Hippocampal and neocortical contributions to memory: advances in the complementary learning systems framework

The complementary learning systems framework provides a simple set of principles, derived from converging biological, psychological and computational constraints, for understanding the differential contributions of the neocortex and hippocampus to learning and memory. The central principles are that the neocortex has a low learning rate and uses overlapping distributed representations to extract the general statistical structure of the environment, whereas the hippocampus learns rapidly using separated representations to encode the details of specific events while minimizing interference. In recent years, we have instantiated these principles in working computational models, and have used these models to address human and animal learning and memory findings, across a wide range of domains and paradigms. Here, we review a few representative applications of our models, focusing on two domains: recognition memory and animal learning in the fear-conditioning paradigm. In both domains, the models have generated novel predictions that have been tested and confirmed.     

Linking working memory and long-term memory: a computational model of the learning of new words

The nonword repetition (NWR) test has been shown to be a good predictor of children's vocabulary size. NWR performance has been explained using phonological working memory, which is seen as a critical component in the learning of new words. However, no detailed specification of the link between phonological working memory and long-term memory (LTM) has been proposed. In this paper, we present a computational model of children's vocabulary acquisition (EPAM-VOC) that specifies how phonological working memory and LTM interact. The model learns phoneme sequences, which are stored in LTM and mediate how much information can be held in working memory. The model's behaviour is compared with that of children in a new study of NWR, conducted in order to ensure the same nonword stimuli and methodology across ages. EPAM-VOC shows a pattern of results similar to that of children: performance is better for shorter nonwords and for wordlike nonwords, and performance improves with age. EPAM-VOC also simulates the superior performance for single consonant nonwords over clustered consonant nonwords found in previous NWR studies. EPAM-VOC provides a simple and elegant computational account of some of the key processes involved in the learning of new words: it specifies how phonological working memory and LTM interact; makes testable predictions; and suggests that developmental changes in NWR performance may reflect differences in the amount of information that has been encoded in LTM rather than developmental changes in working memory capacity.     

工作记忆训练对认知功能和大脑神经系统的影响

工作记忆训练成为近年来提升个体认知绩效的一种有效方式。工作记忆训练主要是指采用工作记忆广度任务、刷新任务以及各种复杂工作记忆任务在计算机上以循序渐进的方式进行训练。近年来的研究发现, 工作记忆训练能提升工作记忆、流体智力、抑制、注意、阅读和数学等认知功能。神经机制的研究发现：工作记忆训练引起大脑额-顶区域激活减弱, 而皮层下结构包括纹状体和尾状核区域的激活增强; 工作记忆训练减少了大脑灰质的数量, 增强了大脑白质的功能连通性; 工作记忆训练引起尾状核上多巴胺受体的变化。未来的研究需要在研究设计、被试人群、研究手段上进一步确认和扩展工作记忆训练的有效性和内在认知神经机制。     

Modeling individual differences in working memory performance: a source activation account

Working memory resources are needed for processing and maintenance of information during cognitive tasks. Many models have been developed to capture the effects of limited working memory resources on performance. However, most of these models do not account for the finding that different individuals show different sensitivities to working memory demands, and none of the models predicts individual subjects' patterns of performance. We propose a computational model that accounts for differences in working memory capacity in terms of a quantity called source activation , which is used to maintain goal-relevant information in an available state. We apply this model to capture the working memory effects of individual subjects at a fine level of detail across two experiments. This, we argue, strengthens the interpretation of source activation as working memory capacity.     

Learning representations in a gated prefrontal cortex model of dynamic task switching

The prefrontal cortex is widely believed to play an important role in facilitating people's ability to switch performance between different tasks. We present a biologically‐based computational model of prefrontal cortex (PFC) that explains its role in task switching in terms of the greater flexibility conferred by activation‐based working memory representations in PFC, as compared with more slowly adapting weight‐based memory mechanisms. Specifically we show that PFC representations can be rapidly updated when a task switches via a dynamic gating mechanism based on a temporal‐differences reward‐prediction learning mechanism. Unlike prior models of this type, the present model develops all of its internal representations via learning mechanisms as shaped by the demands of continuous periodic task switching. This advance opens up a new domain of research into the interactions between working memory task demands and the representations that develop to meet them. Results on a version of the Wisconsin card sorting task are presented for the full model and a number of comparison networks that test the importance of various model features. Furthermore, we show that a lesioned model produces perseverative errors like those seen in frontal patients.     

Neural correlates of learning and working memory in the primate posterior parietal cortex

The posterior parietal cortex has been traditionally associated with coordinate transformations necessary for interaction with the environment and with visual-spatial attention. More recently, involvement of posterior parietal cortex in other cognitive functions such as working memory and task learning has become evident. Neurophysiological experiments in non-human primates and human imaging studies have revealed neural correlates of memory and learning at the single neuron and at the brain network level. During working memory, posterior parietal neurons continue to discharge and to represent stimuli that are no longer present. This activation resembles the responses of prefrontal neurons, although important differences have been identified in terms of the ability to resist stimulation by distracting stimuli, which is more evident in the prefrontal than the posterior parietal cortex. Posterior parietal neurons also become active during tasks that require the organization of information into larger structured elements and their activity is modulated according to learned context-dependent rules. Neural correlates of learning can be observed in the mean discharge rate and spectral power of neuronal spike trains after training to perform new task sets or rules. These findings demonstrate the importance of posterior parietal cortex in brain networks mediating working memory and learning.     

A capacity theory of comprehension: individual differences in working memory.

A theory of the way working memory capacity constrains comprehension is proposed. The theory proposes that both processing and storage are mediated by activation and that the total amount of activation available in working memory varies among individuals. Individual differences in working memory capacity for language can account for qualitative and quantitative differences among college-age adults in several aspects of language comprehension. One aspect is syntactic modularity: The larger capacity of some individuals permits interaction among syntactic and pragmatic information, so that their syntactic processes are not informationally encapsulated. Another aspect is syntactic ambiguity: The larger capacity of some individuals permits them to maintain multiple interpretations. The theory is instantiated as a production system model in which the amount of activation available to the model affects how it adapts to the transient computational and storage demands that occur in comprehension.     

语文学习困难儿童的工作记忆与加工速度

加工速度和工作记忆反应了不同的认知加工过程,在认知发展研究中,加工速度和工作记忆所起的作用仍存在较大的分歧.采用多因素混合实验设计,在严格控制条件下,比较了语文学习困难和控制组儿童的工作记忆和加工速度.结果发现,与控制组相比,语文学习困难儿童在工作记忆和加工速度方面均存在明显的不足,但加工速度不能解释不同能力组之间的差异,语文学习困难儿童的缺陷在于工作记忆能力的下降.工作记忆的缺陷在于言语工作记忆和中央执行功能的不足,与视空间工作记忆能力无关.语文学习困难既存在一般的工作记忆缺陷(中央执行功能)也存在特定的工作记忆(言语工作记忆)能力的不足.     

Neural Representations of Nondeclarative Memories

ABSTRACT— Nondeclarative learning refers to abilities characterized by a lack of awareness of what has been learned and an independence from medial temporal lobe structures that support conscious memories of facts and events. Neuroimaging approaches have been used extensively in two domains of nondeclarative learning—priming and skill learning—to investigate the neural substrates supporting performance. Recent neuroimaging studies have attempted to understand what is being learned in different tasks in order to inform psychological theories of nondeclarative memory. For example, priming may be considered a form of perceptual learning or a form of stimulus–response learning, and correlations between performance and activation patterns in different regions may suggest the nature of the brain changes that support behavior. The attainment of expertise in a skill has been characterized as greater efficiency of processing in the same neural structures that support novice performance or, alternatively, as the recruitment of additional regions. Current research suggests that, within the domains of priming and skill learning, there is much heterogeneity in the underlying brain representations and psychological theories will need to account for these variations.     

The effects of concurrent verbal and visual tasks on category learning

Current theories of category learning posit separate verbal and nonverbal learning systems. Past research suggests that the verbal system relies on verbal working memory and executive functioning and learns rule-defined categories; the nonverbal system does not rely on verbal working memory and learns non-rule-defined categories (E. M. Waldron & F. G. Ashby, 2001; D. Zeithamova & W. T. Maddox, 2006). However, relatively little research has explored the importance of visual working memory or visual processing for either system. The authors investigated the role of working memory (Experiment 1a and 1b), visual processing (Experiment 2), and executive functioning for each system, using a concurrent task methodology. It was found that visual tasks with high executive functioning demands and verbal tasks with high or low executive demands disrupted rule-defined learning, whereas any visual task, regardless of executive functioning demand, disrupted non-rule-defined learning. Taken together, these results confirm the importance of verbal working memory and executive functioning for the verbal system and provide new evidence for the importance of visual processing for the nonverbal system. These results help to clarify understanding of the nonverbal system and have implications for multiple systems theories of category learning (F. G. Ashby, L. A. Alfonso-Reese, A. U. Turken, & E. M. Waldron, 1998).     

Role of Verbal Working Memory in Complex Skill Acquisition

Although research has shown that general cognitive ability is related to performance during the initial stage of skill acquisition, empirical evidence is needed to increase understanding of the primary abilities responsible for the relation. We propose that verbal working memory is related to initial skill acquisition performance based on cognitive activities people enact when learning complex skills. After completing a general cognitive ability measure and a test of verbal working memory, 120 undergraduate students performed a complex computer simulation. The working memory test accounted for a meaningful amount of performance variance after statistically controlling for the general cognitive ability measure. These data suggest that tasks depending heavily on the activation of multiple knowledge structures may be understood in terms of individual differences in working memory.     

The role of interference control in working memory: A study with children at risk of ADHD

The study aimed to test whether the impairment in a working memory task observed in children at risk of ADHD was due to a lack of control of interfering information being processed whilst carrying out the memory task. Two groups of children at risk of ADHD with or without a learning disability (reading impairment) were compared to a control group in a working memory task. Activation of irrelevant items was tested with a lexical decision task presented immediately after the final recall in about half of the trials. Results showed a poor working memory performance in children at risk of ADHD and reading disability associated with a larger activation of irrelevant information than that of control children. Results indicated that the to-be-excluded and interfering items are still highly accessible to working memory in children that fail the working memory task. The examination of working memory and interference control of children at risk of ADHD with a learning disability revealed a counterintuitive picture of children with poor working memory showing better recall/activation of processed information. This picture is consistent with a view of working memory related to an efficient inhibitory control that influences cognitive functioning.     

The role of interference control in working memory: a study with children at risk of ADHD

The study aimed to test whether the impairment in a working memory task observed in children at risk of ADHD was due to a lack of control of interfering information being processed whilst carrying out the memory task. Two groups of children at risk of ADHD with or without a learning disability (reading impairment) were compared to a control group in a working memory task. Activation of irrelevant items was tested with a lexical decision task presented immediately after the final recall in about half of the trials. Results showed a poor working memory performance in children at risk of ADHD and reading disability associated with a larger activation of irrelevant information than that of control children. Results indicated that the to-be-excluded and interfering items are still highly accessible to working memory in children that fail the working memory task. The examination of working memory and interference control of children at risk of ADHD with a learning disability revealed a counterintuitive picture of children with poor working memory showing better recall/activation of processed information. This picture is consistent with a view of working memory related to an efficient inhibitory control that influences cognitive functioning.     

嵌套范式下视空工作记忆对基于规则类别学习

基于COVIS模型与认知加工阶段假设,通过2个实验探讨嵌套范式下, 视空工作记忆对基于规则类别学习的影响。实验1采用类别学习中嵌套视空工作记忆的范式,结果发现视空工作记忆削弱基于规则类别学习成绩,与COVIS模型预测相一致。实验2则采用视空工作记忆中嵌套类别学习任务的范式,结果却发现视空工作记忆对基于规则类别学习的影响消失。实验结果表明嵌套范式下视空工作记忆的位置影响基于规则类别学习,初步验证了类别学习存在多个认知加工阶段的假设,视空工作记忆主要影响基于规则类别学习中规则的发现和检验阶段。     

An account of the relationship between fluid intelligence and complex learning in considering storage capacity and executive attention

Although fluid intelligence and complex learning are conceptualized differently and assessed by apparently different measures, both theoretical accounts and empirical evidence suggest a relationship between the two constructs. In this study, major working memory aspects including the storage capacity and executive attention were proposed to account for the relationship between fluid intelligence and complex learning. A sample of 184 participants completed fluid intelligence and complex learning scales, as well as working memory measures that each included two or three treatment levels differing in the demands on capacity or executive control. The differences among the treatment levels provided a favorable precondition for employing fixed-links models to separate the core processes of storage capacity or executive attention from the auxiliary processes. Results indicated that both storage capacity and executive attention contributed significantly to fluid intelligence and complex learning. A further analysis showed that the two working memory aspects, particularly the storage capacity, accounted for most of the shared variance between fluid intelligence and complex learning.     

Consciousness and working memory: Current trends and research perspectives

Working memory has long been thought to be closely related to consciousness. However, recent empirical studies show that unconscious content may be maintained within working memory and that complex cognitive computations may be performed on-line. This promotes research on the exact relationships between consciousness and working memory. Current evidence for working memory being a conscious as well as an unconscious process is reviewed. Consciousness is shown to be considered a subset of working memory by major current theories of working memory. Evidence for unconscious elements in working memory is shown to come from visual masking and attentional blink paradigms, and from the studies of implicit working memory. It is concluded that more research is needed to explicate the relationship between consciousness and working memory. Future research directions regarding the relationship between consciousness and working memory are discussed.     

Computational principles of working memory in sentence comprehension

Understanding a sentence requires a working memory of the partial products of comprehension, so that linguistic relations between temporally distal parts of the sentence can be rapidly computed. We describe an emerging theoretical framework for this working memory system that incorporates several independently motivated principles of memory: a sharply limited attentional focus, rapid retrieval of item (but not order) information subject to interference from similar items, and activation decay (forgetting over time). A computational model embodying these principles provides an explanation of the functional capacities and severe limitations of human processing, as well as accounts of reading times. The broad implication is that the detailed nature of cross-linguistic sentence processing emerges from the interaction of general principles of human memory with the specialized task of language comprehension.     

工作记忆训练及对数学能力的迁移作用

工作记忆缺陷会影响个体数学能力发展。通过记忆策略、广度、刷新、转换等功能的工作记忆训练,可以改善个体认知功能。然而,工作记忆训练对个体的阅读、数学、流体智力等方面的远迁移效果并不一致。研究表明,工作记忆训练可以改善数感、视觉空间能力、推理能力等数学一般技能;也会通过改善语音工作记忆以及空间能力促进数学计算能力,或者通过改善中央执行系统,提升数学问题表征、模式识别、解题迁移、策略选择等复杂的过程,从而促进数学问题解决能力。因此,区分不同数学任务的认知过程,可以获得工作记忆训练对数学能力迁移效果的进一步证据。今后,神经影像学的证据或许也是未来工作记忆训练对数学能力提高的又一佐证。     

Irrelevant sensory stimuli interfere with working memory storage: Evidence from a computational model of prefrontal neurons

The encoding of irrelevant stimuli into the memory store has previously been suggested as a mechanism of interference in working memory (e.g., Lange & Oberauer, Memory, 13, 333–339, 2005; Nairne, Memory & Cognition, 18, 251–269, 1990). Recently, Bancroft and Servos (Experimental Brain Research, 208, 529–532, 2011) used a tactile working memory task to provide experimental evidence that irrelevant stimuli were, in fact, encoded into working memory. In the present study, we replicated Bancroft and Servos’s experimental findings using a biologically based computational model of prefrontal neurons, providing a neurocomputational model of overwriting in working memory. Furthermore, our modeling results show that inhibition acts to protect the contents of working memory, and they suggest a need for further experimental research into the capacity of vibrotactile working memory.     

工作记忆容量与儿童算术认知加工

工作记忆容量的差别是个体认知差异的一个重要方面。工作记忆容量在多种运算规则下与运算技能、概念性知识等因素交织在一起共同对儿童的算术认知加工产生影响,并且它是数学学习不良儿童学习困难的一个重要影响因素。同时,儿童的工作记忆容量本身也受到多种因素的制约,进而影响儿童的算术认知成绩。