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Population coding of somatic sensations

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Abstract

The somatic sensory system includes a variety of sensory modalities, such as touch, pain, itch, and temperature sensitivity. T The coding of these modalities appears to be best explained by the population-coding theory, which is composed of the following features. First, an individual somatic sensory afferent is connected with a specific neural circuit or network (for simplicity, a sensory-labeled line), whose isolated activation is sufficient to generate one specific sensation under normal conditions. Second, labeled lines are interconnected through local excitatory and inhibitory interneurons. As a result, activation of one labeled line could modulate, or provide gate control of, another labeled line. Third, most sensory fibers are polymodal, such that a given stimulus placed onto the skin often activates two or multiple sensory-labeled lines; crosstalk among them is needed to generate one dominant sensation. Fourth and under pathological conditions, a disruption of the antagonistic interaction among labeled lines could open normally masked neuronal pathways, and allow a given sensory stimulus to evoke a new sensation, such as pain evoked by innocuous mechanical or thermal stimuli and itch evoked by painful stimuli. As a result of this, some sensory fibers operate along distinct labeled lines under normal versus pathological conditions. Thus, a better understanding of the neural network underlying labeled line crosstalk may provide new strategies to treat chronic pain and itch.

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References

  1. Norrsell U, Finger S, Lajonchere C. Cutaneous sensory spots and the “law of specific nerve energies”: history and development of ideas. Brain Res Bull 1999, 48: 457–465.

    Article  PubMed  CAS  Google Scholar 

  2. Torebjörk HE, Ochoa JL. Specific sensations evoked by activity in single identified sensory units in man. Acta Physiol Scand 1980, 110: 445–447.

    Article  PubMed  Google Scholar 

  3. Torebjörk HE, Schady W, Ochoa J. Sensory correlates of somatic afferent fibre activation. Hum Neurobiol 1984, 3: 15–20.

    PubMed  Google Scholar 

  4. Handwerker HO. Sixty years of C-fiber recordings from animal and human skin nerves: historical notes. Prog Brain Res 1996, 113: 39–51.

    Article  PubMed  CAS  Google Scholar 

  5. Alrutz S. On the temperature senses: II. The sensation of ‘hot’. Mind 1898, 7: 141–144.

    Article  Google Scholar 

  6. Head H. Croonian lecture: Release of function in the nervous system. Proceeding of the Royal Society of London. Series B. 1921, 92: 184–209.

    Article  Google Scholar 

  7. Dodt E, Zotterman Y. The discharge of specific cold fibres at high temperatures; the paradoxical cold. Acta Physiol Scand 1952, 26: 358–365.

    Article  PubMed  CAS  Google Scholar 

  8. Simone DA, Baumann TK, LaMotte RH. Dose-dependent pain and mechanical hyperalgesia in humans after intradermal injection of capsaicin. Pain 1989, 38: 99–107.

    Article  PubMed  CAS  Google Scholar 

  9. Lynn B. Capsaicin: actions on C fibre afferents that may be involved in itch. Skin Pharmacol 1992, 5: 9–13.

    Article  PubMed  CAS  Google Scholar 

  10. Shim WS, Oh U. Histamine-induced itch and its relationship with pain. Mol Pain 2008, 4: 29.

    Article  PubMed  Google Scholar 

  11. Imamachi N, Park GH, Lee H, Anderson DJ, Simon MI, Basbaum AI, et al. TRPV1-expressing primary afferents generate behavioral responses to pruritogens via multiple mechanisms. Proc Natl Acad Sci U S A 2009, 106: 11330–11335.

    Article  PubMed  CAS  Google Scholar 

  12. Campbell JN, Meyer RA. Mechanisms of neuropathic pain. Neuron 2006, 52: 77–92.

    Article  PubMed  CAS  Google Scholar 

  13. Costigan M, Scholz J, Woolf CJ. Neuropathic pain: a maladaptive response of the nervous system to damage. Annu Rev Neurosci 2009, 32: 1–32.

    Article  PubMed  CAS  Google Scholar 

  14. Ikoma A, Fartasch M, Heyer G, Miyachi Y, Handwerker H, Schmelz M. Painful stimuli evoke itch in patients with chronic pruritus: central sensitization for itch. Neurology 2004, 62: 212–217.

    PubMed  CAS  Google Scholar 

  15. Hosogi M, Schmelz M, Miyachi Y, Ikoma A. Bradykinin is a potent pruritogen in atopic dermatitis: a switch from pain to itch. Pain 2006, 126: 16–23.

    Article  PubMed  CAS  Google Scholar 

  16. Ishiuji Y, Coghill RC, Patel TS, Dawn A, Fountain J, Oshiro Y, et al. Repetitive scratching and noxious heat do not inhibit histamineinduced itch in atopic dermatitis. Br J Dermatol 2008, 158: 78–83.

    PubMed  CAS  Google Scholar 

  17. McMahon SB, Koltzenburg M. Itching for an explanation. Trends Neurosci 1992, 15: 497–501.

    Article  PubMed  CAS  Google Scholar 

  18. Akiyama T, Merrill AW, Carstens MI, Carstens E. Activation of superficial dorsal horn neurons in the mouse by a PAR-2 agonist and 5-HT: potential role in itch. J Neurosci 2009, 29: 6691–6699.

    Article  PubMed  CAS  Google Scholar 

  19. Campero M, Baumann TK, Bostock H, Ochoa JL. Human cutaneous C fibres activated by cooling, heating and menthol. J Physiol 2009, 587: 5633–5652.

    Article  PubMed  CAS  Google Scholar 

  20. Ma Q. Labeled lines meet and talk: population coding of somatic sensations. J Clin Invest 2010, 120: 3773–3778.

    Article  PubMed  CAS  Google Scholar 

  21. Head H, Sherren J. The consequences of injury to the peripheral nerves in man. Brain 1905, 28: 116–138.

    Article  Google Scholar 

  22. Weddell G, Sinclair DC, Feindel WH. Significance of multiple innervation of cutaneous pain spots in relation to the quality of pain sensibility. Nature 1947, 160: 27.

    Article  PubMed  CAS  Google Scholar 

  23. Noordenbos W. Pain. Amsterdam: Elsevier, 1959.

    Google Scholar 

  24. Melzack R, Wall PD. Pain mechanisms: a new theory. Science 1965, 150: 971–979.

    Article  PubMed  CAS  Google Scholar 

  25. Wall PD. The gate control theory of pain mechanisms. A re-examination and re-statement. Brain 1978, 101: 1–18.

    Article  PubMed  CAS  Google Scholar 

  26. Noordenbos W. Some historical aspects. Pain 1987, 29: 141–150.

    Article  PubMed  CAS  Google Scholar 

  27. Green BG. Temperature perception and nociception. J Neurobiol 2004, 61: 13–29.

    Article  PubMed  Google Scholar 

  28. Mackenzie RA, Burke D, Skuse NF, Lethlean AK. Fibre function and perception during cutaneous nerve block. J Neurol Neurosurg Psychiatry 1975, 38: 865–873.

    Article  PubMed  CAS  Google Scholar 

  29. Fruhstorfer H. Thermal sensibility changes during ischemic nerve block. Pain 1984, 20: 355–361.

    Article  PubMed  CAS  Google Scholar 

  30. Wahren LK, Torebjörk E, Jörum E. Central suppression of coldinduced C fibre pain by myelinated fibre input. Pain 1989, 38: 313–319.

    Article  PubMed  CAS  Google Scholar 

  31. Yarnitsky D, Ochoa JL. Release of cold-induced burning pain by block of cold-specific afferent input. Brain 1990, 113: 893–902.

    Article  PubMed  Google Scholar 

  32. Craig AD, Bushnell MC. The thermal grill illusion: unmasking the burn of cold pain. Science 1994, 265: 252–255.

    Article  PubMed  CAS  Google Scholar 

  33. Price DD, Hu JW, Dubner R, Gracely RH. Peripheral suppression of first pain and central summation of second pain evoked by noxious heat pulses. Pain 1977, 3: 57–68.

    Article  PubMed  CAS  Google Scholar 

  34. LaMotte RH, Thalhammer JG. Response properties of high-threshold cutaneous cold receptors in the primate. Brain Res 1982, 244: 279–287.

    Article  PubMed  CAS  Google Scholar 

  35. Delmas P, Hao J, Rodat-Despoix L. Molecular mechanisms of mechanotransduction in mammalian sensory neurons. Nat Rev Neurosci 2011, 12: 139–153.

    Article  PubMed  CAS  Google Scholar 

  36. Bautista DM, Lumpkin EA. Perspectives on: information and coding in mammalian sensory physiology: probing mammalian touch transduction. J Gen Physiol 2011, 138: 291–301.

    Article  PubMed  CAS  Google Scholar 

  37. Li L, Rutlin M, Abraira VE, Cassidy C, Kus L, Gong S, et al. The functional organization of cutaneous low-threshold mechanosensory neurons. Cell 2011, 147: 1615–1627.

    Article  PubMed  CAS  Google Scholar 

  38. Cervero F, Iggo A, Ogawa H. Nociceptor-driven dorsal horn neurones in the lumbar spinal cord of the cat. Pain 1976, 2: 5–24.

    Article  PubMed  CAS  Google Scholar 

  39. Takazawa T, MacDermott AB. Synaptic pathways and inhibitory gates in the spinal cord dorsal horn. Ann N Y Acad Sci 2010, 1198: 153–158.

    Article  PubMed  CAS  Google Scholar 

  40. Baba H, Ji RR, Kohno T, Moore KA, Ataka T, Wakai A, et al. Removal of GABAergic inhibition facilitates polysynaptic A fibermediated excitatory transmission to the superficial spinal dorsal horn. Mol Cell Neurosci 2003, 24: 818–830.

    Article  PubMed  CAS  Google Scholar 

  41. Torsney C, MacDermott AB. Disinhibition opens the gate to pathological pain signaling in superficial neurokinin 1 receptor-expressing neurons in rat spinal cord. J Neurosci 2006, 26: 1833–1843.

    Article  PubMed  CAS  Google Scholar 

  42. Keller AF, Beggs S, Salter MW, De Koninck Y. Transformation of the output of spinal lamina I neurons after nerve injury and microglia stimulation underlying neuropathic pain. Mol Pain 2007, 3: 27.

    Article  PubMed  Google Scholar 

  43. Miraucourt LS, Moisset X, Dallel R, Voisin DL. Glycine inhibitory dysfunction induces a selectively dynamic, morphine-resistant, and neurokinin 1 receptor-independent mechanical allodynia. J Neurosci 2009, 29: 2519–2527.

    Article  PubMed  CAS  Google Scholar 

  44. Miraucourt LS, Dallel R, Voisin DL. Glycine inhibitory dysfunction turns touch into pain through PKCgamma interneurons. PLoS One 2007, 2: e1116.

    Article  PubMed  Google Scholar 

  45. Drew LJ, MacDermott AB. Neuroscience: Unbearable lightness of touch. Nature 2009, 462: 580–581.

    Article  PubMed  CAS  Google Scholar 

  46. Schmelz M. Itch and pain. Neurosci Biobehav Rev 2010, 34: 171–176.

    Article  PubMed  CAS  Google Scholar 

  47. Davidson S, Giesler GJ. The multiple pathways for itch and their interactions with pain. Trends Neurosci 2010, 33: 550–558.

    Article  PubMed  CAS  Google Scholar 

  48. Patel KN, Dong X. Itch: cells, molecules, and circuits. ACS Chem Neurosci 2011, 2: 17–25.

    Article  PubMed  Google Scholar 

  49. Ross SE. Pain and itch: insights into the neural circuits of aversive somatosensation in health and disease. Curr Opin Neurobiol 2011, 21: 880–887.

    Article  PubMed  CAS  Google Scholar 

  50. Jeffry J, Kim S, Chen ZF. Itch signaling in the nervous system. Physiology (Bethesda) 2011, 26: 286–292.

    Article  CAS  Google Scholar 

  51. Ikoma A, Cevikbas F, Kempkes C, Steinhoff M. Anatomy and neurophysiology of pruritus. Semin Cutan Med Surg 2011, 30: 64–70.

    Article  PubMed  CAS  Google Scholar 

  52. Johanek LM, Meyer RA, Friedman RM, Greenquist KW, Shim B, Borzan J, et al. A role for polymodal C-fiber afferents in nonhistaminergic itch. J Neurosci. 2008, 28: 7659–7669.

    Article  PubMed  CAS  Google Scholar 

  53. Patel KN, Dong X. An itch to be scratched. Neuron 2010, 68: 334–339.

    Article  PubMed  CAS  Google Scholar 

  54. Handwerker HO. Microneurography of pruritus. Neurosci Lett 2010, 470: 193–196.

    Article  PubMed  CAS  Google Scholar 

  55. Perl ER. Ideas about pain, a historical view. Nat Rev Neurosci 2007, 8: 71–80.

    Article  PubMed  CAS  Google Scholar 

  56. Woolf CJ, Ma Q. Nociceptors-noxious stimulus detectors. Neuron 2007, 55: 353–364.

    Article  PubMed  CAS  Google Scholar 

  57. Basbaum AI, Bautista DM, Scherrer G, Julius D. Cellular and molecular mechanisms of pain. Cell 2009, 139: 267–284.

    Article  PubMed  CAS  Google Scholar 

  58. Chen J. History of pain theories. Neurosci Bull 2011, 27: 343–350.

    Article  PubMed  Google Scholar 

  59. Schmelz M, Schmidt R, Bickel A, Handwerker HO, Torebjörk HE. Specific C-receptors for itch in human skin. J Neurosci 1997, 17: 8003–8008.

    PubMed  CAS  Google Scholar 

  60. Andrew D, Craig AD. Spinothalamic lamina I neurons selectively sensitive to histamine: a central neural pathway for itch. Nat Neurosci 2001, 4: 72–77.

    Article  PubMed  CAS  Google Scholar 

  61. Sun YG, Chen ZF. A gastrin-releasing peptide receptor mediates the itch sensation in the spinal cord. Nature 2007, 448: 700–703.

    Article  PubMed  CAS  Google Scholar 

  62. Sun YG, Zhao ZQ, Meng XL, Yin J, Liu XY, Chen ZF. Cellular basis of itch sensation. Science 2009, 325: 1531–1534.

    Article  PubMed  CAS  Google Scholar 

  63. Liu Q, Tang Z, Surdenikova L, Kim S, Patel KN, Kim A, et al. Sensory neuron-specific GPCR Mrgprs are itch receptors mediating chloroquine-induced pruritus. Cell 2009, 139: 1353–1365.

    Article  PubMed  Google Scholar 

  64. Liu Q, Weng HJ, Patel KN, Tang Z, Bai H, Steinhoff M, et al. The distinct roles of two GPCRs, MrgprC11 and PAR2, in itch and hyperalgesia. Sci Signal 2011, 4: ra45.

    Article  PubMed  CAS  Google Scholar 

  65. Wilson SR, Gerhold KA, Bifolck-Fisher A, Liu Q, Patel KN, Dong X, et al. TRPA1 is required for histamine-independent, Mas-related G protein-coupled receptor-mediated itch. Nat Neurosci 2011, 14: 595–602.

    Article  PubMed  CAS  Google Scholar 

  66. Green BG, Shaffer GS. The sensory response to capsaicin during repeated topical exposures: differential effects on sensations of itching and pungency. Pain 1993, 53: 323–334.

    Article  PubMed  CAS  Google Scholar 

  67. Schmelz M, Schmidt R, Weidner C, Hilliges M, Torebjork HE, Handwerker HO. Chemical response pattern of different classes of C-nociceptors to pruritogens and algogens. J Neurophysiol 2003, 89: 2441–2448.

    Article  PubMed  CAS  Google Scholar 

  68. Shim WS, Tak MH, Lee MH, Kim M, Kim M, Koo JY, et al. TRPV1 mediates histamine-induced itching via the activation of phospholipase A2 and 12-lipoxygenase. J Neurosci 2007, 27: 2331–2337.

    Article  PubMed  CAS  Google Scholar 

  69. Costa R, Marotta DM, Manjavachi MN, Fernandes ES, Lima-Garcia JF, Paszcuk AF, et al. Evidence for the role of neurogenic inflammation components in trypsin-elicited scratching behaviour in mice. Br J Pharmacol 2008, 154: 1094–1103.

    Article  PubMed  CAS  Google Scholar 

  70. Namer B, Seifert F, Handwerker HO, Maihöfner C. TRPA1 and TRPM8 activation in humans: effects of cinnamaldehyde and menthol. Neuroreport 2005, 16: 955–959.

    Article  PubMed  CAS  Google Scholar 

  71. Graham DT, Goodell H, Wolff HG. Neural mechanisms involved in itch, itchy skin, and tickle sensations. J Clin Invest 1951, 30: 37–49.

    Article  PubMed  CAS  Google Scholar 

  72. Ward L, Wright E, McMahon SB. A comparison of the effects of noxious and innocuous counterstimuli on experimentally induced itch and pain. Pain 1996, 64: 129–138.

    Article  PubMed  CAS  Google Scholar 

  73. Atanassoff PG, Brull SJ, Zhang J, Greenquist K, Silverman DG, Lamotte RH. Enhancement of experimental pruritus and mechanically evoked dysesthesiae with local anesthesia. Somatosens Mot Res 1999, 16: 291–298.

    Article  PubMed  CAS  Google Scholar 

  74. Akiyama T, Iodi Carstens M, Carstens E. Pruritic and algesic facial stimuli excite partly overlapping populations of primary and second-order trigeminal neurons in mice. J Neurophysiol 2010, 104(5): 2442–2450.

    Article  PubMed  CAS  Google Scholar 

  75. Liu Y, Abdel Samad O, Duan B, Zhang L, Tong Q, Ji RR, et al. VGLUT2-dependent glutamate release from peripheral nociceptors is required to sense pain and suppress itch. Neuron 2010, 68: 543–556.

    Article  PubMed  CAS  Google Scholar 

  76. Sikand P, Shimada SG, Green BG, LaMotte RH. Sensory responses to injection and punctate application of capsaicin and histamine to the skin. Pain 2011, 152: 2485–2494.

    Article  PubMed  CAS  Google Scholar 

  77. Sikand P, Shimada SG, Green BG, LaMotte RH. Similar itch and nociceptive sensations evoked by punctate cutaneous application of capsaicin, histamine and cowhage. Pain 2009, 144: 66–75.

    Article  PubMed  CAS  Google Scholar 

  78. Lagerström MC, Rogoz K, Abrahamsen B, Persson E, Reinius B, Nordenankar K, et al. VGLUT2-dependent sensory neurons in the TRPV1 population regulate pain and itch. Neuron 2010, 68: 529–542.

    Article  PubMed  Google Scholar 

  79. Koga K, Chen T, Li XY, Descalzi G, Ling J, Gu J, et al. Glutamate acts as a neurotransmitter for gastrin releasing peptide-sensitive and insensitive itch-related synaptic transmission in mammalian spinal cord. Mol Pain 2011, 7: 47.

    Article  PubMed  CAS  Google Scholar 

  80. Davidson S, Zhang X, Khasabov SG, Simone DA, Giesler GJ Jr. Relief of itch by scratching: state-dependent inhibition of primate spinothalamic tract neurons. Nat Neurosci 2009, 12: 544–546.

    Article  PubMed  CAS  Google Scholar 

  81. Ross SE, Mardinly AR, McCord AE, Zurawski J, Cohen S, Jung C, et al. Loss of inhibitory interneurons in the dorsal spinal cord and elevated itch in Bhlhb5 mutant mice. Neuron 2010, 65: 886–898.

    Article  PubMed  CAS  Google Scholar 

  82. Woolf CJ. Central sensitization: implications for the diagnosis and treatment of pain. Pain 2011, 152(3 Suppl): S2–15.

    Article  PubMed  Google Scholar 

  83. McMahon SB, Malcangio M. Current challenges in glia-pain biology. Neuron 2009, 64: 46–54.

    Article  PubMed  CAS  Google Scholar 

  84. Kuner R. Central mechanisms of pathological pain. Nat Med 2010, 16: 1258–1266.

    Article  PubMed  CAS  Google Scholar 

  85. Gao YJ, Ji RR. Chemokines, neuronal-glial interactions, and central processing of neuropathic pain. Pharmacol Ther 2010, 126: 56–68.

    Article  PubMed  CAS  Google Scholar 

  86. Liu M, Wood JN. The roles of sodium channels in nociception: implications for mechanisms of neuropathic pain. Pain Med 2011, 12 (Suppl 3): S93–99.

    Article  PubMed  Google Scholar 

  87. Campbell JN, Raja SN, Meyer RA, Mackinnon SE. Myelinated afferents signal the hyperalgesia associated with nerve injury. Pain 1988, 32: 89–94.

    Article  PubMed  CAS  Google Scholar 

  88. Price DD, Bennett GJ, Rafii A. Psychophysical observations on patients with neuropathic pain relieved by a sympathetic block. Pain 1989, 36: 273–288.

    Article  PubMed  CAS  Google Scholar 

  89. Koltzenburg M, Lundberg LE, Torebjörk HE. Dynamic and static components of mechanical hyperalgesia in human hairy skin. Pain 1992, 51: 207–219.

    Article  PubMed  CAS  Google Scholar 

  90. Torebjörk HE, Lundberg LE, LaMotte RH. Central changes in processing of mechanoreceptive input in capsaicin-induced secondary hyperalgesia in humans. J Physiol 1992, 448: 765–780.

    PubMed  Google Scholar 

  91. Price TJ, Cervero F, Gold MS, Hammond DL, Prescott SA. Chloride regulation in the pain pathway. Brain Res Rev 2009, 60: 149–170.

    Article  PubMed  CAS  Google Scholar 

  92. Zhang Z, Cai YQ, Zou F, Bie B, Pan ZZ. Epigenetic suppression of GAD65 expression mediates persistent pain. Nat Med 2011, 17: 1448–1455.

    Article  PubMed  CAS  Google Scholar 

  93. Moore KA, Kohno T, Karchewski LA, Scholz J, Baba H, Woolf CJ. Partial peripheral nerve injury promotes a selective loss of GABAergic inhibition in the superficial dorsal horn of the spinal cord. J Neurosci 2002, 22: 6724–6731.

    PubMed  CAS  Google Scholar 

  94. Polgár E, Todd AJ. Tactile allodynia can occur in the spared nerve injury model in the rat without selective loss of GABA or GABA(A) receptors from synapses in laminae I–II of the ipsilateral spinal dorsal horn. Neuroscience 2008, 156: 193–202.

    Article  PubMed  Google Scholar 

  95. Coull JA, Beggs S, Boudreau D, Boivin D, Tsuda M, Inoue K, et al. BDNF from microglia causes the shift in neuronal anion gradient underlying neuropathic pain. Nature 2005, 438: 1017–1021.

    Article  PubMed  CAS  Google Scholar 

  96. Coull JA, Boudreau D, Bachand K, Prescott SA, Nault F, Sík A, et al. Trans-synaptic shift in anion gradient in spinal lamina I neurons as a mechanism of neuropathic pain. Nature 2003, 424: 938–942.

    Article  PubMed  CAS  Google Scholar 

  97. Beyer C, Roberts LA, Komisaruk BR. Hyperalgesia induced by altered glycinergic activity at the spinal cord. Life Sci 1985, 37: 875–882.

    Article  PubMed  CAS  Google Scholar 

  98. Roberts LA, Beyer C, Komisaruk BR. Nociceptive responses to altered GABAergic activity at the spinal cord. Life Sci 1986, 39: 1667–1674.

    Article  PubMed  CAS  Google Scholar 

  99. Yaksh TL. Behavioral and autonomic correlates of the tactile evoked allodynia produced by spinal glycine inhibition: effects of modulatory receptor systems and excitatory amino acid antagonists. Pain 1989, 37: 111–123.

    Article  PubMed  CAS  Google Scholar 

  100. Belmonte C, Brock JA, Viana F. Converting cold into pain. Exp Brain Res 2009, 196: 13–30.

    Article  PubMed  Google Scholar 

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  1. Dana-Farber Cancer Institute and Department of Neurobiology, Harvard Medical School, Boston, Massachusetts, 02115, USA

    Qiufu Ma

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Ma, Q. Population coding of somatic sensations. Neurosci. Bull. 28, 91–99 (2012). https://doi.org/10.1007/s12264-012-1201-2

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