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Pupil diameter, under constant illumination, is known to reflect individuals' internal states, such as surprise about observation and environmental uncertainty. Despite the growing use of pupillometry in cognitive learning studies as an additional measure for examining internal states, few studies have used pupillometry in human motor learning studies. Here, we provide the first detailed characterization of pupil diameter changes in a short-term reach adaptation paradigm. We measured pupil changes in 121 human participants while they adapted to abrupt, gradual, or switching force field conditions. Sudden increases in movement error caused by the introduction/reversal of the force field resulted in strong phasic pupil dilation during movement accompanied by a transient increase in tonic premovement baseline pupil diameter in subsequent trials. In contrast, pupil responses were reduced when the force field was gradually introduced, indicating that large, unexpected errors drove the changes in pupil responses. Interestingly, however, error-induced pupil responses gradually became insensitive after experiencing multiple force field reversals. We also found an association between baseline pupil diameter and incidental knowledge of the gradually introduced perturbation. Finally, in all experiments, we found a strong co-occurrence of larger baseline pupil diameter with slower reaction and movement times after each rest break. Collectively, these results suggest that tonic baseline pupil diameter reflects one's belief about environmental uncertainty, whereas phasic pupil dilation during movement reflects surprise about a sensory outcome (i.e., movement error), and both effects are modulated by novelty. Our results provide a new approach for nonverbally assessing participants' internal states during motor learning. [ABSTRACT FROM AUTHOR]

Previous studies have shown that cerebellar transcranial direct current stimulation (tDCS) leads to faster adaptation of arm reaching movements to visuomotor rotation and force field perturbations in healthy subjects. The first aim of the present study was to confirm a stimulation-dependent effect on motor adaptation. Second, we investigated whether tDCS effects differ depending on onset, that is, before or at the beginning of the adaptation phase. A total of 120 healthy and right-handed subjects (60 women, mean age 23.2 ± SD 2.7 yr, range 18-31 yr) were tested. Subjects moved a cursor with a manipulandum to one of eight targets presented on a vertically orientated screen. Three baseline blocks were followed by one adaptation block and three washout blocks. Sixty subjects did a force field adaptation task (FF), and 60 subjects did a visuomotor adaptation task (VM). Equal numbers of subjects received anodal, cathodal, or sham cerebellar tDCS beginning either in the third baseline block or at the start of the adaptation block. In FF and VM, tDCS and the onset of tDCS did not show a significant effect on motor adaptation (all P values >0.05). We were unable to support previous findings of modulatory cerebellar tDCS effects in reaching adaptation tasks in healthy subjects. Prior to possible application in patients with cerebellar disease, future experiments are needed to determine which tDCS and task parameters lead to robust tDCS effects. NEW & NOTEWORTHY Transcranial direct current stimulation (tDCS) is a promising tool to improve motor learning. We investigated whether cerebellar tDCS improves motor learning in force field and visuomotor tasks in healthy subjects and what influence the onset of stimulation has. We did not find stimulation effects of tDCS or an effect of onset of stimulation. A reevaluation of cerebellar tDCS in healthy subjects and at the end of the clinical potential in cerebellar patients is demanded. [ABSTRACT FROM AUTHOR]

Recent computational theories and behavioral observations suggest that motor learning is supported by multiple adaptation processes, operating on different timescales, but direct neural evidence is lacking. We tested this hypothesis by applying transcranial magnetic stimulation over motor cortex in 16 human subjects during a validated reach adaptation task. Motor-evoked potentials (MEPs) and cortical silent periods (CSPs) were recorded from the biceps brachii to assess modulations of corticospinal excitability as indices for corticospinal plasticity. Guided by a two-state adaptation model, we show that the MEP reflects an adaptive process that learns quickly but has poor retention, while the CSP correlates with a process that responds more slowly but retains information well. These results provide a physiological link between models of motor learning and distinct changes in corticospinal excitability. Our findings support the relationship between corticospinal gain modulations and the adaptive processes in motor learning. NEW & NOTEWORTHY Computational theories and behavioral observations suggest that motor learning is supported by multiple adaptation processes, but direct neural evidence is lacking. We tested this hypothesis by applying transcranial magnetic stimulation over human motor cortex during a reach adaptation task. Guided by a two-state adaptation model, we show that the motor-evoked potential reflects a process that adapts and decays quickly, whereas the cortical silent period reflects slow adaptation and decay. [ABSTRACT FROM AUTHOR]

To adapt to deterministic force perturbations that depend on the current state of the hand, internal representations are formed to capture the relationships between forces experienced and motion. However, information from multiple modalities travels at different rates, resulting in intermodal delays that require compensation for these internal representations to develop. To understand how these delays are represented by the brain, we presented participants with delayed velocity-dependent force fields, i.e., forces that depend on hand velocity either 70 or 100 ms beforehand. We probed the internal representation of these delayed forces by examining the forces the participants applied to cope with the perturbations. The findings showed that for both delayed forces, the best model of internal representation consisted of a delayed velocity and current position and velocity. We show that participants relied initially on the current state, but with adaptation, the contribution of the delayed representation to adaptation increased. After adaptation, when the participants were asked to make movements with a higher velocity for which they had not previously experienced with the delayed force field, they applied forces that were consistent with current position and velocity as well as delayed velocity representations. This suggests that the sensorimotor system represents delayed force feedback using current and delayed state information and that it uses this representation when generalizing to faster movements. [ABSTRACT FROM AUTHOR]

It is currently unclear whether the brain plans movement kinematics explicitly or whether movement paths arise implicitly through optimization of a cost function that takes into account control and/or dynamic variables. Several cost functions are proposed in the literature that are very different in nature (e.g., control effort, torque change, and jerk), yet each can predict common movement characteristics. We set out to disentangle predictions of the different variables using a combination of modeling and empirical studies. Subjects performed goal-directed arm movements in a force field (FF) in combination with visual perturbations of seen hand position. This FF was designed to have distinct optimal movements for muscle-input and dynamic costs while leaving kinematic cost unchanged. Visual perturbations in turn changed the kinematic cost but left the dynamic and muscle-input costs unchanged. An optimally controlled, physiologically realistic arm model was used to predict movements under the various cost variables. Experimental results were not consistent with a cost function containing any of the control and dynamic costs investigated. Movement patterns of all experimental conditions were adequately predicted by a kinematic cost function comprising both visually and somatosensory perceived jerk. The present study provides clear behavioral evidence that the brain solves kinematic and mechanical redundancy in separate steps: in a first step, movement kinematics are planned; and in a second, separate step, muscle activation patterns are generated. [ABSTRACT FROM AUTHOR]

Pupil diameter, under constant illumination, is known to reflect individuals' internal states, such as surprise about observation and environmental uncertainty. Despite the growing use of pupillometry in cognitive learning studies as an additional measure for examining internal states, few studies have used pupillometry in human motor learning studies. Here, we provide the first detailed characterization of pupil diameter changes in a short-term reach adaptation paradigm. We measured pupil changes in 121 human participants while they adapted to abrupt, gradual, or switching force field conditions. Sudden increases in movement error caused by the introduction/reversal of the force field resulted in strong phasic pupil dilation during movement accompanied by a transient increase in tonic premovement baseline pupil diameter in subsequent trials. In contrast, pupil responses were reduced when the force field was gradually introduced, indicating that large, unexpected errors drove the changes in pupil responses. Interestingly, however, error-induced pupil responses gradually became insensitive after experiencing multiple force field reversals. We also found an association between baseline pupil diameter and incidental knowledge of the gradually introduced perturbation. Finally, in all experiments, we found a strong co-occurrence of larger baseline pupil diameter with slower reaction and movement times after each rest break. Collectively, these results suggest that tonic baseline pupil diameter reflects one's belief about environmental uncertainty, whereas phasic pupil dilation during movement reflects surprise about a sensory outcome (i.e., movement error), and both effects are modulated by novelty. Our results provide a new approach for nonverbally assessing participants' internal states during motor learning. NEW & NOTEWORTHY Pupil diameter is known as a noninvasive window into individuals' internal states. Despite the growing use of pupillometry in cognitive learning studies, it receives little attention in motor learning studies. Here, we characterized the pupil responses in a short-term reach adaptation paradigm by measuring pupil diameter of human participants while they adapted to abrupt, gradual, or switching force field conditions. Our results demonstrate how surprise and uncertainty reflected in pupil diameter develop during motor adaptation.

A compelling idea in cognitive neuroscience links motor control and action observation. Recent work supports the idea that a link exists not just between action observation and action planning, but between observation and motor learning. Several studies support the idea that cortical regions that underlie active motor learning also play a role in motor learning by observing. The goal of the present study was to test whether motor learning by observing is effector dependent (as in active motor learning) or effector independent (as in studies of action observation and mirror neurons). Right-handed human subjects observed a video depicting another individual learning to reach to visual targets in a force field (FF). The video showed reaching in a clockwise FF (CWFF) or a counter-clockwise FF (CCWFF), and depicted an individual reaching with the right or left arm. After observation, all subjects were asked to reach in a CWFF, using their right arm. As in our prior studies, subjects who observed a CWFF prior to the CWFF test performed better than subjects who observed a CCWFF. We show here that this effect was seen both when observers watched others reach using their right arm, and when observers watched others learning to reach using the left arm. These results suggest that information about novel forces learned through observation is represented in an effector-independent coordinate frame, and are consistent with the idea that the brain links not only observation and movement, but motor learning as well, through abstract representations of actions. [ABSTRACT FROM AUTHOR]

Everyday movements often have multiple solutions. Many of these solutions arise from biomechanical redundancies. Often, however, the goal does not require a unique movement. To examine how people exploit task-related redundancy, we observed as participants produced three-dimensional (3-D) reaching movements, moving to one of two rectangular targets that were diagonally oriented in the frontal (x, y) plane. On most trials, the movement was perturbed by a vertical, velocity-dependent force. Since participants were free to move in 3-D space, online corrections could involve movement along the perturbed, vertical dimension, as well as the nonperturbed, horizontal dimension. If the motor system exploits task redundancies, then corrections along the horizontal dimension should depend on the orientation of the target. Consistent with this prediction, participants modified both the horizontal and vertical coordinates of the trajectory over the course of learning, and the horizontal component was sensitive to the orientation of the target. Furthermore, participants produced online corrections with a horizontal component that brought the hand closer to the target. These results suggest that we not only correct for mismatches between expected and experienced forces but also exploit task-specific redundancies to efficiently improve performance. [ABSTRACT FROM AUTHOR]