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Chapter 2. Prehension

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260 THE PHASES OF PREHENSION<br />

might be mediating grip force responses. There was a lack of the<br />

‘tracking response’ with digital anaesthesia as well, confirming the<br />

requirement of continuous afferent input from both thumb and index<br />

finger for grip force regulation in pad opposition. Contrary to their<br />

initial expectations, there appeared to be independent force control for<br />

each finger. The cutaneous strain at digital pulps resulting from<br />

increased shear forces was believed to be the most relevant mechanical<br />

stimulus of the digital mechanoreceptors. Johansson and colleagues<br />

also note extension of the interphalangeal joints and flexion of the<br />

metacarpophalangeal joint of the index finger as possible stimuli for<br />

mechanoreceptors, since these appeared to aid grip force modulation<br />

with digital anesthesia.<br />

To investigate the influence of task instructions on loading or<br />

unloading perturbations, Winstein, Abbs and Petashnick (1991), prior<br />

to each block of trials, verbally told subjects to ‘hold’ or ‘let go’ in<br />

response to any apparent change in object weight. Thus they had a set<br />

to ‘hold’ or ‘let go’. The latencies of evoked grip changes, with a<br />

mean of 66 (+13) ms and as early as 35 ms after perturbation onset,<br />

were unaffected by either instructions or direction of loading. Grip<br />

force responses to unloading were unaffected by instructions; in<br />

contrast, with loading, the magnitude of early grip responses were<br />

27% higher following hold than following let go instructions. In hold<br />

conditions, the post-perturbation grip force level was sustained. In let<br />

go conditions, a second grip force response was observed with<br />

median latencies of 239-401 ms, across subjects. The latency of the<br />

let go response did not vary with unloading or loading conditions.<br />

They noted that the magnitude of the grip force responses were much<br />

more marked (on average 3.6 times higher) to loading than unloading<br />

forces of equivalent magnitude, suggesting this reflects functional<br />

evolution of the hand in a gravity dominant world. Winstein et al.<br />

note the importance of balancing two functional goals: first, to ensure<br />

the object is not dropped or to maintain the object in stable grasp (not<br />

too little gripping force) and second, to ensure the object is not<br />

crushed or broken (not too much gripping force).<br />

In determining the slip ratio in pad opposition, during the time<br />

when individuals med to voluntarily release their grasp, they felt as if<br />

their fingers adhered to the object (Johansson and Westling, 1987,<br />

1990). It is as if the cutaneous afferents and peripheral biomechanics<br />

to grasp are in contradiction to the efferent commands to release. The<br />

effort required to overcome this magnet phenomenon was greater<br />

using sandpaper as the surface structure, and became stronger as the<br />

grip force approached the minimum grip force to prevent slips. They

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