Teaching and Learning Guide for

This guide accompanies the following article(s): Jonathan H. Venezia and Gregory Hickok, ‘Mirror Neurons, the Motor System, and Language – From the Motor Theory to Embodied Cognition and Beyond’, Language and Linguistics Compass 3 (2009) pp. 1403–1416. DOI: 10.1111/j.1749‐818x.2009.00169.x Author’s Introduction Mirror neurons have been invoked to explain a huge number of psychological phenomena including action understanding, imitation, empathy, theory of mind, consciousness, and many more. Indeed, mirror neurons have provided a physiological substrate for motor theories of cognition, which appear elegant, simple, and easy to accept at first glance. A computationally specific motor theory predating the discovery of mirror neurons is the motor theory of speech perception (MTSP). Decades after MTSP had fallen out of favor among speech scientists it enjoyed a renaissance in the age of mirror neurons. However, mirror neurons have done nothing to overcome the arguments leveled against MTSP years ago. We offer this guide as a reminder that we must be cautious when interpreting and drawing inference from new discoveries that seem to explain complex cognitive processes in attractively simple terms. Speech and language draw upon a myriad of sophisticated cognitive processes, distributed throughout cortex, which combine to solve extraordinarily difficult computational problems. However, there is a large body of evidence suggesting that speech perception, specifically, relies heavily on auditory representations in the superior temporal lobes, and we must interpret motor contributions to speech perception in light of this fact. Key Readings 1. Rizzolatti & Craighero. 2004. The mirror‐neuron system. Annual Review of Neuroscience 27. 169–92. doi: 10.1146/annurev.neuro.27.070203.144230 This highly cited review provides appropriate background and motivation for a discussion on motor system contributions in speech perception, as mirror neurons have fueled the emergence of modern motor theories of cognition. The article covers the discovery and basic properties of mirror neurons in the macaque monkey, and outlines a proposed role for mirror neurons in action understanding. Homologies between monkeys and humans are discussed, and neurophysiological evidence for a human mirror neuron system is reviewed. Finally, evidence concerning a role for the human mirror system in speech is highlighted, including a proposed evolutionary connection between gestural communication and verbal speech. Thus, we recommend this article as a comprehensive introduction to the relation between mirror neurons, action, and speech. 2 . Hickok, G. 2009. Eight problems for the mirror neuron theory of action understanding. Journal of Cognitive Neuroscience 21. 1229–43. doi: 10.1162/jocn.2009.21189 We recommend this article as an alternative to the typical interpretation of evidence concerning the mirror neuron theory of action understanding. Critical review of the relevant neurophysiological data in monkeys and humans is offered. Also, a discussion on the relation between mirror neurons and the motor theory of speech is contained. We feel that close examination and careful scrutiny of the mirror neuron theory of action understanding are of critical importance given the purported link between action‐based gestural communication and verbal speech. 3. Galantucci, Fowler, & Turvey. 2006. The motor theory of speech perception reviewed. Psychonomic Bulletin & Review 13. 361–77. http://pbr.psychonomic‐journals.org/content/13/3/361.abstract This is an excellent recent review of the motor theory of speech perception. The major tenets of the motor theory are concisely summarized along with the original findings that have motivated the theory throughout its several iterations. The evidence for each central claim is reviewed and evaluated, and a thorough discussion of the link between perceptual and motor speech systems is offered, including a section on neural evidence that gives mention to mirror neurons. 4. Lotto, A., Hickok, G. & Holt, L. 2009. Reflections on mirror neurons and speech perception. Trends in Cognitive Sciences 13. 110–4. doi: 10.1016/j.tics.2008.11.008 Lotto et al. provide a thorough discussion of the relation between mirror neurons and the motor theory of speech perception. The article establishes that mirror neurons do nothing to solve the computational problems that originally gave rise to the motor theory, nor do they account for decades‐old behavioral and neuropsychological challenges. In fact, the authors argue that the involvement of mirror neurons in speech perception would contradict the central tenets of the theory. 5. Iacoboni, M. 2008. The role of premotor cortex in speech perception: evidence from fMRI and rTMS. Journal of Physiology – Paris 102. 31–34. doi: 10.1016/j.jphysparis.2008.03.003 Iacoboni provides a nice review of the recent evidence for motor system involvement in speech perception, including a historical perspective linking this evidence with the motor theory of speech and with mirror neurons. 6. Wilson, S. & Iacoboni, M. 2004. Listening to speech activates motor areas involved in speech production. Nature Neuroscience 7. 701–2. doi: 10.1038/nn1263 This fMRI study provides unequivocal evidence that passively listening to speech sounds activates the speech motor system. Motor theorists often cite the results as evidence that speech perception is grounded in motor circuits, though extensive activation was observed in superior temporal cortex as well. Moreover, activation measured during speech production extended into regions of premotor cortex that were not active during perception of the same sounds. Indeed, even non‐speech stimuli (noise bursts) activated a premotor region that was active during speech perception and production. Should we conclude that perception of noise bursts, which our motor system cannot produce, is grounded in motor circuits? Thus, we recommend this study as a reminder that we must be careful in interpreting neuroimaging results. 7. Meister, I., Wilson, M., Deblieck, C., Wu, A., & Iacoboni, M. 2007. The essential role of premotor cortex in speech perception. Current Biology 19. 1692–6

Cognition · Cognitive psychology · Cognitive science · Consciousness · Embodied cognition · Empathy · Imitation · Mirror neuron · Motor Cognition · Motor theory of speech perception · Perception · Social cognition · Speech perception · Theory of mind · Action Observation and Synchronization · Computer Science · Neuroscience · Psychology · Artificial Intelligence