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Chapter 2. Prehension

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226 THE PHASES OF PREHENSION<br />

for FA1 unit, a buzzing, wobbling or fluttery sensation was<br />

experienced (Johansson, 1978).<br />

In addition to mechanoreceptors in the skin and subcutaneous<br />

tissues, joint, muscle and tendon receptors are providing<br />

proprioceptive information during contact and manipulation. In<br />

examination of the deep ulnar nerve, Devanandan and colleagues, in<br />

bonnet monkeys (Macaca radiata) found that myelinated sensory fibers<br />

comprised 70% of the total nerve fibers, that 50% of myelinated fibers<br />

in nerves to the intrinsic muscles of the hand are sensory, and further<br />

that 25 - 50% of the fibers in the deep ulnar nerve innervate the joints<br />

of the hand (Devanandan, Ghosh & Simoes, 1980). Followup<br />

anatomical studies demonstrated a paucity of tendon organs in the<br />

intrinsic hand musculature, in contrast to the rich number and density<br />

of muscle spindles (Devanandan, Ghosh & John, 1983). Thus, there<br />

is a large quantum of feedback concerning movement parameters, not<br />

of evolving tension in the intrinsic muscles. They consider the role of<br />

spindles in intrinsic muscles to be, in part, providing rapid feedback in<br />

evolving manipulation so that the intrinsic muscles of the hands can be<br />

appropriately altering the resultant force vectors that act on the<br />

respective digital joints. The muscle spindle afferents in the lumbrical<br />

nerve have been implicated also in perception of direction, distinct<br />

from the perception of movement, given the loss of perception of<br />

direction with injury or administration of local anesthetic<br />

(Devanandan, personal communication, November, 1989).<br />

It appears that there are substantial differences beween<br />

mechanoreceptors in digital joints and more proximal joints, such that<br />

digital joints are specialized particularly for detection of dynamic<br />

mechanical changes, rather than static joint position. In examining the<br />

receptors in the metacarpophalangeal joints of bonnet monkeys and<br />

one male human, Sathian and Devanandan (1983) found a dominance<br />

of dynamic mechanoreceptors: paciniform corpuscles (Type II), a<br />

smaller number of Ruffini endings (Type I) and their restriction to<br />

palmar ligaments, and a complete absence of the static receptors, Golgi<br />

endings (Type 111). Free nerve endings (Type IV) and paciniform<br />

corpuscles were also found close to blood vessels, perhaps to detect<br />

mechanical changes associated with vasodilation. In the<br />

interphalangeal joints, as shown in Figure 6.8, there are some free<br />

nerve endings; however, pacinifondpacinian corpuscles are almost the<br />

exclusive encapsulated mechanoreceptors in the joints. With 3 - 50<br />

lamellae, these were situated in groups, and exclusively in the lateral<br />

and ventral (palmar) aspects of the joint capsule (Babu 8z Devanandan,<br />

1991). A single parent axon may branch and supply endings with

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