Biological Cybernetics

, Volume 72, Issue 3, pp 197–206 | Cite as

A neural network model of phantom limbs

  • Manfred Spitzer
  • Peter Böhler
  • Matthias Weisbrod
  • Udo Kischka


This paper presents a detailed clinical description of phantom limbs and a neuronal network model that provides a comprehensive and parsimonious explanation of otherwise inexplicable or at least unrelated phenomena. Simulations of self-organizing feature maps (Kohonen networks) that had been trained to recognize input patterns were deprived of parts of their input in order to simulate partial deafferentation. This leads to reorganization processes that are shown to be driven by input noise. In patients with an amputated limb, this noise is generated by dorsal root ganglion sensory neurons which are known to fire irregularly upon laceration. According to this model, the long-standing debate concerning non-cortical vs. cortical contributions to the generation of the phenomenon of phantom limbs can be resolved in that it is the peripherally generated noise that causes cortical reorganization. The model can be tested and may have therapeutic implications.


Dorsal Root Ganglion Sensory Neuron Neural Network Model Input Pattern Amputate Limb 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.


Unable to display preview. Download preview PDF.

Unable to display preview. Download preview PDF.


  1. Anderson JA, Pellionisz A, Rosenfeld E (1990) Introduction (to chapters 17,18, and 19). In: Anderson JA, Pellionisz A, Rosenfeld E (eds) Neurocomputing 2. Directions for research. MIT Press, Cambridge, Mass. pp 295–299Google Scholar
  2. Antonini A, Stryker MP (1993) Rapid remodeling of axonal arbors in the visual cortex. Science 260:1819–1821Google Scholar
  3. Bach y Rita P (1990) Brain plasticity as a basis for recovery of function in humans. Neuropsychologia 28:547–554Google Scholar
  4. Bors E (1951) Phantom limbs of patients with spinal cord injury. Arch Neurol Psychiatr 66:610–631Google Scholar
  5. Buchanen DC, Mandel AR (1986) The prevalence of phantom limb experience in amputees. Rehab Psychol 31:183–188Google Scholar
  6. Carlen PL, Wall PD, Nadvorna H, Steinbach T (1978) Phantom limbs and related phenomena in recent traumatic amputations. Neurology 28:211–217Google Scholar
  7. Caudill M, Butler C (1992) Understanding neural networks, Vols 1 and 2. MIT Press, Cambridge, MassGoogle Scholar
  8. Changeux J-P, Heidmann T, Patte P (1984) Learning by selection. In: Marker P, Terrence HS (eds) The biology of learning. Springer, Berlin Heidelberg New York [Quoted from: pp 115–133 Rosenfeld E, Pellionisz A, Anderson JA (eds) (1990) Neurocomputing 2. Directions for research. MIT Press, Cambridge, Mass. pp 300–307Google Scholar
  9. Churchland PS, Koch C, Sejnowski TJ (1990) What is computational neuroscience? In: Schwartz EL (eds) Computational neuroscience. MIT Press, Cambridge, Mass. pp 38–45Google Scholar
  10. Cohen J, Servan-Schreiber D (1992) Context, cortex and dopamine: a connectionist approach to behavior and biology in schizophrenia. Psychol Rev 12:45–77Google Scholar
  11. Creutzfeld DO (1993) Cortex cerebri. Sellstverlag GöttingenGoogle Scholar
  12. Crick F (1988) What mad pursuit. Basic Books, New YorkGoogle Scholar
  13. Crick F (1989) The recent excitement about neural networks. Nature 337:129–132Google Scholar
  14. Cronholm B (1951) Phantom limbs in amputees. Acta Psychiatr Neurol Scand Suppl 72:1–310Google Scholar
  15. Devor M (1984) The pathophysiology and anatomy of damaged nerve. In: Wall PD, Melzak M (eds) Textbook of pain. Churchill-Livingstone, Edinburgh, pp 49–64Google Scholar
  16. Douglas JK, Wilkens L, Pantazelou E, Moss F (1993) Noise enhancement of information transfer in crayfish mechanoreceptors by stochastic resonance. Nature 365:337–340Google Scholar
  17. Edelman GM, Finkel LH (1984) Neuronal group selection in the cerebral cortex. In: Edelman GM, Einar-Gall W, Maxwell W (eds) Dynamic aspects of neocortical function. Wiley-Interscience, New York, pp 653–595 [Quoted from: Anderson, JA, Pellionisz, A Rosenfeld E (ed) (1990) Neurocomputing 2. Directions for research. MIT Press, Cambridge, Mass. pp 308–334Google Scholar
  18. Gilbert CD (1993) Rapid dynamic changes in adult cerebral cortex. Curr Opin Neurobiol 3:100–103Google Scholar
  19. Halligan PW, Marshall JC, Wade DT, Davey J, Morrison D (1993) Thumb in cheek? Sensory reorganization and perceptual plasticity after limb amputation. Neuroreport 4:233–236Google Scholar
  20. Hoffman J (1954) Phantom limb syndrome. A critical review of the literature. J Nerv Ment Dis 119:261–270Google Scholar
  21. Jensen TS, Rasmussen P (1989) Phantom pain and related phenomena after amputation. In: Wall PD, Melzack R (eds) Textbook of pain, 2nd edn Livingstone Churchill, EdinburghGoogle Scholar
  22. Jensen TS, Krebs B, Nielsen J, Rasmussen P (1984) Non-painful phantom limb phenomena in amputees: incidence, clinical characteristics and temporal course. Acta Neurol Scand 70:407–414Google Scholar
  23. Katz J (1992) Psychophysiological contributions to phantom limbs. Can J Psychiatry 37:282–298Google Scholar
  24. Kohonen T (1982) Self-organized formation of topologically correct feature maps. Biol Cybern 43:59–69Google Scholar
  25. Kohonen T (1989) Self-organization and associative memory, 3rd edn Springer, Berlin Heidelberg New YorkGoogle Scholar
  26. Kosslyn S, Koenig O (1992) Wet Mind. The New Cognitive Neuroscience. Macmillan, New York, Toronto, Oxford, Singapore, SydneyGoogle Scholar
  27. Maddox J (1994) Bringing more order out of noisiness. Nature 369:271Google Scholar
  28. Merzenich MM, Kaas JH, Wall J, Nelson RJ, Sur M, Felleman D (1983) Topographic reorganization of somatosensory cortical areas 3b and 1 in adult monkeys following restricted deafferentation. Neuroscience 8:33–55Google Scholar
  29. Merzenich MM, Sameshima K (1993) Cortical plasticity and memory. Curr Opin Neurol 3:187–196Google Scholar
  30. Miikkulainen R (1993) Subsymbolic natural language processing. An integrated model of scripts, lexicon, and memory. MIT Press, Cambridge Mass.Google Scholar
  31. Mitchell SW (1871) Phantom limbs. Lippinocott Mag Pop Lit Sci 8:563–569Google Scholar
  32. Morrison JH, Hof PR (1992) The organization of cerebral cortex: from molecules to circuits. Discuss Neurosci 9:1–80Google Scholar
  33. Neumann von (1958) The computer and the brain. Yale University Press, New HavenGoogle Scholar
  34. O'Leary DDM (1992) Development of connectional diversity and specificity in the mammalian brain by the pruning of collateral projections. Curr Opin Neurobiol 2:70–77Google Scholar
  35. Penfield W, Rasmussen T (1950) The cerebral cortex of man: a clinical study of localization and function. Macmillan, New YorkGoogle Scholar
  36. Poeck K (1963) Zur Psychophysiologie der Phantomerlebnisse. Nervenarzt 34:241–256Google Scholar
  37. Pons TP, Garraghty PE, Ommaya AK, Kaas JH, Taub E, Mishkin M (1991) Massive cortical reorganization after sensory deafferentation in adult macaques. Science 252:1857–1860Google Scholar
  38. Ramachandran VS, Rogers-Ramachandran D, Steward M (1992) Perceptual correlates of massive cortical reorganization. Science 258:1159–1160Google Scholar
  39. Recanzone GH, Jenkins WM, Hradek GT, Merzenich MM (1992a) rogressive improvement in discriminative abilities in adult owl monkeys performing a tactile frequency discrimination task. J Neurophysiol 67:1015–1030Google Scholar
  40. Recanzone GH, Merzenich MM, Jenkins WM, Grajski KA, Dinse HR (1992b) Topographic reorganization of the hand representation in cortical area 3b of owl monkeys trained in a frequency-discrimination task. Neuroiphysiol 67:1031–1056Google Scholar
  41. Recanzone GH, Merzenich MM, Schreiner CE (1992c) Changes in the distributed temporal response properties of SI cortical neurons reflect improvements in performance on a temporally based tactile discrimination task. J Neurophysiol 67:1071–1091Google Scholar
  42. Recanzone GH, Schreiner CE, Merzenich MM (1993) Plasticity in the frequency representation of primary auditory cortex following discrimination training in adult owl monkeys. J Neurosci 13:87–103Google Scholar
  43. Ritter H (1991) Neuronale Netze. Addison-Wesley, ReadingGoogle Scholar
  44. Ritter H, Kohonen T (1989) Self-organizing semantic maps. Biol Cybern 61:241–254Google Scholar
  45. Rumelhart D (1989) The architecture of mind: A connectionist approach. In: Posner M (eds) Foundations of Cognitive Science. MIT Press, Cambridge, Mass. pp 133–159Google Scholar
  46. Schilder P (1923) Über elementate Halluzinationen des Bewegungssehens. Z Neurol Psychiatrie 80:424–431Google Scholar
  47. Shepherd GM (1990) The significance of real neuron architectures for neural network simulations. In: Schwartz EL (ed) Computational neuroscience. MIT Press, Cambridge, Mass. pp 82–96Google Scholar
  48. Spitzer M (1988) Halluzinationen. Springer. Berlin Heidelberg New YorkGoogle Scholar
  49. Spitzer M, Braun U, Hermle L, Maier S (1993) Associative semantic network dysfunction in thought-disordered schizophrenic patients: direct evidence from indirect semantic priming. Biol. Psychiatry 34:864–877Google Scholar
  50. Sutton G, Reggia J, Armentrout S, D'Autrechy C (1994) Cortical map reorganization as a competitive process. Neural Comp 6:1–13Google Scholar
  51. Thomson AM, Deuchars J (1994) Temporal and spatial properties of local circuits in neocortex Trends Neuro Sci 17:119–126Google Scholar
  52. Weinstein S, Sersen EA, Vetter RJ (1964) Phantoms and somatic sensation in cases of congenital aplasia. Neurology 10:905–911Google Scholar
  53. Weiss SA, Fishman S (1963) Extended and telescoped phantom limbs in unilateral amputees. l J Abnorm Soc Psychol 66:489–497Google Scholar
  54. Welk E, Leah JD, Zimmermann M (1990) Characteristics of A- and C-fibers ending in a sensory nerve neuroma in the rat. J. Neurophysiol 63:759–766Google Scholar
  55. Zuk GH (1956) The phantom limb: a proposed theory of unconscious origins. J Nerv Ment Dis 124:510–513Google Scholar

Copyright information

© Springer-Verlag 1995

Authors and Affiliations

  • Manfred Spitzer
    • 1
  • Peter Böhler
    • 1
  • Matthias Weisbrod
    • 1
  • Udo Kischka
    • 1
  1. 1.Section of Experimental PsychopathologyPsychiatrische Universitätsklinik HeidelbergHeidelbergGermany

Personalised recommendations