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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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Foglia, L., & Grush, R. (2010) · Behavioral and Brain Sciences · doi:10.1017/s0140525x10001147 · Cited by 2
Self-citationADAPTIVE CONTROL LOOPS AS AN INTERMEDIATE MIND-BRAIN REDUCTION BASIS
Proust, J. (2009) · De Gruyter eBooks · doi:10.1515/9783110328851.191 · Cited by 2
“…and Euler, were eager to claim, a variational constraint can and should only be explained by an agent's inten24 For a similar view of the mind, see Grush (2004) & Hurley (2008), The grain of truth in enactive theories of perception (Noe, 2004) is that perception is functionally engaged in the…”
Detecting Affordances by Visuomotor SimulationPreprint
Schenck, W., Hasenbein, H., & Möller, R. (2016) · arXiv (Cornell University) · doi:10.48550/arxiv.1611.00274 · Cited by 3
“…theories” were proposed in cognitive science (and related areas) which basically state that simulation is an important foundation for perception and higher–level cognition (Hesslow, 2002; Cruse, 2003; Wolpert et al., 2003; Holland and Goodman, 2003; Grush, 2004; Pezzulo and Castelfranchi, 2009).”
The Concept of Simulation in Control-Theoretic Accounts of Motor Control and Action Perception
Herschbach, M. (2008) · eScholarship (California Digital Library) · Cited by 2
Villalobos, M. (2015) · ERA · Cited by 2
Raffin, E. (2011) · theses.fr (ABES) · doi:10.70675/278b493fzc4b2z450dz83c0z6b2036a820dd · Cited by 2
Understanding rumination as a form of inner speech: probing the role of motor processesPreprint
Nalborczyk, L. (2019) · Thesis Commons (OSF Preprints) · doi:10.31237/osf.io/p6dct · Cited by 4
“In the emulation model proposed by Grush (2004), the emulator is a device that implements the same input-output function as the body (i.e., the musculoskeletal system and relevant sensory systems).2 When the emulator receives a copy of the control signal 2In Grush’s terminology, emulator is used as…”
InfluentialImagined Head Rotations Facilitate Spatial MemoryPreprint
Wyssen, G., Mast, F. W., & Ertl, M. (2022) · Research Square · doi:10.21203/rs.3.rs-1316811/v1 · Cited by 2
Biologically Inspired Robot Body Models and Self-Calibration
Hoffmann, M. (2021) · Springer eBooks · doi:10.1007/978-3-642-41610-1_201-1 · Cited by 2
Impaired prediction of ongoing events in posttraumatic stress disorderPreprint
Eisenberg, M. L., Rodebaugh, T. L., Flores, S. E., & Zacks, J. M. (2022) · doi:10.31234/osf.io/2jpvg · Cited by 2
A Drive to Survive: the free energy principle and the meaning of lifePreprint
Nave, K. (2023) · doi:10.31234/osf.io/ds9mn · Cited by 2
Visual motion estimation and tracking of rigid bodies by physical simulation
Duff, D. J. (2011) · UPT. Syiah Kuala University Library (Syiah Kuala University) · Cited by 2
Buzsáki, G. (2019) · Oxford University Press eBooks · doi:10.1093/oso/9780190905385.003.0003 · Cited by 1
The Neural Mechanisms Underlying Social Norms
Mu, Y., & Gelfand, M. J. (2020) · Cambridge University Press eBooks · doi:10.1017/9781108695374.015 · Cited by 2
Salve, J., Upadhyay, P., Mashood, K. K., & Chandrasekharan, S. (2024) · Science & Education · doi:10.1007/s11191-023-00488-7 · Cited by 1
Nalborczyk, L., Alario, F. ‐., & Longcamp, M. (2024) · Cognition · doi:10.1016/j.cognition.2024.105997 · Cited by 1
“A prominent proposal is that during motor imagery, the sensory consequences of actions may be simulated mentally using pairs of internal models developed for the control of overt actions (e.g., Grush, 2004; Jeannerod, 1994, 2001).”
Knowing that as knowing how: a neurocognitive practicalism
Piccinini, G., & Hetherington, S. (2024) · Synthese · doi:10.1007/s11229-024-04843-1 · Cited by 2
Liu, Y., Wang, Y., Zhao, X., Hu, P., & Hu, Y. (2025) · Brain and Behavior · doi:10.1002/brb3.70826 · Cited by 1
“MI training involves the continuous offline activation of the “emulator” through efferent neural currents, which facilitates the repair or reorganization of impaired motor pathways, boosts synaptic efficiency, and reduces presynaptic inhibition (Grush 2004; F. Li et al. 2018; Ruffino et al. 2017).”
Do low imagers know more words? examining the association between mental imagery and vocabulary size
Yavuz, M., & Nazir, T. A. (2026) · Language Cognition and Neuroscience · doi:10.1080/23273798.2026.2614588 · Cited by 1
Suemitsu, K., & Nambu, I. (2026) · Frontiers in Human Neuroscience · doi:10.3389/fnhum.2025.1713908 · Cited by 1
“Even in the absence of voluntary movement, motor imagery engages internal forward models that generate predicted sensory consequences based on action intention (Jeannerod, 2001; Wolpert et al., 1995; Grush, 2004; Kilteni et al., 2018).”
InfluentialDistance-dependent modulation of sensory attenuation during motor imagery and overt action
Lee, K.-B., Cho, H.-T., Huh, D.-C., & Lee, J.-H. (2026) · Experimental Brain Research · doi:10.1007/s00221-026-07357-z · Cited by 1
“According to the simulation hypothesis (Jeannerod 2001), motor imagery is assumed to rely on internal mod - els that are functionally equivalent to those engaged during overt movement, implying that both forms of action share a common representational structure (Decety 1996 ; Grush 2004).”
A Neurocomputational Model of Anticipation and Sustained Inattentional Blindness in Hierarchies
Morse, A. F., Lowe, R. J., & Tom Ziemke, B. (2009) · Lecture notes in computer science · doi:10.1007/978-3-642-02565-5_9 · Cited by 1
“…it is clear that the ability to predict would be extremely useful to any embodied agent interacting with a complex environment; for example, simulating or anticipating the effects of possible actions for evaluation in an agent-centered way e.g. (Clark & Grush, 1999; Grush, 2004; Hesslow, 2002).”
InfluentialEmbodied Moving-Target Seeking with Prediction and Planning
Oses, N., Hoffmann, M., & Koene, R. A. (2010) · Lecture notes in computer science · doi:10.1007/978-3-642-13803-4_59 · Cited by 1
“At the same time, our scenario is a case for minimalistic model of cognition which is firmly grounded in body dynamics [13,14,15,16,17].”
Internal Simulation of an Agent’s Intentions
Johnsson, M., & Buonamente, M. (2012) · Advances in intelligent systems and computing · doi:10.1007/978-3-642-34274-5_32 · Cited by 1
Piemonte, M. E. P., Kopczynski, M. C., Voos, M. C., Miranda, C. S., & Oliveira, T. D. P. (2015) · Journal of Education and Training Studies · doi:10.11114/jets.v3i3.665 · Cited by 1
“In this situation, a forward sensory model would be used to predict the sensory consequences of the movements (Avanzino et al., 2009; Desmurget et al., 2009; Grush, 2004; Haruno, Wolpert, & Kawato, 2001; Tian & Poeppel, 2012).”
Grounding compositional symbols: no composition without discrimination
Greco, A., & Carrea, E. (2011) · Cognitive Processing · doi:10.1007/s10339-011-0427-7 · Cited by 1
A cognitive robotic model of mental rotation
Seepanomwan, K., Caligiore, D., Baldassarre, G., & Cangelosi, A. (2013) · IEEE Symposium on Computational Intelligence, Cognitive Algorithms, Mind, and Brain (CCMB) · doi:10.1109/ccmb.2013.6609163 · Cited by 1
“Another hypothesis on how premotor areas participate in mental rotation is based on the forward model theory of motor control [20][21][22].”
Accounting for Consciousness: Epistemic and Operational Issues
Peters, F. (2014) · Axiomathes · doi:10.1007/s10516-014-9232-0 · Cited by 1
Morgan, A. D. (2014) · doi:10.7282/t3f47mff · Cited by 1
“When evaluating the significance of Shagrir’s example, it’s important to keep in mind that Ramsey allows that some connectionist theories invoke structural representations; he cites Grush’s (2004) ‘emulation theory’ as an example.”
Consciousness and apparent motion: Paradox resolved
Aksentijevic, A. (2015) · Theory & Psychology · doi:10.1177/0959354315621401 · Cited by 1
“Different solutions have been suggested— from the warping of the mental time line or multiple drafts (Dennett & Kinsbourne, 1992) and time markers (Grush, 2004), to the contraction of an extended interval into a brief moment (Tye, 2003), different processing speeds for location and motion in the…”
Methodological consequences of weak embodied cognition and shared intentionality
Thompson, J. J., Sameen, N., & Racine, T. P. (2016) · New Ideas in Psychology · doi:10.1016/j.newideapsych.2016.03.002 · Cited by 2
Intentionality of Strong Anticipation in Motor Behaviors.
Liu, D. H.-W. (2012) · Cognitive Science · Cited by 1
Eye to I: Males Recognize Own Eye Movements, Females Inhibit Recognition.
Chandrasekharan, S., Date, G., Pande, P., Rahaman, J., Shaikh, R., Srivastava, A., Srivastava, N., & Agrawal, H. (2015) · eScholarship (California Digital Library) · Cited by 1
9 Embedded Cognition and Computation
Rupert, R. D. (2009) · Oxford University Press eBooks · doi:10.1093/acprof:oso/9780195379457.003.0009 · Cited by 1
Conceptualizing Simulation Theory
Goldman, A. I. (2006) · Oxford University Press eBooks · doi:10.1093/0195138929.003.0002 · Cited by 1
8 The Experience of Extension and the Extension of Experience
Rupert, R. D. (2009) · Oxford University Press eBooks · doi:10.1093/acprof:oso/9780195379457.003.0008 · Cited by 1
Modeling of Human Behavior Within the Paradigm of Modern Physics
Lubashevsky, I. (2017) · Understanding complex systems · doi:10.1007/978-3-319-51706-3_6 · Cited by 1
Ezquerro, J., & Iza, M. (2017) · Studies in applied philosophy, epistemology and rational ethics · doi:10.1007/978-3-319-43784-2_4 · Cited by 1
Monitoring prediction errors facilitates cognition in action.
Plass, J., Choi, S. T., Suzuki, S., & Grabowecky, M. (2018) · Journal of Experimental Psychology General · doi:10.1037/xge0000519 · Cited by 1
“…are generated to produce particular environmental effects, motor and premotor systems must predict the outcomes of potential actions to identify motor programs that are most consistent with the organism’s goals or expectations (Kawato & Wolpert, 1998; Grush, 2004; Friston, 2011; Clark, 2013, 2015).”
Effects of emulation training on activities of daily living and job-related skills.
Matthews, K. M., Newcomb, E. T., Morgan, C. A., Chen, T., & Vanderburg, N. M. (2019) · Behavior Analysis Research and Practice · doi:10.1037/bar0000161 · Cited by 1
Causation and Logic in Neuroscience
Buzsáki, G. (2019) · Oxford University Press eBooks · doi:10.1093/oso/9780190905385.003.0002 · Cited by 1
Tracing the Roots of Cognition in Predictive Processing
Pezzulo, G. (2017) · Theoretical Philosophy/MIND Group – JGU Mainz · doi:10.15502/9783958573215 · Cited by 33
“…Piaget’s proposals on the construction of intelligence from sensorimotor experience (Piaget and Cook 1952); but also in many more recent variants (Clark and Grush 1999; Cotterill 1998; Hesslow 2002; Grush 2004) and especially under the theoretical umbrella of action-oriented representation (AOR).”
Influential(2011) · eScholarship (California Digital Library) · Cited by 9
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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