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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
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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Rambusch, J. (2006) · eScholarship (California Digital Library) · Cited by 23
A Schema Based Model of the Praying Mantis
Pezzulo, G., & Calvi, G. (2006) · Lecture notes in computer science · doi:10.1007/11840541_18 · Cited by 23
“The degree of (mis)match between actual and expected stimulus has two functions: (1) Adjustment of Control : the predicted signal can compensate time delays, filter or replace missing or unreliable stimuli; see [7] for a comparison with Kalman filtering; (2) Schema Selection: schemas which predict well gain activation.”
Transformation, encoding and representation
Webb, B. H. (2006) · Current Biology · doi:10.1016/j.cub.2006.02.034 · Cited by 23
Anticipations, Brains, Individual and Social Behavior: An Introduction to Anticipatory Systems
Butz, M. V., Sigaud, O., Pezzulo, G., & Baldassarre, G. (2008) · Lecture notes in computer science · doi:10.1007/978-3-540-74262-3_1 · Cited by 24
“Interactions with the experienced environment are often re-played or projected into the future by means of an internal predictive environmental model [18, 32, 40].”
Menegaldo, L. L. (2017) · Biological Cybernetics · doi:10.1007/s00422-017-0724-z · Cited by 22
“However, state estimation has frequently been regarded as an appropriate theoretical framework for describing several motor and cognitive neuroscience problems (Shadmehr andKrakauer 2008; Grush 2004; Schwartz 2016).”
Fine motor skills and mental imagery: Is it all in the mind?
Martzog, P., & Suggate, S. P. (2019) · Journal of Experimental Child Psychology · doi:10.1016/j.jecp.2019.05.002 · Cited by 29
Pezzulo, G., D’Amato, L., Mannella, F., Priorelli, M., de Maele, T. V., Stoianov, I. P., & Friston, K. J. (2024) · Annals of the New York Academy of Sciences · doi:10.1111/nyas.15118 · Cited by 25
“Various researchers 5,71,134–138 have proposed a plausible transition from pragmatic to more detached and descriptive representations, which can be condensed into two pivotal steps.”
Hughes, R. W., Marsh, J. E., & Jones, D. M. (2011) · Memory & Cognition · doi:10.3758/s13421-011-0116-x · Cited by 22
“From this standpoint, serial rehearsal in STM tasks is akin to a covert Role of Serial Order 21 simulation (or forward model; cf. Grush, 2004) of those bodily movements that are best placed to mimic the presented material.”
Action and Language Integration: From Humans to Cognitive Robots
Borghi, A. M., & Cangelosi, A. (2014) · Topics in Cognitive Science · doi:10.1111/tops.12103 · Cited by 22
van Dijk, L. (2016) · Philosophical Psychology · doi:10.1080/09515089.2016.1213379 · Cited by 22
“…engage in the unfolding acts of walking or talking as evidence that there must be a mechanism prior to these “overt” processes that is their source, such as a “forward model,” “internal model,” or an “emulator” (e.g., Grush, 2004; for criticism see e.g., Chemero, 2009; Dewey, 1896; Ingold, 2000).”
Glazebrook, C. M., Kiernan, D., Welsh, T. N., & Tremblay, L. (2014) · Human Movement Science · doi:10.1016/j.humov.2014.11.005 · Cited by 26
Electrophysiological analysis of a sensorimotor integration task
Velasques, B., Machado, S., Portella, C. E., Silva, J. G., Basile, L. F. H., Cagy, M., Piedade, R., & Ribeiro, P. (2007) · Neuroscience Letters · doi:10.1016/j.neulet.2007.08.061 · Cited by 22
Natural Selection in the Brain
Fernando, C., & Szathmáry, E. (2009) · On thinking · doi:10.1007/978-3-642-03129-8_20 · Cited by 22
“Such forward models feature in influential theories of cognition (Churchland 2002) in the form of emulators (Craik 1943; Grush 2004 ).”
Modeling speech imitation and ecological learning of auditory-motor maps
Canevari, C., Badino, L., D’Ausilio, A., Fadiga, L., & Giorgio Metta, B. (2013) · Frontiers in Psychology · doi:10.3389/fpsyg.2013.00364 · Cited by 23
“Furthermore, the mirror neuron theory suggests that visual (or audio) representations of actions that are part of our motor repertoire can exploit an additional inferential process based on the emulation of analogous motor commands in our brain (Grush, 2004).”
Multi-timescale Nexting in a Reinforcement Learning Robot
Modayil, J., White, A., & Sutton, R. S. (2012) · Lecture notes in computer science · doi:10.1007/978-3-642-33093-3_30 · Cited by 21
A continuum of intentionality: linking the biogenic and anthropogenic approaches to cognition
Sims, M. (2021) · Biology & Philosophy · doi:10.1007/s10539-021-09827-w · Cited by 28
“The idea that cognition is fundamentally anticipatory also follows a long tradition in psychology and cognitive science (Bartlett 1932; Craik 1943; Piaget 1970; Neisser 1976; Drescher 1991; Arbib 1992; Riegler 2001; Grush 2004; Castelfranchi 2005; Bar 2007).”
Dahm, S. F., Weigelt, M., & Rieger, M. (2022) · Psychological Research · doi:10.1007/s00426-022-01645-3 · Cited by 21
“Similarly, it is possible, that during AIP a full-fledged simulation of the action, including motor components, occurs (cf. Davidson & Wolpert, 2005; Grush, 2004; Ingram et al., 2019).”
Farina, M., & Lavazza, A. (2022) · Synthese · doi:10.1007/s11229-022-03963-w · Cited by 22
“To undermine the idea that the best cognitive model capable of explaining cases of smooth coping is the one that draws no firm boundaries between the bodily self and the external resources, Rupert may appeal to the emulator theory (Grush, 2004).”
Concepts as plug & play devices
Shea, N. J. (2022) · Philosophical Transactions of the Royal Society B Biological Sciences · doi:10.1098/rstb.2021.0353 · Cited by 23
“…it is even more useful to be able to sever representations from their worldly correlations, run internal computations that make use of their interconnections, and then restore their input – output profile so that the conclusions that have been worked out offline can be used to guide behaviour.”
Muraki, E. J., Dahm, S. F., & Pexman, P. M. (2023) · Cognition · doi:10.1016/j.cognition.2023.105589 · Cited by 21
The role of accent imitation in sensorimotor integration during processing of intelligible speech
Adank, P., Rueschemeyer, S., & Harold Bekkering, A. (2013) · Frontiers in Human Neuroscience · doi:10.3389/fnhum.2013.00634 · Cited by 21
Computation and Representation in Cognitive Neuroscience
Piccinini, G. (2018) · Minds and Machines · doi:10.1007/s11023-018-9461-x · Cited by 21
How passive is passive listening? Toward a sensorimotor theory of auditory perception
Froese, T., & González-Grandón, X. A. (2019) · Phenomenology and the Cognitive Sciences · doi:10.1007/s11097-019-09641-6 · Cited by 25
“One way of interpreting the possible auditory function of areas normally associated with speech production is in terms of internal forward model structures, or “emulators” as they are known in motor control theory (Grush 2004).”
Tutorial Review of Bio-Inspired Approaches to Robotic Manipulation for Space Debris Salvage
Ellery, A. A. (2020) · Biomimetics · doi:10.3390/biomimetics5020019 · Cited by 22
Sims, M., & Pezzulo, G. (2021) · Synthese · doi:10.1007/s11229-021-03140-5 · Cited by 25
“…models is that they afford forms of internal manipulation and inference—for example, when they are used to steer sensorimotor predictions of action consequences—and for this they are sometimes called “action-oriented” or “embodied representations” (Clark & Grush, 1999; Grush, 2004; Pezzulo, 2008).”
Vecchi, T., & Gatti, D. (2020) · The MIT Press eBooks · doi:10.7551/mitpress/13543.001.0001 · Cited by 31
Corradi‐Dell’Acqua, C., & Tessari, A. (2009) · Neuropsychologia · doi:10.1016/j.neuropsychologia.2009.11.029 · Cited by 22
“own body (holding information about the spatial relation between one’s parts, biomechanical constraints, the orientation of body segments in the external space, and even its kinematics—Wilson & Knoblich, 2005)might be used by the visual system asmodel information for either one’s own body, that of a third person, or e…”
Does the World Leak Into the Mind? Active Externalism, “Internalism” and Epistemology
Dartnall, T. (2005) · Cognitive Science · doi:10.1207/s15516709cog2901_6 · Cited by 21
Tomasi, D. L. (2020) · Springer eBooks · doi:10.1007/978-3-030-35354-4_6 · Cited by 21
The role of imitation in learning to pronounce
Messum, P. (2008) · OpenGrey (Institut de l'Information Scientifique et Technique) · Cited by 21
Errors in Imagined and Executed Typing
Dahm, S., & Rieger, M. (2019) · Vision · doi:10.3390/vision3040066 · Cited by 19
“In MI, assuming that motor commands are generated, forward models may predict the action effects in a similar way [22,23] (Figure 1).”
Mahmoud, S., Billing, E., Svensson, H., & Thill, S. (2022) · Cognitive Systems Research · doi:10.1016/j.cogsys.2022.09.005 · Cited by 20
Representationalism and the phenomenology of mental imagery
Thompson, E. T. (2007) · Synthese · doi:10.1007/s11229-006-9086-0 · Cited by 20
“See Grush (2004).”
Beudel, M., Zijlstra, S., Mulder, T., Zijdewind, I., & de Jong, B. M. (2010) · Human Brain Mapping · doi:10.1002/hbm.21044 · Cited by 20
“By doing this, incoming proprioceptive information can be filtered from information induced by motor commands [Grush, 2004].”
Chemical, Neuronal, and Linguistic Replicators
Fernando, C., & Szathmáry, E. (2010) · The MIT Press eBooks · doi:10.7551/mitpress/9780262513678.003.0009 · Cited by 20
Koordination sportlicher Bewegungen – Sportmotorik
Hossner, E.-J., Müller, H., & Voelcker‐Rehage, C. (2013) · Springer eBooks · doi:10.1007/978-3-642-37546-0_8 · Cited by 23
Typical biomechanical bias in the perception of congenitally absent hands
Vannuscorps, G., & Caramazza, A. (2015) · Cortex · doi:10.1016/j.cortex.2015.02.015 · Cited by 19
“For numerous authors, this demonstrates that the perception of others' bodies and movements is supported by somatosensory and motor representations of our own body (Grush, 2004; Hommel, Müsseler, Aschersleben, & Prinz, 2001; Wilson & Knoblich, 2005).”
A New Imagery Debate: Enactive and Sensorimotor Accounts
Foglia, L., & O'Regan, J. K. (2015) · Review of Philosophy and Psychology · doi:10.1007/s13164-015-0269-9 · Cited by 20
“A final approach involving a form of action is the emulation theory (Grush 1998, 2004).”
Learning a world model and planning with a self-organizing, dynamic neural systemPreprint
Toussaint, M. (2003) · arXiv (Cornell University) · doi:10.48550/arxiv.nlin/0306015 · Cited by 22
“There is clear evidence that nervous systems use such internal models to perform predict ive motor control, imagery, inference, and planning in a way that involves a simulation o f actions and their perceptual implications [1, 2].”
Predictive coding I: Introduction
Sprevak, M. D. (2023) · Philosophy Compass · doi:10.1111/phc3.12950 · Cited by 22
Ptak, R., Doganci, N., & Bourgeois, A. (2021) · Brain Sciences · doi:10.3390/brainsci11121652 · Cited by 24
“Our theory is based on Grush’s emulation theory of mental representation [124].”
Wang, L., Li, X., Zheng, W., Chen, X., Chen, Q., Hu, Y., Cao, L., Ren, J., Qin, W., Lu, J., et al. (2022) · Brain Imaging and Behavior · doi:10.1007/s11682-022-00675-0 · Cited by 19
“The possible mechanisms for MI include psycho-neuromuscular theory (Page et al., 2001) and motor simulation theory (namely the central theory)(Grush, 2004).”
Embodied Social Cognition and Embedded Theory of Mind
Fenici, M. (2012) · Biolinguistics · doi:10.5964/bioling.8921 · Cited by 19
Inference Through Embodied Simulation in Cognitive Robots
Mohan, V., Morasso, P., Sandini, G., & Kasderidis, S. (2013) · Cognitive Computation · doi:10.1007/s12559-013-9205-4 · Cited by 19
“While this perspective has been emphasized in the reviews of Hesslow [32] [33], Grush, [29] among others, more recent developments on the organization of semantic knowledge in the brain (See [16] [48] [49] [57]) provide further insights that help to constrain computational architectures for cognitive agents.”
Zhang, Q., Zhang, P., Lu, S., Yang, Y. J., Yuan, S., Chen, Y., Sun, S., & Bai, X. (2019) · Frontiers in Human Neuroscience · doi:10.3389/fnhum.2019.00080 · Cited by 19
“Grush (2004) proposed that motor imagery drives an emulator of body when the emulator receives an efference copy, it releases an output signal same as an actual sensory feedback signal.”
Visual phenomenology versus visuomotor imagery: How can we be aware of action properties?
Ferretti, G. (2019) · Synthese · doi:10.1007/s11229-019-02282-x · Cited by 19
“Accordingly, philosophers and neuroscientists agree that we can have visual experience of geometrical and shape properties (Siegel 2016; Nanay 2011, 2012a, b, 2013), as well as that we can form and superimpose visuomotor imagery on them (Briscoe 2011, 2018; Grush 2004).”
de Vignemont, F., Pitron, V., & Alsmith, A. (2021) · doi:10.1093/oso/9780198851721.003.0001 · Cited by 19
Gibbs, R. W. (2005) · Cambridge University Press eBooks · doi:10.1017/cbo9780511805844.007 · Cited by 19
The embodied and situated nature of computer game play.
Rambusch, J. (2006) · Cited by 18
Gendron, M., Mesquita, B., & Barrett, L. F. (2020) · Culture, Mind, and Brain · doi:10.1017/9781108695374.010 · Cited by 20
Inferential Role Semantics for Natural Language
Blouw, P. (2017) · eScholarship (California Digital Library) · Cited by 18
Artificial Intelligence in Science and Society: The Vision of USERN
Dorigo, T., Brown, G. D., Casonato, C., Cerdà, A., Ciarrochi, J., Lio, M. D., D’Souza, N., Gauger, N. R., Hayes, S. C., Hofmann, S. G., et al. (2025) · IEEE Access · doi:10.1109/access.2025.3529357 · Cited by 21
4 Innateness and Genetic Information
Godfrey‐Smith, P. (2008) · Oxford University Press eBooks · doi:10.1093/acprof:oso/9780195332834.003.0004 · Cited by 19
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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