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Pleasant Touch 741

Pleasant Touch
A B Vallbo, H Olausson, and J Wessberg, Goteborg University, Goteborg, Sweden
2009 Elsevier Ltd. All rights reserved.

Pleasant Touch A Fundamental Affectionate Variable


It is a universal experience that bodily contact between individuals is a powerful means of nonverbal communication. Skin-to-skin touch may have a very strong emotional impact, immediate as well as long term. The affective potential of bodily contact is perhaps most apparent in erotic and parentchild relations, although touching is a potent means to convey empathy and relieve anxiety in less intimate personal contacts as well. Even a passing touch offered by a friend or an empathic nurse may give great reassurance to a person in fear, distress, or agony. Robin Dunbar stated wisely that a touch may be worth a thousand words. Further, in positive excitement we like to raise the feeling of shared joy, for instance, by hugging the teammates after a goal. On the other hand, the erotic potential of bodily contacts implies a barrier which in many situations makes us abstain from touching another person. It was shown long ago by Harlow and co-workers that pleasant touch is essential to provide affection and security in early childhood. Studies of baby monkeys left in distress because they were deprived of their native mothers led to the conclusion that contact comfort is a very potent factor for the baby to develop affectionate responses, reassurance, and love for the mother. In fact, contact comfort was found to be even more powerful than nutrition. Although a number of sensory mechanisms, as well as psychological factors are probably involved in most situations when we convey positive emotional signals by bodily contact, it is obvious that cutaneous touch is a significant element. The focus of the present account is on highly sensitive cutaneous sense organs which respond to light touch, particularly the unmyelinated tactile (CT) afferents which might have a specific role in this context. With regard to semantics, the term tactile in this account solely refers to the finding that the afferents respond intensely to light touch, whereas the term does not refer to central effects related to haptics.

myelinated Ab-afferents alone. However, in the last decades it has been demonstrated that human skin is also innervated by unmyelinated afferents responding to light touch (CT-afferents). They conduct impulses at a speed of about 1 m s1 whereas Ab-afferents are about 50 times faster (see Figure 1). Six different kinds of Ab-afferents with separate physiological properties have been identified in human skin in addition to the CT-afferents (Table 1). The distinct difference in receptive field size and topography is illustrated in Figure 2 for the main types of tactile afferents encountered in the hairy skin of the forearm. Corresponding illustrations for glabrous skin units may be found in Kandel, Schwartz, and Jessels textbook. Receptive field structure as assessed with an objective method provides convincing evidence for unit identification and separation in distinct groups, although additional observations are essential, for example, conduction velocity and selective sensitivity to hair movements. There is an interesting regional difference with regard to Ab and CT innervation in man. CT-afferents have been demonstrated in face, forearm, hand dorsum, and thigh, suggesting that they are present all over the hairy skin. However, they have not been found in the glabrous skin of the hand in spite of numerous microenurography explorations of this area. Hence, it seems reasonable to conclude that they are lacking in the glabrous skin. Although there is no method to properly assess the proportion of CT-afferents out of total number of tactile units, it is a recurring experience in microneurography experiments that CT-afferents are encountered at least as often as Ab-afferents in nerves innervating the forearm skin. However, there are indications that CT-afferents may be less common in the lower leg. It should be pointed out that unmyelinated tactile afferents were demonstrated long ago in the hairy skin of various animals but they have not been found in the pad skin which corresponds to the glabrous skin in man where CT-afferents are lacking as well. In animals the unmyelinated tactile afferents are often denoted as low threshold mechanoreceptive C-afferents (C-LTM). For long it was assumed that this system of afferents had faded away in the process of evolution from rat, rabbit, and cat to man until microneurogaphy demonstrated CT-afferents in man in the beginning of the 1990s.
Functional Properties of CT-Afferents

Tactile Afferents in Human Skin


The canonical view of human physiology is that light touch is signaled by large and fast conducting

The CT-afferents are equally sensitive to skin deformation as many of the Ab-afferents. They respond to

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Table 1 Mechanoreceptive afferents in human nerves from glabrous and hairy skin Sensory endings and afferent unit acronyms Glabrous skin Hairy skin Merkel SAI Ruffini SA II Pacini FA II (PC) Hair follicle unit Field C-tactile CT (CLTM) Adaptation Slow Slow Fast Fast Fast Intermediate

Merkel SAI Ruffini SAII Pacini FA II (PC) Meissner FAI (RA, QA)

50 g 0.2 s b 50 V SAI CT c 2 ms

Figure 1 Microneurography recording of two single afferents to light touch, that is, a fast conducting Ab-afferent and an unmyelinated CT-afferent. (a) Target point for touch stimulus and microneurography recording site from the lateral antebrachial cutaneous nerve in the upper arm. (b) Microneurography record and time course of tap stimulus showing an early burst of nerve impulses in the Ab-afferent and a late burst in the CT-afferent. Distance between target area and recording site was 247 mm implying a conduction velocity below 1 m s1 of the CT-afferent. The threshold of the CT-afferent as tested with von Frey monofilaments was 1.3 mN. (c) Nerve impulses from the record in (b) displayed on an expanded timescale to highlight the different impulse shape of the Ab-afferent (SAI type) and the CT afferent. Reproduced from Wessberg J, Olausson H, Wiklund Fernstrom K, and Vallbo AB (2003) Receptive field properties of unmyelinated tactile afferents in the human skin. Journal of Neurophysiology 89: 15671575, with permission from The American Physiological Society.

The separate unit types were identified on the basis of microneurography recordings in man while types of end organ were inferred on the basis of physiological and morphological studies in various species. Slow and fast adaptation refers to the presence or lack of sustained response to stationary skin deformation. Most acronyms refer to adaptation properties (fast, rapid, quick, or slow adaptation). Acronyms within parenthesis have mainly been used in studies of nonhuman species. All afferents listed are myelinated and fast conducting (Ab fibers) except CT units which are unmyelinated (C-fibers). The table includes highly sensitive afferents but not nociceptive afferents that may respond weakly to innouous skin deformation. The table entries refer to glabrous skin of the hand and hairy skin of the forearm and are based on recent investigations. However, there are microneurography studies suggesting a somewhat different innervation pattern in particular skin regions, the existence of subgroups, and one or two additional mechanoreceptor types.

indentation forces in the range 0.32.5 mN as tested with von Frey monofilaments and they produce high-frequency trains of action potentials (50100 impulses s1) to pleasant touch stimuli. It should be recalled that maximal impulse frequency recorded from afferent C-fibers is about 100 impulses s1. In contrast, Ab-afferents may fire close to 1000 impulses s1. It is of interest that, in addition to these two groups of sensitive tactile afferents, some of the nociceptive afferents may respond to innocuous touch as well. However, their sensitivity is low and their response to pleasant touch is minimal, usually not more than one or two impulses. While CT-afferents share the high sensitivity with the Ab-afferents, the physiological properties of the two groups differ in most other respects. Ab-afferents fall in two distinct groups with regard to response to steady-state skin deformation (Table 1). The fast adapting units do not respond at all, that is, they lack static sensitivity altogether, while the slowly adapting units provide a continuous discharge for

minutes or more. Actually, many Ruffini units may be firing continuously for life. In contrast, the CT-afferents have intermediate adaptation properties. They respond initially with a burst of high impulse rate, while the rate often falls to zero after a few seconds of sustained indentation. The discharge may then reassume after another few seconds, then wax and wane to finally stop altogether. CT-afferents lack the potential of the Ab-afferents to code fast events. They respond well to slowly moving stimuli but poorly to brisk movements and vibration. Hence, their dynamic response range is limited to low frequencies. Stimuli which are particularly efficient to evoke a massive CT-input are slowly moving, caresslike stimuli over large skin areas with the hand or a broad artist paintbrush. A prominent feature of the CT-afferents is that they are highly fatigable. When identical stimuli are repeated, the first stimulus usually evokes a much larger response than the following. This feature suggests that the system is particularly interested in novel events. When a local skin deformation is released, CT-afferents may produce an after-discharge that may last for many seconds. However, the duration varies a lot even within the individual unit depending on temperature, previous history, and other nondefined factors. An additional and unique feature of the CT-afferents

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SA I SA II CT

0 Hair

10 mm

0 Field

10 mm

0 CT

10 mm

20 mm

20 mm

10 mm

max.

Figure 2 Receptive field topography of five kinds of tactile afferents in the hairy skin. Data from four representative Ab-afferents and two different CTafferents (see Table 1). Note difference in scale for the hair and field units. The colors represent intensity of afferent firing from zero to maximal rate of the particular trial. Peak rates fall in the range 3560 impulses s1. Gray areas which fill up the SAII box represent continuous spontaneous activity of the afferent. (Ab afferents) Adapted from Vallbo AB, Olausson H, Wessberg J, and Kakuda N (1995) Receptive field characteristics of tactile units with myelinated afferents in hairy skin of human subjects. Journal of Physiology 483: 783 795, with permission from Blackwell Publishing. (CT-afferents) Reproduced from Wessberg J, Olausson H, Wiklund Fernstrom K, and Vallbo AB (2003) Receptive field properties of unmyelinated tactile afferents in the human skin. Journal of Neurophysiology 89: 15671575, with permission from The American Physiological Society.

is that the response of the individual afferent is not as consistent and predictable as with individual Abafferents but may vary from one test series to the next for no obvious reason. The response features of the CT-afferents as outlined above all converge toward the interpretation that the system is not designed to code accurately and consistently intensity and time course of skin deformations. These are functions which are essential for discriminative touch. In daily life, the tactile system continually captures details of temporally and spatially variant skin deformations. As a result we can readily perceive more complex features, for instance, the direction of a stimulus moving along the skin and we can discriminate the intensity of component indentations which are closely adjacent

in time and space to assess shape and surface structure of objects. Altogether, it seems evident that the CT-afferents lack response properties required for serving discriminative touch. Significance of conduction velocity The high conduction velocity of the myelinated Ab-afferents is a feature which is essential for discriminative touch, as well as for guidance of motor activities. Hence, the low conduction velocity of the CT-afferents precludes these roles. First, the long delay (0.51.0 s) from stimulus to impulse arrival in the brain is hardly compatible with accurate integration with signals from fast Ab-afferents and other sensory systems, for example, vision and hearing, that is, forms of integration which are often essential for guidance of

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motor activities. Second, the documented interfiber variation in conduction velocity around 1 m s1 implies that the set of impulses in the sample of CTafferents excited by a local touch would be widely dispersed in time when arriving to the brain. As a result, the CT-afferents produce a temporally blurred account of the peripheral event compared to that of the Ab-system. Hence, the slow conduction velocity of the unmyelinated nerve fibers implies an additional argument against the CT-system having a role in discriminative touch.

maps are modified with amputations and other injuries. In relation to a possible role of CT-afferents in this context, it seems significant that the unmyelinated afferents involved in body surface mapping are sensitive to capsaicin. Since human CT-afferents are not sensitive to capsaicin, it seems reasonable to conclude that the CT-afferents are not main candidates for cortical map design.
Affective Touch Hypothesis

CT-Afferents Functional Hypotheses


The physiological properties of the CT-afferents as presented above seem to rule out the possibility that the system plays an essential role in discriminative and motor aspects of cutaneous mechanoreception. Hence, it is of interest to consider what other functions might justify the presence of a slow system with selective sensitivity to light touch. Which functions might afford the deficiencies of the CT-system in coding peripheral events, yet be biologically significant enough to justify the preservation of a parallel tactile system in addition to the fast and accurate Ab-system?
Tickle Hypothesis

Although poor coding accuracy is a serious drawback in discriminative functions, an area where these deficiencies are readily affordable is the limbic realm. Considering the potential roles of light touch in relation to limbic functions, it seems reasonable to single out one function of particular interest, that is, tight physical contact with conspecifics, in the first place with your mother, later in life with your partner and children. We all strongly feel now and then that skin wants skin. These considerations led to the affective touch hypothesis, implying that the essential role of the CT-system is to convey pleasant aspects of light touch, particularly in skin-to-skin contact with affiliative human beings. It should be emphasized that this does not imply that CT-afferents are the only afferents that may contribute or give rise to limbic touch effects, as will be discussed in a later section.

For long, the low threshold mechanoreceptive C-afferents were discussed as a system accounting for the sensation of tickle. This idea proposed by Zotterman in 1939 was largely based on his observation that unmyelinated tactile afferents in the cat may produce long-lasting after-discharges when a skin indentation was released. The tickle hypothesis was then reiterated in a number of successive publications as late as in the 1990s. However, in microneurogaphy studies of human CT-afferents, it has been found that the duration of after-discharge does not match the duration of tickle sensations, nor is the impulse rate of CT-afferents related to the intensity of tickle sensation. Moreover, subjects who lack Ab-afferents but have preserved small fiber functions (see below) are not aware of being tickled in the affected skin areas. These findings indicate that the sensation of tickle is dependent on Ab-afferents rather than on CT-afferents.
Maintenance of Cortical Maps

Responses to Activation of the CT-System


To test the affective touch hypothesis and explore the functional role of the CT-system, it is desirable to activate the CT-afferents selectively without activating Ab-afferents. Unfortunately, this is not possible in normal subjects although stimuli designed to give different proportions of Ab- or CT-input have been tested. It has been demonstrated in proper psychophysical tests that a slowly moving touch stimulus (15 cm s1) which is particularly effective to activate CT-afferents is more pleasant than a fast-moving stimulus. Vibratory stimuli which activate Ab-afferents very efficiently, but CT-afferents only weakly, are rated as less pleasant than slowly moving touch stimuli. These observations are consistent with the hypothesis that CT-afferents are more important for the feeling of pleasant touch than are Ab-afferents. Although findings of this nature are consistent with the affective touch hypothesis, more pertinent data have been obtained from a few subjects, who suffer from a very rare neurological deficiency (sensory neuronopathy syndrome), implying that they lack Ab-afferents from large parts of the body. As a consequence, it is reasonable to assume that their sole tactile afferents are unmyelinated CT-afferents.

Studies in nonhuman subjects (cat and flying fox) have suggested that unmyelinated mechanoreceptive afferents may play an essential role in the complex processes of shaping and maintaining the map of body surface in the somatosensory cortex areas. The hypothesis is based on observations of how these

Pleasant Touch 745 Psychophysical Responses

Two subjects with sensory neuronopathy syndrome are well described in the literature. They have been studied extensively over the years particularly with regard to motor functions. It has also been reported, although more in passing, that they lack tactile sensibility. This observation was consistent with the prevailing view at that time that tactile sensibility is lacking altogether when Ab-afferents are nonfunctioning. This view, in turn, was largely based on experiments in normal subjects which suggested that tactile sensibility is lacking when the Ab-afferents had been selectively blocked by sustained pressure onto the nerve. However, when it became evident that human skin is supplied with a system of unmyelinated afferents sensitive to light touch, it became pertinent to reexamine the tactile sensibility of neuronopathy subjects using more refined approaches. Rigorous psychophysical tests were pursued to study if the neuronopathy subjects were at all able to detect touch stimuli which were particularly designed to evoke massive CT-inputs. It was found, in 2-alternative-force-choice (2-AFC) tests, that subjects lacking Ab-afferents readily detected such stimuli applied to the forearm skin where CT-afferents are abundant. In contrast, they failed altogether to detect the same kind of stimuli applied to the glabrous skin of the hand where CT-afferents are lacking. A difference of similar kind was found between the two skin regions in the neuronopathy subjects when detection threshold was assessed using von Frey monofilament. Relatively weak monofilaments which give a pure sensation of touch in normal subjects were readily detected in the hairy skin but not in the glabrous skin. Here much stronger stimuli (48 times stronger) were required for detection. Moreover, the neuronopathy patients reported that these strong stimuli gave rise to a sharp and marginally painful sensation, suggesting that detection in the palm was based on nociceptive input. In contrast, sensations with threshold von Frey monofilament in the hairy skin lacked all elements of pain or sharpness suggesting that detection was based on afferents sensitive to innocuous stimuli. It should be emphasized that, in normal subjects, there is no difference between the two skin regions in this respect, that is, thresholds for detection of monofilament stimuli are identical in the hairy and glabrous skin. Further, thresholds are generally lower than in the neuronopathy subjects, suggesting detection by Ab-afferents in normal subjects. An additional highly relevant finding is that the neuronopathy subjects are unable to detect vibratory stimuli which are known to a give a poor excitation of CT-afferents but a massive activation of several kinds of Ab-afferents.

Although these observations agree that CT-afferents may well account for detection of touch stimuli in subjects lacking Ab-afferents, it has been more difficult to obtain a clear and consistent apprehension of the quality of the sensation perceived when CT-afferents are selectively stimulated. Still, a number of interesting features emerged, when the qualitative characteristics of the sensation were explored. First, it was obvious that the sensation associated with a massive and selective CT-input was very weak, vague, and inconsistent when slowly stroking a broad and soft artist paintbrush along the forearm of the neuronopathy subjects. In some trials, the subject reported no sensations at all. In others, they reported a sensation of light touch which was barely detectable and difficult to mentally catch and describe in detail. In this context it should be noticed that stimulus detection in a psychophysical 2-AFC test does not prove that the stimulus is very clearly perceived. The weakness and vagueness of the sensation are further illustrated by the fact that conscious perception of CT-stimulus varied from one occasion to the other and between the two subjects as well. In fact, one of the two subjects (GL) reported that she began to feel more touch sensations in daily life, once she had had the experience of touch perception from the affected skin areas and had become aware of the possibility. Although the two neuronopathy subjects were not able to give a clear and detailed description of the sensation evoked by CT-stimulation, some consistent features of the quality emerged, namely that the sensation had no tint of pain, tickle, or itch. Further they reported that the sensation elicited by CT-stimulation was slightly or moderately pleasant.
Brain Responses

When functional magnetic resonance imaging (fMRI) was used to study brain response to touch stimuli (Figure 3) in normal and neuronopathy subjects lacking Ab-afferents, the findings indicated that different sensory areas are activated by the fast (Ab) and the slow (CT) tactile systems from the hairy skin. In normal subjects pleasant touch activated the classical somatosensory areas SI and SII as well as insular cortex, notably the posterior part of the contra-lateral insula. Somatosensory areas SI and SII are known to be significant for discriminative touch whereas insular cortex is a region of great interest in relation to affective mechanisms, because it is considered to house a gateway from sensory systems to the emotional systems of the frontal lobe. When similar stimuli were applied to subjects lacking Ab-afferents, no activation was found in the somatosensory areas whereas the posterior insular region was activated.

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Figure 3 fMRI responses (BOLD) to pleasant touch stimuli. (a) Response in posterior and middle insular region to selective stimulation of CT-afferents in a neuronopathy subject lacking large myelinated Ab-afferents. In this subject light touch produced no BOLD response in somatosensory areas SI and SII, whereas similar stimuli regularly activate these two areas in normal subjects. (b) BOLD response in orbitofrontal cortex to pleasant touch stimuli of the palm in normal subjects (group data). BOLD, blood-oxygen level dependent. (a) Unpublished. (b) Adapted from Rolls ET, ODoherty J, Kringelbach ML, et al. (2003) Representations of pleasant and painful touch in the human orbitofrontal and cingulated cortices. Cerebral Cortex 13: 308317, with permission from Oxford University Press.

The conclusion is that the unmyelinated CT-afferents project toward the emotional systems (insular cortex) but less or not at all toward the discriminative cognitive systems (classical somatosensory areas SI and SII). This observation seems to provide an essential support for the affective touch hypothesis. To what extent the tactile Ab-afferents project toward the emotional systems along with the CTafferents has not been widely explored. However, it seems likely that this may be the case, because pleasant touch stimuli of the palm, where CT-afferents are lacking, give rise to fMRI responses in a target area for insular efferents in orbitofrontal cortex involved in more complex emotional mechanisms. It remains to be explored whether a projection of tactile Abafferents to emotional centers is unique for the palm or is true for other skin areas as well. It may be speculated that Ab-afferents from the palm have obtained an affective role because CT-afferents are lacking here. Anyway, it seems clear that affective aspects of touch are not always dependent on CT-afferents alone. The physiological properties of the CT-afferents as well as the psychophysical and fMRI responses to selective activation of the unmyelinated tactile system converge toward a limbic-emotional role of the system. The affective touch hypothesis implies that the essential role of the CT-system is to provide or support emotional, hormonal, and behavioral responses to skin-to-skin contact with conspecifics. On the other hand, it seems likely that a natural perceptiveemotional response to pleasant touch is dependent on the combination of afferents from the two tactile systems, because selective CT-stimulation fails to evoke anything like a full sensation of pleasant

touch. The combination of CT- and Ab-afferents is obviously required for the complete feeling of pleasant touch in the hairy skin. It is also pertinent to recall a general principle regarding emotional responses. The intensity and even the quality of the emotional response evoked by a particular stimulus is highly dependent on contextual factors, as well as state of deprivation, that is, to what extent your desire with regard to the particular emotion has recently been satisfied or not. Obviously, your sense of pleasure is intense when you hug and touch your partner after weeks of separation, whereas you are not particularly pleased when touched by an unknown person in a crowd. It may be asked why evolution has retained a separate system of unmyelinated afferents for socialemotional touch. Why could this role not be handled by the Ab-afferents? It is difficult to deny that they could and, in fact, they do in the glabrous skin of the hand.A particular reason for preservation of the CT-system is that unmyelinated afferents are parsimonious because they take up little space in the peripheral nerves.

CT-System A Branch of an Interoceptive Afferent System


In a wider perspective, the CT-system may be regarded as a branch of a large afferent system which is basically concerned with your own body rather than external events, as conjectured by Craig. The basic role of the interoceptive system is to continuously watch the condition of body tissues, as well as physiological and chemical variables within the body. Interoceptive afferents have particular access

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to brain centers which control affective, hormonal, autonomic, and behavioral responses with effects to readjust adverse conditions and keep variables within desired ranges. The ultimate aim would be to keep the cell, the individual, and the species going. Included in the interoceptive system are afferents related to perception of pain, itch, temperature, air hunger, vasomotor flush, hunger, thirst, and a range of visceral sensations, as well as afferents which are essential for the subconscious control of physiological variables, such as blood pressure and concentration of blood gases. The peripheral afferents of the interoceptive system consist of small diameter nerve fibers (thin myelinated Ad and C fibers) with projections to the superficial lamina of the dorsal horn in the spinal cord and nucleus of the solitary tract. In this context it is interesting that touch sensitive unmyelinated afferents (C-LTM) have been shown to project to lamina I and II in several mammalian species, consistent with the interpretation that they belong to the interoceptive system. However, apart from this finding, connectivity studies of CT-afferents or homologs in other species are unfortunately lacking altogether in the literature. At higher levels the small fiber afferents are tightly linked to amygdale and hypothalamus and, at cortical levels, to the insular region, primarily the posterior and middle parts. Moreover, there are indications that the anterior insular cortex on the nondominant side constitutes a still higher level housing a re-representation of the interoceptive image of the entire body. This representation of the physical self is postulated to constitute a basic foundation of mental consciousness. The role of the CT-units as an afferent branch of a system guarding the well-being of the body would be to signal pleasure and reassurance as you are close to your parents, lover, kin, or friends. There are indications that pleasant bodily contacts promote hormonal responses, that is, endorphin and oxytocin, which contribute to the feeling of well-being, confidence, and calmness.

Bodily Contact
Although the affective touch hypothesis implies that the particular role of the CT-system is to promote emotional, hormonal, and behavioral responses to pleasant physical contact with conspecifics, it is obvious that a number of other sensory channels are important as well. Interesting observations of social and biological roles of tight bodily contacts in nonhuman species have been reported. A particular kind of bodily contact is grooming. At first sight it appears that grooming has a hygienic

role alone. However, Dunbar has demonstrated in monkeys that it has an affective and social role as well. Obviously, the groomee is very pleased with the procedure even though the handling is often quite tough. Grooming increases the production of endorphin in the groomee. Dunbar has also found that monkeys spend much more time in grooming than required from hygienic point of view. It is particularly interesting that the time spent on grooming is larger, the larger the social group. The bottom line conclusion is that an essential role of grooming is to promote affectionate attachment between individuals and hence to keep the group together. The caring pattern of the mother in handling her young infants has a profound influence on the development of the child. Epidemiological studies in man have demonstrated that adverse conditions in early childhood tend to increase stressful and nonsocial behavior in the adult. Biological mechanisms that might be significant for this phenomenon have been explored in rats by Meaney and co-workers. Individual rat mothers differ with regard to caring pattern. Some are very active, spending a lot of time licking and handling their pups. Others are more passive in that respect. The caring pattern has a profound effect on epigenetic mechanisms related to the expression of stress hormone receptors in the brain. In the end, the offspring of active mothers become less stress sensitive, more curious, and exploratory than the offspring of passive mothers. Interestingly, these differences are permanent and remain in the adult, although they are partially modifiable. Although it remains to be explored to what extent the CT-system is involved in affective response to bodily contacts in a social context, like grooming and mothers care of her baby, there is no question that the CT-system is activated in these situations. The affective touch hypothesis implies that the CT-system is one of several communication channels accounting for responses to bodily contacts which engage perceptive, hormonal, autonomic, and behavioral mechanisms and might range from immediate pleasure, joy, and reassurance in sex to the postnatal development of your childrens stress coping profiles.
See also: Cross-Modal Interactions Between Vision and Touch; Mechanoreceptors; Orbitofrontal Cortex: Visual Functions; Tactile Coding in Peripheral Neural Populations; Tactile Texture.

Further Reading
Calford MB and Tweedale R (1991) C-fibres provide a source of masking inhibition to primary somatosensory cortex.

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Proceedings of the Royal Society, Series B: Biological Sciences 243: 269275. Cole J (1995) Pride and a Daily Marathon. Cambridge, MA: The MIT Press. Cole J, Bushnell MC, McGlone F, et al. (2006) Unmyelinated tactile afferents underpin detection of low-force monofilaments. Muscle Nerve 34: 105107. Craig AD (2002) How do you feel? Interoception: The sense of the physiological condition of the body. Nature Reviews 3: 655666. Dunbar R (1997) Grooming, Gossip and the Evolution of Language. London: Faber & Faber. Harlow HF (1959) Love in infant monkeys. Scientific American 200: 6874. Kandel ER, Schwartz JH, and Jessell TM (eds.) (2000) Principles of Neural Science, 4th edn. New York: McGraw-Hill. Olausson H, Lamarre Y, Backlund, et al. (2002) Unmyelinated tactile afferents signal touch and project to insular cortex. Nature Neuroscience 5: 900904. Rolls ET, ODoherty J, Kringelbach ML, et al. (2003) Representations of pleasant and painful touch in the human orbitofrontal and cingulated cortices. Cerebral Cortex 13: 308317. Sterman AB, Schaumburg HH, and Ashbury AK (1980) The acute sensory neuronopathy syndrome: A distinct clinical entity. Annual Neurology 7: 354358. Uvnas-Moberg K (1997) Physiological and endocrine effects of social contact. In: Carter CS, Lederhendler II, and Kirkpatrick B (eds.) The Integrative Neurobiology of Affiliation (Annals of the New York Academy of Science), vol. 807, pp. 146163. New York: The New York Academy of Sciences. Vallbo AB, Olausson H, and Wessberg J (1999) Unmyelinated afferents constitute a second system coding tactile stimuli of the human hairy skin. Journal of Neurophysiology 81: 27532763. Vallbo AB, Olausson H, Wessberg J, and Kakuda N (1995) Receptive field characteristics of tactile units with myelinated afferents in hairy skin of human subjects. Journal of Physiology 483: 783795. Weaver IC, Cervoni N, Champagne FA, et al. (2004) Epigenetic programming by maternal behaviour. Nature Neuroscience 7: 847854. Wessberg J, Olausson H, Wiklund Fernstrom K, and Vallbo AB (2003) Receptive field properties of unmyelinated tactile afferents in the human skin. Journal of Neurophysiology 89: 15671575.

Relevant Website
http://deafferented.apinc.org Studies on the Deafferent Patient G.L.