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.

This paper concerns how motor actions are neurally represented and coded. Action planning and motor preparation can be studied using a specific type of representational activity, motor imagery. A close functional equivalence between motor imagery and motor preparation is suggested by the positive effects of imagining movements on motor learning, the similarity between the neural structures involved, and the similar physiological correlates observed in both imaging and preparing. The content of motor representations can be inferred from motor images at a macroscopic level, based on global aspects of the action (the duration and amount of effort involved) and the motor rules and constraints which predict the spatial path and kinematics of movements. A more microscopic neural account calls for a representation of object-oriented action. Object attributes are processed in different neural pathways depending on the kind of task the subject is performing. During object-oriented action, a pragmatic representation is activated in which object affordances are transformed into specific motor schemas (independently of other tasks such as object recognition). Animal as well as human clinical data implicate the posterior parietal and premotor cortical areas in schema instantiation. A mechanism is proposed that is able to encode the desired goal of the action and is applicable to different levels of representational organization.

https://www.cambridge.org/core/services/aop-cambridge-core/content/view/S0140525X00034026

The representing brain: Neural correlates of motor intention and imagery

Neural simulation of action: a unifying mechanism for motor cognition.

Grasping objects: the cortical mechanisms of visuomotor transformation

Prehension movements were studied by film in 7 adult subjects. Transportation of the hand to the target-object location had features very similar to any aiming arm movement, that is, it involved a fast-velocity initial phase and a low-velocity final phase. The peak velocity of the movement was highly correlated with its amplitude, although total movement duration tended to remain invariant when target distance was changed. The low-velocity phase consistently began after about 75% of movement time had elapsed. This ration was maintained for different movement amplitudes. Formation of the finger grip occurred during hand transportation. Fingers were first stretched and then began to close in anticipation to contact with the object. The onset of the closure phase was highly correlated to the beginning of the low velocity phase of transportation. This pattern for both transportation and finger grip formation was maintained in conditions whether visual feedback from the moving limb was present or not. Implications of these findings for the central programming of multisegmental movements are discussed.

The Timing of Natural Prehension Movements

An investigation of a subject at the boundaries of psychology and physiology. The author's account is based upon experimental data from human and animal work with particular emphasis on illustrative clinical cases. The organization of goal-directed movements is a problem central to the relations of brain and behaviour in humans and other animals. Movement represents a clue to the study of brain mechanisms and psychological functions. The use of behaving animals in neurophysiological laboratories has allowed direct investigation of the role of discrete neural mechanisms in controlling motor output. The contribution that these mechanisms make can be assessed by reproducing experimentally the motor syndromes observed in man. Kinematic description of motor responses may be directly transferable to theoretical models of motor systems and to their technological applications such as artificial intelligence and robotics. Finally, the use of concepts derived from cognitive psychology in the study of motor control has opened new fields of investigation concerning the mental representation of goal-directed movements.

Marc Jeannerod

Papers

The representing brain: Neural correlates of motor intention and imagery

Neural simulation of action: a unifying mechanism for motor cognition.

Grasping objects: the cortical mechanisms of visuomotor transformation

The Timing of Natural Prehension Movements

The neural and behavioural organization of goal-directed movements