Making order from chaos: the misguided frontal lobe.
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Biomedical subjects
Publications and source records attributed to R Ivry.
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Using an event-related fMRI procedure, we investigated the role of the human cerebellum in sequential finger movements. Subjects performed a delayed sequential finger movement task in which an instructive cue preceded the imperative signal by 16.5 s. Bilateral activation was observed in the cerebellum following both the cue and imperative signals. The activated regions overlapped within the cerebellum, extending across intermediate and lateral regions corresponding to lobules HV-HVII. In contrast, activation in primary motor cortex was primarily restricted to the execution phase and most prominent in the contralateral hemisphere. These results indicate that the cerebellum is bilaterally recruited for the preparation and execution of sequential movements.
The past decade has witnessed a paradigm shift concerning the study of the cerebellum. Results from various studies employing a variety of methodologies suggest that the functional role of this structure is not limited to motor control. The article by Tesche and Karhu appearing in this issue, provides strong evidence that the cerebellum in humans is activated in anticipation of somatosensory events, even when these events do not require overt responses. In their study, the sensory response is observed when the stimuli fail to occur at expected points in time, consistent with the hypothesis that the cerebellum is specialized for representing the temporal relationships between events, motoric or otherwise. Timing and sensory expectancy likely reflect nested hypotheses, and it remains to be seen if one provides a more encompassing yet specific view of cerebellar function.
PET revealed the effects of stimulus characteristics on the neural substrate of motor learning. Right-handed subjects performed a serial reaction time task with colour-coded stimuli to eliminate the potential for learned eye-movements. The task was performed with the right hand under two different conditions. In one condition, subjects simultaneously performed a distractor task. Although they did show behavioural evidence of learning, they were not explicitly aware of the stimulus-response sequence. In the second condition, there was no distractor task, and seven out of the 11 subjects then became explicitly aware of the stimulus sequence. Metabolic correlates of learning were distinct in the two conditions. When learning was implicit under dual-task conditions, learning-related changes were observed in left motor and supplementary motor cortex as well as in the putamen. These regions are similar to those observed in a previous study in which the stimuli were cued by spatial position. Under single-task conditions, metabolic changes were found in the right prefrontal cortex and premotor cortex, as well as in the temporal lobe. A similar shift to the right hemisphere was observed in the spatial study during single-task learning. However, explicit learning of the task with colour stimuli activated more ventral regions. The areas supporting motor-sequence learning are contingent on both stimulus properties and attentional constraints.
Coordinated movement requires the normal operation of a number of different brain structures. Taking a modular perspective, it is argued that these structures provide unique computations that in concert produce coordinated behavior. The coordination problems of patients with cerebellar lesions can be understood as a problem in controlling and regulating the temporal patterns of movement. The timing capabilities of the cerebellum are not limited to the motor domain, but are utilized in perceptual tasks that require the precise representation of temporal information. Patients with cerebellar lesions are impaired in judging the duration of a short auditory stimulus or the velocity of a moving visual stimulus. The timing hypothesis also provides a computational account of the role of the cerebellum in certain types of learning. In particular, the cerebellum is essential for situations in which the animal must learn the temporal relationship between successive events such as in eyeblink conditioning. Modeling and behavioral studies suggest that the cerebellar timing system is best characterized as providing a near-infinite set of interval type timers rather than as a single clock with pacemaker or oscillatory properties. Thus, the cerebellum will be invoked whenever a task requires its timing function, but the exact neural elements that will be activated vary from task to task. The multiple-timer hypothesis suggests an alternative account of neuroimaging results implicating the cerebellum in higher cognitive processes. The activation may reflect the automatic preparation of multiple responses rather than be associated with processes such as semantic analysis, error detection, attention shifting, or response selection.
The role of lateral prefrontal cortex in transducing perception into action was studied in 10 patients with chronic, unilateral lesions. They identified colors in the center of a visual display, while a flanking, distractor color was presented simultaneously in either the ipsilesional or contralesional field. The flanker could be either the same color as the target, or incompatible with the correct response. The effects of compatible and incompatible flankers on reaction time (RT) served as a measure of response channel activation by the flanker. Flankers in the contralesional field influenced RT less than did those in the ipsilesional field. These results suggest that the lateral prefrontal cortex is involved in maintaining stimulus-response channels.
Visual objects are perceived correctly only if their features are identified and then bound together. Illusory conjunctions result when feature identification is correct but an error occurs during feature binding. A new model is proposed that assumes feature binding errors occur because of uncertainty about the location of visual features. This model accounted for data from 2 new experiments better than a model derived from A. M. Treisman and H. Schmidt's (1982) feature integration theory. The traditional method for detecting the occurrence of true illusory conjunctions is shown to be fundamentally flawed. A reexamination of 2 previous studies provided new insights into the role of attention and location information in object perception and a reinterpretation of the deficits in patients who exhibit attentional disorders.
Illusory conjunctions are the incorrect combination of correctly perceived features, such as color and shape. They have been found to occur using a brief exposure (under 200 ms) and a dual task designed to divert attention. The present study investigated the roles of exposure duration and attention in obtaining illusory conjunctions. Several mathematical models of the feature integration task were also assessed. Experiment 1 tested participants' accuracy at combining features using a long exposure and an attention-diverting taks. Experiment 2 compared performance with and without the attention-diverting task. The final experiment compared performance using a brief (0.15 s) and a long (1.5 s) exposure duration without an attention-diverting task. Neither attention nor exposure duration had a significant effect on feature integration.
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This review describes some characteristics of patients with cerebellar lesions, including limb movements, changes in motor planning and disturbances in time-dependent perception. The delay in movement initiation can be explained by a delay in onset of movement-related discharge of neurons in motor cortex. Disorders of movement termination (hypermetria) are accompanied by asymmetric velocity profiles and by prolonged agonist and delayed antagonist EMG activity necessary to brake the movement. During complex movements in three-dimensional space, the cerebellum contributes to timing between single components of a movement, scales the size of muscular action, and coordinates the sequence of agonists and antagonists. The basic structure of motor programs is not generated exclusively within the cerebellum and patients with cerebellar lesions can use precuing information to improve their motor performance. Time-dependent perception in the auditory and visual domains are disturbed in patients with cerebellar lesions.
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This article addresses 2 questions that arise from the finding that visual scenes are first parsed into visual features: (a) the accumulation of location information about objects during their recognition and (b) the mechanism for the binding of the visual features. The first 2 experiments demonstrated that when 2 colored letters were presented outside the initial focus of attention, illusory conjunctions between the color of one letter and the shape of the other were formed only if the letters were less than 1 degree apart. Separation greater than 2 degrees resulted in fewer conjunction errors than expected by chance. Experiments 3 and 4 showed that inside the spread of attention, illusory conjunctions between the 2 letters can occur regardless of the distance between them. In addition, these experiments demonstrated that the span of attention can expand or shrink like a spotlight. The results suggest that features inside the focus of attention are integrated by an expandable focal attention mechanism that conjoins all features that appear inside its focus. Visual features outside the focus of attention may be registered with coarse location information prior to their integration. Alternatively, a quick and imprecise shift of attention to the periphery may lead to illusory conjunctions among adjacent stimuli.
Thirteen patients with bilateral cerebellar disease and 12 patients with unilateral cerebellar disease were instructed to execute movement sequences in response to a simple reaction signal. Each to-be-executed sequence consisted either of a single, two, or three keypress components. Evidence for cerebellar involvement in the execution of programmed responses was sought in the pattern of response onset times and interkeypress times. Patients with mild bilateral cerebellar dysfunction or mild unilateral dysfunction, and neurologically unimpaired subjects showed increases in response onset time as sequence length increased from L = 1 to L = 3. In contrast to this, there were negligible or no effects of sequence length on response onset time in patients with moderate bilateral cerebellar dysfunction and in patients with moderate unilateral cerebellar dysfunction who responded with the hand ipsilateral to the lesion. Furthermore, cerebellar dysfunction was associated with significantly slower interkeypress reaction times. These results support the hypothesis that the translation of a programmed sequence of responses into action involves cerebellar structures which schedule a sequence of ordered responses before onset of movement.
Six experiments investigated the preattentive segregation of line-like patterns composed of discrete elements in a background of distractors. The results indicate that other factors in addition to spatial density influence line segregation. Edge alignment, edge length and principal axis orientation also affect line segregation. Differences in the outputs of Gabor filters fail to account for the perceived segregation of the lines. Possible models of line segregation based on element grouping, feature density and search are briefly discussed.
Different features of stimuli present in the field of view appear to be registered in different cortical maps. How, then, are the features that come from the same object bound together rather than mistakenly assembled with features coming from other simultaneously present objects? One theory supposes that an attentional mechanism intercepts input coming from particular retinal locations at a way station prior to parsing of the features from the same object. Any enhancement (or facilitation) at that stage will cause all the features from that object to be modified simultaneously in the downstream registers. The imposed temporal synchronicity serves as the essential binding cue. Five experiments provided no support for the theory. There is no tendency for synchronicity of features to cause binding unless the features come from the same location. Location, rather than temporal synchronicity, appears to be the essential cue for binding.
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Explore the source record for details and available documents.