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Accurate control of body posture is enforced by a multitude of corrective actions operating over a range of time scales. The earliest correction is the short-latency reflex (SLR) which occurs between 20-45 ms following a sudden displacement of the limb and is generated entirely by spinal circuits. In contrast, voluntary reactions are generated by a highly distributed network but at a significantly longer delay after stimulus onset (greater than 100 ms). Between these two epochs is the long-latency reflex (LLR) (around 50-100 ms) which acts more rapidly than voluntary reactions but shares some supraspinal pathways and functional capabilities. In particular, the LLR accounts for the arm's biomechanical properties rather than only responding to local muscle stretch like the SLR. This paper will review how the LLR accounts for the arm's biomechanical properties and the supraspinal pathways supporting this ability. Relevant experimental paradigms include clinical studies, non-invasive brain stimulation, neural recordings in monkeys, and human behavioral studies. The sum of this effort indicates that primary motor cortex and reticular formation (RF) contribute to the LLR either by generating or scaling its structured response appropriate for the arm's biomechanics whereas the cerebellum scales the magnitude of the feedback response. Additional putative pathways are discussed as well as potential research lines.

Joint movement tracked by a feedback pathway For animals with multi-joint limbs, one of the daunting problems that the nervous system has to solve is how to correctly interpret and respond to sensory input induced by complex combinations of limb movements. For example, one apparently simple displacement of the shoulder could arise from an infinite number of different combinations of forces acting at the shoulder and elbow. Pruszynski et al . use neurophysiological recordings in monkeys and stimulation studies in humans to demonstrate that knowledge of limb mechanics is solved through a feedback pathway involving the primary motor cortex (M1), rather than through the feed-forward processing of motion variables, a view which has been dominant for the past 25 years. The results have implications for the design of humanoid robots and brain–machine interfaces, as well as for understanding and treating patients with motor dysfunctions such as stroke. A basic difficulty for the nervous system is integrating locally ambiguous sensory information to form accurate perceptions about the outside world 1 , 2 , 3 , 4 . This local-to-global problem is also fundamental to motor control of the arm, because complex mechanical interactions between shoulder and elbow allow a particular amount of motion at one joint to arise from an infinite combination of shoulder and elbow torques 5 . Here we show, in humans and rhesus monkeys, that a transcortical pathway through primary motor cortex (M1) resolves this ambiguity during fast feedback control. We demonstrate that single M1 neurons of behaving monkeys can integrate shoulder and elbow motion information into motor commands that appropriately counter the underlying torque within about 50 milliseconds of a mechanical perturbation. Moreover, we reveal a causal link between M1 processing and multi-joint integration in humans by showing that shoulder muscle responses occurring ∼50 milliseconds after pure elbow displacement can be potentiated by transcranial magnetic stimulation. Taken together, our results show that transcortical processing through M1 permits feedback responses to express a level of sophistication that rivals voluntary control; this provides neurophysiological support for influential theories positing that voluntary movement is generated by the intelligent manipulation of sensory feedback 6 , 7 .

Although a limb's motion appears to be similar across unimanual and bimanual movements, here we demonstrate partial, but not complete, transfer of learning across these behavioral contexts, hidden learning that remains intact (but invisible) until the original context is again encountered, and the ability to associate two conflicting force fields simultaneously, one with each context. These results suggest partial, but not complete, overlap in the learning processes involved in the acquisition of unimanual and bimanual skills.

Accurately maintaining a fixed limb posture and quickly moving between postures underlies both everyday skills, including holding and lifting a cup of coffee, and expert skills, such as an Olympic wrestler's holding and throwing an opponent. A fundamental question in limb motor control is whether the brain manages these contrasting goals of posture and movement through a single, robust control process or whether each engages a specialized control process. We addressed this question by examining how individual neurons in the primary motor cortex of macaque monkeys represent mechanical loads during posture and movement tasks. Notably, approximately half of the neurons that expressed load-related activity did so exclusively during either posture only or movement only. Further, those neurons with load-related activity during both tasks randomly switched their magnitude of response between tasks. These random changes in load representation suggest specialized control processes, one for posture and one for movement.

An important window into sensorimotor function is how humans interact and stop moving projectiles, such as stopping a door from closing shut or catching a ball. Previous studies have suggested that humans time the initiation and modulate the amplitude of their muscle activity based on the momentum of the approaching object. However, real-world experiments are constrained by laws of mechanics, which cannot be manipulated experimentally to probe the mechanisms of sensorimotor control and learning. An augmented-reality variant of such tasks allows for experimental manipulation of the relationship between motion and force to obtain novel insights into how the nervous system prepares motor responses to interact with moving stimuli. Existing paradigms for studying interactions with moving projectiles use massless objects and are primarily focused on quantifying gaze and hand kinematics. Here, we developed a novel collision paradigm using a robotic manipulandum where participants mechanically stopped a virtual object moving in the horizontal plane. On each block of trials, we varied the virtual object’s momentum by increasing either its velocity or mass. Participants stopped the object by applying a force impulse that matched the object momentum. We observed that hand force increased as a function of object momentum linked to changes in virtual mass or velocity, similar to results from studies involving catching free-falling objects. In addition, increasing object velocity resulted in later onset of hand force relative to the impending time-to-contact. These findings show that the present paradigm can be used to determine how humans process projectile motion for hand motor control.

Sudden limb displacement evokes a complex sequence of compensatory muscle activity. Following the short-latency reflex and preceding voluntary reactions is an epoch termed the medium-latency reflex (MLR) that could reflect spinal processing of group II muscle afferents. One way to test this possibility is oral ingestion of tizanidine, an alpha-2 adrenergic agonist that inhibits the interneurons transmitting group II signals onto spinal motor neurons. We examined whether group II afferents contribute to MLR activity throughout the major muscles that span the elbow and shoulder. MLRs of ankle muscles were also tested during walking on the same day, in the same participants as well as during sitting in a different group of subjects. In contrast to previous reports, the ingestion of tizanidine had minimal impact on MLRs of arm or leg muscles during motor actions. A significant decrease in magnitude was observed for 2/16 contrasts in arm muscles and 0/4 contrasts in leg muscles. This discrepancy with previous studies could indicate that tizanidine’s efficacy is altered by subtle changes in protocol or that group II afferents do not substantially contribute to MLRs.