The link between brain learning, attention, and consciousness

The processes whereby our brains continue to learn about a changing world in a stable fashion throughout life are proposed to lead to conscious experiences. These processes include the learning of top-down expectations, the matching of these expectations against bottom-up data, the focusing of attention upon the expected clusters of information, and the development of resonant states between bottom-up and top-down processes as they reach an attentive consensus between what is expected and what is there in the outside world. It is suggested that all conscious states in the brain are resonant states and that these resonant states trigger learning of sensory and cognitive representations. The models which summarize these concepts are therefore called Adaptive Resonance Theory, or ART, models. Psychophysical and neurobiological data in support of ART are presented from early vision, visual object recognition, auditory streaming, variable-rate speech perception, somatosensory perception, and cognitive-emotional interactions, among others. It is noted that ART mechanisms seem to be operative at all levels of the visual system, and it is proposed how these mechanisms are realized by known laminar circuits of visual cortex. It is predicted that the same circuit realization of ART mechanisms will be found in the laminar circuits of all sensory and cognitive neocortex. Concepts and data are summarized concerning how some visual percepts may be visibly, or modally, perceived, whereas amodal percepts may be consciously recognized even though they are perceptually invisible. It is also suggested that sensory and cognitive processing in the What processing stream of the brain obey top-down matching and learning laws that are often complementary to those used for spatial and motor processing in the brain's Where processing stream. This enables our sensory and cognitive representations to maintain their stability as we learn more about the world, while allowing spatial and motor representations to forget learned maps and gains that are no longer appropriate as our bodies develop and grow from infanthood to adulthood. Procedural memories are proposed to be unconscious because the inhibitory matching process that supports these spatial and motor processes cannot lead to resonance.     

Augmented visual, auditory, haptic, and multimodal feedback in motor learning: A review

It is generally accepted that augmented feedback, provided by a human expert or a technical display, effectively enhances motor learning. However, discussion of the way to most effectively provide augmented feedback has been controversial. Related studies have focused primarily on simple or artificial tasks enhanced by visual feedback. Recently, technical advances have made it possible also to investigate more complex, realistic motor tasks and to implement not only visual, but also auditory, haptic, or multimodal augmented feedback. The aim of this review is to address the potential of augmented unimodal and multimodal feedback in the framework of motor learning theories. The review addresses the reasons for the different impacts of feedback strategies within or between the visual, auditory, and haptic modalities and the challenges that need to be overcome to provide appropriate feedback in these modalities, either in isolation or in combination. Accordingly, the design criteria for successful visual, auditory, haptic, and multimodal feedback are elaborated.     

The functional role of cross-frequency coupling

Recent studies suggest that cross-frequency coupling (CFC) might play a functional role in neuronal computation, communication and learning. In particular, the strength of phase-amplitude CFC differs across brain areas in a task-relevant manner, changes quickly in response to sensory, motor and cognitive events, and correlates with performance in learning tasks. Importantly, whereas high-frequency brain activity reflects local domains of cortical processing, low-frequency brain rhythms are dynamically entrained across distributed brain regions by both external sensory input and internal cognitive events. CFC might thus serve as a mechanism to transfer information from large-scale brain networks operating at behavioral timescales to the fast, local cortical processing required for effective computation and synaptic modification, thus integrating functional systems across multiple spatiotemporal scales.     

Dissociation of sensory facilitation and decision bias in statistical learning

ABSTRACT

The detection of regularities in the sensory environment, known as statistical learning, is an important brain function that has been observed in many experimental contexts. In these experiments, statistical learning of patterned sensory stimulation leads to improvements in the speed and/or accuracy with which subsequent stimuli are recognized. That is, statistical learning facilitates the transformation of sensory stimuli into motor responses, but the mechanism by which this occurs is unclear. Statistical learning could improve the efficiency of sensory processing, or it could bias responses toward particular outcomes. The distinction is important, as these different hypotheses imply different functions and different neural substrates for statistical learning. Here we address this problem by studying statistical learning as a decision-making process, which allows us to leverage the extensive computational literature on this topic. Specifically we describe a method for applying the Diffusion Decision Model (DDM) to isolate different sensory and cognitive processes associated with decision-making. The results indicate that statistical learning improves performance on a visual learning task in two distinct ways: by altering the efficiency of sensory processing and by introducing biases in the decision-making process. By fitting the parameters of the DDM to data from individual subjects, we find that the prominence of these two factors differed substantially across the population, and that these differences were predictive of individual performance on the psychophysical task. Overall, these results indicate that different cognitive processes can be recruited by statistical learning, and that the DDM is a powerful framework for detecting these influences.     

Different types of environmental enrichment have discrepant effects on spatial memory and synaptophysin levels in female mice

Environmental enrichment paradigms that incorporate cognitive stimulation, exercise, and motor learning benefit memory and synaptic plasticity across the rodent lifespan. However, the contribution each individual element of the enriched environment makes to enhancing memory and synaptic plasticity has yet to be delineated. Therefore, the current study tested the effects of three of these elements on memory and synaptic protein levels. Young female C57BL/6 mice were given 3h of daily exposure to either rodent toys (cognitive stimulation) or running wheels (exercise), or daily acrobatic training for 6 weeks prior to and throughout behavioral testing. Controls were group housed, but did not receive enrichment. Spatial working and reference memory were tested in a water-escape motivated radial arm maze. Levels of the presynaptic protein synaptophysin were then measured in frontoparietal cortex, hippocampus, striatum, and cerebellum. Exercise, but not cognitive stimulation or acrobat training, improved spatial working memory relative to controls, despite the fact that both exercise and cognitive stimulation increased synaptophysin levels in the neocortex and hippocampus. These data suggest that exercise alone is sufficient to improve working memory, and that enrichment-induced increases in synaptophysin levels may not be sufficient to improve working memory in young females. Spatial reference memory was unaffected by enrichment. Acrobat training had no effect on memory or synaptophysin levels, suggesting a minimal contribution of motor learning to the mnemonic and neuronal benefits of enrichment. These results provide the first evidence that different elements of the enriched environment have markedly distinct effects on spatial memory and synaptic alterations.     

Introduction to the special section on theory and data in categorization: Integrating computational, behavioral, and cognitive neuroscience approaches

This special section brings together behavioral, computational, mathematical, and neuroimaging approaches to understand the processes underlying category learning. Over the past decade, there has been growing convergence in research on categorization, with computational-mathematical models influencing the interpretation of brain imaging and neuropsychological data, and with cognitive neuroscience findings influencing the development and refinement of models. Classic debates between single-system and multiple-memory-system theories have become more nuanced and focused. Multiple brain areas and cognitive processes contribute to categorization, but theories differ markedly in whether and when those neurocognitive components are recruited for different aspects of categorization. The articles in this special section approach this issue from several diverse angles.     

基于非侵入性脑刺激的认知增强：方法、伦理和应用

作为人类追求卓越的方式之一,基于非侵入性脑刺激的认知增强成为众多学科和公众关注的问题。首先阐述了非侵入性脑刺激的两种主要技术手段(经颅磁刺激和经颅直流刺激)的技术原理及其在提升健康个体认知功能上的应用,并分析了这两种技术可能带来的安全、自主选择、公平等伦理问题,最后总结了该认知增强技术在体育和军事等两个具体领域的应用。未来可进一步提高技术手段的深部脑区刺激能力及该增强技术的持续和真实效果。     

Exploration in outbred mice covaries with general learning abilities irrespective of stress reactivity, emotionality, and physical attributes

Across multiple learning tasks (that place different sensory, motor, and information processing demands on the animals), we have found that the performance of mice is commonly regulated by a single factor ("general learning") that accounts for 30-40% of the variance across individuals and tasks. Furthermore, individuals' general learning abilities were highly correlated with their propensity to engage in exploration in an open field, a behavior that is potentially stress-inducing. This relationship between exploration in the open field and general learning abilities suggests the possibility that variations in stress sensitivity/responsivity or related emotional responses might directly influence individuals' general learning abilities. Here, the relationship of sensory/motor skills and stress sensitivity/emotionality to animals' general learning abilities were assessed. Outbred (CD-1) mice were tested in a battery of six learning tasks as well as 21 tests of exploratory behavior, sensory/motor function and fitness, emotionality, and stress reactivity. The performances of individual mice were correlated across six learning tasks, and the performance measures of all learning tasks loaded heavily on a single factor (principal component analysis), accounting for 32% of the variability between animals and tasks. Open field exploration and seven additional exploratory behaviors (including those exhibited in an elevated plus maze) also loaded heavily on this same factor, although general activity, sensory/motor responses, physical characteristics, and direct measures of fear did not. In a separate experiment, serum corticosterone levels of mice were elevated in response to a mild environmental stressor (confinement on an elevated platform). Stress-induced corticosterone levels were correlated with behavioral fear responses, but were unsystematically related to individuals' propensity for exploration. In total, these results suggest that although general learning abilities are strongly related to individuals' propensity for exploration, this relationship is not attributable to variations in sensory/motor function or the individuals' physiological or behavioral sensitivity to conditions that promote stress or fear.     

Modeling developmental cognitive neuroscience

In the past few years connectionist models have greatly contributed to formulating theories of cognitive development. Some of these models follow the approach of developmental cognitive neuroscience in exploring interactions between brain development and cognitive development by integrating structural change into learning. We describe two classes of these models. The first focuses on experience-dependent structural elaboration within a brain region by adding or deleting units and connections during learning. The second models the gradual integration of different brain areas based on combinations of experience-dependent and maturational factors. These models provide new theories of the mechanisms of cognitive change in various domains and they offer an integrated framework to study normal and abnormal development, and normal and impaired adult processing.     

Schema theory: a critical appraisal and reevaluation

The authors critically review a number of the constructs and associated predictions proposed in schema theory (R. A. Schmidt, 1975). The authors propose that new control and learning theories should include a reformulated (a) notion of a generalized motor program that is not based on motor program but still accounts for the strong tendency for responses to maintain their relative characteristics; (b) mechanism or processes whereby an abstract movement structure based on proportional principles (e.g., relative timing, relative force) is developed through practice; and (c) explanation for parameter learning that accounts for the benefits of parameter variability but also considers how variability is scheduled. Furthermore, they also propose that new theories of motor learning must be able to account for the consistent findings spawned as a result of the schema theory proposal and must not be simply discounted because of some disfavor with the motor program notion, in general, or schema theory, more specifically.     

A bottom-up view of toddler word learning

A head camera was used to examine the visual correlates of object name learning by toddlers as they played with novel objects and as the parent spontaneously named those objects. The toddlers’ learning of the object names was tested after play, and the visual properties of the head camera images during naming events associated with learned and unlearned object names were analyzed. Naming events associated with learning had a clear visual signature, one in which the visual information itself was clean and visual competition among objects was minimized. Moreover, for learned object names, the visual advantage of the named target over competitors was sustained, both before and after the heard name. The findings are discussed in terms of the visual and cognitive processes that may depend on clean sensory input for learning and also on the sensory–motor, cognitive, and social processes that may create these optimal visual moments for learning.     

The Effect of Erroneous Knowledge of Results on Skill Acquisition when Augmented Information is Redundant

Even though it can be shown that verbal knowledge of results (KR) is redundant with sensory feedback for learning certain motor skills, such findings do not eliminate the possibility that when KR is available it influences underlying learning processes. In order to examine the function of KR more closely, two experiments were designed in which the subjects received conflicting information about their own sensory feedback and the KR presented by the experimenter. In Experiment 1, two erroneous-KR groups, a correct-KR group, and a no-KR group performed 150 practice trials on a simple anticipation timing task and then performed three no-KR retention tests of 30 trials each following intervals of 10 minutes, 1 week, and 1 month. The results supported previous findings that providing correct KR is redundant in anticipation tasks. However, learning was influenced by KR as subjects performed according to the erroneous KR information, thereby ignoring their sensory feedback even after a 1-month interval. In Experiment 2, subjects practised a more complex striking response for the anticipation task for 75 trials and then performed no-KR retention trials either immediately, or 1 day or 1 week later. One of the groups received erroneous KR after 50 practice trials with correct KR. The results confirmed and extended those from Experiment 1, as erroneous KR, even after initial practice with correct KR, influenced retention performance. These results indicate that although KR provides information that is not needed to learn anticipation timing skills, this augmented verbal information is a dominant source of information that influences underlying cognitive processes involved in learning motor skills.     

Solving the "human problem": the frontal feedback model

This paper argues that humans possess unique cognitive abilities due to the presence of a functional system that exists in the human brain that is absent in the non-human brain. This system, the frontal feedback system, was born in the hominin brain when the great phylogenetic expansion of the prefrontal cortex relative to posterior sensory regions surpassed a critical threshold. Surpassing that threshold effectively reversed the preferred direction of information flow in the highest association regions of the neocortex, producing the frontal feedback system. This reversal was from the caudo-rostral bias characteristic of non-human, or pre-human, brain dynamics to a rostro-caudal bias characteristic of modern human brain dynamics. The frontal feedback system works through frontal motor routines, or action schemes, manipulating the release and reconstruction of stored sensory memories in posterior sensory areas. As an obligatory feature of frontal feedback, a central character, or self, emerges within this cortical network that manifests itself as agent in these reconstructions as well as in the experience of sensory perceptions. Dynamical-systems modeling of cortical interactions is combined in the paper with recent neuroimaging studies of "resting-state" brain activity to bridge the gap between microscopic and macroscopic levels of cortical behavior. This synthesis is used to support the proposal of an information flow reversal occurring in the hominin brain and also to explain how such a reversal generates the wide variety of cognitive and experiential phenomena that many consider to be uniquely human.     

Terminal Feedback Outperforms Concurrent Visual,Auditory, and Haptic Feedback in Learning a Complex Rowing-Type Task

Augmented feedback, provided by coaches or displays, is a well-established strategy to accelerate motor learning. Frequent terminal feedback and concurrent feedback have been shown to be detrimental for simple motor task learning but supportive for complex motor task learning. However, conclusions on optimal feedback strategies have been mainly drawn from studies on artificial laboratory tasks with visual feedback only. Therefore, the authors compared the effectiveness of learning a complex, 3-dimensional rowing-type task with either concurrent visual, auditory, or haptic feedback to self-controlled terminal visual feedback. Results revealed that terminal visual feedback was most effective because it emphasized the internalization of task-relevant aspects. In contrast, concurrent feedback fostered the correction of task-irrelevant errors, which hindered learning. The concurrent visual and haptic feedback group performed much better during training with the feedback than in nonfeedback trials. Auditory feedback based on sonification of the movement error was not practical for training the 3-dimensional movement for most participants. Concurrent multimodal feedback in combination with terminal feedback may be most effective, especially if the feedback strategy is adapted to individual preferences and skill level.     

Conscious awareness of motor fluidity improves performance and decreases cognitive effort in sequence learning

Motor skill learning is improved when participants are instructed to judge after each trial whether their performed movements have reached maximal fluidity. Consequently, the conscious awareness of this maximal fluidity can be classified as a genuine learning factor for motor sequences. However, it is unknown whether this effect of conscious awareness on motor learning could be mediated by the increased cognitive effort that may accompany such judgment making. The main aim of this study was to test this hypothesis in comparing two groups with, and without, the conscious awareness of the maximal fluidity. To assess the possible involvement of cognitive effort, we have recorded the pupillary dilation to the task, which is well-known to increase in proportion to cognitive effort. Results confirmed that conscious awareness indeed improved motor sequence learning of the trained sequence specifically. Pupil dilation was smaller during trained than during novel sequence performance, indicating that sequence learning decreased the cognitive cost of sequence execution. However, we found that in the group that had to judge on their maximal fluidity, pupil dilation during sequence production was smaller than in the control group, indicating that the motor improvement induced by the fluidity judgment does not involve additional cognitive effort. We discuss these results in the context of motor learning and cognitive effort theories.     

Conscious thought as simulation of behaviour and perception

A 'simulation' theory of cognitive function can be based on three assumptions about brain function. First, behaviour can be simulated by activating motor structures, as during an overt action but suppressing its execution. Second, perception can be simulated by internal activation of sensory cortex, as during normal perception of external stimuli. Third, both overt and covert actions can elicit perceptual simulation of their normal consequences. A large body of evidence supports these assumptions. It is argued that the simulation approach can explain the relations between motor, sensory and cognitive functions and the appearance of an inner world.     

Learning by observation and learning by doing in Down and Williams syndromes

New skills may be learned by active experience (experiential learning or learning by doing) or by observation of others’ experience (learning by observation). In general, learning by observation reduces the time and the attempts needed to learn complex actions and behaviors. The present research aimed to compare learning by observation and learning by doing in two clinical populations with different etiology of intellectual disability (ID), as individuals with Down syndrome (DS) and individuals with Williams syndrome (WS), with the hypothesis that specific profiles of learning may be found in each syndrome. To this end, we used a mixture of new and existing data to compare the performances of 24 individuals with DS, 24 individuals with WS and 24 typically developing children on computerized tasks of learning by observation or learning by doing. The main result was that the two groups with ID exhibited distinct patterns of learning by observation. Thus, individuals with DS were impaired in reproducing the previously observed visuo‐motor sequence, while they were as efficient as TD children in the experiential learning task. On the other hand, individuals with WS benefited from the observational training while they were severely impaired in detecting the visuo‐motor sequence in the experiential learning task (when presented first). The present findings reinforce the syndrome‐specific hypothesis and the view of ID as a variety of conditions in which some cognitive functions are more disrupted than others because of the differences in genetic profile and brain morphology and functionality. These findings have important implications for clinicians, who should take into account the genetic etiology of ID in developing learning programs for treatment and education.     

Visuomotor Adaptation and Proprioceptive Recalibration

ABSTRACT

Motor learning, in particular motor adaptation, is driven by information from multiple senses. For example, when arm control is faulty, vision, touch, and proprioception can all report on the arm's movements and help guide the adjustments necessary for correcting motor error. In recent years we have learned a lot about how the brain integrates information from multiple senses for the purpose of perception. However, less is known about how multisensory data guide motor learning. Most models of, and studies on, motor learning focus almost exclusively on the ensuing changes in motor performance without exploring the implications on sensory plasticity. Nor do they consider how discrepancies in sensory information (e.g., vision and proprioception) related to hand position may affect motor learning. Here, we discuss research from our lab and others that shows how motor learning paradigms affect proprioceptive estimates of hand position, and how even the mere discrepancy between visual and proprioceptive feedback can affect learning and plasticity. Our results suggest that sensorimotor learning mechanisms do not exclusively rely on motor plasticity and motor memory, and that sensory plasticity, in particular proprioceptive recalibration, plays a unique and important role in motor learning.