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Classical theories of motor learning hypothesize that adaptation is driven by sensorimotor error; this is supported by studies of arm and eye movements that have shown that trial-to-trial adaptation increases with error. Studies of postural control have shown that anticipatory postural adjustments increase with the magnitude of a perturbation. However, differences in adaptation have been observed between the two modalities, possibly due to either the inherent instability or sensory uncertainty in standing posture. Therefore, we hypothesized that trial-to-trial adaptation in posture should be driven by error, similar to what is observed in arm reaching, but the nature of the relationship between error and adaptation may differ. Here we investigated trial-to-trial adaptation of arm reaching and postural control concurrently; subjects made reaching movements in a novel dynamic environment of varying strengths, while standing and holding the handle of a force-generating robotic arm. We found that error and adaptation increased with perturbation strength in both arm and posture. Furthermore, in both modalities, adaptation showed a significant correlation with error magnitude. Our results indicate that adaptation scales proportionally with error in the arm and near proportionally in posture. In posture only, adaptation was not sensitive to small error sizes, which were similar in size to errors experienced in unperturbed baseline movements due to inherent variability. This finding may be explained as an effect of uncertainty about the source of small errors. Our findings suggest that in rehabilitation, postural error size should be considered relative to the magnitude of inherent movement variability.

The motor system retains learning from visuomotor adaptation tasks in the form of “savings” to enable faster readaptation to similar perturbations in the future. Leow et al. ( J Neurophysiol 116: 1603–1614, 2016) suggest that the experience of prior errors during relearning is necessary for savings while repetition of prior actions may not be sufficient. These findings provide novel insight into factors that contribute to visuomotor adaptation and can be applied to future experimental and clinical research.

Long-train electrical stimulation of the motor and premotor cortices of nonhuman primates can produce either hand-to-mouth or grasp-to-inspect movements, depending on the precise location of stimulation. Furthermore, single-neuron recording studies identify discrete neuronal populations in the inferior parietal and ventral premotor cortices that respond uniquely to either grasp-to-eat or grasp-to-place movements, despite their identical mechanistic requirements. These studies demonstrate that the macaque motor cortex is organized around producing functional, goal-oriented movements, rather than simply fulfilling muscular prerequisites of action. In humans, right-handed hand-to-mouth movements have a unique kinematic signature; smaller maximum grip apertures are produced when grasping to eat than when grasping to place identical targets. This is evidence that the motor cortex in humans is also organized around producing functional movements. However, in both macaques and humans, grasp-to-eat/hand-to-mouth movements have always been elicited using edible targets and have (necessarily) been paired with mouth movement. It is therefore unknown whether the kinematic distinction is a natural result of grasping food and/or is simply attributable to concurrent opening of the mouth while grasping. In experiment 1, we used goal-differentiated grasping tasks, directed toward edible and inedible targets, to show that the unique kinematic signature is present even with inedible targets. In experiment 2, we used the same goal-differentiated grasping tasks, either coupled with or divorced from an open-mouth movement, to show that the signature is not attributable merely to a planned opening of the mouth during the grasp. These results are discussed in relation to the role of hand-to-mouth movements in human development, independently of grasp-to-eat behavior.

It is well established that the cortical processing of somatosensory and auditory signals is attenuated when they result from self-generated actions compared with external events. This phenomenon is thought to result from an efference copy of motor commands used to predict the sensory consequences of an action through a forward model. The present work examined whether attenuation also takes place for visual reafferent signals from the moving limb during voluntary reaching movements. To address this issue, EEG activity was recorded in a condition in which visual feedback of the hand was provided in real time and compared with a condition in which it was presented with a 150-ms delay, thus creating a mismatch between the predicted and actual visual consequences of the movement. Results revealed that the amplitude of the N1 component of the visual event-related potential evoked by hand visual feedback over the parietal cortex was significantly smaller when presented in real time compared with when it was delayed. These data suggest that the cortical processing of visual reafferent signals is attenuated when they are correctly predicted, likely as a result of a forward model.

Experimental studies have shown that a continuum of ankle and hip strategies is used to restore posture following an external perturbation. Postural responses can be modeled by feedback control with feedback gains that optimize a specific objective. On the one hand, feedback gains that minimize effort have been used to predict muscle activity during perturbed standing. On the other hand, hip and ankle strategies have been predicted by minimizing postural instability and deviation from upright posture. It remains unclear, however, whether and how effort minimization influences the selection of a specific postural response. We hypothesize that the relative importance of minimizing mechanical work vs. postural instability influences the strategy used to restore upright posture. This hypothesis was investigated based on experiments and predictive simulations of the postural response following a backward support surface translation. Peak hip flexion angle was significantly correlated with three experimentally determined measures of effort, i.e., mechanical work, mean muscle activity and metabolic energy. Furthermore, a continuum of ankle and hip strategies was predicted in simulation when changing the relative importance of minimizing mechanical work and postural instability, with increased weighting of mechanical work resulting in an ankle strategy. In conclusion, the combination of experimental measurements and predictive simulations of the postural response to a backward support surface translation showed that the trade-off between effort and postural instability minimization can explain the selection of a specific postural response in the continuum of potential ankle and hip strategies.

Contains fulltext : 159350.pdf (Publisher’s version ) (Closed access) The natural world continuously presents us with many opportunities for action, thus a process of target selection must precede action execution. While there has been considerable progress in understanding target selection in stationary environments, little is known about target selection when we are in motion. Here we investigated the effect of self-motion signals on saccadic target selection in a dynamic environment. Human subjects were sinusoidally translated (f=0.6 Hz, 30 cm peak-to-peak displacement) along an inter-aural axis using a vestibular sled. During the motion two visual targets were presented asynchronously but equidistantly on either side of fixation. Subjects had to look at one of these targets as quickly as possible. Using an adaptive approach, the time delay between these targets was adjusted until the subject selected both targets equally often. We determined this balanced time delay for different phases of the motion in order to distinguish the effects of body acceleration and velocity on saccadic target selection. Results show that acceleration (or position, as these are indistinguishable during sinusoidal motion), but not velocity, affect target selection for saccades. Subjects preferred to look at targets in the direction of the acceleration - the leftward target was preferred when the sled accelerated to the left, and vice versa. Saccadic reaction times mimicked this selection bias by being reliably shorter to targets in the direction of acceleration. Our results provide evidence that saccade target selection mechanisms are modulated by self-motion signals, which could be derived directly from the otolith system. 9 p.

Weight predictions used to scale lifting forces adapt quickly when repeatedly lifting unusually weighted objects and are readily updated by explicit information provided about weight. In contrast, weight predictions used when making perceptual judgments about weight are more resistant to change and are largely unaffected by explicit information about weight. These observations suggest that distinct memory systems underlie weight prediction when lifting objects and judging their weights. Here we examined whether these weight predictions differ in their reliance on declarative and nondeclarative memory resources by comparing the adaptability of these predictions in older adults, who exhibit relatively impaired declarative memory processes, to those in younger adults. In the size condition, we measured lift forces as participants repeatedly lifted a pair of size-weight inverted objects in alternation. To assess weight judgments, we measured the size-weight illusion every 10 lifts. The material condition was similar, except that we used material-weight inverted objects and measured the material-weight illusion. The strengths of these illusions prior to lifting, and the attenuation of the illusions that arise when lifting inverted objects, were similar for both groups. The magnitude of the change in the illusions was positively correlated with implicit memory performance in both groups, suggesting that predictions used when judging weight rely on nondeclarative memory resources. Updating of lifting forces also did not differ between groups. However, within the older group the success with which lifting forces were updated was positively correlated with working memory performance, suggesting that weight predictions used when lifting rely on declarative memory resources.

Glutamatergic currents play a fundamental role in regulating respiratory motor output and are partially mediated by α-amino-3-hydroxy-5-methyl-isoxazole-propionic acid (AMPA) receptors throughout the premotor and motor respiratory circuitry. Ampakines are pharmacological compounds that enhance glutamatergic transmission by altering AMPA receptor channel kinetics. Here, we examined if ampakines alter the expression of respiratory long-term facilitation (LTF), a form of neuroplasticity manifested as a persistent increase in inspiratory activity following brief periods of reduced O2 [intermittent hypoxia (IH)]. Current synaptic models indicate enhanced effectiveness of glutamatergic synapses after IH, and we hypothesized that ampakine pretreatment would potentiate IH-induced LTF of respiratory activity. Inspiratory bursting was recorded from the hypoglossal nerve of anesthetized and mechanically ventilated mice. During baseline (BL) recording conditions, burst amplitude was stable for at least 90 min (98 ± 5% BL). Exposure to IH (3 × 1 min, 15% O2) resulted in a sustained increase in burst amplitude (218 ± 44% BL at 90 min following final bout of hypoxia). Mice given an intraperitoneal injection of ampakine CX717 (15 mg/kg) 10 min before IH showed enhanced LTF (500 ± 110% BL at 90 min). Post hoc analyses indicated that CX717 potentiated LTF only when initial baseline burst amplitude was low. We conclude that under appropriate conditions ampakine pretreatment can potentiate IH-induced respiratory LTF. These data suggest that ampakines may have therapeutic value in the context of hypoxia-based neurorehabilitation strategies, particularly in disorders with blunted respiratory motor output such as spinal cord injury.

The purpose of the study was to compare the discharge characteristics of single motor units during sustained isometric contractions that required either force or position control in left-handed individuals. The target force for the two sustained contractions (24.9 ± 10.5% maximal force) was identical for each biceps brachii motor unit ( n = 32) and set at 4.7 ± 2.0% of maximal voluntary contraction (MVC) force above its recruitment threshold (range: 0.5–41.2% MVC force). The contractions were not sustained to task failure, but the duration (range: 60–330 s) was identical for each motor unit and the decline in MVC force immediately after the sustained contractions was similar for the two tasks (force: 11.1% ± 13.7%; position: 11.6% ± 9.9%). Despite a greater increase in the rating of perceived exertion during the position task (task × time interaction, P < 0.006), the amplitude of the surface-recorded electromyogram for the agonist and antagonist muscles increased similarly during the two tasks. Nonetheless, mean discharge rate of the biceps brachii motor units declined more during the position task (task × time interaction, P < 0.01) and the variability in discharge times (coefficient of variation for interspike interval) increased only during the position task (task × time interaction, P < 0.008). When combined with the results of an identical study on right-handers (Mottram CJ, Jakobi JM, Semmler JG, Enoka RM. J Neurophysiol 93: 1381–1392, 2005), the findings indicate that handedness does not influence the adjustments in biceps brachii motor unit activity during sustained submaximal contractions requiring either force or position control.

Short-latency afferent inhibition (SAI) is characterized by the suppression of the transcranial magnetic stimulation motor evoked potential (MEP) by the cortical arrival of a somatosensory afferent volley. It remains unknown whether the magnitude of SAI reflects changes in the sensory afferent volley, similar to that observed for somatosensory evoked potentials (SEPs). The present study investigated stimulus-response relationships between sensory nerve action potentials (SNAPs), SAI, and SEPs and their interrelatedness. Experiment 1 ( n = 23, age 23 ± 1.5 yr) investigated the stimulus-response profile for SEPs and SAI in the flexor carpi radialis muscle after stimulation of the mixed median nerve at the wrist using ∼25%, 50%, 75%, and 100% of the maximum SNAP and at 1.2× and 2.4× motor threshold (the latter equated to 100% of the maximum SNAP). Experiment 2 ( n = 20, age 23.1 ± 2 yr) probed SEPs and SAI stimulus-response relationships after stimulation of the cutaneous digital nerve at ∼25%, 50%, 75%, and 100% of the maximum SNAP recorded at the elbow. Results indicate that, for both nerve types, SAI magnitude is dependent on the volume of the sensory afferent volley and ceases to increase once all afferent fibers within the nerve are recruited. Furthermore, for both nerve types, the magnitudes of SAI and SEPs are related such that an increase in excitation within somatosensory cortex is associated with an increase in the magnitude of afferent-induced MEP inhibition.

Removing or adding sensory cues from one sensory system during standing balance causes a change in the contribution of the remaining sensory systems, a process referred to as sensory reweighting. While reweighting changes have been described in many studies under steady-state conditions, less is known about the temporal dynamics of reweighting following sudden transitions to different sensory conditions. The present study changed sensory conditions by periodically adding or removing visual (lights On/Off) or proprioceptive cues (surface sway referencing On/Off) in 12 young, healthy subjects. Evidence for changes in sensory contributions to balance was obtained by measuring the time course of medial-lateral sway responses to a constant-amplitude 0.56-Hz sinusoidal stimulus, applied as support surface tilt (proprioceptive contribution), as visual scene tilt (visual contribution), or as binaural galvanic vestibular stimulation (vestibular contribution), and by analyzing the time course of sway variability. Sine responses and variability of body sway velocity showed significant changes following transitions and were highly correlated under steady-state conditions. A dependence of steady-state responses on upcoming transitions was observed, suggesting that knowledge of impending changes can influence sensory weighting. Dynamic changes in sway in the period immediately following sensory transitions were very inhomogeneous across sway measures and in different experimental tests. In contrast to steady-state results, sway response and variability measures were not correlated with one another in the dynamic transition period. Several factors influence sway responses following addition or removal of sensory cues, partly instigated by but also obscuring the effects of reweighting dynamics.

The process of sensorimotor adaptation is considered to be driven by errors. While sensory prediction errors, defined as the difference between the planned and the actual movement of the cursor, drive implicit learning processes, target errors (e.g., the distance of the cursor from the target) are thought to drive explicit learning mechanisms. This distinction was mainly studied in the context of arm reaching tasks where the position and the size of the target were constant. We hypothesize that in a dynamic reaching environment, where subjects have to hit moving targets and the targets' dynamic characteristics affect task success, implicit processes will benefit from target errors as well. We examine the effect of target errors on learning of an unnoticed perturbation during unconstrained reaching movements. Subjects played a Pong game, in which they had to hit a moving ball by moving a paddle controlled by their hand. During the game, the movement of the paddle was gradually rotated with respect to the hand, reaching a final rotation of 25°. Subjects were assigned to one of two groups: The high-target error group played the Pong with a small ball, and the low-target error group played with a big ball. Before and after the Pong game, subjects performed open-loop reaching movements toward static targets with no visual feedback. While both groups adapted to the rotation, the postrotation reaching movements were directionally biased only in the small-ball group. This result provides evidence that implicit adaptation is sensitive to target errors.

Saccadic adaptation maintains the correct mapping between eye movements and their targets, yet the dynamics of saccadic gain changes in the presence of systematically varying disturbances has not been extensively studied. Here we assessed changes in the gain of saccade amplitudes induced by continuous and periodic postsaccadic visual feedback. Observers made saccades following a sequence of target steps either along the horizontal meridian (Two-way adaptation) or with unconstrained saccade directions (Global adaptation). An intrasaccadic step—following a sinusoidal variation as a function of the trial number (with 3 different frequencies tested in separate blocks)—consistently displaced the target along its vector. The oculomotor system responded to the resulting feedback error by modifying saccade amplitudes in a periodic fashion with similar frequency of variation but lagging the disturbance by a few tens of trials. This periodic response was superimposed on a drift toward stronger hypometria with similar asymptotes and decay rates across stimulus conditions. The magnitude of the periodic response decreased with increasing frequency and was smaller and more delayed for Global than Two-way adaptation. These results suggest that—in addition to the well-characterized return-to-baseline response observed in protocols using constant visual feedback—the oculomotor system attempts to minimize the feedback error by integrating its variation across trials. This process resembles a convolution with an internal response function, whose structure would be determined by coefficients of the learning model. Our protocol reveals this fast learning process in single short experimental sessions, qualifying it for the study of sensorimotor learning in health and disease.

We explored the origins of unintentional changes in performance during accurate force production in isometric conditions seen after turning visual feedback off. The idea of control with referent spatial coordinates suggests that these phenomena could result from drifts of the referent coordinate for the effector. Subjects performed accurate force/moment production tasks by pressing with the fingers of a hand on force sensors. Turning the visual feedback off resulted in slow drifts of both total force and total moment to lower magnitudes of these variables; these drifts were more pronounced in the right hand of the right-handed subjects. Drifts in individual finger forces could be in different direction; in particular, fingers that produced moments of force against the required total moment showed an increase in their forces. The force/moment drift was associated with a drop in the index of synergy stabilizing performance under visual feedback. The drifts in directions that changed performance (non-motor equivalent) and in directions that did not (motor equivalent) were of about the same magnitude. The results suggest that control with referent coordinates is associated with drifts of those referent coordinates toward the corresponding actual coordinates of the hand, a reflection of the natural tendency of physical systems to move toward a minimum of potential energy. The interaction between drifts of the hand referent coordinate and referent orientation leads to counterdirectional drifts in individual finger forces. The results also demonstrate that the sensory information used to create multifinger synergies is necessary for their presence over the task duration.

When a saccade is expected to result in a reward, both neural activity in oculomotor areas and the saccade itself (e.g., its vigor and latency) are altered (compared with when no reward is expected). As such, it is unclear whether the correlations of neural activity with reward indicate a representation of reward beyond a movement representation; the modulated neural activity may simply represent the differences in motor output due to expected reward. Here, to distinguish between these possibilities, we trained monkeys to perform a natural scene search task while we recorded from the frontal eye field (FEF). Indeed, when reward was expected (i.e., saccades to the target), FEF neurons showed enhanced responses. Moreover, when monkeys accidentally made eye movements to the target, firing rates were lower than when they purposively moved to the target. Thus, neurons were modulated by expected reward rather than simply the presence of the target. We then fit a model that simultaneously included components related to expected reward and saccade parameters. While expected reward led to shorter latency and higher velocity saccades, these behavioral changes could not fully explain the increased FEF firing rates. Thus, FEF neurons appear to encode motivational factors such as reward expectation, above and beyond the kinematic and behavioral consequences of imminent reward.

The mammalian superior colliculus (SC) is a midbrain structure that integrates multimodal sensory inputs and computes commands to initiate rapid eye movements. SC neurons burst with the sudden onset of a visual stimulus, followed by persistent activity that may underlie shifts of attention and decision making. Experiments in vitro suggest that circuit reverberations play a role in the burst activity in the SC, but the origin of persistent activity is unclear. In the present study we characterized an afterdepolarization (ADP) that follows action potentials in slices of rat SC. Population responses seen with voltage-sensitive dye imaging consisted of rapid spikes followed immediately by a second distinct depolarization of lower amplitude and longer duration. Patch-clamp recordings showed qualitatively similar behavior: in nearly all neurons throughout the SC, rapid spikes were followed by an ADP. Ionic and pharmacological manipulations along with experiments with current and voltage steps indicated that the ADP of SC neurons arises from Na+ current that either persists or resurges following Na+ channel inactivation at the end of an action potential. Comparisons of pharmacological properties and frequency dependence revealed a clear parallel between patch-clamp recordings and voltage imaging experiments, indicating a common underlying membrane mechanism for the ADP in both single neurons and populations. The ADP can initiate repetitive spiking at intervals consistent with the frequency of persistent activity in the SC. These results indicate that SC neurons have intrinsic membrane properties that can contribute to electrical activity that underlies shifts of attention and decision making.

Several studies have indicated that human observers are very efficient at tracking self-generated hand movements with their gaze, yet it is not clear whether this is simply a by-product of the predictability of self-generated actions or if it results from a deeper coupling of the somatomotor and oculomotor systems. In a first behavioral experiment we compared pursuit performance as observers either followed their own finger or tracked a dot whose motion was externally generated but mimicked their finger motion. We found that even when the dot motion was completely predictable in terms of both onset time and kinematics, pursuit was not identical to that produced as the observers tracked their finger, as evidenced by increased rate of catch-up saccades and by the fact that in the initial phase of the movement gaze was lagging behind the dot, whereas it was ahead of the finger. In a second experiment we recorded EEG in the attempt to find a direct link between the finger motor preparation, indexed by the lateralized readiness potential (LRP) and the latency of smooth pursuit. After taking into account finger movement onset variability, we observed larger LRP amplitudes associated with earlier smooth pursuit onset across trials. The same held across subjects, where average LRP onset correlated with average eye latency. The evidence from both experiments concurs to indicate that a strong coupling exists between the motor systems leading to eye and finger movements and that simple top-down predictive signals are unlikely to support optimal coordination.

The present work clarifies how the nucleus accumbens contributes to action. This region is often assumed to influence behavior “off-line” by evaluating outcomes. Studying rare recordings of local field potentials from the human nucleus accumbens, we observe a perimovement decrease of alpha and beta oscillations in seven of eight individuals, a signal that, in the motor system, is directly related to action preparation. Our results support the idea of an online role of this region for imminent action., The human nucleus accumbens is thought to play an important role in guiding future action selection via an evaluation of current action outcomes. Here we provide electrophysiological evidence for a more direct, i.e., online, role during action preparation. We recorded local field potentials from the nucleus accumbens in patients with epilepsy undergoing surgery for deep brain stimulation. We found a consistent decrease in the power of alpha/beta oscillations (10–30 Hz) before and around the time of movements. This perimovement alpha/beta desynchronization was observed in seven of eight patients and was present both before instructed movements in a serial reaction time task as well as before self-paced, deliberate choices in a decision making task. A similar beta decrease over sensorimotor cortex and in the subthalamic nucleus has been directly related to movement preparation and execution. Our results support the idea of a direct role of the human nucleus accumbens in action preparation and execution.

Various lines of evidence indicate that multiple movements can be prepared in parallel. Here, we show that preparing more than one movement comes with a cost: a movement plan is more variable if it is prepared simultaneously with another plan. This suggests that the representations of movement plans share a common neural resource and implies that the number of alternative plans is constrained by noise., Do movement plans, like representations in working memory, share a limited pool of resources? If so, the precision with which each individual movement plan is specified should decrease as the total number of movement plans increases. To explore this, human participants made speeded reaching movements toward visual targets. We examined if preparing one movement resulted in less variability than preparing two movements. The number of planned movements was manipulated in a delayed response cueing procedure that limited planning to a single target (experiment 1) or hand (experiment 2) or required planning of movements toward two targets (or with two hands). For both experiments, initial movement direction variability was higher in the two-plan condition than in the one-plan condition, demonstrating a cost associated with planning multiple movements, consistent with the limited resource hypothesis. In experiment 3, we showed that the advantage in initial variability of preparing a single movement was present only when the trajectory could be fully specified. This indicates that the difference in variability between one and two plans reflects the specification of full motor plans, not a general preparedness to move. The precision cost related to concurrent plans represents a novel constraint on motor preparation, indicating that multiple movements cannot be planned independently, even if they involve different limbs.

Recent neural and behavioral findings provide support for the influential idea that in situations in which multiple action options are presented simultaneously, we prepare action plans for each competing option before deciding between and executing one of those plans. However, in natural, everyday environments, our available action options frequently change from one moment to the next, and there is often uncertainty as to whether additional options will become available before having to select a particular course of action. Here, with the use of a target-directed reaching task, we show that in this situation, the brain specifies a competing action for each new, sequentially presented potential target and that recently formed action plans can be revisited and updated so as to conform with separate, more newly developed, plans. These findings indicate that the brain forms labile motor plans for sequentially arising target options that can be flexibly restructured to accommodate new motor plans.

We investigated the timing of gait parameter changes (stride length, peak toe velocity, and double-, single-support, and complete step duration) to control gait speed. Eleven healthy participants adjusted their gait speed on a treadmill to maintain a constant distance between them and a fore-aft oscillating cue (a place on a conveyor belt surface). The experimental design balanced conditions of cue modality (vision: eyes-open; manual contact: eyes-closed while touching the cue); treadmill speed (0.2, 0.4, 0.85, and 1.3 m/s); and cue motion (none, ±10 cm at 0.09, 0.11, and 0.18 Hz). Correlation analyses revealed a number of temporal relationships between gait parameters and cue speed. The results suggest that neural control ranged from feedforward to feedback. Specifically, step length preceded cue velocity during double-support duration suggesting anticipatory control. Peak toe velocity nearly coincided with its most-correlated cue velocity during single-support duration. The toe-off concluding step and double-support durations followed their most-correlated cue velocity, suggesting feedback control. Cue-tracking accuracy and cue velocity correlations with timing parameters were higher with the manual contact cue than visual cue. The cue/gait timing relationships generalized across cue modalities, albeit with greater delays of step-cycle events relative to manual contact cue velocity. We conclude that individual kinematic parameters of gait are controlled to achieve a desired velocity at different specific times during the gait cycle. The overall timing pattern of instantaneous cue velocities associated with different gait parameters is conserved across cues that afford different performance accuracies. This timing pattern may be temporally shifted to optimize control. Different cue/gait parameter latencies in our nonadaptation paradigm provide general-case evidence of the independent control of gait parameters previously demonstrated in gait adaptation paradigms.

The role of primary motor cortex (M1) in the control of hand movements is still unclear. Functional magnetic resonance imaging (fMRI) studies of unimanual performance reported a relationship between level of precision of a motor task and additional ipsilateral M1 (iM1) activation. In the present study, we determined whether the demand on accuracy of a movement influences the magnitude of the inhibitory effect between primary motor cortices (IHI). We used transcranial magnetic stimulation (TMS) to measure active IHI (aIHI) of the iM1 on the contralateral M1 (cM1) in the premovement period of a left-hand motor task. Ten healthy participants manipulated a joystick to point to targets of two different sizes. For aIHI, the conditioning stimulus (CS) was applied to iM1, and the test stimulus (TS) to cM1, with an interstimulus interval of 10 ms. The amount of the inhibitory effect of the CS on the motor-evoked potential (MEP) of the subsequent TS was expressed as percentage of the mean MEP amplitude evoked by the single TS. Across different time points of aIHI measurements in the premovement period, there was a significant effect for target size on aIHI. Preparing to point to small targets was associated with weaker aIHI compared with pointing to large targets. The present findings suggest that, during the premovement period, aIHI from iM1 on cM1 is modulated by the demand on accuracy of the motor task. This is consistent with task fMRI findings showing bilateral M1 activation during high-precision movements but only unilateral M1 activity during low-precision movements.

The human speech system exhibits a remarkable flexibility by adapting to alterations in speaking environments. While it is believed that speech motor adaptation under altered sensory feedback involves rapid reorganization of speech motor networks, the mechanisms by which different brain regions communicate and coordinate their activity to mediate adaptation remain unknown, and explanations of outcome differences in adaption remain largely elusive. In this study, under the paradigm of altered auditory feedback with continuous EEG recordings, the differential roles of oscillatory neural processes in motor speech adaptability were investigated. The predictive capacities of different EEG frequency bands were assessed, and it was found that theta-, beta-, and gamma-band activities during speech planning and production contained significant and reliable information about motor speech adaptability. It was further observed that these bands do not work independently but interact with each other suggesting an underlying brain network operating across hierarchically organized frequency bands to support motor speech adaptation. These results provide novel insights into both learning and disorders of speech using time frequency analysis of neural oscillations.

Movements can be learned implicitly in response to new environmental demands or explicitly through instruction and strategy. The former is often studied in an environment that perturbs movement so that people learn to correct the errors and store a new motor pattern. Here, we demonstrate in human walking that implicit learning of foot placement occurs even when an explicit strategy is used to block changes in foot placement during the learning process. We studied people learning a new walking pattern on a split-belt treadmill with and without an explicit strategy through instruction on where to step. When there is no instruction, subjects implicitly learn to place one foot in front of the other to minimize step-length asymmetry during split-belt walking, and the learned pattern is maintained when the belts are returned to the same speed, i.e., postlearning. When instruction is provided, we block expression of the new foot-placement pattern that would otherwise naturally develop from adaptation. Despite this appearance of no learning in foot placement, subjects show similar postlearning effects as those who were not given any instruction. Thus locomotor adaptation is not dependent on a change in action during learning but instead can be driven entirely by an unexpressed internal recalibration of the desired movement.

The presence of a respiratory-related cortical activity during tidal breathing is abnormal and a hallmark of respiratory difficulties, but its detection requires superior discrimination and temporal resolution. The aim of this study was to validate a computational method using EEG covariance (or connectivity) matrices to detect a change in brain activity related to breathing. In 17 healthy subjects, EEG was recorded during resting unloaded breathing (RB), voluntary sniffs, and breathing against an inspiratory threshold load (ITL). EEG were analyzed by the specially developed covariance-based classifier, event-related potentials, and time-frequency (T-F) distributions. Nine subjects repeated the protocol. The classifier could accurately detect ITL and sniffs compared with the reference period of RB. For ITL, EEG-based detection was superior to airflow-based detection ( P < 0.05). A coincident improvement in EEG-airflow correlation in ITL compared with RB ( P < 0.05) confirmed that EEG detection relates to breathing. Premotor potential incidence was significantly higher before inspiration in sniffs and ITL compared with RB ( P < 0.05), but T-F distributions revealed a significant difference between sniffs and RB only ( P < 0.05). Intraclass correlation values ranged from poor (−0.2) to excellent (1.0). Thus, as for conventional event-related potential analysis, the covariance-based classifier can accurately predict a change in brain state related to a change in respiratory state, and given its capacity for near “real-time” detection, it is suitable to monitor the respiratory state in respiratory and critically ill patients in the development of a brain-ventilator interface.

Voluntary limb modifications must be integrated with basic walking patterns during visually guided walking. In this study we tested whether voluntary gait modifications can become more automatic with practice. We challenged walking control by presenting visual stepping targets that instructed subjects to modify step length from one trial to the next. Our sequence learning paradigm is derived from the serial reaction-time (SRT) task that has been used in upper limb studies. Both random and ordered sequences of step lengths were used to measure sequence-specific and sequence-nonspecific learning during walking. In addition, we determined how age (i.e., healthy young adults vs. children) and biomechanical factors (i.e., walking speed) affected the rate and magnitude of locomotor sequence learning. The results showed that healthy young adults (age 24 ± 5 yr, n = 20) could learn a specific sequence of step lengths over 300 training steps. Younger children (age 6–10 yr, n = 8) had lower baseline performance, but their magnitude and rate of sequence learning were the same compared with those of older children (11–16 yr, n = 10) and healthy adults. In addition, learning capacity may be more limited at faster walking speeds. To our knowledge, this is the first study to demonstrate that spatial sequence learning can be integrated with a highly automatic task such as walking. These findings suggest that adults and children use implicit knowledge about the sequence to plan and execute leg movement during visually guided walking.

Paired associative stimulation (PAS) induces changes in the excitability of human sensorimotor cortex that outlast the procedure. PAS typically involves repeatedly pairing stimulation of a peripheral nerve that innervates an intrinsic hand muscle with transcranial magnetic stimulation over the representation of that muscle in the primary motor cortex. Depending on the timing of the stimuli (interstimulus interval of 25 or 10 ms), PAS leads to either an increase (PAS25) or a decrease (PAS10) in excitability. Both protocols, however, have been associated with an increase in excitability of nearby muscle representations not specifically targeted by PAS. Based on these spillover effects, we hypothesized that an additive, excitability-enhancing effect of PAS25 applied to one muscle representation may be produced by simultaneously applying PAS25 or PAS10 to a nearby representation. In different experiments prototypical PAS25 targeting the left thumb representation [abductor pollicis brevis (APB)] was combined with either PAS25 or PAS10 applied to the left little finger representation [abductor digiti minimi (ADM)] or, in a control experiment, with PAS10 also targeting the APB. In an additional control experiment PAS10 targeted both representations. The plasticity effects were quantified by measuring the amplitude of motor evoked potentials (MEPs) recorded before and after PAS. As expected, prototypical PAS25 was associated with an increase in MEP amplitude in the APB muscle. This effect was enhanced when PAS also targeted the ADM representation but only when a different interstimulus timing (PAS10) was used. These results suggest that PAS-induced plasticity is modified by concurrently targeting separate motor cortical representations with excitatory and inhibitory protocols.

The caudal cingulate motor area (CMAc) and the supplementary motor area (SMA) play important roles in movement execution. The present study examined the neural mechanisms underlying these roles by investigating local field potentials (LFPs) from these areas while monkeys pressed buttons with either their left or right hand. During hand movement, power increases in the high-gamma (80–120 Hz) and theta (3–8 Hz) bands and a power decrease in the beta (12–30 Hz) band were observed in both the CMAc and SMA. High-gamma and beta activity in the SMA predominantly represented contralateral hand movements, whereas activity in the CMAc preferentially represented movement of either hand. Theta activity in both brain regions most frequently reflected movement of either hand, but a contralateral hand bias was more evident in the SMA than in the CMAc. An analysis of the relationships of the laterality representations between the high-gamma and theta bands at each recording site revealed that, irrespective of the hand preference for the theta band, the high-gamma band in the SMA preferentially represented contralateral hand movement, whereas the high-gamma band in the CMAc represented movement of either hand. These findings suggest that the input-output relationships for ipsilateral and contralateral hand movements in the CMAc and SMA differ in terms of their functionality. The CMAc may transform the input signals representing general aspects of movement into commands to perform movements with either hand, whereas the SMA may transform the input signals into commands to perform movement with the contralateral hand.

As the major input to the basal ganglia, the striatum is innervated by a wide range of other areas. Overlapping input from these regions is speculated to influence temporal correlations among striatal ensembles. However, the network dynamics among behaviorally related neural populations in the striatum has not been extensively studied. We used large-scale neural recordings to monitor activity from striatal ensembles in mice undergoing Pavlovian reward conditioning. A subpopulation of putative medium spiny projection neurons (MSNs) was found to discriminate between cues that predicted the delivery of a reward and cues that predicted no specific outcome. These cells were preferentially located in lateral subregions of the striatum. Discriminating MSNs were more spontaneously active and more correlated than their nondiscriminating counterparts. Furthermore, discriminating fast spiking interneurons (FSIs) represented a highly prevalent group in the recordings, which formed a strongly correlated network with discriminating MSNs. Spike time cross-correlation analysis showed the existence of synchronized activity among FSIs and feedforward inhibitory modulation of MSN spiking by FSIs. These findings suggest that populations of functionally specialized (cue-discriminating) striatal neurons have distinct network dynamics that sets them apart from nondiscriminating cells, potentially to facilitate accurate behavioral responding during associative reward learning.

Down syndrome cell adherence molecule (DSCAM) contributes to the normal establishment and maintenance of neural circuits. Whereas there is abundant literature regarding the role of DSCAM in the neural patterning of the mammalian retina, less is known about motor circuits. Recently, DSCAM mutation has been shown to impair bilateral motor coordination during respiration, thus causing death at birth. DSCAM mutants that survive through adulthood display a lack of locomotor endurance and coordination in the rotarod test, thus suggesting that the DSCAM mutation impairs motor control. We investigated the motor and locomotor functions of DSCAM2J mutant mice through a combination of anatomical, kinematic, force, and electromyographic recordings. With respect to wild-type mice, DSCAM2J mice displayed a longer swing phase with a limb hyperflexion at the expense of a shorter stance phase during locomotion. Furthermore, electromyographic activity in the flexor and extensor muscles was increased and coactivated over 20% of the step cycle over a wide range of walking speeds. In contrast to wild-type mice, which used lateral walk and trot at walking speed, DSCAM2J mice used preferentially less coordinated gaits, such as out-of-phase walk and pace. The neuromuscular junction and the contractile properties of muscles, as well as their muscle spindles, were normal, and no signs of motor rigidity or spasticity were observed during passive limb movements. Our study demonstrates that the DSCAM mutation induces dystonic hypertonia and a disruption of locomotor gaits.

Interhemispheric interactions through the corpus callosum play an important role in the control of bimanual forces. However, the extent to which physiological connections between primary motor cortices are modulated during increasing levels of bimanual force generation in intact humans remains poorly understood. Here we studied coherence between electroencephalographic (EEG) signals and the ipsilateral cortical silent period (iSP), two well-known measures of interhemispheric connectivity between motor cortices, during unilateral and bilateral 10%, 40%, and 70% of maximal isometric voluntary contraction (MVC) into index finger abduction. We found that EEG-EEG coherence in the alpha frequency band decreased while the iSP area increased during bilateral compared with unilateral 40% and 70% but not 10% of MVC. Decreases in coherence in the alpha frequency band correlated with increases in the iSP area, and subjects who showed this inverse relation were able to maintain more steady bilateral muscle contractions. To further examine the relationship between the iSP and coherence we electrically stimulated the ulnar nerve at the wrist at the alpha frequency. Electrical stimulation increased coherence in the alpha frequency band and decreased the iSP area during bilateral 70% of MVC. Altogether, our findings demonstrate an inverse relation between alpha oscillations and the iSP during strong levels of bimanual force generation. We suggest that interactions between neural pathways mediating alpha oscillatory activity and transcallosal inhibition between motor cortices might contribute to the steadiness of strong bilateral isometric muscle contractions in intact humans.

Power spectral analyses of electrical signals from respiratory nerves reveal prominent oscillations above the primary rate of breathing. Acute exposure to intermittent hypoxia can induce a form of neuroplasticity known as long-term facilitation (LTF), in which inspiratory burst amplitude is persistently elevated. Most evidence indicates that the mechanisms of LTF are postsynaptic and also that high-frequency oscillations within the power spectrum show coherence across different respiratory nerves. Since the most logical interpretation of this coherence is that a shared presynaptic mechanism is responsible, we hypothesized that high-frequency spectral content would be unchanged during LTF. Recordings of inspiratory hypoglossal (XII) activity were made from anesthetized, vagotomized, and ventilated 129/SVE mice. When arterial O2 saturation (SaO2) was maintained >96%, the XII power spectrum and burst amplitude were unchanged for 90 min. Three, 1-min hypoxic episodes (SaO2 = 50 ± 10%), however, caused a persistent (>60 min) and robust (>400% baseline) increase in burst amplitude. Spectral analyses revealed a rightward shift of the signal content during LTF, with sustained increases in content above ∼125 Hz following intermittent hypoxia and reductions in power at lower frequencies. Changes in the spectral content during LTF were qualitatively similar to what occurred during the acute hypoxic response. We conclude that high-frequency content increases during XII LTF in this experimental preparation; this may indicate that intermittent hypoxia-induced plasticity in the premotor network contributes to expression of XII LTF.

Item does not contain fulltext Complex gait (e.g., obstacle avoidance) requires a higher cognitive load than simple steady-state gait, which is a more automated movement. The higher levels of the central nervous system, responsible for adjusting motor plans to complex gait, develop throughout childhood into adulthood. Therefore, we hypothesize that gait strategies in complex gait are likely to mature until adulthood as well. However, little is known about the maturation of complex gait from childhood into adolescence and adulthood. To address this issue, we investigated obstacle avoidance in forty-four 8- to 18-yr-old participants who walked at preferred speed along a 6-m walkway on which a planar obstacle (150% of step length, 1 m wide) was projected. Participants avoided the obstacle by stepping over this projection, while lower body kinematics were recorded. Results showed that step length and speed adjustments during successful obstacle avoidance were similar across all ages, even though younger children modified step width to a greater extent. Additionally, the younger children used larger maximal toe elevations and take-off distances than older children. Moreover, during unsuccessful trials, younger children deployed exaggerated take-off distances, which resulted in obstacle contact upon the consecutive heel strike. These results indicate that obstacle avoidance is not fully matured in younger children, and that the inability to plan precise foot placements is an important factor contributing to failures in obstacle avoidance.

A previous neurophysiological investigation demonstrated an increase in functional projections of expiratory bulbospinal neurons (EBSNs) in the segment above a chronic lateral thoracic spinal cord lesion that severed their axons. We have now investigated how this plasticity might be manifested in thoracic motoneurons by measuring their respiratory drive and the connections to them from individual EBSNs. In anesthetized cats, simultaneous recordings were made intracellularly from motoneurons in the segment above a left-side chronic (16 wk) lesion of the spinal cord in the rostral part of T8, T9, or T10 and extracellularly from EBSNs in the right caudal medulla, antidromically excited from just above the lesion but not from below. Spike-triggered averaging was used to measure the connections between pairs of EBSNs and motoneurons. Connections were found to have a very similar distribution to normal and were, if anything (nonsignificantly), weaker than normal, being present for 42/158 pairs, vs. 55/154 pairs in controls. The expiratory drive in expiratory motoneurons appeared stronger than in controls but again not significantly so. Thus we conclude that new connections made by the EBSNs following these lesions were made to neurons other than α-motoneurons. However, a previously unidentified form of functional plasticity was seen in that there was a significant increase in the excitation of motoneurons during postinspiration, being manifest either in increased incidence of expiratory decrementing respiratory drive potentials or in an increased amplitude of the postinspiratory depolarizing phase in inspiratory motoneurons. We suggest that this component arose from spinal cord interneurons.

Neural synchrony has been suggested as a mechanism for integrating distributed sensorimotor systems involved in coordinated movement. To test the role of corticomuscular and intermuscular coherence in bimanual coordination, we experimentally manipulated the degree of coordination between hand muscles by varying the sensitivity of the visual feedback to differences in bilateral force. In 16 healthy participants, cortical activity was measured using EEG and muscle activity of the flexor pollicis brevis of both hands using high-density electromyography (HDsEMG). Using the uncontrolled manifold framework, coordination between bilateral forces was quantified by the synergy index R V in the time and frequency domain. Functional connectivity was assessed using corticomuscular coherence between muscle activity and cortical source activity and intermuscular coherence between bilateral EMG activity. The synergy index increased in the high coordination condition. R V was higher in the high coordination condition in frequencies between 0 and 0.5 Hz; for the 0.5- to 2-Hz frequency band, this pattern was inverted. Corticomuscular coherence in the beta band (16–30 Hz) was maximal in the contralateral motor cortex and was reduced in the high coordination condition. In contrast, intermuscular coherence was observed at 5–12 Hz and increased with bimanual coordination. Within-subject comparisons revealed a negative correlation between R V and corticomuscular coherence and a positive correlation between R V and intermuscular coherence. Our findings suggest two distinct neural pathways: 1) corticomuscular coherence reflects direct corticospinal projections involved in controlling individual muscles; and 2) intermuscular coherence reflects diverging pathways involved in the coordination of multiple muscles.

The neuropeptide proctolin (RYLPT) plays important roles as both a neurohormone and a cotransmitter in arthropod neuromuscular systems. We used third-instar Drosophila larvae as a model system to differentiate synaptic effects of this peptide from its direct effects on muscle contractility and to determine whether proctolin can work in a cell-selective manner on muscle fibers. Proctolin did not appear to alter the amplitude of excitatory junctional potentials but did induce sustained muscle contractions in preparations where the CNS had been removed and no stimuli were applied to the remaining nerves. Proctolin-induced contractions were dose-dependent, were reduced by knocking down expression of the Drosophila proctolin receptor in muscle tissue, and were larger in some muscle cells than others (i.e., larger in fibers 4, 12, and 13 than in 6 and 7). Proctolin also increased the amplitude of nerve-evoked contractions in a dose-dependent manner, and the magnitude of this effect was also larger in some muscle cells than others (again, larger in fibers 4, 12, and 13 than in 6 and 7). Increasing the intraburst impulse frequency and number of impulses per burst increased the magnitude of proctolin's enhancement of nerve-evoked contractions and decreased the threshold and EC50concentrations for proctolin to enhance nerve-evoked contractions. Reducing proctolin receptor expression decreased the velocity of larval crawling at higher temperatures, and thermal preference in these larvae. Our results suggest that proctolin acts directly on body-wall muscles to elicit slow, sustained contractions and to enhance nerve-evoked contractions, and that proctolin affects muscle fibers in a cell-selective manner.

Cervical spinal cord injury (SCI) in humans typically damages both sides of the spinal cord, resulting in asymmetric functional impairments in the arms. Despite this well-accepted notion and the growing emphasis on the use of bimanual training strategies, how movement of one arm affects the motion of the contralateral arm after SCI remains unknown. Using kinematics and multichannel electromyographic (EMG) recordings we studied unilateral and bilateral reach-to-grasp movements to a small and a large cylinder in individuals with asymmetric arm impairments due to cervical SCI and age-matched control subjects. We found that the stronger arm of SCI subjects showed movement durations longer than control subjects during bilateral compared with unilateral trials. Specifically, movement duration was prolonged when opening and closing the hand when reaching for a large and a small object, respectively, accompanied by deficient activation of finger flexor and extensor muscles. In subjects with SCI interlimb coordination was reduced compared with control subjects, and individuals with lesser coordination between hands were those who showed prolonged times to open the hand. Although the weaker arm showed movement durations during bilateral compared with unilateral trials that were proportional to controls, the stronger arm was excessively delayed during bilateral reaching. Altogether, our findings demonstrate that during bilateral reach-to-grasp movements the more impaired arm has detrimental effects on hand opening and closing of the less impaired arm and that they are related, at least in part, to deficient control of EMG activity of hand muscles. We suggest that hand opening might provide a time to drive bimanual coordination adjustments after human SCI.

Synchronous motor unit firing instances have been attributed to anatomical inputs shared by motoneurons. Yet, there is a lack of empirical evidence confirming the notion that common inputs elicit synchronization under voluntary conditions. We tested this notion by measuring synchronization between motor unit action potential trains (MUAPTs) as their firing rates progressed within a contraction from a relatively low force level to a higher one. On average, the degree of synchronization decreased as the force increased. The common input notion provides no empirically supported explanation for the observed synchronization behavior. Therefore, we investigated a more probable explanation for synchronization. Our data set of 17,546 paired MUAPTs revealed that the degree of synchronization varies as a function of two characteristics of the motor unit firing rate: the similarity and the slope as a function of force. Both are measures of the excitation of the motoneurons. As the force generated by the muscle increases, the firing rate slope decreases, and the synchronization correspondingly decreases. Different muscles have motor units with different firing rate characteristics and display different amounts of synchronization. Although this association is not proof of causality, it consistently explains our observations and strongly suggests further investigation. So viewed, synchronization is likely an epiphenomenon, subject to countless unknown neural interactions. As such, synchronous firing instances may not be the product of a specific design and may not serve a specific physiological purpose. Our explanation for synchronization has the advantage of being supported by empirical evidence, whereas the common input does not.

The goal of this study was to investigate the human ability to stabilize the trunk in space during pelvic tilt. Upper body sway was evoked in kneeling-seated healthy subjects by angular platform perturbations with a rotation around a virtual low-back pivot point between the L4 and L5 vertebrae. To investigate motor control modulation, variations in task instruction (balance naturally or minimize trunk sway), vision (eyes open or closed), and perturbation bandwidth (from 0.2 up to 1, 3, or 10 Hz) were applied. Cocontraction and proprioceptive muscle spindle feedback were associated with minimizing low-back flexion/extension (trunk-on-pelvis stabilization), while vestibular and visual feedback were supposed to contribute to trunk-in-space stabilization. Trunk-in-space stabilization was only observed with the minimize trunk sway task instruction, while the task instruction to balance naturally led to trunk-on-pelvis stabilization with trunk rotations even exceeding the perturbations. This indicates that vestibular feedback is used when minimizing trunk sway but has only a minor contribution during natural trunk stabilization in the sagittal plane. The eyes open condition resulted in reduced global trunk rotations and increased global trunk reflexive responses, demonstrating effective visual contributions to trunk-in-space stabilization. On the other hand, increasing perturbation bandwidth caused a decreased feedback contribution leading to deteriorated trunk-in-space stabilization.

Anatomical studies demonstrate selective compartmental innervation of most human extraocular muscles (EOMs), suggesting the potential for differential compartmental control. This was supported by magnetic resonance imaging (MRI) demonstrating differential lateral rectus (LR) compartmental contraction during ocular counterrolling, differential medial rectus (MR) compartmental contraction during asymmetric convergence, and differential LR, inferior rectus (IR), and superior oblique (SO) compartmental contraction during vertical vergence. To ascertain possible differential compartmental EOM contraction during vertical ductions, surface coil MRI was performed over a range of target-controlled vertical gaze positions in 25 orbits of 13 normal volunteers. Cross-sectional areas and partial volumes of EOMs were analyzed in contiguous, quasi-coronal 2-mm image planes spanning origins to globe equator to determine morphometric features correlating best with contractility. Confirming and extending prior findings for horizontal EOMs during horizontal ductions, the percent change in posterior partial volume (PPV) of vertical EOMs from 8 to 14 mm posterior to the globe correlated best with vertical duction. EOMs were then divided into equal transverse compartments to evaluate the effect of vertical gaze on changes in PPV. Differential contractile changes were detected in the two compartments of the same EOM during infraduction for the IR medial vs. lateral (+4.4%, P = 0.03), LR inferior vs. superior (+4.0%, P = 0.0002), MR superior vs. inferior (−6.0%, P = 0.001), and SO lateral vs. medial (+9.7%, P = 0.007) compartments, with no differential contractile changes in the superior rectus. These findings suggest that differential compartmental activity occurs during normal vertical ductions. Thus all EOMs may contribute to cyclovertical actions.

Intermanual transfer, i.e., generalization of motor learning across hands, is a well-accepted phenomenon of motor learning. Yet, there are open questions regarding the characteristics of this transfer, particularly the intermanual transfer of dynamic learning. In this study, we investigated intermanual transfer in a force field adaptation task concerning the direction and the coordinate frame of transfer as well as the influence of a 24-h consolidation period on the transfer. We tested 48 healthy human subjects for transfer from dominant to nondominant hand, and vice versa. We considered two features of transfer. First, we examined transfer to the untrained hand using force channel trials that suppress error feedback and learning mechanisms to assess intermanual transfer in the form of a practice-dependent bias. Second, we considered transfer by exposing the subjects to the force field with the untrained hand to check for faster learning of the dynamics (interlimb savings). Half of the subjects were tested for transfer immediately after adaptation, whereas the other half were tested after a 24-h consolidation period. Our results showed intermanual transfer both from dominant to nondominant hand and vice versa in extrinsic coordinates. After the consolidation period, transfer effects were weakened. Moreover, the transfer effects were negligible compared with the subjects' ability to rapidly adapt to the force field condition. We conclude that intermanual transfer is a bidirectional phenomenon that vanishes with time. However, the ability to transfer motor learning seems to play a minor role compared with the rapid adaptation processes.

Motor learning during reaching not only recalibrates movement but can also lead to small but consistent changes in the sense of arm position. Studies have suggested that this sensory effect may be the result of recalibration of a forward model that associates motor commands with their sensory consequences. Here we investigated whether similar perceptual changes occur in the lower limbs after learning a new walking pattern on a split-belt treadmill—a task that critically involves proprioception. Specifically, we studied how this motor learning task affects perception of leg speed during walking, perception of leg position during standing or walking, and perception of contact force during stepping. Our results show that split-belt adaptation leads to robust motor aftereffects and alters the perception of leg speed during walking. This is specific to the direction of walking that was trained during adaptation (i.e., backward or forward). The change in leg speed perception accounts for roughly half of the observed motor aftereffect. In contrast, split-belt adaptation does not alter the perception of leg position during standing or walking and does not change the perception of stepping force. Our results demonstrate that there is a recalibration of a sensory percept specific to the domain of the perturbation that was applied during walking (i.e., speed but not position or force). Furthermore, the motor and sensory consequences of locomotor adaptation may be linked, suggesting overlapping mechanisms driving changes in the motor and sensory domains.

How does long-term training affect the neural control of movements? Here we tested the hypothesis that long-term training leading to skilled motor performance alters muscle coordination during challenging, as well as nominal everyday motor behaviors. Using motor module (a.k.a., muscle synergy) analyses, we identified differences in muscle coordination patterns between professionally trained ballet dancers (experts) and untrained novices that accompanied differences in walking balance proficiency assessed using a challenging beam-walking test. During beam walking, we found that experts recruited more motor modules than novices, suggesting an increase in motor repertoire size. Motor modules in experts had less muscle coactivity and were more consistent than in novices, reflecting greater efficiency in muscle output. Moreover, the pool of motor modules shared between beam and overground walking was larger in experts compared with novices, suggesting greater generalization of motor module function across multiple behaviors. These differences in motor output between experts and novices could not be explained by differences in kinematics, suggesting that they likely reflect differences in the neural control of movement following years of training rather than biomechanical constraints imposed by the activity or musculoskeletal structure and function. Our results suggest that to learn challenging new behaviors, we may take advantage of existing motor modules used for related behaviors and sculpt them to meet the demands of a new behavior.

In reaching to grasp an object, the arm transports the hand to the intended location as the hand shapes to grasp the object. Prior studies that tracked arm endpoint and grip aperture have shown that reaching and grasping, while proceeding in parallel, are interdependent to some degree. Other studies of reaching and grasping that have examined the joint angles of all five digits as the hand shapes to grasp various objects have not tracked the joint angles of the arm as well. We, therefore, examined 22 joint angles from the shoulder to the five digits as monkeys reached, grasped, and manipulated in a task that dissociated location and object. We quantified the extent to which each angle varied depending on location, on object, and on their interaction, all as a function of time. Although joint angles varied depending on both location and object beginning early in the movement, an early phase of location effects in joint angles from the shoulder to the digits was followed by a later phase in which object effects predominated at all joint angles distal to the shoulder. Interaction effects were relatively small throughout the reach-to-grasp. Whereas reach trajectory was influenced substantially by the object, grasp shape was comparatively invariant to location. Our observations suggest that neural control of reach-to-grasp may occur largely in two sequential phases: the first determining the location to which the arm transports the hand, and the second shaping the entire upper extremity to grasp and manipulate the object.

Our recent terminal experiments revealed that administration of a single train of repetitive spinal electromagnetic stimulation (sEMS; 35 min) enhanced synaptic plasticity in spinal circuitry following lateral hemisection spinal cord injury. In the current study, we have examined effects of repetitive sEMS applied as a single train and chronically (5 wk, every other day) following thoracic T10 contusion. Chronic studies involved examination of systematic sEMS administration alone and combined with exercise training and transgene delivery of neurotrophin [adeno-associated virus 10-neurotrophin 3 (AAV10-NT3)]. Electrophysiological intracellular/extracellular recordings, immunohistochemistry, behavioral testing, and anatomical tracing were performed to assess effects of treatments. We found that administration of a single sEMS train induced transient facilitation of transmission through preserved lateral white matter to motoneurons and hindlimb muscles in chronically contused rats with effects lasting for at least 2 h. These physiological changes associated with increased immunoreactivity of GluR1 and GluR2/3 glutamate receptors in lumbar neurons. Systematic administration of sEMS alone for 5 wk, however, was unable to induce cumulative improvements of transmission in spinomuscular circuitry or improve impaired motor function following thoracic contusion. Encouragingly, chronic administration of sEMS, followed by exercise training (running in an exercise ball and swimming), induced the following: 1) sustained strengthening of transmission to lumbar motoneurons and hindlimb muscles, 2) better retrograde transport of anatomical tracer, and 3) improved locomotor function. Greatest improvements were seen in the group that received exercise combined with sEMS and AAV-NT3.

How do cats manage to walk so graciously on top of narrow fences or windowsills high above the ground while apparently exerting little effort? In this study we investigated cat full-body mechanics and the activity of limb muscles and motor cortex during walking along a narrow 5-cm path on the ground. We tested the hypotheses that during narrow walking 1) lateral stability would be lower because of the decreased base-of-support area and 2) the motor cortex activity would increase stride-related modulation because of imposed demands on lateral stability and paw placement accuracy. We measured medio-lateral and rostro-caudal dynamic stability derived from the extrapolated center of mass position with respect to the boundaries of the support area. We found that cats were statically stable in the frontal plane during both unconstrained and narrow-path walking. During narrow-path walking, cats walked slightly slower with more adducted limbs, produced smaller lateral forces by hindlimbs, and had elevated muscle activities. Of 174 neurons recorded in cortical layer V, 87% of forelimb-related neurons (from 114) and 90% of hindlimb-related neurons (from 60) had activities during narrow-path walking distinct from unconstrained walking: more often they had a higher mean discharge rate, lower depth of stride-related modulation, and/or longer period of activation during the stride. These activity changes appeared to contribute to control of accurate paw placement in the medio-lateral direction, the width of the stride, rather than to lateral stability control, as the stability demands on narrow-path and unconstrained walking were similar.