Not "just" a coincidence: frontal-striatal interactions in working memory and interval timing

The frontal cortex and basal ganglia play central roles in working memory and in the ability to time brief intervals. We outline recent theoretical and empirical work to suggest that working memory and interval timing rely not only on the same anatomic structures, but also on the same neural representation of a specific stimulus. Specifically, cortical neurons may fire in an oscillatory fashion to form representations of stimuli, and the striatum (a basal ganglia structure) may detect those patterns of cortical firing that occur co-incident to important events. Information about stimulus identity can be extracted from which cortical neurons are involved in the representation, and information about duration can be extracted from their relative phase. The principles derived from these biologically based models also fit well with a family of behaviourally based models that emphasise the importance of time in many working memory phenomena.     

Not “just” a coincidence: Frontal‐striatal interactions in working memory and interval timing

The frontal cortex and basal ganglia play central roles in working memory and in the ability to time brief intervals. We outline recent theoretical and empirical work to suggest that working memory and interval timing rely not only on the same anatomic structures, but also on the same neural representation of a specific stimulus. Specifically, cortical neurons may fire in an oscillatory fashion to form representations of stimuli, and the striatum (a basal ganglia structure) may detect those patterns of cortical firing that occur co‐incident to important events. Information about stimulus identity can be extracted from which cortical neurons are involved in the representation, and information about duration can be extracted from their relative phase. The principles derived from these biologically based models also fit well with a family of behaviourally based models that emphasise the importance of time in many working memory phenomena.     

工作记忆对时间加工的影响

非时间信息加工与时间加工的关系是复杂的, 存在双向干扰或单向干扰, 结果的不一致与任务所需的注意资源及工作记忆都有关。工作记忆的中央执行系统在时间加工中起主要作用, 同时进行的非时间任务对中央执行功能的需求越多, 两种任务之间的干扰程度越大。语言环和视空间模板对时间加工的影响与非时间信息的类型有关。工作记忆作为一个整体以工作记忆容量为指标, 计时成绩也表现出与工作记忆容量有关的年龄、智力等方面的差异。时间加工和工作记忆在额叶、顶叶、基底神经节等皮质存在共同的神经机制。未来应该丰富工作记忆的研究内容, 结合时间加工的分段性探讨工作记忆影响时间加工的具体进程及神经机制, 并力求在应用方面取得一定的成果。     

Working memory modulates the perception of time

Recent research has indicated that reentrant feedback from the contents of working memory can enhance neural representations and the perceptual strengths of matching stimuli in the visual field. However, whether the contents of working memory can also distort conscious experiences of perception remains unclear. Our present results show that the durations of perceptual stimuli matching the nontemporal representations in working memory tend to be perceived as longer than those of mismatching stimuli. This is the first demonstration that working memory can lead to distortions of time perception. Our findings are consistent with the ideas that the perceived duration of a stimulus depends on the magnitude of the neural responses to that stimulus in visual cortex and that there is a common system for representing both temporal and nontemporal magnitudes. We conclude that top-down modulation from the nontemporal contents of working memory distorts the perceptual experience of temporal duration.     

时间认知神经科学研究进展

当前对时间认知的脑机制探讨有三个模型：特异化计时模型、分布网络模型和定域计时模型。在这些模型的框架下,时间认知的神经心理学研究集中探讨了小脑、基底神经节、前额叶在时间信息加工中的作用和大脑两半球在时间认知中的不对称性。小脑作为内部计时系统对时间控制具有重要作用,在周期性动作任务中,小脑对不连续动作计时具有特异性。基底神经节在时间加工任务中与小脑存在明显的作用分离,其具体机制还有待深入研究。前额叶的计时功能可能与注意和工作记忆对时间信息的获得、维持和组织有关。此外,还发现大脑右半球与时问信息的加工关系密切。     

Dedicated and intrinsic models of time perception

Two general frameworks have been articulated to describe how the passage of time is perceived. One emphasizes that the judgment of the duration of a stimulus depends on the operation of dedicated neural mechanisms specialized for representing the temporal relationships between events. Alternatively, the representation of duration could be ubiquitous, arising from the intrinsic dynamics of nondedicated neural mechanisms. In such models, duration might be encoded directly through the amount of activation of sensory processes or as spatial patterns of activity in a network of neurons. Although intrinsic models are neurally plausible, we highlight several issues that must be addressed before we dispense with models of duration perception that are based on dedicated processes.     

事件相关振荡与振荡脑网络

事件相关振荡是伴随认知、情感和行为过程的脑电磁振荡活动,观察到其各类调频、调幅和调相现象,这种介观和宏观尺度上大量神经元的集体活动与微观尺度上神经元平均发放率和发放定时相互影响,共同参与神经信息的编码、表征、通讯和调控。动态细胞集群假说认为大脑认知功能是神经网络通过同步振荡相互作用的结果,在基于振荡的大脑理论指引下,多尺度、跨脑区和跨频率事件相关振荡研究为揭开振荡脑网络的工作原理带来了希望     

Perceived duration decreases with increasing eccentricity

Previous studies examining the influence of stimulus location on temporal perception yield inhomogeneous and contradicting results. Therefore, the aim of the present study is to soundly examine the effect of stimulus eccentricity. In a series of five experiments, subjects compared the duration of foveal disks to disks presented at different retinal eccentricities on the horizontal meridian. The results show that the perceived duration of a visual stimulus declines with increasing eccentricity. The effect was replicated with various stimulus orders (Experiments 1–3), as well as with cortically magnified stimuli (Experiments 4–5), ruling out that the effect was merely caused by different cortical representation sizes. The apparent decreasing duration of stimuli with increasing eccentricity is discussed with respect to current models of time perception, the possible influence of visual attention and respective underlying physiological characteristics of the visual system.     

Inertia and memory in ambiguous visual perception

Perceptual multistability during ambiguous visual perception is an important clue to neural dynamics. We examined perceptual switching during ambiguous depth perception using a Necker cube stimulus, and also during binocular rivalry. Analysis of perceptual switching time series using variance–sample size analysis, spectral analysis and time series shuffling shows that switching times behave as a 1/f noise and possess very long range correlations. The long memory feature contrasts sharply with the traditional satiation models of multistability, where the memory is not incorporated, as well as with recently published models of multistability and neural processing, where memory is excluded. On the other hand, the long memory feature favors the concept of “dynamic core” or coalition of neurons, where neurons form transient coalitions. Perceptual switching then corresponds to replacement of one coalition of neurons by another. The inertia and memory measures the stability of a coalition: a strong and stable coalition has to be won over by another similarly strong and stable coalition, resulting in long switching times. The complicated transient dynamics of competing coalitions of neurons may be addressable using a combination of functional imaging, measurement of frequency-tagged magnetoencephalography and frequency-tagged encephalography, simultaneous recordings of groups of neurons in many areas of the brain, and concepts from statistical mechanics and nonlinear dynamics theory.     

On the Correlation between Synchronized Oscillatory Activities and Consciousness

Recent experiments have shown that the amplitudes of cortical gamma band oscillatory activities that occur during anesthesia are often greater than amplitudes of similar activities that occur without anesthesia. This result is apparently at odds with the hypothesis that synchronized oscillatory activities constitute the neural correlate of consciousness. We argue that while synchronization and oscillatory patterning are necessary conditions for consciousness, they are not sufficient. Based on the results of a binocular rivalry study of Fries et al. (1997), we propose that the degrees of oscillatory strength and synchronization of neuronal activities determine the degree of awareness those activities produce. On the other hand, the overal firing rates of neurons in cortical sensory areas are not correlated with the degree of awareness the activities of those neurons produce. The results of the experiment of Fries et al. (1997) appear to conflict with the results of another binocular rivalry experiment, in which monkeys were trained to pull a lever in order to report which stimulus object was being perceived (Leopold & Logothetis, 1996). In the latter experiment, it was demonstrated that the firing rates of neurons in striate cortex did not change during perceptual alterations, while 90% of neurons in inferior and superior temporal cortices changed their firing rate when the perceived image changed. This result led to the conclusion that activities in temporal cortex are correlated with visual awareness, but those in striate cortex are not. We argue that activities in temporal cortex contribute little, if anything, to perceptual awareness, and that their primary function is computational. Thus the correlation between the firing rates of neurons in these areas and the responses of the monkeys is due to the recognition of a particular stimulus object, which in turn is due to the computations made there.     

Visual working memory contaminates perception

Indirect evidence suggests that the contents of visual working memory may be maintained within sensory areas early in the visual hierarchy. We tested this possibility using a well-studied motion repulsion phenomenon in which perception of one direction of motion is distorted when another direction of motion is viewed simultaneously. We found that observers misperceived the actual direction of motion of a single motion stimulus if, while viewing that stimulus, they were holding a different motion direction in visual working memory. Control experiments showed that none of a variety of alternative explanations could account for this repulsion effect induced by working memory. Our findings provide compelling evidence that visual working memory representations directly interact with the same neural mechanisms as those involved in processing basic sensory events.     

振动触觉频率信息的工作记忆容量及存储机制

工作记忆可以同时保存多个信息并且容量有限, 这一内在机制是工作记忆研究的重点问题。视觉和言语等研究领域都发现工作记忆能够存储多个信息单元, 但对振动触觉工作记忆是否能存储多个频率信息目前尚无相关研究。由于振动触觉频率刺激和视觉刺激具有不同的神经编码机制, 以及振动频率信息是通过躯体感觉产生的模拟的、单维的、参数化信息, 振动触觉工作记忆容量及其加工存储机制的研究也必不可少。首先, 本项目将采用新的实验范式, 探究不同的刺激呈现方式以及不同反应报告方式下, 振动触觉工作记忆的容量及其认知机制。其次, 本项目也将同时运用功能磁共振成像(fMRI)技术, 来阐述振动触觉工作记忆加工存储的神经机制。探究基于触觉频率信息的参数工作记忆容量及其神经机制是完善工作记忆模型的重要补充, 将有助于提高我们对工作记忆系统的理解, 并为视觉、听觉、触觉多模态感知觉信息的跨通道研究奠定基础。     

Working memory retention systems: a state of activated long-term memory

High temporal resolution event-related brain potential and electroencephalographic coherence studies of the neural substrate of short-term storage in working memory indicate that the sustained coactivation of both prefrontal cortex and the posterior cortical systems that participate in the initial perception and comprehension of the retained information are involved in its storage. These studies further show that short-term storage mechanisms involve an increase in neural synchrony between prefrontal cortex and posterior cortex and the enhanced activation of long-term memory representations of material held in short-term memory. This activation begins during the encoding/comprehension phase and evidently is prolonged into the retention phase by attentional drive from prefrontal cortex control systems. A parsimonious interpretation of these findings is that the long-term memory systems associated with the posterior cortical processors provide the necessary representational basis for working memory, with the property of short-term memory decay being primarily due to the posterior system. In this view, there is no reason to posit specialized neural systems whose functions are limited to those of short-term storage buffers. Prefrontal cortex provides the attentional pointer system for maintaining activation in the appropriate posterior processing systems. Short-term memory capacity and phenomena such as displacement of information in short-term memory are determined by limitations on the number of pointers that can be sustained by the prefrontal control systems.     

Remembering the time: a continuous clock

The neural mechanisms for time measurement are currently a subject of much debate. This article argues that our brains can measure time using the same dorsolateral prefrontal cells that are known to be involved in working memory. Evidence for this is: (1) the dorsolateral prefrontal cortex is integral to both cognitive timing and working memory; (2) both behavioural processes are modulated by dopamine and disrupted by manipulation of dopaminergic projections to the dorsolateral prefrontal cortex; (3) the neurons in question ramp their activity in a temporally predictable way during both types of processing; and (4) this ramping activity is modulated by dopamine. The dual involvement of these prefrontal neurons in working memory and cognitive timing supports a view of the prefrontal cortex as a multipurpose processor recruited by a wide variety of tasks.     

Visuospatial working memory influences the interaction between space and time

How do representations of space inform our perception of time? In language, spatial vocabulary is frequently used to describe temporal concepts, and spatial information biases temporal perception even in non-verbal tasks. In contrast, temporal information typically exerts little, if any, influence on the perception of spatial extent. Here, we used spatial and temporal reproduction tasks, both with and without verbal and spatial dual tasks, to investigate the mechanism underlying the asymmetric relation between space and time. Specifically, we tested whether the asymmetric interference between spatial and temporal stimulus attributes arises from interference in verbal or visuospatial working memory. We found that loading visuospatial working memory, but not verbal working memory, eliminated the asymmetric pattern of interference. This suggests that the interference between spatial and temporal representations arises due to processing constraints in visuospatial working memory. These findings are discussed in terms of the load theory of attention and the relative automaticity with which spatial and temporal information is processed.     

言语工作记忆的保持对时间知觉的影响*

采用工作记忆和时间判断的双任务范式,探讨了言语工作记忆的保持过程对时间知觉的影响。研究中操作记忆负荷和任务之间的时间间隔(ISI:记忆刺激消失到时间判断任务开始之间的时间间隔)。结果发现,记忆负荷越大,知觉到的时间越短,韦伯比例越低,反应时间越长;ISI越短,知觉到的时间变短,反应时间越长,这表明知觉时间和反应决策是一致的,不存在累加时间和反应判断之间的trade-off。但是记忆负荷和ISI的交互作用不显著。结果表明记忆负荷和ISI以不同的方式影响时间判断,言语工作记忆的保持和时间加工共享工作记忆资源。     

The time course of response inhibition in masked priming

In two experiments, we studied the temporal dynamics of the response time effects of masked visual prime stimuli, as a function of stimulus eccentricity and size. Experiment 1 factorially varied prime-target congruency, eccentricity, and mask-target stimulus onset asynchrony. Early facilitative and late inhibitory effects of congruency were observed at all eccentricities, with temporal dynamics modulated by eccentricity. To test whether this dependence on eccentricity is due to cortical magnification, Experiment 2 varied stimulus size as well. Response inhibition time courses were influenced by size and eccentricity jointly, with no discernible difference when stimuli were matched for cortical magnification. Analysis of the individual time course data revealed that the timescale of inhibition changes with the strength of the cortical representation of the prime stimulus. This imposes constraints on possible models.     

Timing the “magical number seven”: Presentation rate and regularity affect verbal working memory performance

The informative value of time and temporal structure often remains neglected in cognitive assessments. However, next to information about stimulus identity we can exploit temporal ordering principles, such as regularity, periodicity, or grouping to generate predictions about the timing of future events. Such predictions may improve cognitive performance by optimising adaptation to dynamic stimuli. Here, we investigated the influence of temporal structure on verbal working memory by assessing immediate recall performance for aurally presented digit sequences (forward digit span) as a function of standard (1000 ms stimulus-onset-asynchronies, SOAs), short (700 ms), long (1300 ms) and mixed (700–1300 ms) stimulus timing during the presentation phase. Participant's digit spans were lower for short and mixed SOA presentation relative to standard SOAs. This confirms an impact of temporal structure on the classic “magical number seven,” suggesting that working memory performance can in part be regulated through the systematic application of temporal ordering principles.     

Memory influences on hippocampal and striatal neural codes: effects of a shift between task rules

Interactions with neocortical memory systems may facilitate flexible information processing by hippocampus. We sought direct evidence for such memory influences by recording hippocampal neural responses to a change in cognitive strategy. Well-trained rats switched (within a single recording session) between the use of place and response strategies to solve a plus maze task. Maze and extramaze environments were constant throughout testing. Place fields demonstrated (in-field) firing rate and location-based reorganization [Leutgeb, S., Leutgeb, J. K., Barnes, C. A., Moser, E. I., McNaughton, B. L., & Moser, M. B. (2005). Independent codes for spatial and episodic memory in hippocampal neuronal ensembles. Science, 309, 619-623] after a task switch, suggesting that hippocampus encoded each phase of testing as a different context, or episode. The task switch also resulted in qualitative and quantitative changes to discharge that were correlated with an animal's velocity or acceleration of movement. Thus, the effects of a strategy switch extended beyond the spatial domain, and the movement correlates were not passive reflections of the current behavioral state. To determine whether hippocampal neural responses were unique, striatal place and movement-correlated neurons were simultaneously recorded with hippocampal neurons. Striatal place and movement cells exhibited a response profile that was similar, but not identical, to that observed for hippocampus after a strategy switch. Thus, retrieval of a different memory led both neural systems to represent a different context. However, hippocampus may play a special (though not exclusive) role in flexible spatial processing since correlated firing amongst cell pairs was highest when rats successfully switched between two spatial tasks. Correlated firing by striatal cell pairs increased following any strategy switch, supporting the view that striatum codes change in reinforcement contingencies.     

统计概要表征的内容与机制

人类能够快速提取集合刺激中的统计信息, 并形成以集中趋势和变异性为主要内容的统计概要表征。根据注意资源的使用, 统计概要表征的加工方式可分为分布式注意下的整体加工和聚焦注意下的个体加工。两种加工在工作记忆中相互影响, 共同构建了对集合刺激的表征。统计概要表征适用于多种刺激类型, 发生在低水平和高水平加工中, 因而可能是一种普遍的知觉加工方式。未来研究可关注统计概要表征的神经基础及其对决策的影响。