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Chapter 2. Prehension

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<strong>Chapter</strong> 6 - During Contact 259<br />

50 - 70 ms after onset of load force perturbations, with a silent period<br />

preceding the phasic burst, 30 - 40 ms after perturbation onset (in all<br />

muscles recorded except extensor digitorum communis). Note the<br />

similarity in latency to the ‘automatic’ adjustments in the grip-<br />

force/load-force ratio to slips reported by Johansson and Westling<br />

(1988a). Subjects never made an error in the direction of the<br />

automatic, short latency, response, e.g., a rapid increase in load force<br />

always yielded an increase in grip force. The increase in grip force<br />

averaged 7.2 (f 2) N and was within 6 - 10% of the grip force needed<br />

to lift the object. In contrast to the sustained response to object slip,<br />

the grip force in response to load perturbation steadily decreased (to<br />

the level necessary to maintain grasp). The grip force response to<br />

perturbation was sensitive to the size and velocity of the load force<br />

increase but not the preexisting grip force; thus, Cole and Abbs<br />

concluded that the grip force was at a level to maintain grasp, i.e.,<br />

proportional to the required load force, not at a specified level above<br />

the object’s slip point. They note the impairment with digital<br />

anesthesia, and consider the role of cutaneous, joint and muscle<br />

mechanoreceptors in signalling the load changes, with emphasis on<br />

cutaneous mechanoreceptors detecting shearing forces on the skin of<br />

the finger pads.<br />

Johansson and colleagues (Johansson et al., 1992a,b,c)<br />

performed a series of load perturbation experiments, using pad<br />

opposition. In the absence of instructions to initiate grip changes, they<br />

showed that the latency of the initial grip force responses, termed<br />

‘catch-up response’ was dependent on load force rate (e.g., 8W,<br />

108+13, 138f27 and 174f39 ms for the 32, 8, 4, and 2 N/s rates<br />

respectively). The latency of grip force response to different loads<br />

(with a constant load force rate of 4 N/s) was 140 (f30) ms. The<br />

amplitude of the grip force response was a function of both load force<br />

and load force rate, but its time course was similar over all rates of<br />

perturbing load force. For longer lasting loading phases, the catch-up<br />

response was followed by a ‘tracking’ response, in which grip force<br />

increased in parallel with load force, to maintain a slip ratio.<br />

Interestingly, with digital anaesthesia, subjects had to voluntarily<br />

attend to the task (usually automatic, without anesthesia), the grip<br />

force latencies were prolonged (to several hundred milliseconds), with<br />

less modulation of grip force, absence of grip force responses (absent<br />

from 25 - 93% of trials, depending on subject), and there was a lack<br />

of dependence of the ‘catch-up’ grip force responses on load force<br />

rate. With digital anaesthesia, there were large individual differences,<br />

suggesting that for some subjects, more proximal mechanoreceptors

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