Differential effects of radiant and mechanically applied thermal stimuli on human C-tactile afferent firing patterns.

C-tactile (CT) afferents respond to gentle tactile stimulation, but only a handful of studies in humans and animals have investigated whether their firing is modified by temperature. We describe the effects of radiant thermal stimuli, and of stationary and very slowly moving mechanothermal stimuli, on CT afferent responses. We find that CT afferents are primarily mechanoreceptors, as they fired little during radiant thermal stimuli, but they exhibited different patterns of firing during combined mechano-cool stimulation compared with warming.

Microneurography: how it started and how it works.

In the first section, this historical review describes endeavors to develop the method for recording normal nerve impulse traffic in humans, designated microneurography. The method was developed at the Department of Clinical Neurophysiology of the Academic Hospital in Uppsala, Sweden. Microneurography involves the impalement of a peripheral nerve with a tungsten needle electrode.

Somatotopic organization of gentle touch processing in the posterior insular cortex.

A network of thin (C and A delta) afferents relays various signals related to the physiological condition of the body, including sensations of gentle touch, pain, and temperature changes. Such afferents project to the insular cortex, where a somatotopic organization of responses to noxious and cooling stimuli was recently observed. To explore the possibility of a corresponding body-map topography in relation to gentle touch mediated through C tactile (CT) fibers, we applied soft brush stimuli to the right forearm and thigh of a patient (GL) lacking A beta afferents, and six healthy subjects during functional magnetic resonance imaging (fMRI).

The neurophysiology of unmyelinated tactile afferents.

CT (C tactile) afferents are a distinct type of unmyelinated, low-threshold mechanoreceptive units existing in the hairy but not glabrous skin of humans and other mammals. Evidence from patients lacking myelinated tactile afferents indicates that signaling in these fibers activate the insular cortex. Since this system is poor in encoding discriminative aspects of touch, but well-suited to encoding slow, gentle touch, CT fibers in hairy skin may be part of a system for processing pleasant and socially relevant aspects of touch.

Unmyelinated tactile afferents have opposite effects on insular and somatosensory cortical processing.

A previous functional magnetic resonance imaging (fMRI) study of an A-beta deafferented subject (GL) showed that stimulation of tactile C afferents (CT) activates insular cortex whereas no activation was seen in somatosensory cortices. Psychophysical studies suggested that CT afferents contribute to affective but not to discriminative aspects of tactile stimulation. We have now examined cortical processing following CT stimulation in a second similarly deafferented subject (IW), as well as revisited the data from GL.

Discriminative touch and emotional touch.

Somatic sensation comprises four main modalities, each relaying tactile, thermal, painful, or pruritic (itch) information to the central nervous system. These input channels can be further classified as subserving a sensory function of spatial and temporal localization, discrimination, and provision of essential information for controlling and guiding exploratory tactile behaviours, and an affective function that is widely recognized as providing the afferent neural input driving the subjective experience of pain, but not so widely recognized as also providing the subjective experience of affiliative or emotional somatic pleasure of touch. The discriminative properties of tactile sensation are mediated by a class of fast-conducting myelinated peripheral nerve fibres--A-beta fibres--whereas the rewarding, emotional properties of touch are hypothesized to be mediated by a class of unmyelinated peripheral nerve fibres--CT afferents (C tactile)--that have biophysical, electrophysiological, neurobiological, and anatomical properties that drive the temporally delayed emotional somatic system.

Functional role of unmyelinated tactile afferents in human hairy skin: sympathetic response and perceptual localization.

In addition to A-beta fibres the human hairy skin has unmyelinated (C) fibres responsive to light touch. Previous functional magnetic resonance imaging (fMRI) studies in a subject with a neuronopathy who specifically lacks A-beta afferents indicated that tactile C afferents (CT) activate insular cortex, whereas no response was seen in somatosensory areas 1 and 2. Psychophysical tests suggested that CT afferents give rise to an inconsistent perception of weak and pleasant touch.

Unmyelinated tactile afferents underpin detection of low-force monofilaments.

Human hairy but not glabrous skin has unmyelinated (C) tactile (CT) afferents that project to insular cortex. We studied two subjects with the rare sensory neuronopathy syndrome who lack A-beta fibers but have relatively preserved C-fiber function. Weak monofilaments were detected on hairy skin alone.

Microneurography: how the technique developed and its role in the investigation of the sympathetic nervous system.

A historical review is given of the development of microneurography and its application for studies of sympathetic nerve activity in humans.

Receptive field properties of unmyelinated tactile afferents in the human skin.

We recorded, with the microneurography technique, single-unit impulses from nine cutaneous mechanoreceptive afferents with conduction velocities in the C range and receptive fields in the hairy skin of the forearm. The units responded with high impulse rates to light touch and had low monofilament thresholds. The geography of receptive fields was explored with a scanning method: a lightweight probe with a small and rounded tip was made to scan the field area in a series of closely adjacent tracks while single-unit activity was recorded.

Unmyelinated tactile afferents signal touch and project to insular cortex.

There is dual tactile innervation of the human hairy skin: in addition to fast-conducting myelinated afferent fibers, there is a system of slow-conducting unmyelinated (C) afferents that respond to light touch. In a unique patient lacking large myelinated afferents, we found that activation of C tactile (CT) afferents produced a faint sensation of pleasant touch. Functional magnetic resonance imaging (fMRI) analysis during CT stimulation showed activation of the insular region, but not of somatosensory areas S1 and S2.

Proprioceptive feedback is reduced during adaptation to a visuomotor transformation: preliminary findings.

Adapting movements in relation to visual feedback is a ubiquitous characteristic of sensorimotor control and involves the integration of multiple sources of sensory information. We recorded sensory feedback from muscle spindle afferents during visuomotor adaptation while subjects performed an aiming task to investigate whether the activity of the muscle spindles was modulated by the fusimotor system under these learning conditions. None of the muscle spindles showed an increase in activity, rather in 83% of the trials the firing rates were decreased.

Directional tuning of human forearm muscle afferents during voluntary wrist movements.

1. Single unit activity was recorded with the microneurography technique from sixteen spindle afferents and one Golgi tendon organ afferent originating from the forearm extensor muscles. Impulse rates were studied while subjects performed unobstructed aiming movements at the wrist in eight different directions 45 deg apart.

Unmyelinated afferents constitute a second system coding tactile stimuli of the human hairy skin.

Impulses were recorded from unmyelinated afferents innervating the forearm skin of human subjects using the technique of microneurography. Units responding to innocuous skin deformation were selected. The sample (n = 38) was split into low-threshold units (n = 27) and high-threshold units (n = 11) on the basis of three distinctive features, i.

Is human muscle spindle afference dependent on perceived size of error in visual tracking?

Impulses of 16 muscle spindle afferents from finger extensor muscles were recorded from the radial nerve along with electromyographic (EMG) activity and kinematics of joint movement. Twelve units were classified as Ia and 4 as II spindle afferents. Subjects were requested to perform precision movements at a single metacarpophalangeal joint in an indirect visual tracking task.

Pulsatile motor output in human finger movements is not dependent on the stretch reflex.

1. Stretch perturbations were delivered during slow voluntary finger movements with the aim of exploring the role of the stretch reflex in generating the 8-10 Hz discontinuities that characterize these movements. Afferent activity from muscle spindle primary endings in the finger extensor muscles was recorded from the radial nerve, along with the EMG activity of these muscles, and kinematics of the relevant metacarpo-phalangeal joint.

Fusimotor and skeletomotor activities are increased with precision finger movement in man.

1. Impulses of eighteen muscle spindle afferents from finger extensor muscles were recorded from the radial nerve while subjects performed single joint finger movements of two kinds, i.e.

Coding of pulsatile motor output by human muscle afferents during slow finger movements.

1. Impulse activities of thirty-eight muscle spindle and tendon organ afferents from the finger extensor muscles were recorded in the radial nerve of human subjects while the subjects performed voluntary flexion and extension finger movements at a single metacarpophalangeal joint. 2.

Receptive field characteristics of tactile units with myelinated afferents in hairy skin of human subjects.

1. Impulses in single nerve fibres from the lateral antebrachial cutaneous nerve were recorded using the microneurography technique in human subjects. 2.

Human muscle spindle afferent activity in relation to visual control in precision finger movements.

1. Impulse activities of muscle spindle afferents from the finger extensor muscles were recorded in the radial nerve of human subjects. In addition to single unit activity, surface EMG was recorded as well as finger joint position and angular velocity.

The discharge behaviour of single vasoconstrictor motoneurones in human muscle nerves.

1. The discharge behaviour of fourteen single sympathetic vasoconstrictor efferents was studied using a tungsten microelectrode inserted percutaneously into a motor fascicle of the radial or peroneal nerve in eight awake supine subjects. Units were classified as vasoconstrictor because their firing properties correlated appropriately to changes in cardiac interval and arterial pressure.

A system of unmyelinated afferents for innocuous mechanoreception in the human skin.

It is generally held that tactile mechanisms in the human skin are served by fast-conducting myelinated nerve fibres, whereas touch-sensitive afferents with unmyelinated axons are lacking in man, in contrast to other mammals. In the present study we found evidence that sensitive mechanoreceptive afferents with unmyelinated fibres are quite common and widespread in the hairy skin of human subjects. Their biological role remains an enigma which might attract more attention now that their existence in man has been demonstrated.

Organization of motor output in slow finger movements in man.

1. Slow finger movements were analysed in normal human subjects with regard to kinematics and EMG activity of the long finger muscles. Surface EMG from the finger extensor and flexor muscles on the forearm was recorded along with angular position and angular velocity during voluntary ramp movements at single metacarpophalangeal joints.