribute multiplicatively to phi; (2) phi = 0 or phi = pi regardless of K when delta omega = 0; (3) phi approximately 0 or phi approximately pi regardless of delta omega when K is large (relative to delta omega); (4) results (1) to (3) hold identically for both in phase and antiphase coordination. The results also indicate that the relevant frequency is omega c/omega v rather than omega c.(ABSTRACT TRUNCATED AT 250 WORDS)

Adult↗