We review evidence for partially segregated networks of brain areas that carry out different attentional functions. One system, which includes parts of the intraparietal cortex and superior frontal cortex, is involved in preparing and applying goal-directed (top-down) selection for stimuli and responses. This system is also modulated by the detection of stimuli. The other system, which includes the temporoparietal cortex and inferior frontal cortex, and is largely lateralized to the right hemisphere, is not involved in top-down selection. Instead, this system is specialized for the detection of behaviourally relevant stimuli, particularly when they are salient or unexpected. This ventral frontoparietal network works as a 'circuit breaker' for the dorsal system, directing attention to salient events. Both attentional systems interact during normal vision, and both are disrupted in unilateral spatial neglect.

Control of goal-directed and stimulus-driven attention in the brain

Co-Planar Stereotaxic Atlas of the Human Brain—3-Dimensional Proportional System: An Approach to Cerebral Imaging, J. Talairach, P. Tournoux. Georg Thieme Verlag, New York (1988), 122 pp., 130 figs. DM 268

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.

The anterior insular cortex (AIC) is implicated in a wide range of conditions and behaviours, from bowel distension and orgasm, to cigarette craving and maternal love, to decision making and sudden insight. Its function in the re-representation of interoception offers one possible basis for its involvement in all subjective feelings. New findings suggest a fundamental role for the AIC (and the von Economo neurons it contains) in awareness, and thus it needs to be considered as a potential neural correlate of consciousness.

How do you feel--now? The anterior insula and human awareness.

Prior to the twentieth century, theories of knowledge were inherently perceptual. Since then, developments in logic, statis- tics, and programming languages have inspired amodal theories that rest on principles fundamentally different from those underlying perception. In addition, perceptual approaches have become widely viewed as untenable because they are assumed to implement record- ing systems, not conceptual systems. A perceptual theory of knowledge is developed here in the context of current cognitive science and neuroscience. During perceptual experience, association areas in the brain capture bottom-up patterns of activation in sensory-motor areas. Later, in a top-down manner, association areas partially reactivate sensory-motor areas to implement perceptual symbols. The stor- age and reactivation of perceptual symbols operates at the level of perceptual components - not at the level of holistic perceptual expe- riences. Through the use of selective attention, schematic representations of perceptual components are extracted from experience and stored in memory (e.g., individual memories of green, purr, hot). As memories of the same component become organized around a com- mon frame, they implement a simulator that produces limitless simulations of the component (e.g., simulations of purr). Not only do such simulators develop for aspects of sensory experience, they also develop for aspects of proprioception (e.g., lift, run) and introspec- tion (e.g., compare, memory, happy, hungry). Once established, these simulators implement a basic conceptual system that represents types, supports categorization, and produces categorical inferences. These simulators further support productivity, propositions, and ab- stract concepts, thereby implementing a fully functional conceptual system. Productivity results from integrating simulators combinato- rially and recursively to produce complex simulations. Propositions result from binding simulators to perceived individuals to represent type-token relations. Abstract concepts are grounded in complex simulations of combined physical and introspective events. Thus, a per- ceptual theory of knowledge can implement a fully functional conceptual system while avoiding problems associated with amodal sym- bol systems. Implications for cognition, neuroscience, evolution, development, and artificial intelligence are explored.

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Perceptual symbol systems.

Receptive field response of LGB neurons during vestibular stimulation in awake cats.

Brains Top Down:Is Top-Down Causation Challenging Neuroscience?

How do we become aware of our own actions? This classical question is still a matter of debate: does consciousness depend on central efferent signals or derive from peripheral information? In this paper, we had the opportunity to study a haptically deafferented patient using a well-tested experimental paradigm where a cognitive conflict is produced between motor intention, proprioception and visual feedback. Our results show that the patient was able to solve the conflict and to generate accurate movements to a target in the absence of proprioceptive feedback and with very limited visual feedback from her movements. Yet, she could not report any conscious perception of the conflict and showed no conscious knowledge of her actual performance. We suggest that information derived from efferent processes cannot in themselves be a source for conscious experience about our own actions.

Lack of conscious knowledge about one's own actions in a haptically deafferented patient

The influence of deprivation procedures on the development of motion detection mechanisms has been studied in twenty-two kittens. Superior colliculus neurons did not acquire direction selectivity and normal ocular dominance in animals reared in the dark or in stroboscopic light. Neuron immaturity persisted in spite of a five week additional recovery period in normal conditions. Exposure to unidirectional visual motion for 10 h during the fifth week of postnatal age produced an asymmetric development of the two superior colliculi. Finally, unilateral neonatal ablation of visual cortex permanently impaired development of the ipsilateral superior colliculus. In the same or in different animals, development of optokinetic nystagmus, a typical visuomotor response, was similarly influenced by the global or selective deprivation procedures. These results suggest that motion detection mechanisms (both afferent and efferent) strongly depend upon constraints imposed by the visual world during the first weeks of life.

Marc Jeannerod

Papers

Receptive field response of LGB neurons during vestibular stimulation in awake cats.

Brains Top Down:Is Top-Down Causation Challenging Neuroscience?

Lack of conscious knowledge about one's own actions in a haptically deafferented patient

Developmental constraints of motion detection mechanisms in the kitten.

New Issues in Spatial Neglect