Vg. Macefield et Rs. Johansson, CONTROL OF GRIP FORCE DURING RESTRAINT OF AN OBJECT HELD BETWEEN FINGER AND THUMB - RESPONSES OF MUSCLE AND JOINT AFFERENTS FROM THE DIGITS, Experimental Brain Research, 108(1), 1996, pp. 172-184
Pulling or pushing forces applied to an object gripped between finger
and thumb excite tactile afferents in the digits in a manner awarding
these afferents probable roles in triggering the reactive increases in
grip force and in scaling the changes in grip force to the changes in
applied load-force. In the present study we assessed the possible con
tributions from slowly adapting afferents supplying muscles involved i
n the generation of grip forces and from digital joint afferents. Impu
lses were recorded from single afferents via tungsten microelectrodes
inserted percutaneously into the median or ulnar nerves of awake human
subjects. The subject held a manipulandum with a precision grip betwe
en the receptor related digit (index finger, middle finger, ring finge
r or thumb) and an opposing digit (thumb or index finger). Ramp-and-ho
ld load forces of various amplitudes (0.5-2.0 N) and ramp rates (2-32
N/s) were delivered tangential to the parallel grip surfaces in both t
he distal (pulling) and the proximal (pushing) directions. Afferents f
rom the long flexors of the digits (n=19), regardless of their muscle-
spindle or tendon-organ origin, did not respond to the load forces bef
ore the onset of the automatic grip response, even with the fastest ra
mp rates. Their peak discharge closely followed the peak rate of incre
ase in grip force. During the hold phase of the load stimulus, the aff
erents sustained a tonic discharge. The discharge rates were significa
ntly lower with proximally directed loads despite the mean grip-force
being similar in the two directions. This disparity could be explained
by the differing contributions of these muscles to the finger-tip for
ces necessary to restrain the manipulandum in the two directions. Most
afferents from the short flexors of the digits (n=17), including the
lumbricals, dorsal interossei, opponens pollicis, and flexor pollicis
brevis, did not respond at all, even with the fastest ramps. Furthermo
re, the ensemble pattern from the joint afferents (n=6) revealed no si
gnificant encoding of changes in finger-tip forces before the onset of
the increase in grip force. We conclude that mechanoreceptors in the
flexors of the digits and in the interphalangeal joints cannot be awar
ded a significant role in triggering the automatic changes in grip for
ce. Rather, their responses appeared to reflect the reactive forces ge
nerated by the muscles to restrain the object. Hence, it appears that
tactile afferents of the skin in contact with the object are the only
species of receptor in the hand capable of triggering and initially sc
aling an appropriate change in grip force in response to an imposed ch
ange in load force, but that muscle and joint afferents may provide in
formation related to the reactive forces produced by the subject.