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Hall Of Fames Human Beings that you must know!

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Old 18-05-2009, 05:19 PM   #1
Human Primate Social Groomer and Neuroelastician
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Default Hakan Olausson

I think this Swedish researcher deserves a place in our Hall of Fame. For some strange reason, he became interested in, and has worked tirelessly for decades humbly studying, humble unmyelinated afferents, tracing them to the ends of their lines in the brain (mostly insular cortex, right and left) and their behaviour when excited. There is a list of 36 publications for "Olausson H" in Pubmed. But pubmed won't let me bring you a simple link to the list. No....

His work is likely some of the most relevant to what we do with our hands on other peoples' skin. As far as I'm concerned at least - lateral skin stretch and all. When our hands are on the skin we really are affecting the brain, through these unmyelinated afferents.

I now have a better appreciation for what the organism/ectoderm is trying to do, when it kicks off neural crest, just before it turns into skin. It's as though it were trying to integrate itself throughout the entire organism; not content with just being related to brain - it tries to re-invade brain as well as mesoderm. All I can say is wow.

My pet project for the summer (if I can manage it) will be to carefully read through all these and think about how all of it fits together. Here is a blogpost I wrote yesterday on apotemnophilia, after feeling a few adjacent neurons go pop in my own brain. (Entire article by Colapinto on Ramachandran can be found here. Once more, Deric at Mingblog activated my cortex.)

Here is an old news story about his research, from CBC, called "Researchers 'feel out' pleasure nerves in humans." It has a little picture of him bending over an MRI machine.

Unfortunately, he seems to not have his own website, and the only other news story I can find is called Pleasurable Skin Contact (a very shallow treatment indeed!) of his work.

I think there might be more than one H Olausson, unless the one I'm interested in has two or three main lines of research.. bones and fetuses and unmyelinated afferents..

Anyway, here is a list of abstracts (my bolds throughout) relating to afferents, oldest papers first, so that I can add newer ones to the list at the bottom as they come along:

1. Human, tactile, directional sensibility and its peripheral origins. 1992 (I can't access this.)
Tactile directional sensibility is probably functionally important and deserves attention as it is known to be sensitive to many different disturbances of the somatosensory system. Therefore, the ability of healthy adults to determine the direction of motion of a light tactile stimulus travelling proximally or distally along a straight line on depilated, hairy skin of the forearm was examined with two-alternative, forced-choice technique. The aim was to investigate the relative importance of different types of afferent information which may be used for this purpose. A test was started with the moving stimulus covering a distance of no less than 2.5 mm, which was subsequently increased until the subject could report the direction of motion reliably. Afterwards, the distance was decreased until the subject could no longer do so. Three different stimulation conditions were used and for a point stimulator touching the skin it was found that the necessary distance decreased to 2.5 mm after a moderate increase of the vertical contact load. No such decrease was found when a frictionless air-stream point stimulator was used instead. The distances which had to be covered by the point stimulator touching the skin increased to values which were comparable to those obtained with the air-stream stimulator after the lateral extensibility of the skin had been diminished. This was achieved by attaching a surgical sticky plaster around the stimulated skin area. The present findings consequently indicated that optimal, tactile, directional sensitivity depends on peripheral afferent messages which signal the direction of lateral stretching of the skin.
2. Observations on human tactile directional sensibility. 1993 (Link to full text)
1. The ability to tell the direction of a motion across the skin deserve attention for being an easily observed function which provides a sensitive test for disturbances of the peripheral and central nervous systems. The mode of operation, on the other hand, of this tactile directional sensibility is still uncertain. 2. The dependence of directional sensibility on the contact load and distance of movement of a blunt metal tip, has now been determined for the skin of the forearm of normal subjects with the two-alternative forced-choice method. The testing was done under two conditions: elbow bent or straight. Straightening of the arm always reduced the accuracy of the directional sensibility. It also caused measurable changes of cutaneous mechanical properties, which presumably decreased the reliability of afferent information about lateral distension. 3. The average accuracy of the directional sensibility was found to be correlated linearly to the logarithm of the contact load, and straightening of the arm decreased the accuracy for each load by corresponding amounts. Similar relationships were found between the accuracy and the distance of movement. 4. Straightening of the arm did not cause any significant average reduction of the contact threshold for point stimulation of the same receptive field. A consistently lowered contact sensitivity, however, was observed for some of the subjects, which may have contributed to the reduction of the directional sensibility in these cases. 5. Correct directional estimations of the movement of the metal tip were obtained for a distance which was a fifth of the shortest distance for a corresponding estimation of the movement of a frictionless stimulus. The findings thus indicated that the friction between a moving object and the underlying skin, which can be mediated via stretch-sensitive cutaneous receptors, is critical for the determination of its direction of motion. 6. The present observations and previous observations by various authors are suggested to indicate that typical tactile directional sensibility depends on parallel processing of direction-selective data, and spatial data expressed as a function of time.
3. A system of unmyelinated afferents for innocuous mechanoreception in the human skin. 1993 (Link to paper)
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.
4. The influence of spatial summation on human tactile directional sensibility. 1994 (Can't get this one)
Spatial summation is known to influence the magnitude of sensation for stationary cutaneous stimuli. Yet analysis of moving stimuli may also be pertinent, since most stimuli that attract our attention involve movements over the skin surface. The present investigation dealt with the importance of spatial summation for the appreciation of the direction of motion for moving stimuli. The ability to detect the direction of motion was tested on the radial surface of the forearm with the two-alternative forced-choice method. Stimulation was performed with a rolling wheel, in order to exclude friction-generated activation of stretch receptors. Each subject was tested with two wheels with the same radius but different widths, 1 mm and 15 mm. On average, the subjects performed better with the wide wheel than with the narrow one for stimulation distances > or = 16 mm. This value also probably exceeds the threshold distance for directional discrimination for the narrow wheel, which indicates that spatial summation improves suprathreshold performance.
5. Spatial cues serving the tactile directional sensibility of the human forearm]Receptive field characteristics of tactile units with myelinated afferents in hairy skin of human subjects.. 1994 (see article here)
1. Tactile directional sensibility is considered to rely on the parallel processing of direction-contingent sensory data that depend on skin stretching caused by friction, and spatial cues that vary with time. A temperature-controlled airstream stimulus that prevented the activation of stretch receptors was used to investigate directional sensibility for the skin of the forearm. 2. The dependence on contact load and distance of movement was determined for normal subjects with a two-alternative forced-choice method. Testing was performed under two conditions, elbow bent or straight. Bracing the skin by straightening the arm did not alter the accuracy of the directional sensibility, in contrast to previous findings with stimuli that caused friction. 3. The accuracy of directional sensibility was correlated linearly to the logarithm of the distance of movement of the air jet. No correlation was found between accuracy and contact load, unlike findings with stimuli that cause friction. 4. Measurements were made with different subjects to determine the threshold distance at constant load. On average, subjects were able to distinguish direction with movements of < or = 8 mm. This acuity is sharper than has been reported with static stimuli. There was no correlation between subjects' threshold distances for judging direction and spatial acuity measured with absolute point localization. 5. The ability to distinguish direction was poor for the airstream stimulus compared with stimuli causing frictional contact with hairy skin. Nevertheless, the present findings are consistent with the suggestion that cutaneous spatial acuity is better for dynamic than for static stimuli.
6. 1995 (full text free access)
1. Impulses in single nerve fibres from the lateral antebrachial cutaneous nerve were recorded using the microneurography technique in human subjects. 2. In a sample of fifty-five mechanoreceptive units with fast-conducting nerve fibres, five types were identified, i.e. SAI (slowly adapting type I, Merkel), SAII (slowly adapting type II, Ruffini), hair units, field units and Pacinian-type units. The latter three unit types were all rapidly adapting. 3. The detailed structure of thirty-five receptive fields of SAI, SAII, hair and field units was explored with a method which was objective and independent of the experimenter's skill and experience. A lightweight probe was used to scan the receptive field area in a series of tracks 0.23 mm apart while single-unit activity was recorded. 4. SAI fields were small and composed of two to four well-separated high-sensitivity spots and often, in addition, one minor spot of lower sensitivity. SAII units typically fired spontaneously at a low and regular rate. Most fields consisted of one single spot of high sensitivity with diffuse borders. The hair units innervated ten to thirty-three (or more) hairs, which were evenly distributed over a large area. The field units were characterized by a number of small and closely packed high-sensitivity spots with diffuse borders. A conservative estimate indicated eleven spots per unit. 5. The findings indicate that the sheet of mechanoreceptors on the skin of the forearm is distinctly different from that on the dorsum of the hand and in the face. It seems reasonable to assume that the former is more representative for the hairy skin covering the main parts of the body.
7. Directional sensibility for quantification of tactile dysfunction. 1997 (Link to paper)
Examination of tactile directional sensibility, i.e., the ability to tell the direction of an object's motion across the skin, has been recommended by several authors for examination of patients with somatosensory disorders. Recent findings about the physiological mechanisms underlying directional sensibility suggested possibilities to further improve the test. In the present investigation a test was constructed that allowed a semiquantification of the directional sensibility of six body areas within 20 min. Normal values were obtained by testing healthy subjects (n = 40), and the normal values were compared to those obtained in a group of patients with tactile symptoms (n = 20). Ten of the patients had abnormal sensory conduction in one or several nerves, and they also had abnormal directional sensibility. Hence, examination of directional sensibility, according to the present protocol, provides a semiquantitative test that appears to be as sensitive as electrophysiological measurement of conduction in detecting dysfunction in tactile nerves.
8. Remarkable capacity for perception of the direction of skin pull in man. 1998 (Link to paper)
We determined the ability to appreciate the direction of a skin pull caused by a moving pin that was glued to the forearm skin. A majority of the subjects were able to tell the direction of pin movements with an excursion of 0.13 mm (>/=66% correct responses, p<0.05). Local skin anaesthesia showed that stretch sensitive receptors located over 15 mm in front and behind the pin correctly signalled the direction of these minute skin pulls. It was concluded that information about patterns of skin stretch is an important component of the somatosensory system that may contribute not only to kinaesthetic, but also to cutaneous sensations.
9. Unmyelinated afferents constitute a second system coding tactile stimuli of the human hairy skin. 1999 (I can't get this one.)
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.e., thresholds to skin deformation, size of response to innocuous skin deformation, and differential response to sharp and blunt stimuli. The low-threshold units provisionally were denoted tactile afferents on the basis of their response properties, which strongly suggest that they are coding some feature of tactile stimuli. They exhibited, in many respects, similar functional properties as described for low-threshold C-mechanoreceptive units in other mammals. However, a delayed acceleration, not previously demonstrated, was observed in response to long-lasting innocuous indentations. It was concluded that human hairy skin is innervated by a system of highly sensitive mechanoreceptive units with unmyelinated afferents akin to the system previously described in other mammals. The confirmation that the system is present in the forearm skin and not only in the face area where it first was identified suggests a largely general distribution although there are indications that the tactile C afferents may be lacking in the very distal parts of the limbs. The functional role of the system remains to be assessed although physiological properties of the sense organs invite to speculations that the slow tactile system might have closer relations to limbic functions than to cognitive and motor functions.
10. Our second touch system: Receptive field properties of unmyelinated tactile afferents in man. 1999 (no abstract) (Paper here)

11. Tactile directional sensibility: peripheral neural mechanisms in man. 2000 (Link to paper)
Tactile directional sensibility, i.e. the ability to tell the direction of an object's motion across the skin, is an easily observed sensory function that is highly sensitive to disturbances of the somatosensory system. Based on previous psychophysical experiments on healthy subjects it was concluded that directional sensibility depends on two kinds of information from cutaneous mechanoreceptors; spatio-temporal information and information about friction-induced changes in skin stretch. In the present study responses to similar probe movements as in the psychophysical experiments were recorded from human single mechanoreceptors in the forearm skin. All slowly adapting type 2 (SA2) units were spontaneously active, and with increasing force of friction their discharge rates were modified by probe movements at increasing distances from the Ruffini end-organ, reflecting the high stretch-sensitivity of these units. Slowly adapting type 1 (SA1) and field units responded to the moving probe within well-defined skin areas directly overlying the individual receptor terminals, and compared to the SA2 units their response properties were less dependent on the force of friction. The results suggest that SA1 and field units have the capacity to signal spatio-temporal information, whereas a population of SA2 units have the capacity to signal direction-specific information about changes in lateral skin stretch.
12. Cortical activation by tactile and painful stimuli in hemispherectomized patients. 2001 (Link to full text online)
Hemispherectomized patients are able to perceive tactile and painful stimuli on their nonparetic as well as paretic body halves. We have used functional MRI to study the cortical mechanisms underlying this preserved somatosensory capacity. Nonpainful brushing and painful heat were applied to the skin of the legs in four hemispherectomized patients and, for comparison, in four normal subjects. Cortical activation was studied with a 1.5 T scanner using a BOLD (blood oxygen level dependent) protocol. All patients rated both the brushing and the heat pain as almost equally intense on each leg and the ratings were similar to those in normals. Brushing on the nonparetic leg activated primary and secondary somatosensory cortices (S1 and S2) in all patients, similar to findings in normals. Brushing on the paretic leg activated S1 in two patients and S2 in one of these patients. Heat pain activated S2, insular cortex and anterior cingulate cortex to a similar degree for both legs, but the activation was weaker in the patients than in the normals. For the individual patient, there was generally no obvious correlation between cortical activation as studied with the BOLD technique and psychophysical performance. The findings from tactile stimulation of the nonparetic leg, that the activation was similar to the contralateral activation in normals, suggest that tactile information processing in the hemisphere contralateral to the stimulation is independent of the corpus callosum. In contrast, the pain activation for the nonparetic leg was weaker than in normals, suggesting that pain activation in the hemisphere contralateral to the stimulation is dependent on transcallosal information processing. The latter finding was corroborated by a subnormal capacity for pain localization on the nonparetic foot in two of the patients. The findings from stimulation of the paretic leg show that areas typically involved in the processing of tactile and painful stimuli can be activated by ipsilateral pathways directly from the periphery. The tactile-evoked ipsilateral S1 activation may be due to subcortical reorganization, since it was not observed in the normal subjects.
13. Central pain in a hemispherectomized patient. 2001 (Link to paper here)
We have examined a hemispherectomized patient who complained of touch-evoked pricking and burning pain in her paretic hand, especially when the hand was cold. Psychophysical examination showed that for the paretic side she confused cool and warm temperatures, and confirmed that she had a robust allodynia to brush stroking that was enhanced at a cold ambient temperature. Functional magnetic resonance imaging (fMRI) showed that during brush-evoked allodynia, brain structures implicated in normal pain processing (viz. posterior part of the anterior cingulate cortex, secondary somatosensory cortex, and prefrontal cortices) were activated. The fMRI findings thus indicate that the central pain in this patient was served by brain structures implicated in normal pain processing. Possible pathophysiological mechanisms include plasticity as well as thalamic disinhibition.
14. Tactile directional sensibility and diabetic neuropathy. 2001 (Link to paper here)
Five different procedures used to diagnose neuropathy were compared in a "blind" study with diabetic patients. The aim was to evaluate tests of tactile directional sensibility. Three matched groups were examined, two groups with type I diabetes, either with or without suspected neuropathy, and one of healthy controls. Testing consisted of: (1) examination by a specialist in neurology, (2) electrophysiologic measurement of nerve conduction velocity and determination of cool sensitivity, and (3) determination of directional sensibility in two stages, with categorical and quantitative techniques. Abnormal test results were obtained for both groups of diabetic patients. Quantitatively measured directional sensibility had the highest sensitivity (89%) and specificity (85%) when calculated for patients who had received a diagnosis of neuropathy from the neurologist, despite one case of abnormal directional sensibility among the healthy controls. Conduction velocity testing was almost comparably sensitive (80%) and cool sensitivity, comparably specific (85%) when calculated in the same manner.
15. Unmyelinated tactile afferents signal touch and project to insular cortex. 2002 (Link to paper)
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. These findings identify CT as a system for limbic touch that may underlie emotional, hormonal and affiliative responses to caress-like, skin-to-skin contact between individuals.
16. Receptive field properties of unmyelinated tactile afferents in the human skin. 2003 (Link to free paper online)
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. The fields of the nine units varied considerably in size as well as complexity. The individual field consisted of one to nine small responsive spots distributed over an area of 1-35 mm(2) when explored with a moving indentation of 5 mN. The fields were roughly round or oval in shape with no preferred orientation. The size of the response differed between individual sensitive spots in a field, suggesting a highly nonuniform terminal organization. The properties of the fields seem consistent with a role of tactile C afferents to provide information about pleasant touch and skin-to-skin contacts to central structures controlling emotions and affiliative behavior.
16. Sumatriptan (5-HT1B/1D-agonist) causes a transient allodynia. 2004 (Link to paper)
Unpleasant sensory symptoms are commonly reported in association with the use of 5-HT1B/1D-agonists, i.e. triptans. In particular, pain/pressure symptoms from the chest and neck have restricted the use of triptans in the acute treatment of migraine. The cause of these triptan induced side-effects is still unidentified. We have now tested the hypothesis that sumatriptan influences the perception of tactile and thermal stimuli in humans in a randomized, double-blind, placebo-controlled cross-over study. Two groups were tested; one consisted of 12 (mean age 41.2 years, 10 women) subjects with migraine and a history of cutaneous allodynia in association with sumatriptan treatment. Twelve healthy subjects (mean age 38.7 years, 10 women) without migraine served as control group. During pain- and medication-free intervals tactile directional sensibility, perception of dynamic touch (brush) and thermal sensory and pain thresholds were studied on the dorsal side of the left hand. Measurements were performed before, 20, and 40 min after injection of 6 mg sumatriptan or saline. Twenty minutes after injection, sumatriptan caused a significant placebo-subtracted increase in brush-evoked feeling of unpleasantness in both groups (P < 0.01), an increase in brush-evoked pain in migraineurs only (P = 0.021), a reduction of heat pain threshold in all participants pooled (P = 0.031), and a reduction of cold pain threshold in controls only (P = 0.013). At 40 min after injection, no differences remained significant. There were no changes in ratings of brush intensity, tactile directional sensibility or cold or warm sensation thresholds. Thus, sumatriptan may cause a short-lasting allodynia in response to light dynamic touch and a reduction of heat and cold pain thresholds. This could explain at least some of the temporary sensory side-effects of triptans and warrants consideration in the interpretation of studies on migraine-induced allodynia.
17. Tactile functions after cerebral hemispherectomy. 2005 (Link to paper)
Patients that were hemispherectomized due to brain lesions early in life sometimes have remarkably well-preserved tactile functions on their paretic body half. This has been attributed to developmental neuroplasticity. However, the tactile examinations generally have been fairly crude, and subtle deficits may not have been revealed. We investigated monofilament detection and three types of tactile directional sensibility in four hemispherectomized patients and six healthy controls. Patients were examined bilaterally on the face, forearm and lower leg. Normal subjects were examined unilaterally. Following each test of directional sensibility, subjects were asked to rate the intensity of the stimulation. On the nonparetic side, results were almost always in the normal range. On the paretic side, the patients' capacity for monofilament detection was less impaired than their directional sensibility. Despite the disturbed directional sensibility on their paretic side the patients rated tactile sensations evoked by the stimuli, on both their paretic and nonparetic body halves, as more intense than normals. Thus, mechanisms of plasticity seem adequate for tactile detection and intensity coding but not for more complex tactile functions such as directional sensibility. The reason for the high vulnerability of tactile directional sensibility may be that it depends on spatially and temporally precise afferent information processed in a distributed cortical network.
Feelings of warmth correlate with neural activity in right anterior insular cortex
. 2005 (Link to paper)
The neural coding of perception can differ from that for the physical attributes of a stimulus. Recent studies suggest that activity in right anterior insular cortex may underlie thermal perception, particularly that of cold. We now examine whether this region is also important for the perception of warmth. We applied cutaneous warm stimuli on the left leg (warmth) in normal subjects (n = 7) during functional magnetic resonance imaging (fMRI). After each stimulus, subjects rated their subjective intensity of the stimulus using a visual analogue scale (VAS), and correlations were determined between the fMRI signal and the VAS ratings. We found that intensity ratings of warmth correlated with the fMRI signal in the right (contralateral to stimulation) anterior insular cortex. These results, in conjunction with previous reports, suggest that the right anterior insular cortex is important for different types of thermal perception.
19. Tactile directional sensitivity and postural control. 2005 (Link to paper)
People are good at telling the direction of a moving tactile stimulus and this capacity provides a sensitive clinical test of somatosensory disturbances. Tactile directional sensitivity depends on two different kinds of somatosensory information, i.e. spatiotemporal information and information about friction-induced changes in skin stretch. The objective of this study was to compare the relative contribution to postural control of these two types of information for both glabrous and hairy skin. Postural sway amplitudes and sway paths were recorded, with or without access to tactile and/or visual stabilizing stimuli. Subjects were standing on two types of surface, either solid metal or 50 mm foam plastic. Two types of stimulus were used to generate sway-related tactile information. One was a thin air-stream that was used to assess the contribution by spatiotemporal information, and the second was a narrow steel rod that was glued to the skin to assess the contribution by skin-stretch information. The stimuli were applied to the hairy skin of the forearm and to the glabrous skin of the fingertip. In addition, we studied the ability to tell the direction of movement of an air-stream stimulus on glabrous and hairy skin. The air-stream caused significant sway reductions when applied to glabrous, but not hairy skin. The weak effect on hairy skin reflected the perceptually poor directional sensitivity for the air-stream stimulus in this cutaneous area. In contrast, the glued rod reduced sway when applied to both glabrous and hairy skin reflecting the tactile afferents' high sensitivity to skin stretch in these areas. Both types of tactile stimulus reduced sway amplitudes more than sway paths for both hairy and glabrous skin. The visual cue, on the other hand, tended to reduce sway paths more than amplitudes. The two types of tactile receptive surface seem to influence postural control in the same manner, despite anatomical and physiological differences. The results invite speculation that patients with poor directional sensitivity might have reduced postural stability compared with healthy individuals.
20. Unmyelinated tactile afferents underpin detection of low-force monofilaments. 2006 (here it is)
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. Hence, the ability to detect light touch does not depend entirely on the A-beta somatosensory system; CT afferents may contribute to the detection of weak monofilaments.
21. Activation of the cortical pain network by soft tactile stimulation after injection of sumatriptan. 2007 (Link to paper)
The anti-migraine drug sumatriptan often induces unpleasant somatosensory side effects, including a dislike of being touched. With a double-blind cross-over design, we studied the effects of sumatriptan and saline on perception (visual analogue scale) and cortical processing (functional magnetic resonance imaging) of tactile stimulation in healthy subjects. Soft brush stroking on the calf (n=6) was less pleasant (p<0.04) and evoked less activation of posterior insular cortex in the sumatriptan compared to the saline condition. Soft brushing activated pain processing regions (anterior insular, lateral orbitofrontal, and anterior cingulate cortices, and medial thalamus) only in the sumatriptan condition, whereas activation of somatosensory cortices was similar in both conditions. Soft brush stroking on the palm (n=6) was equally pleasant in both conditions. One possible mechanism for the activation of pain processing regions by brush stroking is sensitization of nociceptors by sumatriptan. Another possibility is inhibition of a recently discovered system of low-threshold unmyelinated tactile (CT) afferents that are present in hairy skin only, project to posterior insular cortex, and serve affective aspects of tactile sensation. An inhibition of impulse transmission in the CT system by sumatriptan could disinhibit nociceptive signalling and make light touch less pleasant. This latter alternative is consistent with the observed reduction in posterior insular cortex activation and the selective effects of stimulation on hairy compared to glabrous skin, which are not explained by the nociceptor sensitization account.
22. Functional role of unmyelinated tactile afferents in human hairy skin: sympathetic response and perceptual localization. 2008 (Link to paper)
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. By examining two neuronopathy subjects as well as control subjects we have now demonstrated that CT stimulation can elicit a sympathetic skin response. Further, the neuronopathy subjects' ability to localize stimuli which activate CT afferents was very poor but above chance level. The findings support the interpretation that the CT system is well suited to underpin affective rather than discriminative functions of tactile sensations.
23. Discriminative touch and emotional touch. 2007 (Link to the paper)
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. CT afferents have not been found in the glabrous skin of the hand in spite of numerous electrophysiological explorations of this area. Hence, it seems reasonable to conclude that they are lacking in the glabrous skin. A full understanding of the behavioural and affective consequences of the differential innervation of CT afferents awaits a fuller understanding of their function.
24. Cortical processing of lateral skin stretch stimulation in humans. 2008 (Link to paper)
Direction discrimination of a moving tactile stimulus requires intact dorsal columns and provides a sensitive clinical test of somatosensory dysfunction. Cortical mechanisms are poorly understood. We have applied tangential skin pulls to the right lower leg during functional magnetic resonance imaging. Healthy subjects judged the direction of the skin pulls (task experiment, n = 7) or received skin pulls passively (no task experiment, n = 8). Second somatosensory cortex (S2) was activated in the task as well as no task experiment, and there was no significant difference in cortical activation between the two experiments. Within S2 nearly all subjects had prominent activations in the caudal and superficial part, i.e., in the opercular parietal (OP) area 1. S1 was activated in only one of the subjects. Thus, S2 and especially OP 1 seems to be important for processing of lateral skin stretch stimulation. The finding suggests that a lesion of this area might cause a disturbance in tactile direction discrimination which should be relevant for clinical testing.
25. The neurophysiology of unmyelinated tactile afferents. 2008 (Can't get this one)
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. CT fiber activation may also have a role in pain inhibition. This review outlines the growing evidence for unique properties and pathways of CT afferents.
26. Unmyelinated tactile afferents have opposite effects on insular and somatosensory cortical processing. 2008 (Link to paper)

AB 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. The results in IW showed similar activation of posterior insular cortex following CT stimulation as in GL and so strengthen the view that CT afferents underpin emotional aspects of touch. In addition, CT stimulation evoked significant fMRI deactivation in somatosensory cortex in both subjects supporting the notion that CT is not a system for discriminative touch.

27. Coding of pleasant touch by unmyelinated afferents in humans. 2009 (Link to paper)
Pleasant touch sensations may begin with neural coding in the periphery by specific afferents. We found that during soft brush stroking, low-threshold unmyelinated mechanoreceptors (C-tactile), but not myelinated afferents, responded most vigorously at intermediate brushing velocities (1-10 cm s(-1)), which were perceived by subjects as being the most pleasant. Our results indicate that C-tactile afferents constitute a privileged peripheral pathway for pleasant tactile stimulation that is likely to signal affiliative social body contact.
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"In times of change, learners inherit the earth, while the learned find themselves beautifully equipped to deal with a world that no longer exists" ~Roland Barth

"Doubt is not a pleasant mental state, but certainty is a ridiculous one."~Voltaire

Last edited by Diane; 19-05-2009 at 04:41 AM.
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