Cited by 1,387Embodied and Extended CognitionMotor Control and AdaptationAction Observation and Synchronization

The emulation theory of representation: Motor control, imagery, and perception

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Grush, Rick·2004·Behavioral and Brain Sciences

Abstract

The emulation theory of representation is developed and explored as a framework that can revealingly synthesize a wide variety of representational functions of the brain. The framework is based on constructs from control theory (forward models) and signal processing (Kalman filters). The idea is that in addition to simply engaging with the body and environment, the brain constructs neural circuits that act as models of the body and environment. During overt sensorimotor engagement, these models are driven by efference copies in parallel with the body and environment, in order to provide expectations of the sensory feedback, and to enhance and process sensory information. These models can also be run off-line in order to produce imagery, estimate outcomes of different actions, and evaluate and develop motor plans. The framework is initially developed within the context of motor control, where it has been shown that inner models running in parallel with the body can reduce the effects of feedback delay problems. The same mechanisms can account for motor imagery as the off-line driving of the emulator via efference copies. The framework is extended to account for visual imagery as the off-line driving of an emulator of the motor-visual loop. I also show how such systems can provide for amodal spatial imagery. Perception, including visual perception, results from such models being used to form expectations of, and to interpret, sensory input. I close by briefly outlining other cognitive functions that might also be synthesized within this framework, including reasoning, theory of mind phenomena, and language.

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  • Predictive coding models are well-established in accounting for various features of perception, cognition, and motor control.

    “Predictive coding models are now well-established in accounting for various features of perception (Rao and Ballard, 1999; Yuille and Kersten, 2006), cognition (Grush, 2004), and motor control (Wolpert and Ghahramani, 2000)”

    From An Interoceptive Predictive Coding Model of Conscious Presence (2012) · cites this paper as Grush, 2004

    The citation to Grush (2004) is attached to cognition. Section 6.2, “Cognition” (page unavailable), connects anticipatory mental models to cognition, but says the emulation framework “might be a part of our eventual understanding” of it. That tentative account does not support describing predictive models as already well-established in accounting for cognition. (LLM, conf=0.87)

    Experimental — accuracy not yet measured
  • Grush proposes that the proprioceptive and kinesthetic feelings experienced in motor imagery arise from emulating intended motor commands.

    “Grush (2004) further proposes, based on the notion of a Kalman filter, that the proprioceptive and kinesthetic sensations produced during motor imagery are the consequences of emulation of planned motor commands.”
  • Common features of predictive processing and emulator frameworks include the appeal to forward models and mechanisms for estimating uncertainty and flexibly balancing prior expectations and driving sensory inputs.

    “Common features include the appeal to forward models and the provision of mechanisms (such as Kalman fi ltering -see Friston 2002; Grush 2004; Rao & Ballard 1999) for estimating uncertainty and (thus) fl exibly balancing the in fl uence of prior expectations and driving sensory inputs.”
  • Grush's account of perception as 'environmental emulation' looks highly congruent with the depiction of perception as reconstructing the hidden causes structuring the sensory signal.

    “In addition, Grush ' s account of perception as ' environmental emulation ' (see section 5.2 of Grush 2004) looks highly congruent with the depiction (Friston 2003 and elsewhere) of perception as reconstructing the hidden causes structuring the sensory signal.”
  • For couples, familiarity can lead to greater neural synchrony when familiar actions and communications are incorporated as neural predictions.

    “For our couples, who have been in a committed relationship for some time, such familiarity can lead to greater neural synchrony, when familiar actions56 and communications57 are incorporated as neural predictions and pave the way for greater brain coordination.”

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  1. Dartnall, T. H. (2003) Externalism extended. In: Proceedings of the Joint Fourth International Conference on Cognitive Science and the Seventh Australasian Society for Cognitive Science Conference, University of New South Wales, Sydney, Australia. July 2003, ed. P. Slezak, pp. 94 -99. The University of New South Wales Press.
  2. Dretske, F. I. (1988) Explaining behavior. MIT Press.
  3. Droulez, J. & Darlot, C. (1989) The geometric and dynamic implications of the coherence constraints in three-dimensional sensorimotor interactions. In: Attention and performance, vol. XIII, ed. M. Jeannerod, pp. 495 -526. Erlbaum.
  4. Eccles, J. C. (1979) Introductory remarks. In: Cerebro-cerebellar interactions, ed. J. Massion & K. Sasaki, pp. 10 -18. Elsevier.
  5. Fadiga, L., Fogassi, L., Gallese, V. & Rizzolatti, G. (2000) Visuomotor neurons: Ambiguity of the discharge or "motor" perception? International Journal of Psychophysiology 35:165 -77.
  6. Feynman, R. P. (2001) The pleasure of finding things out, 1st edition. Penguin Books.
  7. Freides, D. (1974) Human information processing and sensory modality: Cross- modal functions, information complexity and deficit. Psychological Bulletin 81:284 -310.
  8. Graziano, M. S. A. (1999) Where is my arm? Relative role of vision and proprioception in the neural representation of limb position. Proceedings of the National Academy of Sciences USA 96:10418 -21.
  9. Hanakawa, T., Immisch, I., Toma, K., Dimyan, M. A., van Gelderen, P. & Hallett, M. (2003c) Functional properties of brain areas associated with motor execution and imagery. Journal of Neurophysiology 89(2):989 -1002.
  10. Heisenberg, M. & Wolf, R. (1988) Reafferent control of optomotor yaw torque in Drosophila melongaster. Journal of Comparative Physiology A163:373-88.
  11. W. T. Miller, R. S. Sutton & P. J. Werbos. MIT Press.
  12. Hurley, S. (forthcoming a) Active perception and perceiving action: The shared circuits hypothesis. In: Perceptual experience, ed. T. Gendler & J. Hawthorne. Oxford University Press.
  13. Hurley, S. (forthcoming b) The shared circuits hypothesis: A unified functional architecture for control, imitation, and simulation. In: Perspectives on Imitation: From mirror neurons to memes, ed. S. Hurley & N. Chater. MIT Press.
  14. Johnson, M. (1987) The body in the mind. University of Chicago Press.
  15. 2003) The embodiment of intentionality In: Dynamical systems approaches to embodied cognition, ed. W. Tschacher, pp. 201-27. Springer Verlag.
  16. Karni, A., Meyer, G., Rey-Hipolito, C., Jezzard, P., Adams, M. M., Turner, R. & Ungerleider, L. G. (1998) The acquisition of skilled motor performance: Fast and slow experience-driven changes in primary motor cortex. Proceedings of the National Academy of Sciences USA 95:861-68.
  17. Kosslyn, S. M. (1994) Image and brain. MIT Press.
  18. Kosslyn, S. M., Ball, T. M. & Reiser, B. J. (1978) Visual imagers preserve metric spatial transformation: Evidence from studies of images scanning. Journal of Experimental Psychology: Human Perception and Performance 4:47-60.
  19. M. S. Gazzaniga, pp. 1035 -42. MIT Press. [arRG, OW]
  20. Llinas, R. & Pare, D. (1991) On dreaming and wakefulness. Neuroscience 44(3):521-35.
  21. Matthews, P. B. C. (1959) The dependence of tension upon extension in the stretch reflex of the soleus of the decerebrate cat. Journal of Physiology 47:521-46.
  22. Mel, B. W. (1986) A connectionist learning model for 3-d mental rotation, zoom, and pan. In: Proceedings of the Eighth Annual Conference of the Cognitive Science Society, pp. 562-71. Erlbaum. [aRG] (1988)
  23. Merfeld, D. M., Young, L., Oman, C. & Shelhamer, M. (1993) A multi-dimensional model of the effect of gravity on the spatial orientation of the monkey. Journal of Vestibular Research 3:141-61.
  24. Merfeld, D. M. & Zupan, L. H. (2002) Neural processing of gravitoinertial cues in humans. III. Modeling tilt and translation responses. Journal of Neurophysiology 87(2):819 -33.
  25. Milner, A. D. & Goodale, M. A. (1996) Visual brain in action. Oxford University Press.
  26. Naito, E., Ehrsson, H. H., Geyer, S., Zilles, K. & Roland, P. E. (1999) Illusory arm movements activate cortical motor areas: A PET study. Journal of Neuroscience 19:6134 -44.
  27. Niedenthal, P. M., Barsalou, L. W., Winkielman, P., Krauth-Gruber, S. & Ric, F. (in press) Embodiment in attitudes, social perception, and emotion. Personality and Social Psychology Review.
  28. O'Reilly, R. & Munakata, Y. (2000) Computational explorations in cognitive neuroscience: Understanding the mind by simulating the brain. MIT Press.
  29. Pisella, L., Grea, H., Tilikete, C., Vighetto, A., Desmurget, M., Rode, G., Boisson, D. & Rossetti, Y. (2000) An "automatic pilot" for the hand in human posterior parietal cortex: Toward reinterpreting optic ataxia. Nature Neuroscience 3(7):729 -36.
  30. Reed, C. (2002a) Chronometric comparisons of imagery to action: Visualizing versus physically performing springboard dives. Memory and Cognition 30(8):1169 -78.
  31. OD, CLR] (2002b) What is the body schema? In: The imitative mind, ed. A. Meltzoff & W. Prinz, pp. 233 -46. Cambridge University Press.
  32. Reisberg, D. & Chambers, D. (1991) Neither pictures nor propositions: What can we learn from a mental image? Canadian Journal of Psychology 45:366-52.
  33. Calvert, C. Spence & B. Stein, pp. 703-709. MIT Press.
  34. Shadmehr, R. & Wise, S. P. (2003) Motor learning and memory for reaching and pointing. In: The new cognitive neurosciences, 3rd edition, ed.
  35. M. S. Gazzaniga. MIT Press.
  36. Sperry, R. W. (1950) Neural basis of the spontaneous optokinetic response produced by vision inversion. Journal of Comparative and Physiological Psychology 43:482-89. [DMM, BW]
  37. Stein, B. E. & Meredith, M. A. (1993) Merging of the senses. MIT Press.
  38. Stojanov, G. (1997) Expectancy theory and interpretation of electroexpectograms (EXG) curves in the context of biological and machine intelligence. Ph.D. Thesis, Electrical Engineering Faculty, Saints Cyril and Methodius University, Skopje, Macedonia.
  39. Stojanov, G., Bozinovski, S. & Trajkovski, G. (1997a) Interactionist expectative view on agency and learning. IMACS Journal of Mathematics and Computers in Simulation 44:295 -310.
  40. 1997b) The status of representation in behaviour based robotic systems: The problem and a solution. Paper presented at the IEEE Conference on Systems, Man, and Cybernetics, Orlando, FL, 1997.
  41. Stojanov, G., Stefanovski, S. & Bozinovski, S. (1995) Expectancy based emergent environment models for autonomous agents. Proceedings of the 5th International Symposium on Automatic Control and Computer Science, Iasi, Romania 1:217-21.
  42. Tomasino, B., Borroni, P., Isaja, A., Baldiserra, F. & Rumiati, R. I. (in press) The primary motor cortex subserves not only movements but also their imagination. Cognitive Neuropsychology.
  43. Tversky, B. (2000) Remembering spaces. In: The Oxford handbook of memory, ed.
  44. E. Tulving & F. I. M. Craik. Oxford University Press.
  45. van Sommers, P. (1984) Drawing and cognition. Cambridge University Press.
  46. von Holst, E. & Mittelstädt, H. (1950/1973) Das Reafferenzprinzip: Wechselwirkungen zwischen Zentralnerven-system und Peripherie. Naturwissenschaften 37:467-76. (Original German publication, 1950.) En- glish translation, 1973: The reafference principle. In: The behavioral physiology of animals and man. The collected papers of Erich von Holst, trans. R. Martin, pp. 139 -73. University of Miami Press. [JSJ, MLL, DMM, BW]
  47. Wachholder, K. & Altenburger, H. (1927/2002) Do our limbs have only one rest length. Simultaneously a contribution to the measurement of elastic forces in active and passive movements. Pflüger's Archive für die gesamte Physiologie 215:627-40. (English translation by D. Sternard, 2002.) [see trans. in Sternad 2002]. [MLL] (2002) Foundational experiments for current hypotheses on equilibrium point control in voluntary movements. Motor Control 6:299 -318. (English translation by D. Sternard, 2002.)
  48. Wang, H., Johnson, T. R. & Zhang, J. (2001) The mind's views of space. In: Proceedings of the Third International Conference of Cognitive Science. Beijing.
  49. Wraga, M., Church, J. & Badre, D. (2002) Event-related fMRI study of imaginal self and object rotations. Journal of Cognitive Neuroscience 104:144.
  50. R. Alterman & D. Kirsch, pp. 1275-80. Lawrence Erlbaum Associates. [HW]

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Authors
Grush, R.
Affiliations
University of California San Diego (US)
Corresponding author
Grush, Rick
Venue
Behavioral and Brain Sciences
Published
2004-06
Volume
27
Issue
3
Pages
377-396
ISSN
  • 0140-525X (ISSN-L)
  • 1469-1825
OpenAlex
W2123786976
Cited by
1,387
References
312
Citations by year
2026: 32 · 2025: 39 · 2024: 65 · 2023: 39 · 2022: 59 · 2021: 47 · 2020: 136 · 2019: 66 · 2018: 55 · 2017: 49 · 2016: 38 · 2015: 74 · 2014: 66 · 2013: 97 · 2012: 50
FWCI
19.81
Type
journal article
Language
en
Open access
Locations
  • Behavioral and Brain Sciences (publishedVersion)
  • PubMed (publishedVersion)
  • eScholarship (California Digital Library) (submittedVersion)
Funders
  • Japan Society for the Promotion of Science (15700257, 15016113)
  • National Institute of Neurological Disorders and Stroke
Topics
  • Life Sciences › Neuroscience › Cognitive Neuroscience › Embodied and Extended Cognition
  • Life Sciences › Neuroscience › Cognitive Neuroscience › Motor Control and Adaptation
  • Social Sciences › Psychology › Social Psychology › Action Observation and Synchronization
Keywords
forward model, Kalman filter, efference copy, sensorimotor control, motor imagery, visual imagery, perception, feedback delay, motor planning, theory of mind, language, reasoning
MeSH terms
Brain, Environment, Humans, Imagination, Language, Models, Theoretical, Perception, Psychomotor Performance, Psychophysiology, Visual Perception

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