In the previous paper regarding the somatosensory control of the human precision grip, we concluded that the elicited automatic grip force adjustments are graded by the amplitude of the imposed loads when restraining an 'active' object subjected to unpredictable pulling forces (Johansson et al. 1992a). Using the same subjects and apparatus, the present study examines the capacity to respond to imposed load forces applied at various rates. Grip and load forces (forces normal and tangential to the grip surfaces) and the position of the object in the pulling direction (distal) were recorded. Trapezoidal load force profiles with plateau amplitudes of 2 N were delivered at the following rates of loading and unloading in an unpredictable sequence: 2 N/s, 4 N/s or 8 N/s. In addition, trials with higher load rate (32 N/s) at a low amplitude (0.7 N) were intermingled. The latencies between the start of the loading and the onset of the grip force response increased with decreasing load force rate. They were 80 +/- 9 ms, 108 +/- 13 ms, 138 +/- 27 ms and 174 +/- 39 ms for the 32, 8, 4 and 2 N/s rates, respectively. These data suggested that the grip response was elicited after a given minimum latency once a load amplitude threshold was exceeded. The amplitude of the initial rapid increase of grip force (i.e., the 'catch-up' response) was scaled by the rate of the load force, whereas its time course was similar for all load rates. This response was thus elicited as a unit, but its amplitude was graded by afferent information about the load rate arising very early during the loading. The scaling of the catch-up response was purposeful since it facilitated a rapid reconciliation of the ratio between the grip and load force to prevent slips. In that sense it apparently also compensated for the varying delays between the loading phase and the resultant grip force responses. However, modification of the catch-up response may occur during its course when the loading rate is altered prior to the grip force response or very early during the catch-up response itself. Hence, afferent information may be utilized continuously in updating the response although its motor expression may be confined to certain time contingencies. Moreover, this updating may take place after an extremely short latency (45-50 ms).(ABSTRACT TRUNCATED AT 400 WORDS)
In manipulating 'passive' objects, for which the physical properties are stable and therefore predictable, information essential for the adaptation of the motor output to the properties of the current object is principally based on 'anticipatory parameter control' using sensorimotor memories, i.e., an internal representation of the object's properties based on previous manipulative experiences. Somatosensory afferent signals only intervene intermittently according to an 'event driven' control policy. The present study is the first in a series concerning the control of precision grip when manipulating 'active' objects that exert unpredictable forces which cannot be adequately represented in a sensorimotor memory. Consequently, the manipulation may be more reliant on a moment-to-moment sensory control. Subjects who were prevented from seeing the hand used the precision grip to restrain a manipulandum with two parallel grip surfaces attached to a force motor which produced distally directed (pulling) loads tangential to the finger tips. The trapezoidal load profiles consisted of a loading phase (4 N/s), plateau phase and an unloading phase (4 N/s) returning the load force to zero. Three force amplitudes were delivered in an unpredictable sequence; 1 N, 2 N and 4 N. In addition, trials with higher load rate (32 N/s) at a low amplitude (0.7 N), were superimposed on various background loads. The movement of the manipulandum, the load forces and grip forces (normal to the grip surfaces) were recorded at each finger. The grip force automatically changed with the load force during the loading and unloading phases. However, the grip responses were initiated after a brief delay. The response to the loading phase was characterized by an initial fast force increase termed the 'catch-up' response, which apparently compensated for the response delay--the grip force adequately matched the current load demands by the end of the catch-up response. In ramps with longer lasting loading phases (amplitude greater than or equal to 2 N) the catch-up response was followed by a 'tracking' response, during which the grip force increased in parallel with load force and maintained an approximately constant force ratio that prevented frictional slips. The grip force during the hold phase was linearly related to the load force, with an intercept close to the grip force used prior to the loading. Likewise, the grip force responses evoked by the fast loadings superimposed on existing loads followed the same linear relationship.(ABSTRACT TRUNCATED AT 400 WORDS)
1. Single units in striate cortex were studied in alert macaques while they viewed a ganzfeld. Of the 385 well-isolated units studied for 10 min to 2 h, 24% gave "luxotonic" responses, i.e., their rate of discharge for 1 min or more in diffuse, featureless, wideangle illumination (20-450 cd/m2) was at least double that during a comparable period in darkness, or vice versa, and not attributable to eye movements of blinking. Those discharging faster in the light, "photergic" units, outnumber those responding to darkness, "scotergic" units 1 by 4:1. 2. In the lateral geniculate nucleus, on the other hand, among 46 units studied, 28% were luxotonic, but scotergic units were the more common. Both types were present in both magno- and parvocellular laminae. 3. For striate cortex two-thirds of the luxotonic units were binocular. Some showed highly similar response for either eye alone, and essentially no summation binocularly; others had grossly differing responses from each eye, and complex binocular interaction. 4. Many units of all types at striate cortex showed significant modulation of their activity consequent to saccadic eye movements made in darkness, whereas comparable modulation was not observed at the lateral geniculate nucleus. 5. On the basis of these and other findings it is concluded that luxotonic cortical activity is prominent probably only in alert primates, and that this is a consequence of the fact that all retinal ganglion cells in primates synapse in the lateral geniculate nucleus (Ref. 9). Possible functions range from mere trophic input to providing a veridical image or a scaling factor for maintenance of perceptual constancy in the face of varying levels of general illumination.
Activity from muscle afferents regarding ankle kinesthesia was recorded using cuff electrodes in a rabbit preparation in which tactile input from the foot was eliminated. The purpose was to determine if such activity can provide information useful in controlling functional electrical stimulation (FES) systems that restore mobility in spinal injured man. The rabbit's ankle was passively flexed and extended while the activity of the tibial and peroneal nerves was recorded. Responses to trapezoidal stimulus profiles were investigated for excursions from 10 to 60 using velocities from 5 /s to 30 /s and different initial ankle positions. The recorded signals mainly reflect activity from primary and secondary muscle afferents. Dorsiflexion stretched the ankle extensors and produced velocity dependent activity in the tibial nerve, and this diminished to a tonic level during the stimulus plateau. The peroneal nerve was silent during dorsiflexion, but was activated by stretch of the peroneal muscles during ankle extension. The responses of the two nerves behaved in a reciprocal manner, but exhibited considerable hysteresis, since motion that relaxed the stretch to the driving muscle produced an immediate cessation of the prior stretch induced activity. A noted difference between the tibial and peroneal nerve responses is that the range of joint position change that activated the flexor afferents was greater then for the extensor afferents. Ankle rotation at higher velocities increased the dynamic stretch evoked responses during the stimulus ramp but showed no effect on the tonic activity during the stimulus plateau. Prestretching the muscles by altering the initial position increased the response to the ramp movement, however, for the peroneal nerve, when the prestretch brought the flexor muscles near to their maximal lengths, the response to additional stretch provided by the ramp movement was diminished. The results indicate that the whole nerve recorded muscle afferent activity may be useful for control of FES assisted standing, because it can indicate the direction of rotation of the passively moved ankle joint, along with coarse information regarding the rate of movement and static joint position. Index Terms-Functional electrical stimulation (FES), muscle afferents, natural sensors, nerve cuff recordings. I. INTRODUCTION F UNCTIONAL electrical stimulation (FES) can be used to restore function in individuals with paralysis [48]. While the forces generated in muscles activated using FES can be Manuscript
This paper is part of a project whose aim is the implementation of closed-loop control of ankle angular position during functional electrical stimulation (FES) assisted standing in paraplegic subjects using natural sensory information. In this paper, a neural fuzzy (NF) model is implemented to extract angular position information from the electroneurographic signals recorded from muscle afferents using cuff electrodes in an animal model. The NF model, named dynamic nonsingleton fuzzy logic system is a Mamdani-like fuzzy system, implemented in the framework of recurrent neural networks. The fuzzification procedure implemented was the nonsingleton technique which has been shown in previous works to be able to take into account the uncertainty in the data. The proposed algorithm was tested in different situations and was able to predict reasonably well the ankle angular trajectories especially for small excursions (as during standing) and when the stimulation sites are far from the registration sites. This suggests it may be possible to use activity from muscle afferents recorded with cuff electrodes for FES closed-loop control of ankle position during quite standing.
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