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The emulation theory of representation: Motor control, imagery, and perception
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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
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“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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Radical Embodied Cognitive Science
Chemero, A. P. (2013) · Review of General Psychology · doi:10.1037/a0032923 · Cited by 282
“The representations they call on are indexical functional (Agre & Chapman, 1987), pushmi pullyu (Millikan, 1995), action-oriented (Clark, 1997), or emulator representations (Grush, 2004).”
An expanded role for the dorsal auditory pathway in sensorimotor control and integration
Rauschecker, J. P. (2010) · Hearing Research · doi:10.1016/j.heares.2010.09.001 · Cited by 294
Mental imagery of speech and movement implicates the dynamics of internal forward models
Tian, X. (2010) · Frontiers in Psychology · doi:10.3389/fpsyg.2010.00166 · Cited by 259
“Decety, J., Jeannerod, M., and Prablanc, C. (1989).”
InfluentialVartanian, O. A., Navarrete, G., Chatterjee, A. K., Fich, L. B., González–Mora, J. L., Leder, H., Modroño, C., Nadal, M., Rostrup, N., & Skov, M. (2014) · Journal of Environmental Psychology · doi:10.1016/j.jenvp.2014.11.006 · Cited by 306
“In fact, to the extent that this exploration is coupled with an intention to approach (or avoid) a space, it would also be accompanied by activation in brain regions implicated in motor imagery and/or planning of voluntary motor movement (Crammond, 1997; Decety, 1996; Dieber et al., 1998; Grush, 2004; Hanakawa, Dimyan…”
New directions in predictive processing
Hohwy, J. (2020) · Mind & Language · doi:10.1111/mila.12281 · Cited by 283
“Pioneering work from Chris Eliasmith (2003) and Rick Grush (2004) did not appeal to Friston’s approach but independently provided influential philosophical frameworks congenial to PP.”
Compromised motor control in children with DCD: A deficit in the internal model?—A systematic review
Adams, I. L. J., Lust, J. M., Wilson, P. H., & Steenbergen, B. (2014) · Neuroscience & Biobehavioral Reviews · doi:10.1016/j.neubiorev.2014.08.011 · Cited by 255
Body Schema in Robotics: A Review
Hoffmann, M., Marques, H. G., Arieta, A. H., Sumioka, H., Lungarella, M., & Pfeifer, R. (2010) · IEEE Transactions on Autonomous Mental Development · doi:10.1109/tamd.2010.2086454 · Cited by 253
“joint angle positions) that interact according to the laws of dynamics and mechanics [61].”
InfluentialEvolving intentions for social interaction: from entrainment to joint action
Knoblich, G., & Sebanz, N. (2008) · Philosophical Transactions of the Royal Society B Biological Sciences · doi:10.1098/rstb.2008.0006 · Cited by 269
“The second component consists of simulation mechanisms tapping into the observer's motor system (Blakemore & Decety 2001; Grush 2004; Wilson & Knoblich 2005).”
Information processing, computation, and cognition
Piccinini, G., & Scarantino, A. (2010) · Journal of Biological Physics · doi:10.1007/s10867-010-9195-3 · Cited by 252
“, [111–115]).”
Roles of supplementary motor areas in auditory processing and auditory imagery
Lima, C. F., Krishnan, S., & Scott, S. K. (2016) · Trends in Neurosciences · doi:10.1016/j.tins.2016.06.003 · Cited by 227
“These ideas resonate with the ‘emulation’ framework discussed in relation to action control, and generalized to motor imagery, perception, and prediction [112,113].”
Brain Activation during Anticipation of Sound Sequences
Leaver, A. M., Lare, J. V., Zielinski, B. A., Halpern, A. R., & Rauschecker, J. P. (2009) · Journal of Neuroscience · doi:10.1523/jneurosci.4921-08.2009 · Cited by 238
“This prospective signal may be related to the “efference copy” proposed in previous models (von Holst and Mittelstaedt, 1950; Troyer and Doupe, 2000), including so-called emulator (or “forward”) models (Grush, 2004; Callan et al., 2007; Ghasia et al., 2008).”
After the Philosophy of Mind: Replacing Scholasticism with Science*
Chemero, A. P., & Silberstein, M. (2008) · Philosophy of Science · doi:10.1086/587820 · Cited by 228
Gesture as simulated action: Revisiting the framework
Hostetter, A. B., & Alibali, M. W. (2018) · Psychonomic Bulletin & Review · doi:10.3758/s13423-018-1548-0 · Cited by 250
“Simulation has been contrasted with motor imagery (e.g., O’Shea & Moran, 2017; Willems, Toni, Hagoort, & Casasanto, 2009) and with emulation (e.g., Grush, 2004), and there is debate about the roles of conscious representations and internal models in simulation (e.g., Hesslow, 2012; Pezzulo,…”
Neural Computation and the Computational Theory of Cognition
Piccinini, G., & Bahar, S. (2012) · Cognitive Science · doi:10.1111/cogs.12012 · Cited by 241
“There is evidence that nervous systems contain and manipulate analog models of their environment (Grush, 2004; Waskan, 2003).”
Neural mechanisms for prediction: do insects have forward models?
Webb, B. H. (2004) · Trends in Neurosciences · doi:10.1016/j.tins.2004.03.004 · Cited by 221
“This article considers recent evidence from invertebrates that demonstrates the predictive modulation of sensory processes by motor output, and discusses to what extent this supports the conclusion that insect nervous systems also implement forward models.”
Nánay, B. (2013) · doi:10.1093/acprof:oso/9780199695379.001.0001 · Cited by 225
How Beat Perception Co-opts Motor Neurophysiology
Cannon, J. J., & Patel, A. D. (2020) · Trends in Cognitive Sciences · doi:10.1016/j.tics.2020.11.002 · Cited by 232
Embodied spatial transformations: "Body analogy" for the mental rotation of objects.
Amorim, M., Isableu, B., & Jarraya, M. (2006) · Journal of Experimental Psychology General · doi:10.1037/0096-3445.135.3.327 · Cited by 228
“This resonance would be part of an emulation process (Grush, 2004) that plays a role in representing and understanding the behavior of conspecifics (Wilson & Knoblich, 2005).”
The current status of the simulation theory of cognition
Hesslow, G. (2011) · Brain Research · doi:10.1016/j.brainres.2011.06.026 · Cited by 226
“…a combination of ideas with a long history, going back to associationist philosophers such as David Hume (1739) and Alexander Bain (1868) and similar ideas have been suggested by several modern authors (too numerous to be credited here but see for instance Grush (2004) or Decety and Grèzes (2006)).”
Pezzulo, G., Barsalou, L. W., Cangelosi, A., Fischer, M. H., McRae, K., & Spivey, M. J. (2013) · Frontiers in Psychology · doi:10.3389/fpsyg.2012.00612 · Cited by 229
“…central aim is realizing complete robotic architectures that, on the one hand, include principles and constraints derived from animal and human cognition and, on the other hand, have realistic embodiment, sensors and effectors, and learn to act autonomously in complex, open-ended (and social).”
Cerebral correlates of motor imagery of normal and precision gait
Bakker, M., de Lange, F. P., Helmich, R. C. G., Scheeringa, R., Bloem, B. R., & Toni, I. (2008) · NeuroImage · doi:10.1016/j.neuroimage.2008.03.020 · Cited by 205
“This issue is an instance of the general distinction that has been drawn between processes leading to the generation of a motor plan (that include predictions of the sensory consequences of the action), and processes related to the evaluation of sensory feedback (Blakemore and Sirigu, 2003; Grush, 2004; Wolpert et al.…”
Perceptual Processing Affects Conceptual Processing
van Dantzig, S., Pecher, D., Zeelenberg, R., & Barsalou, L. W. (2008) · Cognitive Science · doi:10.1080/03640210802035365 · Cited by 206
“Embodied views of cognition suggest that these internal representations are grounded in the systems of perception and action that are used to interact with the world (e.g., Barsalou, 1999; Glenberg, 1997; Grush, 2004; Pecher & Zwaan, 2005).”
How something can be said about telling more than we can know: On choice blindness and introspection
Johansson, P., Hall, L., Sikström, S., Tärning, B., & Lind, A. (2006) · Consciousness and Cognition · doi:10.1016/j.concog.2006.09.004 · Cited by 224
The Dorsal Frontoparietal Network: A Core System for Emulated Action
Ptak, R., Schnider, A., & Fellrath, J. (2017) · Trends in Cognitive Sciences · doi:10.1016/j.tics.2017.05.002 · Cited by 218
“We use the term ‘emulation’ to denote this core function of the dFPN [11,12].”
Searching for Roots of Entrainment and Joint Action in Early Musical Interactions
Phillips-Silver, J., & Keller, P. E. (2012) · Frontiers in Human Neuroscience · doi:10.3389/fnhum.2012.00026 · Cited by 231
“…motor/perceptual resonance and action simulation: While resonance is driven exogenously and automatically (i.e., preattentively) by the perception of external events, simulation may be generated endogenously in the absence of external stimuli (cf. Grush, 2004) and may be modulated by attention.”
Scrutinizing visual images: The role of gaze in mental imagery and memory
Laeng, B., Bloem, I. M., D’Ascenzo, S., & Tommasi, L. (2014) · Cognition · doi:10.1016/j.cognition.2014.01.003 · Cited by 217
“’’ Within this account, imagery could be an anticipatory phase of perception like a ‘‘disposition to see’’ (see also Freyd, 1987; Grush, 2004; Kosslyn & Sussman, 1994; Ryle, 1949) that takes place all the time; only when the perceptual pickup of information is either interrupted or delayed, imagery becomes subjectivel…”
Vancouver, J. B., Weinhardt, J. M., & Schmidt, A. M. (2010) · Journal of Applied Psychology · doi:10.1037/a0020628 · Cited by 212
A cognitive architecture that combines internal simulation with a global workspace
Shanahan, M. (2005) · Consciousness and Cognition · doi:10.1016/j.concog.2005.11.005 · Cited by 206
Neural Dissociations between Action Verb Understanding and Motor Imagery
Willems, R. M., Toni, I., Peter Hagoort, A., & Casasanto, D. J. (2009) · Journal of Cognitive Neuroscience · doi:10.1162/jocn.2009.21386 · Cited by 205
“Like overt motor execution, motor imagery may entail the generation of an action plan (inverse model) as well as a prediction of the actionʼ sensory consequences (forward model) (e.g., Grush, 2004; Wolpert & Ghahramani, 2000).”
Motor imagery: A window into the mechanisms and alterations of the motor system
de Lange, F. P., Karin Roelofs, A., & Toni, I. (2007) · Cortex · doi:10.1016/j.cortex.2007.09.002 · Cited by 192
“…(de Lange et al., 2006; Vargas et al., 2004) studies have showed that there is a clear proprioceptive influence on motor imagery performance in healthy subjects, favouring the view that motor imagery relies on the generation of a complete motor plan that is prevented from operating on the body.”
Few believe the world is flat: How embodiment is changing the scientific understanding of cognition.
Glenberg, A. M. (2015) · Canadian Journal of Experimental Psychology/Revue canadienne de psychologie expérimentale · doi:10.1037/cep0000056 · Cited by 202
Toward a Neurobiologically Plausible Model of Language-Related, Negative Event-Related Potentials
Bornkessel‐Schlesewsky, I., & Schlesewsky, M. (2019) · Frontiers in Psychology · doi:10.3389/fpsyg.2019.00298 · Cited by 196
“This can be operationalized via a Kalman filter (Grush, 2004), which informally speaking determines to what extent the existing model is “trusted” over the current prediction error or vice versa.”
Interactions Between Posterior Gamma and Frontal Alpha/Beta Oscillations During Imagined Actions
de Lange, F. D., Jensen, O., Bauer, M., & Toni, I. (2008) · Frontiers in Human Neuroscience · doi:10.3389/neuro.09.007.2008 · Cited by 456
“To address this issue, it is important to disentangle planningrelated activity from the effects of motor execution and sensory feedback (Blakemore and Sirigu, 2003; Davidson and Wolpert, 2003; Grush, 2004).”
Stepp, N., & Turvey, M. T. (2010) · Cognitive Systems Research · doi:10.1016/j.cogsys.2009.03.003 · Cited by 202
Macuga, K. L., & Frey, S. H. (2011) · NeuroImage · doi:10.1016/j.neuroimage.2011.09.083 · Cited by 175
“It has been suggested that during imagery, the brain internally simulates the predicted sensory consequences of a movement via a forward model (Grush, 2004), and the cerebellum seems well suited to support this function (Miall and King, 2008; Wolpert et al.”
InfluentialPiccinini, G. (2020) · doi:10.1093/oso/9780198866282.001.0001 · Cited by 181
Tian, X., & Poeppel, D. (2013) · Journal of Cognitive Neuroscience · doi:10.1162/jocn_a_00381 · Cited by 172
“Importantly, imagery tasks are typically argued to be mediated by internal simulation and prediction processes (Tian & Poeppel, 2010, 2012; Grush, 2004; Miall & Wolpert, 1996; Sirigu et al., 1996; Jeannerod, 1994, 1995).”
InfluentialKinematics fingerprints of leader and follower role-taking during cooperative joint actions
Sacheli, L. M., Tidoni, E., Pavone, E. F., Aglioti, S. M., & Candidi, M. (2013) · Experimental Brain Research · doi:10.1007/s00221-013-3459-7 · Cited by 178
““Simulative” processes (e.g., “emulation,” Grush 2004) are likely to be recruited to achieve better temporal and spatial predictions of a partner’s actions.”
Ridderinkhof, K. R., & Braß, M. (2015) · Journal of Physiology-Paris · doi:10.1016/j.jphysparis.2015.02.003 · Cited by 191
“External signals are assessed by exteroceptive inference of the causes of these signals; that is, by comparing desired to anticipated and actual sensory states (Grush, 2004).”
InfluentialNREM and REM Sleep: Complementary Roles in Recovery after Wakefulness
Vyazovskiy, V. V., & Delogu, A. (2014) · The Neuroscientist · doi:10.1177/1073858413518152 · Cited by 169
“This notion is largely consistent with the tenets of the neural emulation theory, according to which inner models of the body and the environment can be run offline to produce “imagery, estimate outcomes of different actions, and evaluate and develop motor plans” (Churchland 2002; Grush 2004).”
Tian, X., & Poeppel, D. (2012) · Frontiers in Human Neuroscience · doi:10.3389/fnhum.2012.00314 · Cited by 161
“Mental imagery has been linked to the concept of internal forward models by the argument that the subjective feeling in mental imagery is the result of the internal estimation of the perceptual consequences following the internal simulation of an action (Grush, 2004).”
Moving Ourselves, Moving Others
Zlatev, J., Racine, T. P., Lüdtke, U. M., & Foolen, A. (2012) · Consciousness & emotion book series · doi:10.1075/ceb.6 · Cited by 161
Recruitment of the motor system during music listening: An ALE meta-analysis of fMRI data
Gordon, C. L., Cobb, P. R., & Balasubramaniam, R. (2018) · PLoS ONE · doi:10.1371/journal.pone.0207213 · Cited by 183
Measuring motor imagery ability: A review
McAvinue, L. P., & Robertson, I. H. (2008) · The European Journal of Cognitive Psychology · doi:10.1080/09541440701394624 · Cited by 186
“Motor imagery has been at the centre of many theoretical discussions and has been conceptualised in terms of motor representations (Jeannerod, 1994, 1995, 1997), action prototypes coupled with memory processes (Annett, 1996) and forward models (Grush, 2004; Schwoebel, Boronat, & Coslett, 2002; Wilson, 2003; Wolpert &…”
Stich, S. G. W. (2008) · doi:10.1093/acprof:oso/9780195332834.001.0001 · Cited by 160
MENTAL IMAGERY AND THE VARIETIES OF AMODAL PERCEPTION
BRISCOE, R. E. (2011) · Pacific philosophical quarterly · doi:10.1111/j.1468-0114.2011.01393.x · Cited by 157
Huang, J. Y., & Bargh, J. A. (2014) · Behavioral and Brain Sciences · doi:10.1017/s0140525x13000290 · Cited by 173
Functional coupling underlying motor and cognitive functions of the dorsal premotor cortex.
Abe, M., & Hanakawa, T. (2009) · Behavioural Brain Research · doi:10.1016/j.bbr.2008.10.046 · Cited by 155
A touch with words: Dynamic synergies between manual actions and language
García, A. M., & Ibáñez, A. M. (2016) · Neuroscience & Biobehavioral Reviews · doi:10.1016/j.neubiorev.2016.04.022 · Cited by 157
Rauschecker, J. P. (2017) · Cortex · doi:10.1016/j.cortex.2017.10.020 · Cited by 159
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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