Physiological characteristics of low-threshold mechanoreceptors in joints, muscle and skin in human subjects.

1. The development of microneurography, in which an insulated tungsten microelectrode is inserted into an accessible peripheral or cranial nerve in awake human subjects, has allowed detailed analyses of the signalling capacities of single mechanoreceptive afferents from the skin, muscles and joints. For example, we know much about how the two classes of rapidly adapting (Meissner and Pacinian) and two classes of slowly adapting (Merkel and Ruffini) cutaneous mechanoreceptors encode forces applied normal or tangential to the skin of the hand and the similarities and differences in glabrous versus non-glabrous skin (and receptors associated with hairs). We also know about stretch- and force-sensitive endings in muscle (the muscle spindle and Golgi tendon organ, respectively) and how they behave during passive or active movements or during isometric contractions. In addition, we have characterized the firing properties of mechanoreceptors in the joint capsules of the fingers. However, we know little about sensory nerves in the periosteum, other than that nociceptors and Pacinian corpuscles exist. 2. In addition to studies on the physiology of sensory endings in human subjects, microstimulation through the recording microelectrode has revealed how the brain deals with the sensory information conveyed by a single afferent. From this work, we know that there is specificity in the sensory channels: electrical stimulation of a single Meissner or Pacinian corpuscle generates frequency dependent illusions of 'flutter' or 'vibration', whereas microstimulation of a single Merkel afferent can produce a percept of 'pressure' and stimulation of a single joint afferent can evoke a sensation of 'joint rotation'. Interestingly, the input from a single Ruffini ending in the skin cannot be perceived and the same is true of muscle spindle afferents. So, where does this leave us with osseoperception from the mouth? Given that the periodontal receptors in the vicinity have been lost, which mechanoreceptive endings could encode forces applied to a bone-implanted prosthesis? 3. Meissner and Merkel endings have very small receptive fields and respond only to local forces. Pacinian corpuscles have an exquisite sensitivity to brisk mechanical events and could respond to such stimuli transmitted through the bone to a remote receptor, but would not be able to encode sustained forces. Ruffini endings also respond to forces applied remote to the receptive field and, unlike the Pacinian corpuscles, respond in a sustained fashion, but would their signals be perceived? Like muscle spindles, it is possible that the coactivation of many Ruffini endings could provide meaningful information. Finally, as we have seen, the input from a single joint receptor can be perceived, but they mostly respond at the limits of joint rotation, so it is unlikely that any associated with the temporomandibular joint could contribute to osseoperception.