Stimulus-Specific Delay Activity in Human Primary Visual Cortex

ABSTRACT— Working memory (WM) involves maintaining information in an on-line state. One emerging view is that information in WM is maintained via sensory recruitment , such that information is stored via sustained activity in the sensory areas that encode the to-be-remembered information. Using functional magnetic resonance imaging, we observed that key sensory regions such as primary visual cortex (V1) showed little evidence of sustained increases in mean activation during a WM delay period, though such amplitude increases have typically been used to determine whether a region is involved in on-line maintenance. However, a multivoxel pattern analysis of delay-period activity revealed a sustained pattern of activation in V1 that represented only the intentionally stored feature of a multifeature object. Moreover, the pattern of delay activity was qualitatively similar to that observed during the discrimination of sensory stimuli, suggesting that WM representations in V1 are reasonable "copies" of those evoked during pure sensory processing.     

Keeping track of where we are: Spatial working memory in navigation

Spatial working memory (WM) seems to include two types of spatial information: locations and relations. However, this distinction has been based on small-scale tasks. Here, we used a virtual navigation paradigm to examine whether WM for locations and relations applies to the large-scale spatial world. We found that navigators who successfully learned two routes and also integrated them were superior at maintaining multiple locations and multiple relations in WM. However, over the entire spectrum of navigators, WM for spatial relations, but not locations, was specifically predictive of route integration performance. These results lend further support to the distinction between these two forms of spatial WM and point to their critical role in individual differences in navigation proficiency.     

Evidence for working memory storage operations in perceptual cortex

Isolating the short-term storage component of working memory (WM) from the myriad of associated executive processes has been an enduring challenge. Recent efforts have identified patterns of activity in visual regions that contain information about items being held in WM. However, it remains unclear (1) whether these representations withstand intervening sensory input and (2) how communication between multimodal association cortex and the unimodal perceptual regions supporting WM representations is involved in WM storage. We present evidence that the features of a face held in WM are stored within face-processing regions, that these representations persist across subsequent sensory input, and that information about the match between sensory input and a memory representation is relayed forward from perceptual to prefrontal regions. Participants were presented with a series of probe faces and indicated whether each probe matched a target face held in WM. We parametrically varied the feature similarity between the probe and target faces. Activity within face-processing regions scaled linearly with the degree of feature similarity between the probe face and the features of the target face, suggesting that the features of the target face were stored in these regions. Furthermore, directed connectivity measures revealed that the direction of information flow that was optimal for performance was from sensory regions that stored the features of the target face to dorsal prefrontal regions, supporting the notion that sensory input is compared to representations stored within perceptual regions and is subsequently relayed forward. Together, these findings indicate that WM storage operations are carried out within perceptual cortex.     

Assessing the concreteness of relational representation

Research has shown that people's ability to transfer abstract relational knowledge across situations can be heavily influenced by the concrete objects that fill relational roles. This article provides evidence that the concreteness of the relations themselves also affects performance. In 3 experiments, participants viewed simple relational patterns of visual objects and then identified these same patterns under a variety of physical transformations. Results show that people have difficulty generalizing to novel concrete forms of abstract relations, even when objects are unchanged. This suggests that stimuli are initially represented as concrete relations by default. In the 2nd and 3rd experiments, the number of distinct concrete relations in the training set was increased to promote more abstract representation. Transfer improved for novel concrete relations but not for other transformations such as object substitution. Results indicate that instead of automatically learning abstract relations, people's relational representations preserve all properties that appear consistently in the learning environment, including concrete objects and concrete relations.     

Functional connectivity during working memory maintenance

Neurophysiological experiments with monkeys have demonstrated that working memory (WM) is associated with persistent neural activity in multiple brain regions, such as the prefrontal cortex (PFC), the parietal cortex, and posterior unimodal association areas. WM maintenance is believed to require the coordination of these brain regions, which do not function in isolation but, rather, interact to maintain visual percepts that are no longer present in the environment. However, single-unit physiology studies and traditional univariate analyses of functional brain imaging data cannot evaluate interactions between distant brain regions, and so evidence of regional integration during WM maintenance is largely indirect. In this study, we utilized a recently developed multivariate analysis method that allows us to explore functional connectivity between brain regions during the distinct stages of a delayed face recognition task. To characterize the neural network mediating the on-line maintenance of faces, the fusiform face area (FFA) was defined as a seed and was then used to generate whole-brain correlation maps. A random effects analysis of the correlation data revealed a network of brain regions exhibiting significant correlations with the FFA seed during the WM delay period. This maintenance network included the dorsolateral and ventrolateral PFC, the premotor cortex, the intraparietal sulcus, the caudate nucleus, the thalamus, the hippocampus, and occipitotemporal regions. These findings support the notion that the coordinated functional interaction between nodes of a widely distributed network underlies the active maintenance of a perceptual representation.     

Domain-dependent activation during spatial and nonspatial auditory working memory

Visual system has been proposed to be divided into two, the ventral and dorsal, processing streams. The ventral pathway is thought to be involved in object identification whereas the dorsal pathway processes information regarding the spatial locations of objects and the spatial relationships among objects. Several studies on working memory (WM) processing have further suggested that there is a dissociable domain-dependent functional organization within the prefrontal cortex for processing of spatial and nonspatial visual information. Also the auditory system is proposed to be organized into two domain-specific processing streams, similar to that seen in the visual system. Recent studies on auditory WM have further suggested that maintenance of nonspatial and spatial auditory information activates a distributed neural network including temporal, parietal, and frontal regions but the magnitude of activation within these activated areas shows a different functional topography depending on the type of information being maintained. The dorsal prefrontal cortex, specifically an area of the superior frontal sulcus (SFS), has been shown to exhibit greater activity for spatial than for nonspatial auditory tasks. Conversely, ventral frontal regions have been shown to be more recruited by nonspatial than by spatial auditory tasks. It has also been shown that the magnitude of this dissociation is dependent on the cognitive operations required during WM processing. Moreover, there is evidence that within the nonspatial domain in the ventral prefrontal cortex, there is an across-modality dissociation during maintenance of visual and auditory information. Taken together, human neuroimaging results on both visual and auditory sensory systems support the idea that the prefrontal cortex is organized according to the type of information being maintained in WM.     

Minds, persons, and space: an fMRI investigation into the relational complexity of higher-order intentionality

Mental state reasoning or theory-of-mind has been the subject of a rich body of imaging research. Although such investigations routinely tap a common set of regions, the precise function of each area remains a contentious matter. With the help of functional magnetic resonance imaging (fMRI), we sought to determine which areas are involved when processing mental state or intentional metarepresentations by focusing on the relational aspect of such representations. Using non-intentional relational representations such as spatial relations between persons and between objects as a contrast, the results ascertained the involvement of the precuneus, the temporal poles, and the medial prefrontal cortex in the processing of intentional representations. In contrast, the anterior superior temporal sulcus and the left temporo-parietal junction were implicated when processing representations that refer to the presence of persons in relational contexts in general. The right temporo-parietal junction, however, was specifically activated for persons entering spatial relations. The level of representational complexity, a previously unexplored factor, was also found to modulate the neural response in some brain regions, such as the medial prefrontal cortex and the right temporo-parietal junction. These findings highlight the need to take into account the critical roles played by an extensive network of neural regions during mental state reasoning.     

Language, perception, and the schematic representation of spatial relations

Schemas are abstract nonverbal representations that parsimoniously depict spatial relations. Despite their ubiquitous use in maps and diagrams, little is known about their neural instantiation. We sought to determine the extent to which schematic representations are neurally distinguished from language on the one hand, and from rich perceptual representations on the other. In patients with either left hemisphere damage or right hemisphere damage, a battery of matching tasks depicting categorical spatial relations was used to probe for the comprehension of basic spatial concepts across distinct representational formats (words, pictures, and schemas). Left hemisphere patients underperformed right hemisphere patients across all tasks. However, focused residual analyses using voxel-based lesion-symptom mapping (VLSM) suggest that (1) left hemisphere deficits in the representation of categorical spatial relations are difficult to distinguish from deficits in naming these relations and (2) the right hemisphere plays a special role in extracting schematic representations from richly textured pictures.     

Spatial relational memory in 9-month-old macaque monkeys

This experiment assesses spatial and nonspatial relational memory in freely moving 9-mo-old and adult (11-13-yr-old) macaque monkeys (Macaca mulatta). We tested the use of proximal landmarks, two different objects placed at the center of an open-field arena, as conditional cues allowing monkeys to predict the location of food rewards hidden in one of two sets of three distinct locations. Monkeys were tested in two different conditions: (1) when local visual cues marked the two sets of potentially baited locations, so that monkeys could use both local and spatial information to discriminate these locations from never-baited locations; and (2) when no local visual cues marked the two sets of potentially baited locations, so that monkeys had to rely on a spatial relational representation of the environment to discriminate these locations. No 9-mo-old or adult monkey associated the presence of the proximal landmarks, at the center of the arena, with the presence of food in one set of three distinct locations. All monkeys, however, discriminated the potentially baited locations in the presence of local visual cues, thus providing evidence of visual discrimination learning. More importantly, all 9-mo-old monkeys tested discriminated the potentially baited locations in absence of the local visual cues, thus exhibiting evidence of spatial relational learning. These findings indicate that spatial memory processes characterized by a relational representation of the environment are present as early as 9 mo of age in macaque monkeys.     

Development of an abstract category representation for the spatial relation between in 6- to 10-month-old infants

Nine experiments examined the formation of an abstract category representation for the spatial relation between by 6- to 10-month-old infants. The experiments determined that 9- to 10-month-olds, but not 6- to 7-month-olds, could form an abstract category representation for between when performing in an object-variation version of the between categorization task. The results also demonstrated that 6- to 7-month-olds could form category representations for between in the object-variation version of the between categorization task but that such representations were specific to the particular objects presented. The evidence confirms that representations for different spatial relations emerge at different points during development, and suggests that each representation undergoes its own period of development from concrete to abstract.     

Flexible working memory representation of the relationship between an object and its location as revealed by interactions with attention

Working memory (WM) selectively maintains a limited amount of currently relevant information in an active state to influence future perceptual processing, thought, and behavior. The representation of the information held in WM is unknown, particularly the degree of separation between the representation of an object’s identity and its location. The present experiments examined the flexibility of object and location WM representations by measuring reaction times on a visual discrimination task during the delay period of a WM recognition task for object identities, locations, or both. The results demonstrate that during WM delay periods, attention is biased toward information that matches the current contents of WM. Attention is not biased toward information that was present in the encoded memory sample but not relevant for the memory recognition test. This specificity of the interaction between WM and attention applies to both the identity and the location of the remembered stimulus and to the relationship between an object and its location. The results suggest that when this relationship is necessary for task performance, WM represents an object and its identity in an integrated manner. However, if this relationship is not task relevant, the object and location information are represented in WM separately.     

Rehearsal in Spatial Working Memory: Evidence From Neuroimaging

A variety of biological evidence has identified a frontal-parietal circuit underlying spatial working memory for visual stimuli. But the question remains, how do these neural regions accomplish the goal of maintaining location information on-line? We tested the hypothesis that the active rehearsal of spatial information in working memory is accomplished by means of focal shifts of spatial selective attention to memorized locations. Spatial selective attention has been shown to cause changes in the early visual processing of stimuli that appear in attended locations. Thus, the hypothesis of attention-based rehearsal predicts similar modulations of visual processing at memorized locations. We used functional magnetic resonance imaging to observe posterior visual activations during the performance of a spatial working memory task. In line with the hypothesis, spatial rehearsal led to enhanced activation in the early visual areas contralateral to the memorized locations.     

Infants lost in (peripersonal) space?

A significant challenge in developing spatial representations for the control of action is one of multisensory integration. Specifically, we require an ability to efficiently integrate sensory information arriving from multiple modalities pertaining to the relationships between the acting limbs and the nearby external world (i.e. peripersonal space), across changes in body posture and limb position. Evidence concerning the early development of such spatial representations points towards the independent emergence of two distinct mechanisms of multisensory integration. The earlier-developing mechanism achieves spatial correspondence by representing body parts in their typical or default locations, and the later-developing mechanism does so by dynamically remapping the representation of the position of the limbs with respect to external space in response to changes in postural information arriving from proprioception and vision.     

Binding serial order to representations in working memory: a spatial/verbal dissociation

Verbal information is coded naturally as ordered representations in working memory (WM). However, this may not be true for spatial information. Accordingly, we used memory span tasks to test the hypothesis that serial order is more readily bound to verbal than to spatial representations. Removing serial-order requirements improved performance more for spatial locations than for digits. Furthermore, serial order was freely reproduced twice as frequently for digits as for locations. When participants reordered spatial sequences, they minimized the mean distance between items. Participants also failed to detect changes in serial order more frequently for spatial than for verbal sequences. These results provide converging evidence for a dissociation in the binding of serial order to spatial versus verbal representations. There may be separable domain-specific control processes responsible for this binding. Alternatively, there may be fundamental differences in how effectively temporal information can be bound to different types of stimulus features in WM.     

The role of body-based sensory information in the acquisition of enduring spatial representations

Although many previous studies have shown that body-based sensory modalities such as vestibular, kinesthetic, and efferent information are useful for acquiring spatial information about one's immediate environment, relatively little work has examined how these modalities affect the acquisition of long-term spatial memory. Three groups of participants learned locations along a 146 m indoor route, and subsequently pointed to these locations, estimated distances between them, and constructed maps of the environment. One group had access to visual, proprioceptive, and inertial information, another had access to matched visual and matched inertial information, and another had access only to matched visual information. In contrast to previous findings examining transient, online spatial representations, our results showed very few differences among groups in the accuracy of the spatial memories acquired. The only difference was the improved pointing accuracy of participants who had access to proprioceptive information relative to that of participants in the other conditions. Results are discussed in terms of differential sensory contributions to transient and enduring spatial representations.     

Working memory as internal attention: Toward an integrative account of internal and external selection processes

Working memory (WM) and attention have been studied as separate cognitive constructs, although it has long been acknowledged that attention plays an important role in controlling the activation, maintenance, and manipulation of representations in WM. WM has, conversely, been thought of as a means of maintaining representations to voluntarily guide perceptual selective attention. It has more recently been observed, however, that the contents of WM can capture visual attention, even when such internally maintained representations are irrelevant, and often disruptive, to the immediate external task. Thus, the precise relationship between WM and attention remains unclear, but it appears that they may bidirectionally impact one another, whether or not internal representations are consistent with the external perceptual goals. This reciprocal relationship seems, further, to be constrained by limited cognitive resources to handle demands in either maintenance or selection. We propose here that the close relationship between WM and attention may be best described as a give-and-take interdependence between attention directed toward either actively maintained internal representations (traditionally considered WM) or external perceptual stimuli (traditionally considered selective attention), underpinned by their shared reliance on a common cognitive resource. Put simply, we argue that WM and attention should no longer be considered as separate systems or concepts, but as competing and influencing one another because they rely on the same limited resource. This framework can offer an explanation for the capture of visual attention by irrelevant WM contents, as well as a straightforward account of the underspecified relationship between WM and attention.     

Bayesian analogy with relational transformations

How can humans acquire relational representations that enable analogical inference and other forms of high-level reasoning? Using comparative relations as a model domain, we explore the possibility that bottom-up learning mechanisms applied to objects coded as feature vectors can yield representations of relations sufficient to solve analogy problems. We introduce Bayesian analogy with relational transformations (BART) and apply the model to the task of learning first-order comparative relations (e.g., larger, smaller, fiercer, meeker) from a set of animal pairs. Inputs are coded by vectors of continuous-valued features, based either on human magnitude ratings, normed feature ratings (De Deyne et al., 2008), or outputs of the topics model (Griffiths, Steyvers, & Tenenbaum, 2007). Bootstrapping from empirical priors, the model is able to induce first-order relations represented as probabilistic weight distributions, even when given positive examples only. These learned representations allow classification of novel instantiations of the relations and yield a symbolic distance effect of the sort obtained with both humans and other primates. BART then transforms its learned weight distributions by importance-guided mapping, thereby placing distinct dimensions into correspondence. These transformed representations allow BART to reliably solve 4-term analogies (e.g., larger:smaller::fiercer:meeker), a type of reasoning that is arguably specific to humans. Our results provide a proof-of-concept that structured analogies can be solved with representations induced from unstructured feature vectors by mechanisms that operate in a largely bottom-up fashion. We discuss potential implications for algorithmic and neural models of relational thinking, as well as for the evolution of abstract thought.     

无关长时记忆表征能否引导视觉注意选择?

与任务相关的长时记忆表征在引导视觉注意选择的过程中扮演着重要的角色,它可以使人们在熟悉的视觉情境中快速搜索目标刺激,并偏离干扰刺激。但当长时记忆表征与任务无关时,还能否引导视觉注意选择?目前还不清楚。实验1采用眼动追踪技术直接比较无关工作记忆表征与无关长时记忆表征在视觉搜索阶段对视觉注意的捕获效应,行为反应时与首次注视点百分率的结果都发现,当无关工作记忆表征在视觉搜索中再次出现时能引导视觉注意偏向到与之匹配的干扰刺激,但无关长时记忆表征并没有表现出类似的注意引导效应;实验2探讨记忆表征由工作记忆系统转移到长时记忆系统的过程中对视觉注意的引导效应,结果发现,随着记忆表征的转移,注意引导效应消失了,实验3排除工作记忆表征的干扰后,依然没有发现无关长时记忆表征对注意的引导效应。以上结果表明,无关长时记忆表征并不能像工作记忆表征一样引导视觉注意选择,工作记忆表征和长时记忆表征对视觉注意的引导属于两个不同的认知过程。     

抽象概念表征的具身认知观

抽象概念是否通过感知经验来表征以及如何被感知经验表征是具身认知面临的一大问题.在抽象概念表征是否具有感知经验基础的问题上,具身认知理论认为抽象概念通过情境模拟或隐喻与感知经验发生联系.在抽象概念如何与感知经验表征发生联系的问题上,概念模拟理论强调情景或运动模拟在抽象概念表征中的直接作用;概念隐喻理论则侧重具体经验或具体经验与抽象概念之间的共同结构关系在抽象概念表征中的间接作用.未来研究应改变概念表征的稳定的心理实体观,从语言和抽象表征的关系、正常儿童和特殊群体的抽象概念表征差异入手,整合不同的具身认知观点.     

Roles of egocentric and allocentric spatial representations in locomotion and reorientation

Four experiments investigated the nature of spatial representations used in locomotion. Participants learned the layout of several objects and then pointed to the objects while blindfolded in 3 conditions: before turning (baseline), after turning to a new heading (updating), and after disorientation (disorientation). The internal consistency of pointing in the disorientation condition was relatively high and equivalent to that in the baseline and updating conditions, when the layout had salient intrinsic axes and the participants learned the locations of the objects on the periphery of the layout. The internal consistency of pointing was disrupted by disorientation when participants learned the locations of objects while standing amid them and the layout did not have salient intrinsic axes. It was also observed that many participants retrieved spatial relations after disorientation from the original learning heading. These results indicate that people form an allocentric representation of object-to-object spatial relations when they learn the layout of a novel environment and use that representation to locate objects around them. Egocentric representations may be used to locate objects when allocentric representations are not of high fidelity.