A category of stimuli of great importance for primates, humans in particular, is that formed by actions done by other individuals. If we want to survive, we must understand the actions of others. Furthermore, without action understanding, social organization is impossible. In the case of humans, there is another faculty that depends on the observation of others' actions: imitation learning. Unlike most species, we are able to learn by imitation, and this faculty is at the basis of human culture. In this review we present data on a neurophysiological mechanism--the mirror-neuron mechanism--that appears to play a fundamental role in both action understanding and imitation. We describe first the functional properties of mirror neurons in monkeys. We review next the characteristics of the mirror-neuron system in humans. We stress, in particular, those properties specific to the human mirror-neuron system that might explain the human capacity to learn by imitation. We conclude by discussing the relationship between the mirror-neuron system and language.

The mirror-neuron system.

� Abstract A category of stimuli of great importance for primates, humans in particular, is that formed by actions done by other individuals. If we want to survive, we must understand the actions of others. Furthermore, without action understanding, social organization is impossible. In the case of humans, there is another faculty that depends on the observation of others’ actions: imitation learning. Unlike most species, we are able to learn by imitation, and this faculty is at the basis of human culture. In this review we present data on a neurophysiological mechanism—the mirror-neuron mechanism—that appears to play a fundamental role in both action understanding and imitation. We describe first the functional properties of mirror neurons in monkeys. We review next the characteristics of the mirror-neuron system in humans. We stress, in particular, those properties specific to the human mirror-neuron system that might explain the human capacity to learn by imitation. We conclude by discussing the relationship between the mirror-neuron system and language.

/pdf/the-mirror-neuron-system-4coc36y6dq.pdf

The mirror neuron system.

The parieto-frontal cortical circuit that is active during action observation is the circuit with mirror properties that has been most extensively studied. Yet, there remains controversy on its role in social cognition and its contribution to understanding the actions and intentions of other individuals. Recent studies in monkeys and humans have shed light on what the parieto-frontal cortical circuit encodes and its possible functional relevance for cognition. We conclude that, although there are several mechanisms through which one can understand the behaviour of other individuals, the parieto-frontal mechanism is the only one that allows an individual to understand the action of others 'from the inside' and gives the observer a first-person grasp of the motor goals and intentions of other individuals.

/pdf/the-functional-role-of-the-parieto-frontal-mirror-circuit-1kxsl17xok.pdf

The functional role of the parieto-frontal mirror circuit: interpretations and misinterpretations.

https://www.cns.nyu.edu/~david/courses/sm12/Readings/Wandell-Neuron2007.pdf

Visual Field Maps in Human Cortex

http://www.hegde.us/PIIS1364661312002471.pdf

The ventral visual pathway: an expanded neural framework for the processing of object quality

Though other species of primates also use tools, humans appear unique in their capacity to understand the causal relationship between tools and the result of their use. In a comparative fMRI study, we scanned a large cohort of human volunteers and untrained monkeys, as well as two monkeys trained to use tools, while they observed hand actions and actions performed using simple tools. In both species, the observation of an action, regardless of how performed, activated occipitotemporal, intraparietal, and ventral premotor cortex, bilaterally. In humans, the observation of actions done with simple tools yielded an additional, specific activation of a rostral sector of the left inferior parietal lobule (IPL). This latter site was considered human-specific, as it was not observed in monkey IPL for any of the tool videos presented, even after monkeys had become proficient in using a rake or pliers through extensive training. In conclusion, while the observation of a grasping hand activated similar regions in humans and monkeys, an additional specific sector of IPL devoted to tool use has evolved in Homo sapiens, although tool-specific neurons might reside in the monkey grasping regions. These results shed new light on the changes of the hominid brain during evolution.

https://www.jneurosci.org/content/jneuro/29/37/11523.full.pdf

The Representation of Tool Use in Humans and Monkeys: Common and Uniquely Human Features

Using functional magnetic resonance imaging (fMRI), we mapped the retinotopic organization throughout the visual cortex of fixating monkeys. The retinotopy observed in areas V1, V2, and V3 was completely consistent with the classical view. V1 and V3 were bordered rostrally by a vertical meridian representation, and V2 was bordered by a horizontal meridian. More anterior in occipital cortex, both areas V3A and MT-V5 had lower and upper visual field representations split by a horizontal meridian. The rostral border of dorsal V4 was characterized by the gradual transition of a representation of the vertical meridian (dorsally) to a representation of the horizontal meridian (more ventrally). Central and ventral V4, on the other hand, were rostrally bordered by a representation of the horizontal meridian. The eccentricity lines ran perpendicular to the ventral V3-V4 border but were parallel to the dorsal V3-V4 border. These results indicate different retinotopic organizations within dorsal and ventral V4, suggesting that the latter regions may not be merely the lower and upper visual field representations of a single area. Moreover, because the present fMRI data are in agreement with previously published electrophysiological results, reported distinctions in the retinotopic organization of human and monkey dorsal V4 reflect genuine species differences that cannot be attributed to technical confounds. Finally, aside from dorsal V4, the retinotopic organization of macaque early visual cortex (V1, V2, V3, V3A, and ventral V4) is remarkably similar to that observed in human fMRI studies. This finding indicates that early visual cortex is mostly conserved throughout hominid evolution.

http://www.cnbc.cmu.edu/~tai/readings/v2andAll/v4monkeyfMRI.pdf

The Retinotopic Organization of Primate Dorsal V4 and Surrounding Areas: A Functional Magnetic Resonance Imaging Study in Awake Monkeys

Early time points perfusion imaging

Representation of observed actions in macaque occipitotemporal and parietal cortex

Enhanced fMRI using iron oxide contrast agent in awake, behaving primates

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