Deficient internal models for planning hand-object interactions in apraxia.

Motor imagery (MI) has been associated with planning stages of motor production, and in particular, with internal models that predict the sensory consequences of motor commands and specify the motor commands required to achieve a given outcome. In this study we investigated several predictions derived from the hypothesis that ideomotor apraxia (IM), a deficit in pantomime and imitation of skilled actions, may be attributable in part to deficits in internal models for planning object-related actions, in the face of relatively intact on-line, feedback-driven control of action. This hypothesis predicts that in IM, motor imagery should be (a) strongly correlated with other motor tasks not providing strong visual, tactile, and proprioceptive feedback from objects, i.e., object-related pantomime and imitation; (b) poorly correlated with performance tasks providing strong environmental feedback about the locations of effectors and targets, i.e., actual interaction with objects; and (c) particularly deficient in conditions that are computationally difficult for the motor planning system. Eight left fronto-parietal stroke patients with IM, five stroke patients without IM, and six healthy matched controls imagined grasping dowels and widgets presented at varying orientations, and actually grasped the same objects. The experimental predictions were confirmed. In addition, patients with IM and motor imagery deficits were significantly more likely than the non-apraxic group to have lesions in the intraparietal sulcus, a region previously implicated in imagery for hand-object interactions. The findings suggest a principled explanation for the deficits of IM patients in object-related gesture pantomime, imitation, and learning of new object-related gestures.

A distributed left hemisphere network active during planning of everyday tool use skills.

Determining the relationship between mechanisms involved in action planning and/or execution is critical to understanding the neural bases of skilled behaviors, including tool use. Here we report findings from two fMRI studies of healthy, right-handed adults in which an event-related design was used to distinguish regions involved in planning (i.e. identifying, retrieving and preparing actions associated with a familiar tools' uses) versus executing tool use gestures with the dominant right (experiment 1) and non-dominant left (experiment 2) hands. For either limb, planning tool use actions activates a distributed network in the left cerebral hemisphere consisting of: (i) posterior superior temporal sulcus, along with proximal regions of the middle and superior temporal gyri; (ii) inferior frontal and ventral premotor cortices; (iii) two distinct parietal areas, one located in the anterior supramarginal gyrus (SMG) and another in posterior SMG and angular gyrus; and (iv) dorsolateral prefrontal cortex (DLFPC). With the exception of left DLFPC, adjacent and partially overlapping sub-regions of left parietal, frontal and temporal cortex are also engaged during action execution. We suggest that this left lateralized network constitutes a neural substrate for the interaction of semantic and motoric representations upon which meaningful skills depend.

The neural bases of complex tool use in humans.

The behaviors involved in complex human tool use cut across boundaries traditionally drawn between social, cognitive, perceptual and motor processes. Longstanding neuropsychological evidence suggests a distinction between brain systems responsible for representing: (1) semantic knowledge about familiar tools and their uses, and (2) the acquired skills necessary for performing these actions. Contemporary findings in functional neuroimaging support and refine this distinction by revealing the distributed neural systems that support these processes and the conditions under which they interact. Together, these findings indicate that behaviors associated with complex tool use arise from functionally specialized networks involving temporal, parietal and frontal areas within the left cerebral hemisphere.

Stimulation through simulation? Motor imagery and functional reorganization in hemiplegic stroke patients.

A key factor influencing reorganization of function in damaged neural networks of the adult brain is stimulation. How to stimulate motor areas of patients with paralyses is a formidable challenge. One possibility is to use internal movement simulations, or motor imagery, as an alternative to conventional therapeutic interventions that require voluntary limb movements. Before this alternative can be entertained, two preliminary issues must be resolved. First, do internal movement simulations involve the same neural circuits as comparable overt actions? Second, are motor-impaired populations capable of imagining movements they can no longer perform? Here, I show that under specific conditions, answers to these questions are affirmative. Further, I discuss preliminary evidence that internally simulating movements may induce functional reorganization of the contralesional hand representation of a chronic, densely hemiplegic, cerebral vascular accident (CVA) patient.

Functional imaging of face and hand imitation: towards a motor theory of empathy.

Empathy requires the ability to map the feelings of others onto our own nervous system. Until recently, there was no plausible mechanism to explain how such a mapping might occur. The discovery of mirror neurons, however, suggests that the nervous system is capable of mapping the observed actions of others onto the premotor cortex of the self, at least for reaching and grasping movements. Is there a mirroring system for emotive actions, such as facial expression? Subjects (N = 15; all right-handed; eight men, seven women) watched movies of facial expressions (smile or frown) and hand movements (move index or middle finger) while brain activity was imaged using functional magnetic resonance imaging (fMRI). Subjects watched the movies under three different conditions: passive viewing, active imitation, and an active motor control. Subjects also performed a verb generation task to functionally identify language-processing areas. We found evidence for a common cortical imitation circuit for both face and hand imitation, consisting of Broca's area, bilateral dorsal and ventral premotor areas, right superior temporal gyrus (STG), supplementary motor area, posterior temporo-occipital cortex, and cerebellar areas. For faces, passive viewing led to significant activation in the right ventral premotor area, whereas imitation produced bilateral activation. This result is consistent with evidence for right hemisphere (RH) dominance for emotional processing, and suggests that there may be a right hemisphere mirroring system that could provide a neural substrate for empathy.

Actions or hand-object interactions? Human inferior frontal cortex and action observation.

Cells in macaque ventral premotor cortex (area F5c) respond to observation or production of specific hand-object interactions. Studies in humans associate the left inferior frontal gyrus, including putative F5 homolog pars opercularis, with observing hand actions. Are these responses related to the realized goal of a prehensile action or to the observation of dynamic hand movements? Rapid, event-related fMRI was used to address this question. Subjects watched static pictures of the same objects being grasped or touched while performing a 1-back orienting task. In all 17 subjects, bilateral inferior frontal cortex was differentially activated in response to realized goals of observed prehensile actions. Bilaterally, precentral gyrus was most frequently activated (82%) followed by pars triangularis (73%) and pars opercularis (65%).

What's so special about human tool use?

Evidence suggests homologies in parietofrontal circuits involved in object prehension among humans and monkeys. Likewise, tool use is known to induce functional reorganization of their visuotactile limb representations. Yet, humans are the only species for whom tool use is a defining and universal characteristic. Why? Comparative studies of chimpanzee tool use indicate that critical differences are likely to be found in mechanisms involved in causal reasoning rather than those implementing sensorimotor transformations. Available evidence implicates higher-level perceptual areas in these processes.