It is well established clinically that rhythmic auditory cues can improve gait and other motor behaviors in Parkinson's disease (PD) and other disorders. However, the neural systems underlying this therapeutic effect are largely unknown. To investigate this question we scanned people with PD and age‐matched healthy controls using functional magnetic resonance imaging (fMRI). All subjects performed a rhythmic motor behavior (right hand finger tapping) with and without simultaneous auditory rhythmic cues at two different speeds (1 and 4 Hz). We used spatial independent component analysis (ICA) and regression to identify task‐related functional connectivity networks and assessed differences between groups in intra‐ and inter‐network connectivity. Overall, the control group showed greater intra‐network connectivity in perceptual and motor related networks during motor tapping both with and without rhythmic cues. The PD group showed greater inter‐network connectivity between the auditory network and the executive control network, and between the executive control network and the motor/cerebellar network associated with the motor task performance. We interpret our results as indicating that the temporal rhythmic auditory information may assist compensatory mechanisms through network‐level effects, reflected in increased interaction between auditory and executive networks that in turn modulate activity in cortico‐cerebellar networks.
Concept learning, the ability to extract commonalities and highlight distinctions across a set of related experiences to build organized knowledge, is a critical aspect of cognition. Previous reviews have focused on concept learning research as a means for dissociating multiple brain systems. The current review surveys recent work that uses novel analytical approaches, including the combination of computational modeling with neural measures, focused on testing theories of specific computations and representations that contribute to concept learning. We discuss in detail the roles of the hippocampus, ventromedial prefrontal, lateral prefrontal, and lateral parietal cortices, and how their engagement is modulated by the coherence of experiences and the current learning goals. We conclude that the interaction of multiple brain systems relating to learning, memory, attention, perception, and reward support a flexible concept-learning mechanism that adapts to a range of category structures and incorporates motivational states, making concept learning a fruitful research domain for understanding the neural dynamics underlying complex behaviors.
Effective generalization in a multiple-category situation involves both assessing potential membership in individual categories and resolving conflict between categories while implementing a decision bound. We separated generalization from decision bound implementation using an information integration task in which category exemplars varied over two incommensurable feature dimensions. Human subjects first learned to categorize stimuli within limited training regions, and then, during fMRI scanning, they also categorized transfer stimuli from new regions of perceptual space. Transfer stimuli differed both in distance from the training region prototype and distance from the decision bound, allowing us to independently assess neural systems sensitive to each. Across all stimulus regions, categorization was associated with activity in the extrastriate visual cortex, basal ganglia, and the bilateral intraparietal sulcus. Categorizing stimuli near the decision bound was associated with recruitment of the frontoinsular cortex and medial frontal cortex, regions often associated with conflict and which commonly coactivate within the salience network. Generalization was measured in terms of greater distance from the decision bound and greater distance from the category prototype (average training region stimulus). Distance from the decision bound was associated with activity in the superior parietal lobe, lingual gyri, and anterior hippocampus, whereas distance from the prototype was associated with left intraparietal sulcus activity. The results are interpreted as supporting the existence of different uncertainty resolution mechanisms for uncertainty about category membership (representational uncertainty) and uncertainty about decision bound (decisional uncertainty).
We identified dynamic changes in recruitment of neural connectivity networks across three phases of a flexible rule learning and set-shifting task similar to the Wisconsin Card Sort Task: switching, rule learning via hypothesis testing, and rule application. During fMRI scanning, subjects viewed pairs of stimuli that differed across four dimensions (letter, color, size, screen location), chose one stimulus, and received feedback. Subjects were informed that the correct choice was determined by a simple unidimensional rule, for example “choose the blue letter.” Once each rule had been learned and correctly applied for 4-7 trials, subjects were cued via either negative feedback or visual cues to switch to learning a new rule. Task performance was divided into three phases: Switching (first trial after receiving the switch cue), hypothesis testing (subsequent trials through the last error trial), and rule application (correct responding after the rule was learned). We used both univariate analysis to characterize activity occurring within specific regions of the brain, and a multivariate method, constrained principal component analysis for fMRI (fMRI-CPCA), to investigate how distributed regions coordinate to subserve different processes. As hypothesized, switching was subserved by a limbic network including the ventral striatum, thalamus, and parahippocampal gyrus, in conjunction with cortical salience network regions including the anterior cingulate and frontoinsular cortex. Activity in the ventral striatum was associated with switching regardless of how switching was cued; visually cued shifts were associated with additional visual cortical activity. After switching, as subjects moved into the hypothesis testing phase, a broad fronto-parietal-striatal network (associated with the cognitive control, dorsal attention, and salience networks) increased in activity. This network was sensitive to rule learning speed, with greater extended activity for the slowest learning speed late in the time course of learning. As subjects shifted from hypothesis testing to rule application, activity in this network decreased and activity in the somatomotor and default mode networks increased.
Categorization and memory for specific items are fundamental processes that allow us to apply knowledge to novel stimuli. This study directly compares categorization and memory using delay match to category (DMC) and delay match to sample (DMS) tasks. In DMC participants view and categorize a stimulus, maintain the category across a delay, and at the probe phase view another stimulus and indicate whether it is in the same category or not. In DMS, a standard item working memory task, participants encode and maintain a specific individual item, and at probe decide if the stimulus is an exact match or not. Constrained Principal Components Analysis was used to identify and compare activity within neural networks associated with these tasks, and we relate these networks to those that have been identified with resting state-fMRI. We found that two frontoparietal networks of particular interest. The first network included regions associated with the dorsal attention network and frontoparietal salience network; this network showed patterns of activity consistent with a role in rapid orienting to and processing of complex stimuli. The second uniquely involved regions of the frontoparietal central-executive network; this network responded more slowly following each stimulus and showed a pattern of activity consistent with a general role in role in decision-making across tasks. Additional components were identified that were associated with visual, somatomotor and default mode networks.
Contemporary models of categorization typically tend to sidestep the problem of how information is initially encoded during decision-making. Instead, a focus of this work has been to investigate how, through selective attention, stimulus representations are contorted such that behaviourally-relevant dimensions are accentuated (or "stretched"), and representations of irrelevant dimensions are ignored (or "compressed"). In high-dimensional real-world environments, it is computationally infeasible to sample all available information, and human decision-makers selectively sample information from sources expected to provide relevant information. To address these and other shortcomings, we develop an active sampling model, Sampling Emergent Attention (SEA), which sequentially and strategically samples information sources until the expected cost of information exceeds the expected benefit. The model specifies the interplay of two components, one involved in determining the expected utility of different information sources and the other in representing knowledge and beliefs about the environment. These two components interact such that knowledge of the world guides information sampling, and what is sampled updates knowledge. Like human decision-makers, the model displays strategic sampling behaviour, such as terminating information search when sufficient information has been sampled and adaptively adjusting the search path in response to previously sampled information. The model also shows human-like failure modes. For example, when information exploitation is prioritized over exploration, the bidirectional influences between information-sampling and learning can lead to the development of beliefs that systematically differ from reality.
Through selective attention, decision-makers can learn to ignore behaviorally irrelevant stimulus dimensions. This can improve learning and increase the perceptual discriminability of relevant stimulus information. Across cognitive models of categorization, this is typically accomplished through the inclusion of attentional parameters, which provide information about the importance assigned to each stimulus dimension by each participant. The effect of these parameters on psychological representation is often described geometrically, such that perceptual differences over relevant psychological dimensions are accentuated (or stretched), and differences over irrelevant dimensions are down-weighted (or compressed). In sensory and association cortex, representations of stimulus features are known to covary with their behavioral relevance. Although this implies that neural representational space might closely resemble that hypothesized by formal categorization theory, to date, attentional effects in the brain have been demonstrated through powerful experimental manipulations (e.g., contrasts between relevant and irrelevant features). This approach sidesteps the role of idiosyncratic conceptual knowledge in guiding attention to useful information sources. To bridge this divide, we used formal categorization models, which were fit to behavioral data, to make inferences about the concepts and strategies used by individual participants during decision-making. We found that when greater attentional weight was devoted to a particular visual feature (e.g., “color”), its value (e.g., “red”) was more accurately decoded from occipitotemporal cortex. We also found that this effect was sufficiently sensitive to reflect individual differences in conceptual knowledge, indicating that occipitotemporal stimulus representations are embedded within a space closely resembling that formalized by classic categorization theory.
Through its connections with widespread cortical areas and with dopaminergic midbrain areas, the basal ganglia are well situated to integrate patterns of cortical input with the dopaminergic reward signal originating in the midbrain. In this review, we consider the functions of the basal ganglia in relation to its gross and cellular anatomy, and discuss how these mechanisms subserve the thresholding and selection of motor and cognitive processes. We also discuss how the dopaminergic reward signal enables flexible task learning through modulation of striatal plasticity, and how reinforcement learning models have been used to account for various aspects of basal ganglia activity. Specifically, we will discuss the important role of the basal ganglia in instrumental learning, cognitive control, sequence learning, and categorization tasks. Finally, we will discuss the neurobiological and cognitive characteristics of Parkinson's disease, Huntington's disease and addiction to illustrate the relationship between the basal ganglia and cognitive function. WIREs Cogn Sci 2013, 4:135-148. doi: 10.1002/wcs.1217 For further resources related to this article, please visit the WIREs website.
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