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      Why does a cooled object feel heavier? Psychophysical investigations into the Weber’s Phenomenon

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          Abstract

          Background

          It has long been known that a concomitantly cooled stimulus is perceived as heavier than the same object at a neutral temperature—termed Weber’s Phenomenon (WP). In the current study, we re-examined this phenomenon using well-controlled force and temperature stimuli to explore the complex interplay between thermal and tactile systems, and the peripheral substrates contributing to these interactions. A feedback-controlled apparatus was constructed using a mechanical stimulator attached to a 5- × 5-mm thermode. Force combinations of 0.5 and 1 N (superimposed on 1-N step) were applied to the ulnar territory of dorsal hand. One of the forces had a thermal component, being cooled from 32 to 28 °C at a rate of 2 °C/s with a 3-s static phase. The other stimulus was thermally neutral (32 °C). Participants were asked to report whether the first or the second stimulus was perceived heavier. These observations were obtained in the all-fibre-intact condition and following the preferential block of myelinated fibres by compression of ulnar nerve.

          Results

          In normal condition, when the same forces were applied, all subjects displayed a clear preference for the cooled tactile stimulus as being heavier than the tactile-only stimulus. The frequency of this effect was augmented by an additional ~17% when cooling was applied concurrently with the second stimulus. Following compression block, the mean incidence of WP was significantly reduced regardless of whether cooling was applied concurrently with the first or the second stimulus. However, while the effect was abolished in case of former (elicited in <50% of trials), the compression block had little effect in four out of nine participants in case of latter who reported WP in at least 80% of trials (despite abolition of vibration and cold sensations).

          Conclusions

          WP was found to be a robust tactile–thermal interaction in the all-fibre-intact condition. The emergence of inter-individual differences during myelinated block suggests that subjects may adopt strategies, unbeknownst to them, that focus on the dominant input (myelinated fibres, hence WP abolished by block) or the sum of convergent inputs (myelinated and C fibres, hence WP preserved during block) in order to determine differences in perceived heaviness.

          Electronic supplementary material

          The online version of this article (doi:10.1186/s12868-016-0322-3) contains supplementary material, which is available to authorized users.

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          Most cited references 31

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          Central changes in processing of mechanoreceptive input in capsaicin-induced secondary hyperalgesia in humans.

          1. Capsaicin, the algesic substance in chilli peppers, was injected intradermally in healthy human subjects. A dose of 100 micrograms given in a volume of 10 microliters caused intense pain lasting for a few minutes after injection and resulted in a narrow area of hyperalgesia to heat and a wide surrounding area of hyperalgesia to mechanical stimuli (stroking) lasting for 1-2 h. 2. Nerve compression experiments with selective block of impulse conduction in myelinated (A) but not in unmyelinated (C) fibres indicated that afferent signals in C fibres contributed to pain from capsaicin injection and to heat hyperalgesia, whereas conduction in afferent A fibres was necessary for the perception of mechanical hyperalgesia. 3. Electrical intraneural microstimulation normally eliciting non-painful tactile sensations was accompanied by pain when the sensation was projected to skin areas within the region of mechanical hyperalgesia induced by capsaicin injection. 4. The threshold for pain evoked by intraneural microstimulation was reversibly lowered and pain from suprathreshold stimulation was exaggerated during the period of mechanical hyperalgesia, regardless of lidocaine anaesthesia of the cutaneous innervation territory of the stimulated fibres. 5. The results indicate that hyperalgesia to stroking on a skin area surrounding a painful intradermal injection of capsaicin is due to reversible changes in the central processing of mechanoreceptive input from myelinated fibres which normally evoke non-painful tactile sensations.
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            The structure and function of a slowly adapting touch corpuscle in hairy skin.

            1. Slowly adapting cutaneous mechanoreceptors, in the cat and primates, have been studied by histological and neurophysiological methods.2. Each touch corpuscle is a dome-shaped elevation of the epidermis, whose deepest layer contains up to fifty specialized tactile cells.3. Nerve plates, enclosed by the tactile cell (Merkel cells), are connected to a single myelinated axon in the dense collagenous core of the corpuscle.4. The corpuscle generated > 1000 impulses/sec when excited by vertical surface pressure. The response was highly localized and showed a low mechanical threshold, the frequency being dependent upon the velocity and amplitude of the displacement. There was a period of rapid adaptation before a sustained response which might continue for > 30 min.5. A quantitative analysis of the responses to excitation by displacements of differing amplitude, velocity and duration is included.6. The discharge of touch corpuscle units evoked by a mechanical stimulus was temperature-sensitive, and was enhanced by a fall in skin temperature.
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              Low-threshold mechanoreceptive and nociceptive units with unmyelinated (C) fibres in the human supraorbital nerve.

               M. Nordin (1990)
              1. In recordings from the human supraorbital nerve with tungsten microelectrodes, eleven afferent units with unmyelinated (C) axons were identified on the basis of their conduction velocities (0.6-1.4 m/s). 2. Eight units had low mechanical thresholds (less than or equal to 0.23 g) and could be activated up to their maximal firing rates of about 100 impulses/s by weak tactile stimuli, whereas three units had higher thresholds (5.5 g) and responded vigorously to noxious stimuli only. 3. During a skin indentation the low-threshold units adapted to an irregular low-frequency discharge, and release of the stimulus elicited a prominent off-response often ending with an after-discharge. Slow stroking was a particularly effective stimulus, even when done with cotton wool, whereas rapid stroking reduced the response. All types of stroking stimuli were occasionally followed by after-discharges. Repeated mechanical stimulation at short intervals resulted in a decline of the response, indicating receptor fatigue. For two units a response to skin cooling was observed. 4. The above low-threshold C units have all the main characteristics of the C mechanoreceptors known from the cat and primates but not previously proven to exist in man. The high-threshold C units are similar to the polymodal nociceptors found in other human skin areas.
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                Author and article information

                Contributors
                James.Dunn@westernsydney.edu.au
                D.Mahns@westernsydney.edu.au
                S.Nagi@westernsydney.edu.au
                Journal
                BMC Neurosci
                BMC Neurosci
                BMC Neuroscience
                BioMed Central (London )
                1471-2202
                3 January 2017
                3 January 2017
                2017
                : 18
                Affiliations
                [1 ]School of Medicine, Western Sydney University, Locked Bag 1797, Penrith, NSW 2751 Australia
                [2 ]Department of Clinical and Experimental Medicine, Center for Social and Affective Neuroscience, Linköping University, 58183 Linköping, Sweden
                Article
                322
                10.1186/s12868-016-0322-3
                5209941
                © The Author(s) 2017

                Open AccessThis article is distributed under the terms of the Creative Commons Attribution 4.0 International License ( http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The Creative Commons Public Domain Dedication waiver ( http://creativecommons.org/publicdomain/zero/1.0/) applies to the data made available in this article, unless otherwise stated.

                Funding
                Funded by: Western Sydney University Early Career Research Grant
                Award ID: P00021752
                Award Recipient :
                Categories
                Research Article
                Custom metadata
                © The Author(s) 2017

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