The capacity to use tools is a fundamental evolutionary achievement. Its essence stands in the capacity to transfer a proximal goal (grasp a tool) to a distal goal (e.g., grasp food). Where and how does this goal transfer occur? Here, we show that, in monkeys trained to use tools, cortical motor neurons, active during hand grasping, also become active during grasping with pliers, as if the pliers were now the hand fingers. This motor embodiment occurs both for normal pliers and for ''reverse pliers,'' an implement that requires finger opening, instead of their closing, to grasp an object. We conclude that the capacity to use tools is based on an inherently goal-centered functional organization of primate cortical motor areas.neurophysiology ͉ tool use ͉ goal coding ͉ motor act
Most of the research on cortical processing of taste has focused on either the primary gustatory cortex (GC) or the orbitofrontal cortex (OFC). However, these are not the only areas involved in taste processing. Gustatory information can also reach another frontal region, the medial prefrontal cortex (mPFC), via direct projections from GC. mPFC has been studied extensively in relation to its role in controlling goal-directed action and reward-guided behaviors, yet very little is known about its involvement in taste coding. The experiments presented here address this important point and test whether neurons in mPFC can significantly process the physiochemical and hedonic dimensions of taste. Spiking responses to intraorally delivered tastants were recorded from rats implanted with bundles of electrodes in mPFC and GC. Analysis of single-neuron and ensemble activity revealed similarities and differences between the two areas. Neurons in mPFC can encode the chemosensory identity of gustatory stimuli. However, responses in mPFC are sparser, more narrowly tuned, and have a later onset than in GC. Although taste quality is more robustly represented in GC, taste palatability is coded equally well in the two areas. Additional analysis of responses in neurons processing the hedonic value of taste revealed differences between the two areas in temporal dynamics and sensitivities to palatability. These results add mPFC to the network of areas involved in processing gustatory stimuli and demonstrate significant differences in taste-coding between GC and mPFC.
Evidence from a large number of brain imaging studies has shown that, in humans, the insula, and especially its anterior part, is involved in emotions and emotion recognition. Typically, however, these studies revealed that, besides the insula, a variety of other cortical and subcortical areas are also active. Brain imaging studies are correlative in nature, and, as such, they cannot give indications about the necessary contribution of the different centers involved in emotions. In the present study, we aimed to define more clearly the role of the insula in emotional and social behavior of the monkey by stimulating it electrically. Using this technique, one may determine whether direct activation of the insula can produce specific emotional or social behaviors and exactly which parts of this structure are responsible for these behaviors. The results showed that two emotional behaviors, a basic one (disgust) and a social one (affiliative state), were easily elicited by electrical stimulation of specific parts of the insula. Both behaviors were characterized by specific motor and vegetative responses and by a dramatic change in the monkey's responsiveness to external stimuli.
We recorded motor-evoked potentials (MEPs) to transcranial magnetic stimulation from the right opponens pollicis (OP) muscle while participants observed an experimenter operating two types of pliers: pliers opened by the extension of the fingers and closed by their flexion ("normal pliers") and pliers opened by the flexion of the fingers and closed by their extension ("reverse pliers"). In one experimental condition, the experimenter merely opened and closed the pliers; in the other, he grasped an object with them. In a further condition, the participants imagined themselves operating the normal and reverse pliers. During the observation of actions devoid of a goal, the MEP amplitudes, regardless of pliers used,reflectedthemuscularpatterninvolvedintheexecutionoftheobservedaction.Incontrast,duringtheobservationofgoal-directedactions, the MEPs from OP were modulated by the action goal, increasing during goal achievement despite the opposite hand movements necessary to obtain it. During motor imagery, the MEPs recorded from OP reflected the muscular pattern required to perform the imagined action. We propose that covert activity in the human motor cortex may reflect different aspects of motor behavior. Imagining oneself performing tool actions or observing tool actions devoid of a goal activates the representation of the hand movements that correspond to the observed ones. In contrast, the observation of tool actions with a goal incorporates the distal part of the tool in the observer's body schema, resulting in a higher-order representation of the meaning of the motor act.
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