Stretch reflexes contribute to arm impedance and longer-latency stretch reflexes exhibit increased sensitivity during interactions with compliant or unstable environments. This increased sensitivity is consistent with a regulation of arm impedance to compensate for decreased stability of the environment, but the specificity of this modulation has yet to be investigated. Many tasks, such as tool use, compromise arm stability along specific directions, and stretch reflexes tuned to those directions could present an efficient mechanism for regulating arm impedance in a task-appropriate manner. To be effective, such tuning should adapt not only to the mechanical properties of the environment but to those properties in relation to the arm, which also has directionally specific mechanical properties. The purpose of this study was to investigate the specificity of stretch reflex modulation during interactions with mechanical environments that challenge arm stability. The tested environments were unstable, having the characteristics of a negative stiffness spring. These were either aligned or orthogonal to the direction of maximal endpoint stiffness for each subject. Our results demonstrate preferential increases in reflexes, elicited within 50-100 ms of perturbation onset, to perturbations applied specifically along the direction of the destabilizing environments. This increase occurred only when the magnitude of the environmental instability exceeded endpoint stiffness along the same direction. These results are consistent with task-specific reflex modulation tuned to the mechanical properties of the environment relative to those of the human arm. They demonstrate a highly adaptable, involuntary mechanism that may be used to modulate limb impedance along specific directions.

BackgroundThe human motor system is highly redundant, having more kinematic degrees of freedom than necessary to complete a given task. Understanding how kinematic redundancies are utilized in different tasks remains a fundamental question in motor control. One possibility is that they can be used to tune the mechanical properties of a limb to the specific requirements of a task. For example, many tasks such as tool usage compromise arm stability along specific directions. These tasks only can be completed if the nervous system adapts the mechanical properties of the arm such that the arm, coupled to the tool, remains stable. The purpose of this study was to determine if posture selection is a critical component of endpoint stiffness regulation during unconstrained tasks.Methodology/Principal FindingsThree-dimensional (3D) estimates of endpoint stiffness were used to quantify limb mechanics. Most previous studies examining endpoint stiffness adaptation were completed in 2D using constrained postures to maintain a non-redundant mapping between joint angles and hand location. Our hypothesis was that during unconstrained conditions, subjects would select arm postures that matched endpoint stiffness to the functional requirements of the task. The hypothesis was tested during endpoint tracking tasks in which subjects interacted with unstable haptic environments, simulated using a 3D robotic manipulator. We found that arm posture had a significant effect on endpoint tracking accuracy and that subjects selected postures that improved tracking performance. For environments in which arm posture had a large effect on tracking accuracy, the self-selected postures oriented the direction of maximal endpoint stiffness towards the direction of the unstable haptic environment.Conclusions/SignificanceThese results demonstrate how changes in arm posture can have a dramatic effect on task performance and suggest that postural selection is a fundamental mechanism by which kinematic redundancies can be exploited to regulate arm stiffness in unconstrained tasks.

The often studied stretch reflex is fundamental to the involuntary control of posture and movement. Nevertheless, there remains controversy regarding its functional role. Many studies have demonstrated that stretch reflexes can be modulated in a task appropriate manner. This review focuses on modulation of the long latency stretch reflex, thought to be mediated, at least in part, by supraspinal pathways. For example, this component of the stretch reflex increases in magnitude during interactions with compliant environments, relative to the sensitivity during interactions with rigid environments. This suggests that reflex sensitivity increases to augment limb stability when that stability is not provided by the environment. However, not all results support the stabilizing role of stretch reflexes. Some studies have demonstrated that involuntary responses within the time period corresponding to the long latency reflex can destabilize limb posture. We propose that this debate stems from the fact that multiple perturbation-sensitive pathways can contribute to the long latency stretch reflex and that these pathways have separate functional roles. The presented studies suggest that neural activity occurring within the period normally ascribed to the long latency stretch reflex is highly adaptable to current task demands and possibly should be considered more intelligent than "reflexive."

To manipulate an object, we must simultaneously control the contact forces exerted on the object and the movements of our hand. Two alternative views for manipulation have been proposed: one in which motions and contact forces are represented and controlled by separate neural processes, and one in which motions and forces are controlled jointly, by a single process. To evaluate these alternatives, we designed three tasks in which subjects maintained a specified contact force while their hand was moved by a robotic manipulandum. The prescribed contact force and hand motions were selected in each task to induce the subject to attain one of three goals: (1) exerting a regulated contact force, (2) tracking the motion of the manipulandum, and (3) attaining both force and motion goals concurrently. By comparing subjects' performances in these three tasks, we found that behavior was captured by the summed actions of two independent control systems: one applying the desired force, and the other guiding the hand along the predicted path of the manipulandum. Furthermore, the application of transcranial magnetic stimulation impulses to the posterior parietal cortex selectively disrupted the control of motion but did not affect the regulation of static contact force. Together, these findings are consistent with the view that manipulation of objects is performed by independent brain control of hand motions and interaction forces.