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The emulation theory of representation: Motor control, imagery, and perception
Open accessAbstract
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 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
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“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.”
From Mental imagery of speech and movement implicates the dynamics of internal forward models (2010) · cites this paper as Grush, 2004
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“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.”
From Whatever next? Predictive brains, situated agents, and the future of cognitive science (2013) · cites this paper as Grush 2004
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“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.”
From Whatever next? Predictive brains, situated agents, and the future of cognitive science (2013) · cites this paper as Grush 2004
In the citing paper
“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.”
From Brain-to-Brain Synchrony during Naturalistic Social Interactions (2017) · cites this paper as [56]
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Neural Simulation of Actions for Serpentine Robots
Morasso, P. (2024) · Biomimetics · doi:10.3390/biomimetics9070416 · Cited by 3
“…is the consequence of virtual force fi elds applied to an internal representation of the body, or generative body schema. cognitive agent to achieve prospection is an internal representation of the body or body schema, supported by a unifying simulation/emulation theory of action [31–33].”
Mechanisms of Change and Creativity in Nature and Culture
Dietrich, A. (2025) · Cambridge University Press eBooks · doi:10.1017/9781009091701 · Cited by 2
Chandrasekharan, S., Sinha, R., & Date, G. (2025) · Synthese · doi:10.1007/s11229-025-05081-9 · Cited by 3
“Such ‘off-line’ activation of forward model networks contributes to mental processes (Grush, 2004) – including motor imagery (Vingerhoets et al., 2002 ; Schubotz & Von Cramon, 2004; Fiorio et al. 2006), estimation of outcomes of different possible actions (Bosbach et al., 2005; Chandrasekharan et…”
The Nature of Mental Imagery: Beyond a Basic View
Shepherd, J. (2025) · Analysis · doi:10.1093/analys/anaf024 · Cited by 3
Scott, M. W. (2025) · Brain and Cognition · doi:10.1016/j.bandc.2025.106350 · Cited by 3
“The idea that MI is predictive in nature is not new to the literature (e. g., Blakemore & Sirigu, 2003), and has served as the basis of previous theories for MI (see Grush, 2004; Rieger et al., 2024 for examples).”
Explorando a trajetória espácio-temporal da representação dinâmica de projéteis
Teixeira, N. A. D. S., & Oliveira, A. M. (2013) · Psicologia Reflexão e Crítica · doi:10.1590/s0102-79722013000400012 · Cited by 3
“Este ponto está longe de ser displicente pois é hoje aceite que, aquando da visualização de um evento dinâmico, este é de alguma forma simulado ou emulado representacionalmente (e.g., Grush, 2004).”
Ganesh, G., & Ikegami, T. (2015) · Springer tracts in advanced robotics · doi:10.1007/978-3-319-25739-6_7 · Cited by 3
“In an inter-personal context, this theory believes that the observed actions can be understood by simulating them in one’s own brain using the same (or similar) internal models [93–96].”
Why Imagining Requires Content
Roelofs, L. (2018) · Thought A Journal of Philosophy · doi:10.1002/tht3.393 · Cited by 3
Worthman, C. M. (2020) · Cambridge University Press eBooks · doi:10.1017/9781108695374.007 · Cited by 3
Mental Imagery: Greasing the Mind's Gears
Cavedon‐Taylor, D. (2023) · Philosophers Imprint · doi:10.3998/phimp.1289 · Cited by 3
Agency at a distance: learning causal connections
Gärdenfors, P., & Lombard, M. (2024) · Phenomenology and the Cognitive Sciences · doi:10.1007/s11097-024-09992-9 · Cited by 3
“In control-theoretic terms, this model is called a feed-forward system since the simulator predicts what is going to happen in the real process (Grush, 2004; Kawato, 1999 ; Wolpert et al., 2003 , Gärd - cenfors, 2007b).”
Embodied simulation and knowledge of possibilities
Jones, M., & Schoonen, T. (2024) · Philosophical Psychology · doi:10.1080/09515089.2024.2417990 · Cited by 3
“However, perception is an active and dynamic process, which rarely, if ever, delivers us with anything like a static image (cf. Noë, 2002, 2004; O’Regan & Noë, 2001).”
Kusano, K., Hayashi, M., Iwama, S., & Ushiba, J. (2025) · Frontiers in Neural Circuits · doi:10.3389/fncir.2024.1510324 · Cited by 3
“Grush’s Emulation theory suggests that afferent feedback is involved in creating mental images of movement (Naito et al., 2002; Grush, 2004; Mizuguchi et al., 2009, 2012).”
Bai, Y., Brillinger, M., Karlinsky, A. D., Poliakoff, E., Welsh, T. N., & Gowen, E. (2025) · Scientific Reports · doi:10.1038/s41598-025-97036-w · Cited by 3
“Like action perception, MI is thought to occur via simulation involving the sub-threshold activation of perception-action codes for a specific movement 12,36,37 .”
The Perception–Action Hierarchy and its Implementation Using Binons (Binary Neurons)
Martensen, B. (2020) · Procedia Computer Science · doi:10.1016/j.procs.2020.02.220 · Cited by 3
The First Computational Theory of Cognition
Piccinini, G. (2020) · Oxford University Press eBooks · doi:10.1093/oso/9780198866282.003.0006 · Cited by 3
Künzell, S. (2021) · Springer eBooks · doi:10.1007/978-3-662-53386-4_56-1 · Cited by 3
The Perception of Bodily Expressions of Emotion and the Implications for Computing
Winand, H., & Anthony, P. (2008) · doi:10.5772/6181 · Cited by 3
“Work on motor control indicates ways in which such simulations may be computationally instantiated, in the form of forward models, inverse models, or both (Miall, 2003; Grush, 2004; Wolpert et al., 2003).”
Machine Consciousness in CiceRobot, a Museum Guide Robot.
Chella, A., & Macaluso, I. (2007) · National Conference on Artificial Intelligence · Cited by 3
A Modelling Framework for Functional Imagination
Marques, H. G., Holland, O. E., & Newcombe, R. A. (2008) · Cited by 3
Uncovering a Differentiated Theory of Mind in Children with Autism and Asperger Syndrome
Tine, M. T. (2009) · Cited by 3
The modal breadth of consciousness
Trestman, M. (2013) · Philosophical Psychology · doi:10.1080/09515089.2013.776476 · Cited by 3
“…action-oriented—hence oriented toward a range of possible, but non-compossible, futures—in more subtle ways as well.6 This is because the perception of an object is at least partly constituted by expectations of how else the object might appear (Grush, 2004; Noe, 2004; Zahavi, 1999, pp. 92–103).”
Cortical lateralization in stroke patients measured by event-related potentials during motor imagery
Gong, W., ZHANG, T., & Shan, L. (2013) · Molecular Medicine Reports · doi:10.3892/mmr.2013.1705 · Cited by 4
Henningsen, H. P. (2010) · Neurocomputing · doi:10.1016/j.neucom.2009.10.032 · Cited by 3
Wie wir unser motorisches System nutzen, um Ereignisse vorherzusagen
Schubotz, R. I. (2010) · Zeitschrift für Sportpsychologie · doi:10.1026/1612-5010/a000017 · Cited by 4
Vision-sensorimotor abstraction and imagination towards exploring robot’s inner world
Alnajjar, F. S. K., Hafiz, A. R., Zin, I. B. M., & Murase, K. (2008) · International Joint Conference on Neural Networks (IJCNN) · doi:10.1109/ijcnn.2008.4634134 · Cited by 3
Anticipating Action Effects with Different Attention Foci is Reflected in Brain Activation
Bischoff, M., Zentgraf, K., Pilgramm, S., Krüger, B., Balser, N., Sauerbier, I., Stark, R. M., & Munzert, J. (2015) · Perceptual and Motor Skills · doi:10.2466/22.24.pms.120v10x7 · Cited by 3
“It has been proposed that a predictive forward simulation in the human brain estimates the sensory eff ects of a movement, and that this prediction is based on one's own movement experience ( JohnsonLaird, 1983 ; Wolpert, et al. , 1995 ; Grush, 2004 ).”
Clark, A. (2008) · Oxford University Press eBooks · doi:10.1093/acprof:oso/9780195333213.003.0002 · Cited by 3
Motor Imagery and Mental Training in Older Adults
Kalicinski, M., Thomas, M., & Lobinger, B. (2016) · Elsevier eBooks · doi:10.1016/b978-0-12-803377-7.00008-9 · Cited by 3
Embodiment und Sense of Agency
Rieger, M., & Wenke, D. (2016) · Springer eBooks · doi:10.1007/978-3-642-53898-8_21 · Cited by 3
Computational mental simulation: A review
Polceanu, M., & Buche, C. (2016) · Computer Animation and Virtual Worlds · doi:10.1002/cav.1732 · Cited by 3
Welshon, R. (2019) · Oxford University Press eBooks · doi:10.1093/oso/9780190854614.003.0004 · Cited by 3
The Auditory Cortex of Primates Including Man With Reference to Speech
Rauschecker, J. P. (2020) · Elsevier eBooks · doi:10.1016/b978-0-12-805408-6.00029-4 · Cited by 3
Body models in humans, animals, and robots: mechanisms and plasticity
Hoffmann, M. (2021) · Oxford University Press eBooks · doi:10.1093/oso/9780198851721.003.0010 · Cited by 5
“This resembles the emulation theory of representation proposed by (Grush, 2004) who uses the Kalman filter 2 metaphor: the amodal, long-term body model is a central representation, which also includes its relationship with individual sensory modalities.”
AlphaGo, Locked Strategies, and Eco-cognitive Openness
Magnani, L. (2021) · Cognitive systems monographs · doi:10.1007/978-3-030-81447-2_3 · Cited by 3
Buzsáki, G. (2019) · Oxford University Press eBooks · doi:10.1093/oso/9780190905385.003.0001 · Cited by 4
Hohlefeld, F. U. (2011) · Universitätsbibliothek der FU Berlin Hochschulschriftenstelle u. Dokumentenserver · doi:10.17169/refubium-10445 · Cited by 3
“It is assumed that proprioception is simulated by using the efference copy of an issued motor command (forward-modeling; Grush, 2004; Wolpert & Gahramani, 2000; Wolpert et al., 2001).”
InfluentialTowards the Adoption of a Perception-Driven Perspective in the Design of Complex Robotic Systems
Chella, A., Cossentino, M., & Seidita, V. (2009) · Zenodo (CERN European Organization for Nuclear Research) · Cited by 3
Sensorimotor experience in speech perception
Schuerman, W. L. (2017) · Radboud Repository (Radboud University) · Cited by 3
HIGHER-ORDER ROBOT PERCEPTION LOOP
Chella, A., & Irene Macaluso, B. (2008) · Nova Science Publishers (Nova Science Publishers, Inc.) · Cited by 3
Gravity highlights a dual role of the insula in internal modelsPreprint
Rousseau, C., Barbiero, M., Pozzo, T., Papaxanthis, C., & White, O. (2019) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/659870 · Cited by 3
“This computational perspective assumes that actual and mental movements trigger similar motor representations (simulation Theory, Jeannerod, 2001) and similar predicted sensory consequences (emulation theory, Grush, 2004).”
Ecological Semantics: Programming Environments for Situated Language UnderstandingPreprint
Tamari, R., Stanovsky, G., Shahaf, D., & Tsarfaty, R. (2020) · arXiv (Cornell University) · doi:10.48550/arxiv.2003.04567 · Cited by 3
Displacement or Augmentation? The Effects of AI Innovation on Workforce Dynamics and Firm ValuePreprint
Chen, M. A., & Wang, X. (2025) · SSRN Electronic Journal · doi:10.2139/ssrn.5246388 · Cited by 4
Hossner, E.-J. (2024) · Sport und Gesellschaft · doi:10.1515/sug-2024-2012 · Cited by 2
Amodal completion in vision and audition: A relationship between perception and mental imagery
Bona, E. D. (2025) · Analysis · doi:10.1093/analys/anaf006 · Cited by 2
Even with exposure to errors, motor imagery cannot update internal models
Rowe, J. M., Cramer, B. C., & Boe, S. G. (2025) · Psychological Research · doi:10.1007/s00426-025-02138-9 · Cited by 2
“This interpretation aligns with the key principles of Motor Emulation Theory, particularly the idea that motor imagery has no capacity to influence the predictions of the internal model (or the ‘motor emulator’ as described by Grush), meaning that it cannot update internal models (Grush, 2004).”
InfluentialWilken, S., Böttcher, A., Beste, C., Raab, M., & Hoffmann, S. F. (2025) · Cerebral Cortex · doi:10.1093/cercor/bhaf158 · Cited by 5
Metadata
- Authors
- Grush, R.
- Affiliations
- University of California San Diego (US)
- Corresponding author
- Grush, Rick
- ORCID
- Grush, Rick
- Venue
- Behavioral and Brain Sciences
- Publisher
- Cambridge University Press (CUP)
- 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
- green
- www.escholarship.org
- 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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