Investigation of the Role of Skin and Muscle Receptors in Proprioception at the Ankle Joint in Humans.

This thesis is an investigation of the role of skin and muscle receptors in proprioception at the ankle joint in humans. Somatosensory afferents provide the central nervous system with cues that code for movement and position of the segments, senses collectively known as proprioception. Intramuscular receptors, in particular muscle spindles, code for length and movement velocity of muscles surrounding a joint and therefore play an integral role in proprioception. The role of cutaneous receptors is less clear. When activated via skin stretch they create illusory movements of the limbs, and they are capable of responding to movements of the joints. To what extent these cues are utilized over and above input from muscle spindles remains unknown. In addition, there is evidence that cutaneous receptors may influence the sensitivity of muscle spindles by modulating their level of fusimotor activation. The aim of this thesis was to further examine the role of skin in proprioception and to determine whether or not skin of the foot and ankle is capable of modulating fusimotor drive to muscle spindles of the lower limb. The current thesis is comprised of three studies. The first experiment utilized a matching task at the ankle joint and determined that skin from the dorsum of the foot and ankle is necessary for accurate proprioception. The remaining two experiments used the technique of microneurography to record from single nerve afferents in awake, human participants. Initially, cutaneous afferents were isolated and recorded to determine the efficacy of using cooling over their receptive field as a method to decrease their sensitivity and output. Once cooling was established as an effective tool, the final experiment isolated and recorded from muscle spindles in response to passive, ramp and hold movements at the ankle. It was determined that a reduction in skin input (via cooling) altered the firing response of a portion of spindles. It is likely that this change in firing was due to modulation of fusimotor drive to the spindles. Collectively, the current work contributes the novel findings that skin on the dorsum of the foot is necessary for accurate proprioception at the ankle and that this is largely due to the role of skin as an independent proprioceptive channel. In addition, we have shown for the first time that a reduction in skin input from the foot dorsum is capable of modulating spindle discharge during a passive ramp and hold movement at the ankle, demonstrating a minor role for this interaction in proprioception. A secondary finding of the thesis was that cooling with ice is an effective tool for reducing input from all four classes of cutaneous mechanoreceptors.