Subcortical loops through the basal ganglia and the cerebellum form computationally powerful distributed processing modules (DPMs). This paper relates the computational features of a DPM's loop through the basal ganglia to experimental results for two kinds of natural action selection. First, functional imaging during a serial order recall task was used to study human brain activity during the selection of sequential actions from working memory. Second, microelectrode recordings from monkeys trained in a step-tracking task were used to study the natural selection of corrective submovements. Our DPM-based model assisted in the interpretation of puzzling data from both of these experiments. We come to posit that the many loops through the basal ganglia each regulate the embodiment of pattern formation in a given area of cerebral cortex. This operation serves to instantiate different kinds of action (or thought) mediated by different areas of cerebral cortex. We then use our findings to formulate a model of the aetiology of schizophrenia.
The ability to anticipate predictable stimuli allows faster responses. The predictive saccade (PRED) task has been shown to quickly induce such anticipatory behavior in humans. In a PRED task subjects track a visual target jumping back and forth between fixed positions at a fixed time interval. During this task, saccade latencies drop from approximately 200 ms to <80 ms as subjects anticipate target appearance. This change in saccade latency indicates that subjects' behavior shifts from being sensory driven to being memory driven. We conducted functional magnetic resonance imaging studies with 10 healthy adults performing the PRED task using a standard block design. We compared the PRED task with a visually guided saccade (VGS) task using unpredictable targets matched for number, direction and amplitude of required saccades. Our results show greater activation during the PRED task in the prefrontal, pre-supplementary motor and anterior cingulate cortices, hippocampus, mediodorsal thalamus, striatum and cerebellum. The VGS task elicited greater activation in the cortical eye fields and occipital cortex. These results demonstrate the important dissociation between sensory and predictive neural control of similar saccadic eye movements. Anticipatory behavior induced by the PRED task required less sensory-related processing activity and was subserved by a distributed cortico-subcortical memory system including prefronto-striatal circuitry.
BackgroundProprioception plays important roles in planning and control of limb posture and movement. The impact of proprioceptive deficits on motor function post-stroke has been difficult to elucidate due to limitations in current tests of arm proprioception. Common clinical tests only provide ordinal assessment of proprioceptive integrity (eg. intact, impaired or absent). We introduce a standardized, quantitative method for evaluating proprioception within the arm on a continuous, ratio scale. We demonstrate the approach, which is based on signal detection theory of sensory psychophysics, in two tasks used to characterize motor function after stroke.MethodsHemiparetic stroke survivors and neurologically intact participants attempted to detect displacement- or force-perturbations robotically applied to their arm in a two-interval, two-alternative forced-choice test. A logistic psychometric function parameterized detection of limb perturbations. The shape of this function is determined by two parameters: one corresponds to a signal detection threshold and the other to variability of responses about that threshold. These two parameters define a space in which proprioceptive sensation post-stroke can be compared to that of neurologically-intact people. We used an auditory tone discrimination task to control for potential comprehension, attention and memory deficits.ResultsAll but one stroke survivor demonstrated competence in performing two-alternative discrimination in the auditory training test. For the remaining stroke survivors, those with clinically identified proprioceptive deficits in the hemiparetic arm or hand had higher detection thresholds and exhibited greater response variability than individuals without proprioceptive deficits. We then identified a normative parameter space determined by the threshold and response variability data collected from neurologically intact participants. By plotting displacement detection performance within this normative space, stroke survivors with and without intact proprioception could be discriminated on a continuous scale that was sensitive to small performance variations, e.g. practice effects across days.ConclusionsThe proposed method uses robotic perturbations similar to those used in ongoing studies of motor function post-stroke. The approach is sensitive to small changes in the proprioceptive detection of hand motions. We expect this new robotic assessment will empower future studies to characterize how proprioceptive deficits compromise limb posture and movement control in stroke survivors.
To accurately estimate the state of the body, the nervous system needs to account for delays between signals from different sensory modalities. To investigate how such delays may be represented in the sensorimotor system, we asked human participants to play a virtual pong game in which the movement of the virtual paddle was delayed with respect to their hand movement. We tested the representation of this new mapping between the hand and the delayed paddle by examining transfer of adaptation to blind reaching and blind tracking tasks. These blind tasks enabled to capture the representation in feedforward mechanisms of movement control. A Time Representation of the delay is an estimation of the actual time lag between hand and paddle movements. A State Representation is a representation of delay using current state variables: the distance between the paddle and the ball originating from the delay may be considered as a spatial shift; the low sensitivity in the response of the paddle may be interpreted as a minifying gain; and the lag may be attributed to a mechanical resistance that influences paddle’s movement. We found that the effects of prolonged exposure to the delayed feedback transferred to blind reaching and tracking tasks and caused participants to exhibit hypermetric movements. These results, together with simulations of our representation models, suggest that delay is not represented based on time, but rather as a spatial gain change in visuomotor mapping.
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