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 well-established in accounting for various features of perception, cognition, and motor control.
“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
The citation to Grush (2004) is attached to cognition. Section 6.2, “Cognition” (page unavailable), connects anticipatory mental models to cognition, but says the emulation framework “might be a part of our eventual understanding” of it. That tentative account does not support describing predictive models as already well-established in accounting for cognition. (LLM, conf=0.87)
Experimental — accuracy not yet measuredGrush proposes that the proprioceptive and kinesthetic feelings experienced in motor imagery arise from emulating intended motor commands.
“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
Common features of predictive processing and emulator frameworks include the appeal to forward models and mechanisms for estimating uncertainty and flexibly balancing prior expectations and driving sensory inputs.
“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
Grush's account of perception as 'environmental emulation' looks highly congruent with the depiction of perception as reconstructing the hidden causes structuring the sensory signal.
“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
For couples, familiarity can lead to greater neural synchrony when familiar actions and communications are incorporated as neural predictions.
“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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Rauschecker, J. P., & Scott, S. K. (2009) · Nature Neuroscience · doi:10.1038/nn.2331 · Cited by 1,766 · see citing works
“More recently, forward models have been used to describe the predictive nature of perception and imager”
Goldman, A. I. (2006) · doi:10.1093/0195138929.001.0001 · Cited by 1,490 · see citing works
Embodiment and cognitive science
Gibbs, R. W. (2007) · Choice Reviews Online · doi:10.5860/choice.45-2314 · Cited by 1,264 · see citing works
Neural reuse: A fundamental organizational principle of the brain
Anderson, M. L. (2010) · Behavioral and Brain Sciences · doi:10.1017/s0140525x10000853 · Cited by 1,212 · see citing works
Radical embodied cognitive science
Chemero, A. P. (2010) · Choice Reviews Online · doi:10.5860/choice.47-3480 · Cited by 1,144 · see citing works
Consensus Paper: The Cerebellum's Role in Movement and Cognition
Koziol, L. F., Budding, D. E., Andreasen, N. C., D’Arrigo, S., Bulgheroni, S., Imamizu, H., Ito, M., Manto, M. U., Marvel, C. L., Parker, K. L., et al. (2013) · The Cerebellum · doi:10.1007/s12311-013-0511-x · Cited by 1,136 · see citing works
“It has been proposed that bodily internal models can be reenacted and used in simulation [199, 200], emulation [201], or other (Bayesian) generative schemes [202] to support higher cognition in individual and social domains.”
An integrated theory of language production and comprehension
Pickering, M. J., & Garrod, S. (2013) · Behavioral and Brain Sciences · doi:10.1017/s0140525x12001495 · Cited by 1,569 · see citing works
Simulation, situated conceptualization, and prediction
Barsalou, L. W. (2009) · Philosophical Transactions of the Royal Society B: Biological Sciences · doi:10.1098/rstb.2008.0319 · Cited by 1,017 · see citing works
An Interoceptive Predictive Coding Model of Conscious Presence
Seth, A. K., Suzuki, K., & Critchley, H. D. (2012) · Frontiers in Psychology · doi:10.3389/fpsyg.2011.00395 · Cited by 950 · see citing works
“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) (see Bubic et al., 2010 for a review).”
Visible embodiment: Gestures as simulated action
Hostetter, A. B., & Alibali, M. W. (2008) · Psychonomic Bulletin & Review · doi:10.3758/pbr.15.3.495 · Cited by 875 · see citing works
“Grush, R. (2004). The emulation theory of representation: Motor”
The dynamics of perception and action.
Warren, W. H. (2006) · Psychological Review · doi:10.1037/0033-295x.113.2.358 · Cited by 866 · see citing works
“For example, it is conceivable that a dynamical perceptual– motor process might become detached from its original informational and behavioral context to form an emergent representation, which could then be deployed in mental practice, imagery, off-line planning, or strategic reasoning (cf. Barsalou, 1999; Grush, 2004…”
The Case for Motor Involvement in Perceiving Conspecifics.
Wilson, M., & Knoblich, G. (2005) · Psychological Bulletin · doi:10.1037/0033-2909.131.3.460 · Cited by 814 · see citing works
Inner Speech: Development, Cognitive Functions, Phenomenology, and Neurobiology
Alderson-Day, B., & Fernyhough, C. (2015) · Psychological Bulletin · doi:10.1037/bul0000021 · Cited by 808 · see citing works
“…inputs: for example, producing overt speech is thought to involve the sending of an “efference copy” of the speech motor plan to speech perception areas, forming the basis for a predictive model of what the utterance will sound like, and inhibiting the ensuing auditory response (Grush, 2004).”
Failure to Detect Mismatches Between Intention and Outcome in a Simple Decision Task
Johansson, P., Hall, L., Sikström, S., & Olsson, A. (2005) · Science · doi:10.1126/science.1111709 · Cited by 683 · see citing works
Action-based language: A theory of language acquisition, comprehension, and production
Glenberg, A. M., & D, V. G. M. (2011) · Cortex · doi:10.1016/j.cortex.2011.04.010 · Cited by 675 · see citing works
“We add to the Wolpert and Kawato scheme two features addressed by Grush (2004) and Hurley (2004).”
Active Inference, homeostatic regulation and adaptive behavioural control
Pezzulo, G., Rigoli, F., & Friston, K. (2015) · Progress in Neurobiology · doi:10.1016/j.pneurobio.2015.09.001 · Cited by 650 · see citing works
“of the same perceptual-motor loops – and internally generated brain dynamics – might realize increasingly complex cognitive functions such as planning, imagery, and conscious thought (Buzsáki et al., 2014; Grush, 2004; Hesslow, 2002; Jeannerod, 2001; Pezzulo, 2011; Pezzulo and Castelfranchi, 2009).”
Prediction, cognition and the brain
Bubić, A. (2010) · Frontiers in Human Neuroscience · doi:10.3389/fnhum.2010.00025 · Cited by 658 · see citing works
“…of prediction have much greater value (Friston and Stephan, 2007), as they may signal unsuccessful learning, a major change in the surroundings or noise and smaller changes in the body or the environment, corresponding to normal (plant or world) drifts which typically occur over time (Grush, 2004).”
InfluentialHaidt, J., & Joseph, C. (2008) · The Innate Mind, Volume 3 · doi:10.1093/acprof:oso/9780195332834.003.0019 · Cited by 650 · see citing works
Motor imagery and action observation: cognitive tools for rehabilitation
Mulder, T. (2007) · Journal of Neural Transmission · doi:10.1007/s00702-007-0763-z · Cited by 652 · see citing works
“Grush (2004) takes a related standpoint, but goes one step further by stating that motor imagery results from the control centers of the brain driving an emulator of the body, whereby the normal efferent flow is disengaged from the muscle system.”
Do people use language production to make predictions during comprehension?
Pickering, M. J., & Garrod, S. (2007) · Trends in Cognitive Sciences · doi:10.1016/j.tics.2006.12.002 · Cited by 633 · see citing works
Oberman, L. M., & Ramachandran, V. S. (2007) · Psychological Bulletin · doi:10.1037/0033-2909.133.2.310 · Cited by 630 · see citing works
“An emulator is defined as " a device that mimics the input/output operation of a target system " (such as the motor system; Grush, 2004, p. 387).”
InfluentialDevelopmental robotics: a survey
Lungarella, M., Giorgio Metta, B., Pfeifer, R., & Sandini, G. (2003) · Connection Science · doi:10.1080/09540090310001655110 · Cited by 626 · see citing works
“Forward models are ‘neural simulators’ of the musculo-skeletal system and the environment (Clark and Grush 1999, Grush 2003, Wolpert et al. 2003), and thus allow prediction of the future state of the system given the present state and a certain input (a state specifies a particular body…”
Hallucinations and strong priors
Corlett, P. R., Horga, G., Fletcher, P. C., Alderson-Day, B., Schmack, K., & Powers, A. R. (2019) · Trends in Cognitive Sciences · doi:10.1016/j.tics.2018.12.001 · Cited by 534 · see citing works
“In particular, this renders perception similar to hallucination [1,10].”
Hanakawa, T., Dimyan, M. A., & Hallett, M. (2008) · Cerebral Cortex · doi:10.1093/cercor/bhn036 · Cited by 565 · see citing works
“From the standpoint of the emulation theory of representation (Grush 2004), what is present in motor imagery, but not in the ISrelated processes, is probably willed generation of efference copy and analysis of ''mock'' sensory (mainly proprioceptive and partly visual) signals.”
Prediction of external events with our motor system: towards a new framework
Schubotz, R. I. (2007) · Trends in Cognitive Sciences · doi:10.1016/j.tics.2007.02.006 · Cited by 550 · see citing works
Hurley, S. (2008) · Behavioral and Brain Sciences · doi:10.1017/s0140525x07003123 · Cited by 512 · see citing works
“Cf. Wolpert (1997), Wolpert and Kawato (1998), Haruno et al. (2001), and Wolpert et al. (2003) on selection among forward models; Grush (2004) on continuous combination of real and simulated input; and Grush (1995) linking motor control and mindreading.”
InfluentialWhere's the action? The pragmatic turn in cognitive science
Engel, A. K., Maÿe, A., Kurthen, M., & Peter König, B. (2013) · Trends in Cognitive Sciences · doi:10.1016/j.tics.2013.03.006 · Cited by 435 · see citing works
Shared Neural Mechanisms of Visual Perception and Imagery
Dijkstra, N., Bosch, S., & van Gerven, M. A. J. (2019) · Trends in Cognitive Sciences · doi:10.1016/j.tics.2019.02.004 · Cited by 441 · see citing works
Brain-to-Brain Synchrony during Naturalistic Social Interactions
Kinreich, S., Djalovski, A., Kraus, L., Louzoun, Y., & Feldman, R. (2017) · Scientific Reports · doi:10.1038/s41598-017-17339-5 · Cited by 402 · see citing works
Keller, P. E., Novembre, G., & Hove, M. J. (2014) · Philosophical Transactions of the Royal Society B: Biological Sciences · doi:10.1098/rstb.2013.0394 · Cited by 466 · see citing works
Cognitive Systems and the Extended Mind
Rupert, R. D. (2009) · doi:10.1093/acprof:oso/9780195379457.001.0001 · Cited by 410 · see citing works
Impact of contour on aesthetic judgments and approach-avoidance decisions in architecture
Vartanian, O. A., Navarrete, G., Chatterjee, A. K., Fich, L. B., Leder, H., Modroño, C., Nadal, M., Rostrup, N., & Skov, M. (2013) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1301227110 · Cited by 417 · see citing works
Horizons for the Enactive Mind: Values, Social Interaction, and Play
Paolo, E. A. D., Rohde, M., & Jaegher, H. D. (2010) · The MIT Press eBooks · doi:10.7551/mitpress/9780262014601.003.0003 · Cited by 405 · see citing works
Joint Action, Interactive Alignment, and Dialog
Garrod, S., & Pickering, M. J. (2009) · Topics in Cognitive Science · doi:10.1111/j.1756-8765.2009.01020.x · Cited by 402 · see citing works
Bruineberg, J., Kiverstein, J., & Rietveld, E. (2018) · Synthese · doi:10.1007/s11229-016-1239-1 · Cited by 383 · see citing works
“…dynamics have been at the heart of a large number of influential and very diverse theories of mind, brain and skilled behavior more generally (Helmholtz 1860/1962; Gregory 1980; Grush 2004; Turvey 2010; Noë 2004; Stepp and Turvey 2010; Port and Van Gelder 1995; Thompson 2007; Chemero 2009).”
From the Revolution to Embodiment
Glenberg, A. M., Witt, J. K., & Metcalfe, J. (2013) · Perspectives on Psychological Science · doi:10.1177/1745691613498098 · Cited by 377 · see citing works
“Critically, the forward model can also be used imaginatively, in mental practice, to evaluate outcomes without performing them (e.g., Grush, 2004).”
Navigating the Affordance Landscape: Feedback Control as a Process Model of Behavior and Cognition
Pezzulo, G., & Cisek, P. (2016) · Trends in Cognitive Sciences · doi:10.1016/j.tics.2016.03.013 · Cited by 419 · see citing works
Free-Energy Minimization and the Dark-Room Problem
Friston, K., Thornton, C., & Clark, A. (2012) · Frontiers in Psychology · doi:10.3389/fpsyg.2012.00130 · Cited by 367 · see citing works
“that we can predict (cf., emulation theory; Grush, 2004); ensuring we keep to well-trodden paths in the space of all the physical and physiological variables that underwrite our existence.”
Buzsáki, G. (2019) · doi:10.1093/oso/9780190905385.001.0001 · Cited by 358 · see citing works
How we know our own minds: The relationship between mindreading and metacognition
Carruthers, P. (2009) · Behavioral and Brain Sciences · doi:10.1017/s0140525x09000545 · Cited by 610 · see citing works
The Routledge Companion to Landscape Studies
Howard, P., Thompson, I., & Waterton, E. (2013) · Routledge eBooks · doi:10.4324/9780203096925 · Cited by 331 · see citing works
Challenges and opportunities for grounding cognition.
Barsalou, L. W. (2020) · Journal of Cognition · doi:10.5334/joc.116 · Cited by 319 · see citing works
Multiple roles of motor imagery during action observation
Vogt, S., Rienzo, F. D., Collet, C., Collins, A. F., & Guillot, A. (2013) · Frontiers in Human Neuroscience · doi:10.3389/fnhum.2013.00807 · Cited by 301 · see citing works
“…proposed by Jeannerod (1994, 2001, 2006), and more recently motor simulation, along with prediction as its most prominent cognitive function, has become a commonly accepted framework for a wide range of cognitive domains (Grush, 2004; Kilner et al., 2007; Bubic et al., 2010; Pezzulo et al., 2013).”
Song and speech: brain regions involved with perception and covert production
Callan, D. E., Tsytsarev, V., Hanakawa, T., Callan, A. M., Katsuhara, M., Fukuyama, H., & Turner, R. (2006) · NeuroImage · doi:10.1016/j.neuroimage.2006.01.036 · Cited by 287 · see citing works
“A recent model (Grush, 2004) proposes that there should be domain-general type imagery, called supramodal or amodal imagery.”
Mental Training as a Tool in the Neuroscientific Study of Brain and Cognitive Plasticity
Slagter, H. A., Davidson, R. J., & Lutz, A. (2011) · Frontiers in Human Neuroscience · doi:10.3389/fnhum.2011.00017 · Cited by 288 · see citing works
Motor imagery involves predicting the sensory consequences of the imagined movement
Kilteni, K., Andersson, B. J., Houborg, C., & Ehrsson, H. H. (2018) · Nature Communications · doi:10.1038/s41467-018-03989-0 · Cited by 293 · see citing works
Radical Embodied Cognitive Science
Chemero, A. P. (2013) · Review of General Psychology · doi:10.1037/a0032923 · Cited by 282 · see citing works
“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 273 · see citing works
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 · see citing works
“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 277 · see citing works
“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 256 · see citing works
“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 · see citing works
Body Schema in Robotics: A Review
Hoffmann, M., Marques, H., Arieta, A., Sumioka, H., Lungarella, M., & Pfeifer, R. (2010) · IEEE Transactions on Autonomous Mental Development · doi:10.1109/tamd.2010.2086454 · Cited by 247 · see citing works
“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 244 · see citing works
“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 241 · see citing works
“, [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 · see citing works
“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 229 · see citing works
“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 · see citing works
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 233 · see citing works
“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 225 · see citing works
“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 · see citing works
“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 220 · see citing works
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 215 · see citing works
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 214 · see citing works
“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 213 · see citing works
“…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 210 · see citing works
“…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 · see citing works
“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 · see citing works
“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 210 · see citing works
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 204 · see citing works
“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 202 · see citing works
“…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 202 · see citing works
“’’ 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 199 · see citing works
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 195 · see citing works
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 194 · see citing works
“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 · see citing works
“…(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 188 · see citing works
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 187 · see citing works
“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 · see citing works
“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 · see citing works
Macuga, K. L., & Frey, S. H. (2011) · NeuroImage · doi:10.1016/j.neuroimage.2011.09.083 · Cited by 175 · see citing works
“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 176 · see citing works
Kinematics 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 166 · see citing works
““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 172 · see citing works
“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 · see citing works
“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).”
Horizons for the Enactive Mind: Values, Social Interaction, and Play
Paolo, E. A. D., Rohde, M., & Jaegher, H. D. (2010) · The MIT Press eBooks · doi:10.7551/mitpress/8629.003.0003 · Cited by 162 · see citing works
Tian, X., & Poeppel, D. (2012) · Frontiers in Human Neuroscience · doi:10.3389/fnhum.2012.00314 · Cited by 161 · see citing works
“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 160 · see citing works
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 160 · see citing works
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 173 · see citing works
“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 158 · see citing works
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 · see citing works
Huang, J. Y., & Bargh, J. A. (2014) · Behavioral and Brain Sciences · doi:10.1017/s0140525x13000290 · Cited by 157 · see citing works
Functional coupling underlying motor and cognitive functions of the dorsal premotor cortex
Abe, M., & Hanakawa, T. (2008) · Behavioural Brain Research · doi:10.1016/j.bbr.2008.10.046 · Cited by 157 · see citing works
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 152 · see citing works
Rauschecker, J. P. (2017) · Cortex · doi:10.1016/j.cortex.2017.10.020 · Cited by 145 · see citing works
- 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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