Developmental emergence of self-referential and inhibition mechanisms of body movements underling felicitous behaviors

In young infants, activation or inhibition of body movements on perception of environmental events is important to enable them to act on the world or understand the world. To reveal the development of this ability, we observed movement patterns in all four limbs under the two experimental conditions. Infants assigned to the interaction condition were provided a connection between their arm and a toy, which allowed movements of the critical arm to produce movements in the overhead mobile. Infants assigned to the stimulation condition were presented with similar movements of the toy emulated by the experimenter. In 3-month-old infants, the arm movements increased under the interaction condition, whereas they decreased under the stimulation condition. Such condition-dependent behavior dissociation was not observed in 2-month-old infants. These results suggest that the distinct behavior of an "observer" of environmental events was differentiated from that of a "player" interacting with the surroundings between 2 and 3 months of age. The emergence of differentiation of behaviors points toward the development of self-referential and inhibition mechanisms in the perceptual-motor and attention/evaluation system in the cortico-subcortical network.     

Toy-oriented changes in hand and joint kinematics during the emergence of purposeful reaching

Infants first consistently reach for objects between 3 and 5 months of age. In the months before reaching, infants produce a variety of arm movements. The relationship between these early arm movements and the emergence of purposeful reaching is still unclear. The purpose of the present study was to determine how groups of non-reaching, nearly reaching, and newly reaching infants changed the kinematics of their spontaneous arm movements in the presence of a toy. Five infants in each of these groups were observed with a high-speed motion capture system during trials with and without a toy present. Kinematic analyses examined 3D hand, shoulder, and elbow motions. Our results suggest that with a toy present, non-reachers altered their movement quantity whereas near- and new-reachers altered their movement quality through spatio-temporal dissociation and reorientation of the arm. When comparing the changes across groups we observed three preliminary patterns of toy-oriented changes. Our results join other studies to strengthen the relationship between early arm movements and purposeful reaching. Future longitudinal studies are now required to begin to fully understand the complex process by which infants adapt their early arm movements for purposeful behaviors.     

Matching of Movements Made Independently by the Two Arms in Normal Humans

Comparisons were made of voluntary movements of the right and left arms in normal human subjects. A series of movements of different amplitudes, made at the subject’s own speed, was performed with one limb. After a rest period, the same series was repeated with the contralateral limb. The relation between movement peak velocity and movement amplitude was linear and was the same for both arms. With repeated testing over periods up to two months, the slope of the peak velocity—amplitude relation decreased during the first week, thereafter remaining unchanged. In a second series of experiments, six normal subjects continuously wore a 1 lb (0.45 kg) weight strapped to their left (non-dominant) forearm for up to 1 week. This resulted in an increase in the slope of the peak-velocity/amplitude relation in this arm. A parallel change occurred in movements made independently by the right (non-loaded) arm. A similar matching of movement performance of the two limbs was seen following removal of the weight. The data is interpreted as providing support for the hypothesis that there is a single movement “command” which is applied to both limbs. The interaction of this command with the limbs which have similar second-order mechanical properties yields similar movements even when they are made independently.     

The coordination of limb movements with different kinematic patterns

The principles underlying the coordination of limb movements with different spatiotemporal features were explored. After an initial training session in which the same unidirectional movement had to be performed with both upper limbs, subjects attempted to coordinate two different movements in a second session, i.e., the learned unidirectional movement in the left limb and a new double reversal movement in the right limb. The findings uncovered a wide variety in patterns of interlimb dependence among and within subjects, going from a high degree of dependence to relative independence. The relationship between limbs was studied by means of a detailed analysis of the displacement and acceleration patterns and the electromyographic activity of the major muscles involved. The general underlying principle that appeared to account for the diversity in movement organization was this: higher independence between limb movements is achieved when subjects initiate the movements to be coordinated successively. This asynchrony in movement onset can possibly be viewed as an attempt to safeguard against interference.     

Interlimb coordination following stroke

Studies investigating whether simultaneous bilateral movements can facilitate performance of the impaired limb(s) of stroke patients have returned mixed results. In the present study we compared unilateral limb performance (amplitude, cycle duration) with performance during an interlimb coordination task involving both homologous (both arms, both legs) and non-homologous (one arm, one leg) limbs in stroke participants (n=7) and healthy age-matched controls (n=7). In addition, the effect of on-line augmented visual feedback on interlimb coordination was investigated. Participants performed cyclical flexion-extension movements of the arms and legs in the sagittal plane paced by an auditory metronome (1 Hz). Movement amplitudes were larger and cycle durations shorter during homologous limb coordination than non-homologous coordination. Compared with unilateral movements both groups had reduced movement amplitudes and the stroke group increased cycle duration when interlimb coordination tasks were performed. These effects were most evident during non-homologous (arm and leg) coordination. No evidence of facilitation of the impaired limb(s) was found in any of the interlimb coordination conditions. Augmented visual feedback had minimal effect on the movements of control participants but lead to an increase of cycle duration for stroke participants.     

Matching of movements made independently by the two arms in normal humans

Comparisons were made of voluntary movements of the right and left arms in normal human subjects. A series of movements of different amplitudes, made at the subject' own speed, was performed with one limb. After a rest period, the same series was repeated with the contralateral limb. The relation between movement peak velocity and movement amplitude was linear and was the same for both arms. With repeated testing over periods up to two months, the slope of the peak velocity-amplitude relation decreased during the first week, thereafter remaining unchanged. In a second series of experiments, six normal subjects continuously wore a 1 lb (0.45 kg) weight strapped to their left (non-dominant) forearm for up to 1 week. This resulted in an increase in the slope of the peak-velocity/amplitude relation in this arm. A parallel change occurred in movements made independently by the right (non-loaded) arm. A similar matching of movement performance of the two limbs was seen following removal of the weight. The data is interpreted as proving support for the hypothesis that there is a single movement "command" which is applied to both limbs. The interaction of this command with the limbs which have similar second-order mechanical properties yields similar movements even when they are made independently.     

Additional weight influences the reaching behavior of low-risk preterm infants

The authors investigated the influence of additional weight in the reaching behavior of preterm infants. Nine low-risk preterm and 10 full-term infants age 5-7 months participated. A toy was presented in two procedures: (a) baseline and (b) additional weight (bracelet with weight of 20% of the infant's upper limb mass). The additional weight reduced the straightness index at 5 months, increased the mean velocity and decreased the movement units at all months, and increased unsuccessful grasps at 5 and 7 months in preterm infants. Preterm infants showed less movement units at all months and less unsuccessful grasps at 6 months compared with full-term infants. Weight disturbance seems to change the intrinsic dynamics involved in reaching movements of 5-7-month-old preterm infants.     

Kinetic attraction during bimanual coordination

Two experiments examined the effects of independent variations in kinetic and kinematic requirements on interlimb coupling during a bimanual task. The goal of the investigation was to provide preliminary evidence regarding one general class of physical variables that constrains discrete bimanual movements. Subjects attempted to execute a smooth unidirectional movement with the left arm, along with a three-segment reversal movement with the right arm. The first experiment manipulated the torque required to produce the reversal action, while movement duration and average angular velocity were held constant for both limbs. Several indications of increased interlimb coupling, due to the kinetic variation, were evident. The converse manipulation was used in the second experiment, with movement time and kinematics (velocity, acceleration) changed independently of joint torque requirements for the reversal limb. No clear effect of kinematics on coupling strength was noted. The results suggest that one variable influencing interlimb attraction toward common spatiotemporal trajectories may be kinetic in nature.     

Kinetic Attraction During Bimanual Coordination

Two experiments examined the effects of independent variations in kinetic and kinematic requirements on interlimb coupling during a bimanual task. The goal of the investigation was to provide preliminary evidence regarding one general class of physical variables that constrains discrete bimanual movements. Subjects attempted to execute a smooth unidirectional movement with the left arm, along with a three-segment reversal movement with the right arm. The first experiment manipulated the torque required to produce the reversal action, while movement duration and average angular velocity were held constant for both limbs. Several indications of increased interlimb coupling, due to the kinetic variation, were evident. The converse manipulation was used in the second experiment, with movement time and kinematics (velocity, acceleration) changed independently of joint torque requirements for the reversal limb. No clear effect of kinematics on coupling strength was noted. The results suggest that one variable influencing interlimb attraction toward common spatiotemporal trajectories may be kinetic in nature.     

Toy-oriented changes during early arm movements IV: shoulder-elbow coordination

Our recent work on the initial emergence of reaching identified a mosaic of developmental changes and consistencies within the hand and joint kinematics of arm movements across the pre-reaching period. The purpose of this study was to test hypotheses regarding the coordination of hand and joint kinematics over this same pre-reaching period. Principal component analysis (PCA) was conducted on hand, shoulder, and elbow kinematic data from 15 full-term infants observed biweekly from 8 weeks of age through the week of reach onset. Separate PCAs were calculated for spatial variables and for velocity variables in trials with a toy and without a toy. From the PCA results, we constructed ‘variance profiles’ to reflect the coordinative structure of the hand, shoulder, and elbow. By coordinative structure is meant here the relative contribution of each joint to the factors revealed by the PCA. Shifts in these profiles, which reflected coordination changes, were compared across the hand and joints within each pre-reaching phase (Early, Mid, Late) as well as across phases and trial conditions (no-toy and toy). Results identified both surprising consistencies and important developmental changes in coordination. First, over development, spatial coordination changed in different ways for the shoulder and elbow. Between the Early and Late phases, spatial coordination at the shoulder showed more adult-like coordination during both spontaneous movements and movements with a toy present. In contrast, elbow spatial coordination became more adult-like only during movements with a toy and less adult-like during spontaneous movements. Second, over development, velocity coordination became more adult-like at both joints in movements with and without a toy present. We propose that the features of coordination that changed over development suggest explanations for the differential roles and developmental trajectories of the control of arm movements between the shoulder and elbow. We propose that features that remained consistent over development suggest the presence of developmentally important constraints inherent in arm biomechanics, which may simplify arm control for reaching. Taken together, these findings highlight the critical role of spontaneous arm movements in the emergence of purposeful reaching.     

Effects of velocity and limb loading on the coordination between limb movements during walking

The authors investigated the effects of velocity (increasing from 0.5 to 5.0 km/hr in steps of 0.5 km/hr) and limb loading on the coordination between arm and leg movements during treadmill walking in 7 participants. Both the consistency of the individual limb movements and the stability of their coordination increased with increasing velocity; the frequency coordination between arm and leg movements was 2:1 at the lower velocities and 1:1 at the higher velocities. The mass manipulation affected the individual limb movements but not their coordination, indicating that a stable walking pattern was preserved. The results differed qualitatively from those obtained in studies on bimanual interlimb coordination, implying that the dynamical principles identified therein are not readily applicable to locomotion.     

Patterns of movement in the first 6 months of life: new directions

The field of infant perceptual development has relied heavily on the preferential-looking and habituation paradigms. Despite the obvious advantages of employing standardized methodologies, there is a need to study how perceptual variables give rise to and guide action beginning in the neonatal period. It is argued here that, in the first 5 months of life, it is important to distinguish reflexive patterns of movement from voluntary and cortically controlled patterns. Some criteria are defined. This paper describes three experiments which utilized movement counters to record developmental changes in the frequency and synchrony of limb movements in infants under 5 months. The results show that there is a general increase in frequency of limb movements from 1 to 5 months, but with a plateau or even a decrease in activity from the middle of the third month to the end of the fourth month. It was also demonstrated that synchrony scores (co-activation) for pairs of limbs showed significant increases from 1 month to 5 months. Finally, in all three experiments the presentation of an attractive toy resulted in suppression of activity. The results of the three experiments are interpreted to measure a gradual increase in cortically controlled movement patterns.     

On the space-time structure of human interlimb co-ordination

In three experiments we show, using behavioural measures of movement outcome, as well as movement trajectory information and resultant kinematic profiles, that there is a strong tendency for the limbs to be co-ordinated as a unitary structure even under conditions where the movements are of disparate difficulty. Environmental constraints (an obstacle placed in the path of one limb, but not in the other) are shown to modulate the space-time behaviour of both limbs (Experiment II). Our results obtain for symmetrical (Experiment I) as well as asymmetrical movements that involve non-homologous muscle groups (Experiment III). These findings suggest that in multi-joint limb movements, the many degrees of freedom are organised to function temporarily as a single coherent unit that is uniquely specific to the task demands placed on it. For movements in general, and two-handed movements in particular, such units are revealed in a partitioning of the relevant force demands for each component (a force scaling characteristic) and a preservation of the internal “topology” of the action, as indexed by the relative timing among components. These features, as well as systematic deviations from perfect synchrony between the limbs can be rationalised by a model that assumes the limbs behave qualitatively like non-linear oscillators.     

Limb and gender differences in the development of coordination in early infancy

Young infants produce a variety of spontaneous arm and leg movements in the first few months of life. Coordination of leg joints has been extensively investigated, whereas arm joint coordination has mainly been investigated in the sitting position in the context of early reaching and grasping. The current study investigated arm and leg joint coordination of movements produced in the supine position in 10 fullterm infants aged 6, 12 and 18 weeks. Longitudinal comparisons within limbs (intralimb) as well as between limbs (interlimb, ipsilateral and contralateral) were made as well as an exploration of differences in the development for boys and girls. The relationship between the joint angles was examined by measuring pair-wise cross-correlation functions for the angular displacement curves of the leg (hip, knee and ankle) and arm (shoulder, elbow and wrist) joints of both the right and left side. Both the arms and legs were found to follow a similar pattern of intralimb coordination, although the leg joints were more tightly coupled than the arm joints, particularly the proximal with the middle joint. In support of earlier findings, differences in the development of the right and left side were identified. In addition, gender differences in joint coordination were found for both intralimb and interlimb coordination. This contrasts with the view that gender differences in motor development may be primarily a result of environmental influences.     

Perception of the symmetrical patterning of human gait by infants

Two experiments tested 3- and 5-month-old infants' sensitivity to properties of point-light displays of human gait. In Experiment 1, infants were tested for discrimination of point-light displays of a walker and a runner, which, although they differed in many ways, were equivalent with regard to the phasing of limb movements. Results revealed that 3-month-old, but not 5-month-old, infants discriminated these displays. In Experiment 2, the symmetrical phase-patterning of the runner display was perturbed by advancing two of its limbs by 25% of the gait cycle. Both 3- and 5-month-old infants discriminated the walker display from this new phase-shifted runner display. These findings suggest that 3-month-old infants respond to the absolute and relative motions within a single limb, whereas 5-month-old infants respond primarily to the relations between limbs and, in particular, to the bilateral symmetry between the limbs.     

Spatial Error Detection and Assimilation Effects in Rapid Single and Bimanual Aiming Movements

In 2 experiments, spatial error detection capability and movement accuracy were investigated in both single and bimanual rapid aiming movements. In both experiments, right-handed college-age participants (N = 40 [Experiment 1]; N = 24 [Experiment 2]) used light, aluminum levers to make quick single and dual reversal movements in the sagittal plane in a time to reversal of 210 ms to either the same or different target locations involving identical (Experiment 1) or mirror-image (Experiment 2) movements. In Experiment 1, the shorter-distance limb overshot the target by 15-23&percent; when paired with a limb traveling at least 20 degrees farther, but no spatial assimilations were shown when movements differed by 20 degrees or less. In Experiment 2, the shorter-distance limb overshot 22-29&percent; when paired with a limb traveling 20 degrees farther, but spatial assimilations were not mitigated when both limbs moved to the same target position. Participants underestimated movement amplitude in all dual conditions but particularly when spatial assimilations were noted. Correlations between actual and estimated errors decreased from single to dual trials in both experiments. The findings suggest that spatial assimilations are caused by bimanual differences in movement amplitude, regardless of movement direction, and that individuals have greater difficulty identifying errors in simultaneous actions, especially when spatial assimilations are present, than identifying errors in single-limb actions.     

Toy-oriented changes in early arm movements III: constraints on joint kinematics

The purpose of this study was to identify invariant features of shoulder and elbow kinematics during prereaching arm movements with and without a toy present. Invariant movement features may reflect the presence of constraints that reduce the complexity of learning to reach and provide a link between early arm movements and reaching. Joint excursion and smoothness were consistently greater at the shoulder than the elbow suggesting strong organismal constraints on prereaching movements. Speed became greater in the shoulder than the elbow only with a toy present during the 4 weeks leading up to reach onset suggesting the introduction of task related constraints. We propose that organismal constraints on joint coordination throughout the prereaching period provide a foundation for the overlay of task related constraints closer to reach onset. We also suggest that the coordinative structures of early arm movements and later reaching may be much more similar than currently thought. This similarity would significantly reduce the elements needing to be actively controlled, and simplify the learning process.     

Postural fluctuations during pointing from a unilateral or bilateral stance

An experiment was conducted to compare the effects of bilateral and unilateral stance on postural fluctuations and intralimb coordination during active balance control. Fifteen participants stood bilaterally and unilaterally while conducting a pointing task with an outstretched arm. Excursion of center of foot pressure (CoP) and limb movements were recorded with a force plate and eight dual-axis accelerometers, respectively. Compared to bilateral stance, unilateral stance resulted in wider CoP trajectories and greater postural fluctuations, especially in the lower limbs. The limb-dependent postural fluctuations during unilateral stance were associated with an increased coupling between the upper limb segments and a decreased coupling between the segments of the stance leg. Unilateral stance further resulted in greater regularity and spectral changes in postural fluctuations of the trunk and lower limb due to increased central oscillations (8-15 Hz). The observed structural differences in postural fluctuations between unilateral and bilateral stance strongly suggested that the postural control system modulates joint stiffness in a stance-dependent manner. Probably, in unilateral stance, attentive control was shifted to the stance leg at the expense of increasing arm stiffness to reduce movement redundancy.     

Blocking Perturbations and Multilimb Control in Rapid-Aiming Movements

The effect of an unexpected mechanical block on the control of multilimbaiming movements was studied in two experiments. In the first experiment, subjects (N = 10) attempted to push two hand levers 9 cm forward in 200 ms without vision. In the second experiment, subjects (N = 9) attempted to push hand levers and foot pedals forward 9 cm in 200 ms. After a practice period, five attempts at blocking the limb movements were made on the left lever and the right lever (Experiment 1) and on both levers (Experiment 2 only) during randomly selected trials. When one hand was blocked in Experiment 1, the other hand undershot the target on the first blocked trial, with slight reductions in movement time. When one hand was blocked in Experiment 2, the contralateral limb undershot the target on all blocked trials, but this had little effect on the lower limbs. The lower limbs undershot the target when both up- per limbs were blocked. Discrete movement corrections were made on more of the blocked trials relative to the unblocked control trials. Interlimb correlations decreased following the block, suggesting that subjects dissociated the limbs in an effort to minimize the effect of the block.     

Influence of target uncertainty on reaching movements while standing in stroke

Stroke individuals frequently have balance problems and impaired arm movements that affect their daily activities. We investigated the influence of target uncertainty and the side of the brain lesion on the performance of arm movements and postural adjustments during reaching in a standing position by stroke individuals. Participants stood on force plates and reached a target displayed on the center of a monitor screen under conditions differentiated by the prior knowledge of the target location at the beginning of the movement. Individuals who had a stroke in the right side of the brain performed the tasks with the ipsilesional, right upper limb while the individuals with a left stroke performed with the ipsilesional, left upper limb. Healthy individuals performed with right and left limbs, which data were later averaged for statistical analysis. Kinematic analysis of the arm and lower limb joints and displacements of the center of pressure of each lower limb were compared between target conditions and groups. Stroke individuals showed larger center of pressure displacements of the contralesional compared to the ipsilesional limb while these displacements were symmetrical between lower limbs for the healthy individuals, regardless of the target condition. The target uncertainty affected both the characteristics of the arm movements and postural adjustments before movement onset. Right stroke individuals used more ankle joint movements under the uncertain compared to the certain condition. The uncertainty in target location affects the arm reaching in upright standing, but the effects depend on the side of the brain lesion.