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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
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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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“These results are in accordance with theories that emphasize the importance of sensorimotor contingency between the subject and the sources of sensory stimulation [16, 24, 25].”
Svensson, H., Thill, S., & Ziemke, T. (2013) · Adaptive Behavior · doi:10.1177/1059712313491295 · Cited by 18
“, perception (Möller, 1999), mental imagery (Jeannerod, 1994; Moulton & Kosslyn, 2009; Thomas, 1999), mental representation (Grush, 2004), concepts (Barsalou, 1999) or language (Zwaan, 2004).”
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“Another hypothesis on how motor areas might participate in mental rotation comes from the theories (Grush, 2004), neuroscientific evidence (Miall, 2003), and computational architectures (Wolpert & Kawato, 1998) on motor control based on forward models.”
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“In this way, motor imagery could be defined as an internal dynamical state related to participants’ ability to use internal models to predict the sensory consequences of an action (Gabbard, Caçola, & Bobbio, 2011; Grush, 2004; Papaxanthis, Pozzo et al., 2002).”
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“…useful in VPT as they provide information about what other people may see (for example, head orientation lets you know which direction a person is looking), and motor representation of the body is also thought of as the first step in performing an egocentric spatial transformation (Grush, 2004).”
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“It has been reported that motor imagery is used to predict the proprioceptive consequences of movement and then contributes to movement planning (Grush 2004).”
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“During planning, we very often picture the course of actions as an imaginary episode, from taking the plane to visiting the first museum, in a process that has been called imagination-based planning, model-based planning, mental simulations or emulation, each term carrying somehow different meanings [35, 36, 37, 38, 3…”
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“Covert phonological processes (inner speech) might work as emulators of the to-be-recognized perceptual situation [7–9]; see also [10].”
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“This distinction may be useful in explaining how offline forms of motor planning— resembling earlier theories of motor simulation (Grush 2004)—evolved progressively as elaborations on earlier sensorimotor control loops (Pezzulo 2012).”
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Young, A. (2011) · Wiley eBooks · doi:10.1002/9781444343359.ch7 · Cited by 16
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“…Bubic, von Cramon, and Schubotz 2010; Llinás 2001); an understanding of mental representations as structural models (Cummins 1989; Gładziejewski 2015; Grush 2004; Ryder 2004; Shea 2014; Williams and Colling 2017); and an increased focus on regulation and predictive control as principles of brain…”
Saimpont, A., Malouin, F., Durand, A., Mercier, C., Rienzo, F. D., Saruco, E., Collet, C., Guillot, A., & Jackson, P. L. (2021) · Scientific Reports · doi:10.1038/s41598-021-93240-6 · Cited by 15
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“Among other things, these processes would support the simulation (Jeannerod, 2001; Hesslow, 2002) and emulation (Grush, 2004) of bodily states, as tacitly assumed by recent models characterizing the prediction and understanding of external events (Schubotz, 2007).”
Attentional Reinforcement Learning in the Brain
Yamakawa, H. (2020) · New Generation Computing · doi:10.1007/s00354-019-00081-z · Cited by 18
Effect of Physical Fatigue Elicited by Continuous and Intermittent Exercise on Motor Imagery Ability
Kanthack, T. F. D., Guillot, A., Clémençon, M., Debarnot, U., & Rienzo, F. D. (2020) · Research Quarterly for Exercise and Sport · doi:10.1080/02701367.2019.1691709 · Cited by 15
Neurological and behavioral features of locomotor imagery in the blind
Amemiya, K., Morita, T., Hirose, S., Ikegami, T., Hirashima, M., & Naito, E. (2020) · Brain Imaging and Behavior · doi:10.1007/s11682-020-00275-w · Cited by 15
“These are a substitute for the sensory feedback that would normally arise in association with an overt action, probably by accessing memories of the experienced action (Annett 1995, 1996; Grush 2004; Guillot et al. 2009; Hanakawa 2016; Naito et al. 2002).”
Culture, Mind, and Brain: Emerging Concepts, Models, and Applications
Kirmayer, L. J., Worthman, C. M., Kitayama, S., Lemelson, R., & Cummings, C. A. (2020) · Cited by 15
Multiple Levels of Heuristic Reasoning Processes in Scientific Model Construction
Clement, J. J. (2022) · Frontiers in Psychology · doi:10.3389/fpsyg.2022.750713 · Cited by 18
“Future approaches will likely need to include the role of elemental action components such as anticipatory predictions from schemas in motor control theory (e.g., Schmidt, 1982; Grush, 2004; Barsalou, 2009; Kühn et al., 2011).”
Attentional loads associated with interlimb interactions underlying rhythmic bimanual coordination
Ridderikhoff, A., Peper, C. E., & Beek, P. J. (2008) · Cognition · doi:10.1016/j.cognition.2008.10.002 · Cited by 15
“In contrast, motor control constitutes a form of cognition in the sense that every coordinated action requires the planning of well-orchestrated activity patterns and associated correction processes verifying the outcome ( Grush, 2004; Wolpert & Ghahramani, 2000).”
The Instrumental Structure of Actions
Valaris, M. (2014) · The Philosophical Quarterly · doi:10.1093/pq/pqu054 · Cited by 15
Rahaman, J., Agrawal, H., Srivastava, N., & Chandrasekharan, S. (2018) · Cognitive Sciences · doi:10.1111/cogs.12518 · Cited by 16
Zhang, T., Li, M., Zhang, L., Biswal, B. B., Yao, D., & Xu, P. (2018) · IEEE Access · doi:10.1109/access.2018.2875492 · Cited by 15
“The results are in line with those of many other previous studies [12], [21], [58], [59] and”
The effects of visual impairment on motor imagery in children and adolescents
Schott, N., Beach, P., Knöpfle, I., & Neuberger, V. (2021) · Research in Developmental Disabilities · doi:10.1016/j.ridd.2020.103835 · Cited by 15
Gładziejewski, P. (2015) · New Ideas in Psychology · doi:10.1016/j.newideapsych.2015.01.005 · Cited by 15
“…composed of elements whose dynamics of interactions are meant to model the dynamics of interactions between parts of the musculoskeletal system (see Grush 2004)then they are successful in performing their action-guiding function to the extent to which the way the emulator works structurally or…”
6 Poverty of Stimulus Arguments Concerning Language and Folk Psychology
Segal, G. (2008) · Oxford University Press eBooks · doi:10.1093/oso/9780195332827.003.0006 · Cited by 16
Moore, J. W., & Haggard, P. (2006) · Consciousness and Cognition · doi:10.1016/j.concog.2006.09.003 · Cited by 15
“CBP. In contrast to our exploratory work on choice blindness, the research on agency they present is carried by a strong theoretical framework developed within the field of computational motor control (e.g., Wolpert & Ghahramani, 2004, and taken to its limit as a general model of cognition by Grush, 2004 ).”
The implicit effect of action mental simulation on action evaluation
Ianì, F., Limata, T., Bucciarelli, M., & Mazzoni, G. A. L. (2022) · Quarterly Journal of Experimental Psychology · doi:10.1177/17470218221091096 · Cited by 14
“These mental representations of a sequence of events as they unfold in time are akin to the idea of perceptual emulators (Grush, 2004; see also Wilson & Knoblich, 2005).”
The Hard Problem of Content is Neither
Ramsey, W. (2023) · Review of Philosophy and Psychology · doi:10.1007/s13164-023-00714-9 · Cited by 14
Invariant representations in abstract concept grounding – the physical world in grounded cognition
Friedrich, J., Fischer, M. H., & Raab, M. (2024) · Psychonomic Bulletin & Review · doi:10.3758/s13423-024-02522-3 · Cited by 17
“Importantly, models can be detached, i.e., allow for covert simulation, allowing representation of objects in their absence via internal loops (Grush, 2004; Pezzulo, 2011; Wolpert et al., 2003).”
Gregoromichelaki, E., Kempson, R., Howes, C., & Eshghi, A. (2013) · Advances in interaction studies · doi:10.1075/ais.6.04gre · Cited by 14
The attachment control system and computational modeling: Origins and prospects.
Petters, D. (2018) · Developmental Psychology · doi:10.1037/dev0000647 · Cited by 17
“The final article will be available, upon publication, via its DOI: 10.1037/dev0000647 the system” (Bowlby,1969, p .43) Grush (2004) provides a broader review of forward models, describing their wide use in diverse theories of human cognition.”
InfluentialNakano, H., Murata, S., Shiraiwa, K., & Nonaka, K. (2020) · Brain Sciences · doi:10.3390/brainsci10060332 · Cited by 15
“These are the defining features of simulation theory—specifically, that motor imagery is the psychological counterpart of the offline operation of the efferent motor areas [2].”
Guilbert, J., Fernandez, J., Molina, M., Morin, M., & Alamargot, D. (2020) · Psychological Research · doi:10.1007/s00426-020-01399-w · Cited by 14
“Consequently, MI could be viewed as an internal dynamic state, during which individuals use internal models to consciously predict the sensory consequences of an action (Gabbard, Caçola, & Bobbio, 2011; Grush, 2004; Papaxanthis, Pozzo, Skoura, & Schieppati, 2002).”
Colling, L. J., & Williamson, K. (2014) · Frontiers in Human Neuroscience · doi:10.3389/fnhum.2014.00754 · Cited by 14
“Running the forward model offline—that is, without producing motor output—can also be used to internally simulate limb movements; that is, to engage in motor imagery (Grush, 2004).”
Mental imagery of gravitational motion
Gravano, S., Zago, M., & Lacquaniti, F. (2017) · Cortex · doi:10.1016/j.cortex.2017.08.005 · Cited by 13
Hierarchical minds and the perception/cognition distinction
Williams, D. (2019) · Inquiry · doi:10.1080/0020174x.2019.1610045 · Cited by 26
“…aspects of intelligence: unsupervised learning (Kiefer and Hohwy 2018), reinforcement learning (Garrison, Erdeniz, and Done 2013), perception (Clark 2013), action and motor control (Grush 2004), planning (Bubic, von Cramon, and Schubotz 2010), and even memory (Schacter, Addis, and Buckner 2007).”
Putting the Pieces Together Again
Sloman, A. (2001) · Cambridge University Press eBooks · doi:10.1017/cbo9780511816772.031 · Cited by 14
“So perhaps the primary form of representation for visual information should be representations of processes extended in time and space, whereas many researchers (an exception being Grush, 2004), assume that vision produces information about objects, properties and relationships.”
Fekete, T. (2009) · Minds and Machines · doi:10.1007/s11023-009-9166-2 · Cited by 14
Extending cognitive architectures with spatial and visual imagery mechanisms
Laird, J. E., & Lathrop, S. D. (2008) · Deep Blue (University of Michigan) · Cited by 14
The Brain Evolved to Guide Action
Anderson, M. L., & Chemero, A. P. (2016) · Wiley eBooks · doi:10.1002/9781118316757.ch1 · Cited by 14
Distinct neural mechanisms support inner speaking and inner hearingPreprint
Nalborczyk, L., Longcamp, M., Bonnard, M., Serveau, V., Spieser, L., & Alario, F. ‐. (2021) · PsyArXiv (OSF Preprints) · doi:10.1016/j.cortex.2023.09.007 · Cited by 18
“…are compatible with the models developed in Lœvenbruck et al. (2018) and Grandchamp et al. (2019), as well as other models of motor imagery, such as Grush (2004), in which the role of the primary motor cortex during inner speech is to issue motor commands from which predictions of sensory…”
Pouw, W. T. J. L., & Hostetter, A. B. (2016) · MPG.PuRe (Max Planck Society) · doi:10.12832/83918 · Cited by 14
Neural representations unobserved—or: a dilemma for the cognitive neuroscience revolution
Facchin, M. (2023) · Synthese · doi:10.1007/s11229-023-04418-6 · Cited by 14
“The first—and more widespread—variant is ultimately based on the analysis of the behavior of a large class of neurocomputational models (cf. Churchland, 1995; O’Brien and Opie 2004; Grush, 2004; Shagrir, 2012; Williams, 2017; Wiese, 2016, 2017).”
Auditory verbal hallucinations: Social, but how?
Alderson‐Day, B., & Fernyhough, C. (2016) · PubMed · Cited by 14
Perini, I., Čeko, M., Cerliani, L., van Ettinger-Veenstra, H., Minde, J., & Morrison, I. (2020) · Cerebral Cortex · doi:10.1093/cercor/bhaa078 · Cited by 13
“Further, the general perspective that action goals influence sensory perception has become widely accepted in recent decades, for example, in the interplay between vision and action (e.g., Prinz, 1997; Nöe, 2004; Grush, 2004).”
O’Shea, H. (2022) · Frontiers in Human Neuroscience · doi:10.3389/fnhum.2022.984053 · Cited by 13
“…proposed to drive perceptual-cognitive-motor processes, including association (Barsalou, 1999, 2008; Hesslow, 2012; Pulvermüller, 2018), emulation (Grush, 2004), simulation (Barsalou, 1999, 2008; Jeannerod, 2001, 2006), perception-action coding (Hommel et al., 2001; Hommel, 2019), or predictive…”
InfluentialNeural Pathways of Embodied Simulation
Svensson, H., Morse, A. F., & Tom Ziemke, B. (2009) · Lecture notes in computer science · doi:10.1007/978-3-642-02565-5_6 · Cited by 13
Neuroscientific implications for situated and embodied artificial intelligence
Downing, K. L. (2007) · Connection Science · doi:10.1080/09540090701192584 · Cited by 13
Representations in Dynamical Embodied Agents: Re‐Analyzing a Minimally Cognitive Model Agent
Mirolli, M. (2012) · Cognitive Science · doi:10.1111/j.1551-6709.2012.01233.x · Cited by 13
“Hence, our minimalist simulations seem to demonstrate that it is possible to have an agent possessing ''full-blooded'' representations (in the strong sense of Clark and Grush) without having coupled inverse and forward models , as the emulator theory of representations suggests (Grush, 2004).”
Schema-Based Design and the AKIRA Schema Language: An Overview
Pezzulo, G., & Calvi, G. (2007) · SAB ABiALS · doi:10.1007/978-3-540-74262-3_8 · Cited by 14
“According to Grush [26], simulations can nest to produce increasingly abstract levels of description, in which the criteria for 'matching' are increasingly distant from perceptual matching, although they remain grounded on (actual or simulated) sensorimotor interaction.”
InfluentialFuruta, T., Morita, T., Miura, G., & Naito, E. (2024) · Scientific Reports · doi:10.1038/s41598-024-68425-4 · Cited by 12
“The brain emulates various sensory experiences (feedback) that would be expected (predicted) if the imagined movements were actually performed by referring memories of previous enactments 2,5,36,49,64–66 .”
Bodily structure and body representation
Alsmith, A. (2019) · Synthese · doi:10.1007/s11229-019-02200-1 · Cited by 13
“This in turn might be the basis for cognitive processes such as planning and imagination, in so far as these can also concern distant objects, properties and events that are absent in a spatial sense (Clark and Grush 1999; Grush 2004).”
Multimodal interaction with Loki
Bustos, P., Martínez-Gómez, J., García-Varea, I., Bachiller, P., Calderita, L. V., Manso, L. J., Sánchez, A., & Balderas, A. (2013) · Cited by 12
Brain Responses to Hypnotic Verbal Suggestions Predict Pain Modulation
Desmarteaux, C., Streff, A., Chen, J.-I., Houzé, B., Piché, M., & Rainville, P. (2021) · Frontiers in Pain Research · doi:10.3389/fpain.2021.757384 · Cited by 19
“The notion of emulation refers not only to a sensorimotor preparation, but also to the ability of the brain to run an off-line model in order to produce mental imagery (60), a conscious access to simulations of emulators, being engaged in hypnosis (2).”
The “eye” in imagination: The role of eye movements in a reciprocal aiming task
Pathak, A., Patel, S., Karlinsky, A. D., Taravati, S., & Welsh, T. N. (2022) · Behavioural Brain Research · doi:10.1016/j.bbr.2022.114261 · Cited by 12
Mannella, F., & Tummolini, L. (2022) · Philosophical Transactions of the Royal Society B Biological Sciences · doi:10.1098/rstb.2021.0370 · Cited by 14
“compensating for delay or noise in sensory signals), and that they may have later become available to enable more complex forms of goal-directed or intentional control, which, by inverting the process, use these predictions to select the actions that will cause the predicted sensory effects [30,36,73].”
Mental representation of space: Insights from an oblique distribution of hallucinations
Girard, T. A., Martius, D., & Cheyne, J. A. (2006) · Neuropsychologia · doi:10.1016/j.neuropsychologia.2006.10.002 · Cited by 12
Dynamic Simulation and Static Matching for Action Prediction: Evidence From Body Part Priming
Springer, A., Brandstädter, S., & Prinz, W. (2013) · Cognitive Science · doi:10.1111/cogs.12044 · Cited by 13
“This corresponds to the notion that sensorimotor simulations enable for extrapolating future actions (Grush, 2004; Thornton & Knoblich, 2006).”
Anticipation by multi-modal association through an artificial mental imagery process
Gaona, W., Escobar, E., Hermosillo, J., & Lara, B. (2014) · Connection Science · doi:10.1080/09540091.2014.956289 · Cited by 12
“In the quest of a basic simulation mechanism, forward and inverse models have been proposed (e.g. Grush, 2004; Miall & Wolpert, 1996; Wolpert, Ghahramani, & Flanagan, 2001).”
Molina, M., Kudlinski, C., Guilbert, J., Spruijt, S., Steenbergen, B., & Jouen, F. (2014) · Research in Developmental Disabilities · doi:10.1016/j.ridd.2014.10.053 · Cited by 12
“MI would be used to predict the proprioceptive consequences of an action and then contribute to movement planning (Grush, 2004; Papaxanthis, Pozzo, Skoura, & Schieppati, 2002).”
Brattan, V., Baker, D. H., & Tipper, S. P. (2015) · Journal of Experimental Psychology Human Perception & Performance · doi:10.1037/xhp0000079 · Cited by 13
Top Mysteries of the Mind: Insights From the Default Space Model of Consciousness
Jerath, R., & Beveridge, C. (2018) · Frontiers in Human Neuroscience · doi:10.3389/fnhum.2018.00162 · Cited by 14
“…neurology and psychology that what we experience is actually heavily influenced by our executive expectations, past experiences, and biases instead of a direct experience of external stimuli (Driver and Vuilleumier, 2001; Smythies, 2003; Grush, 2004; Revonsuo, 2006; Maldonato and Dell’Orco, 2012).”
CPG Control for Harmonic Motion of Assistive Robot With Human Motor Control Identification
Zhao, J.-X., & Iwasaki, T. (2020) · IEEE Transactions on Control Systems Technology · doi:10.1109/tcst.2019.2910160 · Cited by 12
“An improved feedback model is discussed in [26], where an inner observer is added to the control loop.”
Tonic dopamine, uncertainty and basal ganglia action selection
Gilbertson, T., & Steele, J. D. (2021) · Neuroscience · doi:10.1016/j.neuroscience.2021.05.010 · Cited by 15
Building human-like communicative intelligence: A grounded perspective
Dubova, M. (2021) · Cognitive Systems Research · doi:10.1016/j.cogsys.2021.12.002 · Cited by 15
“This proposal has been developed in L. Barsalou’s Theory of Perceptual Symbols (Barsalou, 1999) and in other views arguing for modal simulations underlying language understanding (e.g. Zwaan & Radvansky, 1998; Glenberg & Robertson, 1999; Grush, 2004; Gallese, 2008; Zwaan, 2016).”
Time course of learning sequence representations in action imagery practice
Dahm, S. F., & Rieger, M. (2022) · Human Movement Science · doi:10.1016/j.humov.2022.103050 · Cited by 12
“We therefore conclude that implicit sequence learning is based on similar mechanisms in AIP and AEP (Grush, 2004; Jeannerod, 2001).”
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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