The vision of Hsiao on somatosensation.

The goal of this review is to start to consolidate and distill the substantial body of research that comprises the published work of the late Professor Steven S. Hsiao. The studies of Hsiao began by demonstrating the receptive field properties of somatosensory neurons, progressed to describing cortical feature selectivity, and then eventually elevated the field to hopes of tapping into natural neural codes with artificial somatosensory feedback.

Predictive mechanisms in the control of contour following.

In haptic exploration, when running a fingertip along a surface, the control system may attempt to anticipate upcoming changes in curvature in order to maintain a consistent level of contact force. Such predictive mechanisms are well known in the visual system, but have yet to be studied in the somatosensory system. Thus, the present experiment was designed to reveal human capabilities for different types of haptic prediction.

Patterns of muscle activity for digital coarticulation.

Although piano playing is a highly skilled task, basic features of motor pattern generation may be shared across tasks involving fine movements, such as handling coins, fingering food, or using a touch screen. The scripted and sequential nature of piano playing offered the opportunity to quantify the neuromuscular basis of coarticulation, i.e.

Hand kinematics of piano playing.

Dexterous use of the hand represents a sophisticated sensorimotor function. In behaviors such as playing the piano, it can involve strong temporal and spatial constraints. The purpose of this study was to determine fundamental patterns of covariation of motion across joints and digits of the human hand.

Multiple Factors Underlying Haptic Perception of Length and Orientation.

Information about the shape and spatial orientation of an object can be gathered during exploratory hand and arm movements, and then must be synthesized into a unified percept. During the robotically guided exploration of virtual polygons or triangles, the perception of the lengths of two adjoining segments is not always geometrically consistent with the perception of the internal angles between these segments. The present study further characterized this established inconsistency, and also found that subjects' internal angle judgments were influenced by the spatial orientations of the segments, especially the segment that was explored last in the sequence.

Coordination of hand shape.

The neural control of hand movement involves coordination of the sensory, motor, and memory systems. Recent studies have documented the motor coordinates for hand shape, but less is known about the corresponding patterns of somatosensory activity. To initiate this line of investigation, the present study characterized the sense of hand shape by evaluating the influence of differences in the amount of grasping or twisting force, and differences in forearm orientation.

What is the biological basis of sensorimotor integration?

This Prospects presents the problems that must be solved by the vertebrate nervous system in the process of sensorimotor integration and motor control. The concepts of efference copy and inverse model are defined, and multiple biological mechanisms are described, including those that form the basis of integration, extrapolation, and comparison/cancellation operations. Open questions for future research include the biological basis of continuous and distributed versus modular control, and somatosensory-motor coordination.

Somatosensory comparison during haptic tracing.

Active sensing involves memory retrieval and updating as well as mechanisms that trigger corrections to the ongoing exploratory movement. The present study examined this process in a task where human subjects moved the index fingertip clockwise around the circumference of a virtual sphere created by a robotic device. The fingertip pressed into the sphere during the movement, and the subjects were to report slight differences in sphere size (or surface curvature), which occurred from trial to trial.

Effects of object compliance on three-digit grasping.

Compared with rigid objects, grasping and lifting compliant objects presents additional uncertainties. For any static grasp, forces at the fingertips depend on factors including the locations of the contact points and the contact forces must be coordinated to maintain equilibrium. For compliant objects, the locations and orientations of the contact surfaces change in a force-dependent manner, thus changing the force requirements.

Factors Influencing Haptic Perception of Complex Shapes.

Exploration of an object by arm movement and somatosensation is a serial process that relies on memories and expectations. The present experiments tested the hypothesis that this process involves breaking the object into component shapes (primitives). This was tested by having human subjects explore shapes composed of semicircular arcs, as well as quarter circles or quarter ellipses.

Sensorimotor control of contact force.

Interacting with objects in the environment introduces several new challenges for motor control: the potential for instability, external constraints on possible motions and novel dynamics. Grasping and manipulating objects provide the most elaborate examples of such motor tasks. We review each of these topics and suggest that when sensory feedback is reliable, it is used to adapt the motion to the requirements imposed by the object.

Extrapolation of visual motion for manual interception.

A frequent goal of hand movement is to touch a moving target or to make contact with a stationary object that is in motion relative to the moving head and body. This process requires a prediction of the target's motion, since the initial direction of the hand movement anticipates target motion. This experiment was designed to define the visual motion parameters that are incorporated in this prediction of target motion.

Multi-digit control of contact forces during rotation of a hand-held object.

Rotation of an object held with three fingers is produced by modulation of force amplitude and direction at one or more contact points. Changes in the moment arm through which these forces act can also contribute to the modulation of the rotational moment. Therefore force amplitude and direction as well as the center of pressure on each contact surface must be carefully coordinated to produce a rotation.

Multidigit control of contact forces during transport of handheld objects.

When an object is lifted vertically, the normal force increases and decreases in tandem with tangential (load) force to safely avoid slips. For horizontal object transport, horizontal forces at the contact surfaces can be decomposed into manipulation forces (producing acceleration/deceleration) and grasping forces. Although the grasping forces must satisfy equilibrium constraints, it is not clear what determines their modulation across time, nor the extent to which they result from active muscle contraction or mechanical interactions of the digits with the moving object.

Intrinsic Hand Muscle Activation for Grasp and Horizontal Transport.

During object manipulation, the hand and arm muscles produce internal forces on the object (grasping forces) and forces that result in external translation or rotation of the object in space (transport forces). The present study tested whether the intrinsic hand muscles are actively involved in transport as well as grasping. Intrinsic hand muscle activity increased with increasing demands for grasp stability, but also showed the timing and directional tuning patterns appropriate for actively transmitting external forces to the object, during the translational acceleration and deceleration of object transport.

Timing of muscle activation in a hand movement sequence.

Recent studies have described muscle synergies as overlapping, multimuscle groups defined by synchronous covariation in activation intensity. A different approach regards a synergy as a fixed temporal sequence of bursts of activity across groups of motoneurons. To pursue this latter definition, the present study used a principal component (PC) analysis tailored to reveal the across-muscle temporal synergies of human hand movement.

Oculomotor responses to gradual changes in target direction.

Smooth pursuit tracking of targets moving linearly (in one dimension) is well characterized by a model where retinal image motion drives eye acceleration. However, previous findings suggest that this model cannot be simply extended to two-dimensional (2D) tracking. To examine 2D pursuit, in the present study, human subjects tracked a target that moved linearly and then followed the arc of a circle.

Planning and drawing complex shapes.

Arm and hand movements are generally controlled using a combination of sensory-based and memory-based guidance mechanisms. This study examined similarities and differences in visually-guided and memory-guided arm movements, and sought to determine as to what extent certain control principles apply to each type of movement. In particular, the 2/3 power law is a principle that appears to govern the formation of complex, curved hand trajectories; it specifies that the tangential velocity should be proportional to the radius of curvature raised to an exponent of 1/3.

Manual tracking in three dimensions.

Little is known about the manual tracking of targets that move in three dimensions. In the present study, human subjects followed, with the tip of a hand-held pen, a virtual target moving four times (period 5 s) around a novel, unseen path. Two basic types of target paths were used: a peanut-shaped Cassini ellipse and a quasi-spherical shape where four connected semicircles lay in orthogonal planes.

Haptic feature extraction.

This study examined the process by which the shape of a haptically explored object is synthesized from the geometric characteristics of simpler constituent elements, such as arcs and ellipses. Subjects traced the outlines of virtual objects by means of whole arm movements. Each object consisted of the union of a large central ellipse and two smaller circles, extending upward and outward from the top left and right sides of the base.

Effect of compliance on haptic perception of curvature.

The haptic sense of geometric properties such as the curvature of a contour is derived from somatosensory cues about the motions and forces experienced during exploratory actions. This study addressed the question of whether compliance, the relationship between force and displacement, influences haptic perception of curvature. Subjects traced a curved 30 cm long compliant contour by grasping the handle of a manipulandum and reported whether the contour curved towards or away from them.

Functional somatotopy in sensorimotor cortex.

In an effort to understand the highly distributed somatotopy of primary motor cortex, this review draws on principles derived from studies of auditory, visual and somatosensory cortical areas. In each case, a behaviorally important feature or function is overlaid in multiple locations on an underlying topographic map of the peripheral sensory surface. Recent studies of hand muscle synergies suggest the types of two-dimensional functional axes that might reasonably be mapped to the two-dimensional surface of the primary motor cortex.

Distortions in the visual perception of shape.

It is known that visual illusions lead to a distorted perception of the length and orientation of lines, but it is not clear how these illusions affect the appreciation of the shape of closed forms. In this study two experiments were performed to characterize distortions in the visual perception of the shape of quadrilaterals and the extent to which these distortions were similar to the distortions of haptically sensed shapes. In the first experiment human subjects were presented with two quadrilaterals side by side on a computer monitor.

Smooth pursuit tracking of an abrupt change in target direction: vector superposition of discrete responses.

The directional control of smooth pursuit eye movements was studied by presenting human subjects with targets that moved in a straight line at a constant speed and then changed direction abruptly and unpredictably. To minimize the probability of saccadic responses in the interval following the target's change in direction, target position was offset so as to eliminate position error after the reaction time. Smooth pursuit speed declined at a latency of 90 ms, whereas the direction of smooth pursuit began to change later (130 ms).

Muscular and postural synergies of the human hand.

Because humans have limited ability to independently control the many joints of the hand, a wide variety of hand shapes can be characterized as a weighted combination of just two or three main patterns of covariation in joint rotations, or "postural synergies." The present study sought to align muscle synergies with these main postural synergies and to describe the form of membership of motor units in these postural/muscle synergies. Seventeen joint angles and the electromyographic (EMG) activities of several hand muscles (both intrinsic and extrinsic muscles) were recorded while human subjects held the hand statically in 52 specific shapes (i.