Information In Visual Short-term Memory Research Articles

Capacity and precision in an animal model of visual short-term memory

Temporary storage of information in visual short-term memory (VSTM) is a key component of many complex cognitive abilities. However, it is highly limited in capacity. Understanding the neurophysiological nature of this capacity limit will require a valid animal model of VSTM. We used a multiple-item color change detection task to measure macaque monkeys' VSTM capacity. Subjects' performance deteriorated and reaction times increased as a function of the number of items in memory. Additionally, we measured the precision of the memory representations by varying the distance between sample and test colors. In trials with similar sample and test colors, subjects made more errors compared to trials with highly discriminable colors. We modeled the error distribution as a Gaussian function and used this to estimate the precision of VSTM representations. We found that as the number of items in memory increases the precision of the representations decreases dramatically. Additionally, we found that focusing attention on one of the objects increases the precision with which that object is stored and degrades the precision of the remaining. These results are in line with recent findings in human psychophysics and provide a solid foundation for understanding the neurophysiological nature of the capacity limit of VSTM.

Topographic Contribution of Early Visual Cortex to Short-Term Memory Consolidation: A Transcranial Magnetic Stimulation Study

The neural correlates for retention of visual information in visual short-term memory are considered separate from those of sensory encoding. However, recent findings suggest that sensory areas may play a role also in short-term memory. We investigated the functional relevance, spatial specificity, and temporal characteristics of human early visual cortex in the consolidation of capacity-limited topographic visual memory using transcranial magnetic stimulation (TMS). Topographically specific TMS pulses were delivered over lateralized occipital cortex at 100, 200, or 400 ms into the retention phase of a modified change detection task with low or high memory loads. For the high but not the low memory load, we found decreased memory performance for memory trials in the visual field contralateral, but not ipsilateral to the side of TMS, when pulses were delivered at 200 ms into the retention interval. A behavioral version of the TMS experiment, in which a distractor stimulus (memory mask) replaced the TMS pulses, further corroborated these findings. Our findings suggest that retinotopic visual cortex contributes to the short-term consolidation of topographic visual memory during early stages of the retention of visual information. Further, TMS-induced interference decreased the strength (amplitude) of the memory representation, which most strongly affected the high memory load trials.

Splitting attention across the two visual fields in visual short-term memory

Humans have the ability to attentionally select the most relevant visual information from their extrapersonal world and to retain it in a temporary buffer, known as visual short-term memory (VSTM). Research suggests that at least two non-contiguous items can be selected simultaneously when they are distributed across the two visual hemifields. In two experiments, we show that attention can also be split between the left and right sides of internal representations held in VSTM. Participants were asked to remember several colors, while cues presented during the delay instructed them to orient their attention to a subset of memorized colors. Experiment 1 revealed that orienting attention to one or two colors strengthened equally participants’ memory for those colors, but only when they were from separate hemifields. Experiment 2 showed that in the absence of attentional cues the distribution of the items in the visual field per se had no effect on memory. These findings strongly suggest the existence of independent attentional resources in the two hemifields for selecting and/or consolidating information in VSTM.

The impact of extrafoveal information on visual short-term memory for object position

The role of extrafoveal information in visual short-term memory has been investigated relatively little, and, in most existing studies, using verbalisable stimuli susceptible to the recruitment of long-term memory (LTM). In addition, little is known about the impact of extrafoveal information available pre- and posttarget foveation, as it is typical to provide extrafoveal information prior to the foveation of memory targets. In this study, two object-position recognition experiments were conducted (each with two conditions) to establish the impact of extrafoveal information provided before and after the foveation of memory targets. Stimuli comprised 1/f noise discs that minimised the recruitment of LTM by eliminating verbal and semantic cues. Overall, a greater hit rate was found where extrafoveal information was available; however, performance analyses in which extrafoveal information was considered relative to the temporal lag at which target stimuli were foveated reveals both costs and benefits. A beneficial effect arose only where extrafoveal information was provided after the target had been foveated, but not prior to target foveation. Findings are discussed in terms of recency and extrafoveal perception effects, incorporating a postfoveation object-file refresh mechanism.

Contralateral delay activity tracks object identity information in visual short term memory

Previous studies suggested that ERP component contralateral delay activity (CDA) tracks the number of objects containing identity information stored in visual short term memory (VSTM). Later MEG and fMRI studies implied that its neural source lays in superior IPS. However, since the memorized stimuli in previous studies were displayed in distinct spatial locations, hence possibly CDA tracks the object-location information instead. Moreover, a recent study implied the activation in superior IPS reflected the location load. The current research thus explored whether CDA tracks the object-location load or the object-identity load, and its neural sources. Participants were asked to remember one color, four identical colors or four distinct colors. The four-identical-color condition was the critical one because it contains the same amount of identity information as that of one color while the same amount of location information as that of four distinct colors. To ensure the participants indeed selected four colors in the four-identical-color condition, we also split the participants into two groups (low- vs. high-capacity), analyzed late positive component (LPC) in the prefrontal area, and collected participant's subjective-report. Our results revealed that most of the participants selected four identical colors. Moreover, regardless of capacity-group, there was no difference on CDA between one color and four identical colors yet both were lower than 4 distinct colors. Besides, the source of CDA was located in the superior parietal lobule, which is very close to the superior IPS. These results support the statement that CDA tracks the object identity information in VSTM.

Mental rotation impairs attention shifting and short-term memory encoding: Neurophysiological evidence against the response-selection bottleneck model of dual-task performance

Dual tasks and their associated delays have often been used to examine the boundaries of processing in the brain. We used the dual-task procedure and recorded event-related potentials (ERPs) to investigate how mental rotation of a first stimulus (S1) influences the shifting of visual-spatial attention to a second stimulus (S2). Visual-spatial attention was monitored by using the N2pc component of the ERP. In addition, we examined the sustained posterior contralateral negativity (SPCN) believed to index the retention of information in visual short-term memory. We found modulations of both the N2pc and the SPCN, suggesting that engaging mechanisms of mental rotation impairs the deployment of visual-spatial attention and delays the passage of a representation of S2 into visual short-term memory. Both results suggest interactions between mental rotation and visual-spatial attention in capacity-limited processing mechanisms indicating that response selection is not pivotal in dual-task delays and all three processes are likely to share a common resource like executive control.

Object-based representations govern both the storage of information in visual short-term memory and the retrieval of information from it

Interest is growing in how information is retained in visual short-term memory (VSTM). We describe an experiment that assesses VSTM within the context of multidimensional signal detection theory. On every trial, participants were presented with a 250-ms display containing four colored shapes. They were then probed 900 ms later with a colored shape and made separate old/new judgments about the color and the shape. In any particular trial, one, both, or neither of the probed features had been presented. Performance differed according to whether both probed features belonged to a single object or to two different objects. When both probed features belonged to the same object, featural retrieval was better than would be predicted by independent feature storage. When both probed features belonged to two different objects, featural retrieval was worse than would be predicted by independent feature storage. We conclude that storage in and retrieval from VSTM operate on the basis of object-based representations.

Mental Rotation Requires Visual Short-term Memory: Evidence from Human Electric Cortical Activity

The purpose of the present study was to seek evidence that mental rotation is accomplished by transforming a representation held in visual short-term memory (VSTM). In order to accomplish this goal, we utilized the sustained posterior contralateral negativity (SPCN), an electrophysiological index of the maintenance of information in VSTM. We hypothesized that if mental rotation is accomplished by transforming a representation held in VSTM, then the duration that this representation is maintained in VSTM should be related to the degree to which the representation must be rotated to reach the desired orientation. Therefore, the SPCN should offset at progressively longer latencies as the degree of rotation required increases. We tested this prediction in two experiments utilizing rotated alphanumeric characters. Experiment 1 utilized a normal versus mirror discrimination task that is known to require mental rotation. Experiment 2 utilized a letter versus digit discrimination, a task that does not require mental rotation. In Experiment 1, the offset latency of the SPCN wave increased with increases in the angle of rotation of the target. This effect indicates that targets were maintained in VSTM for longer durations as the angle of rotation increased. Experiment 2 revealed that target orientation did not affect SPCN offset latency when subjects did not adopt a mental rotation strategy, confirming that the effects on the SPCN latency effects observed in Experiment 1 were not due to the mere presentation of rotated patterns. Thus, these two experiments provide clear evidence that mental rotation involves representations maintained in VSTM.

Priming the semantic neighbourhood during the attentional blink.

BackgroundWhen two targets are presented in close temporal proximity amongst a rapid serial visual stream of distractors, a period of disrupted attention and attenuated awareness lasting 200–500 ms follows identification of the first target (T1). This phenomenon is known as the “attentional blink” (AB) and is generally attributed to a failure to consolidate information in visual short-term memory due to depleted or disrupted attentional resources. Previous research has shown that items presented during the AB that fail to reach conscious awareness are still processed to relatively high levels, including the level of meaning. For example, missed word stimuli have been shown to prime later targets that are closely associated words. Although these findings have been interpreted as evidence for semantic processing during the AB, closely associated words (e.g., day-night) may also rely on specific, well-worn, lexical associative links which enhance attention to the relevant target.Methodology/Principal FindingsWe used a measure of semantic distance to create prime-target pairs that are conceptually close, but have low word associations (e.g., wagon and van) and investigated priming from a distractor stimulus presented during the AB to a subsequent target (T2). The stimuli were words (concrete nouns) in Experiment 1 and the corresponding pictures of objects in Experiment 2. In both experiments, report of T2 was facilitated when this item was preceded by a semantically-related distractor.Conclusions/SignificanceThis study is the first to show conclusively that conceptual information is extracted from distractor stimuli presented during a period of attenuated awareness and that this information spreads to neighbouring concepts within a semantic network.

Exogenous attention influences visual short‐term memory in infants

Two experiments examined the hypothesis that developing visual attentional mechanisms influence infants' Visual Short-Term Memory (VSTM) in the context of multiple items. Five- and 10-month-old infants (N = 76) received a change detection task in which arrays of three differently colored squares appeared and disappeared. On each trial one square changed color and one square was cued; sometimes the cued item was the changing item, and sometimes the changing item was not the cued item. Ten-month-old infants exhibited enhanced memory for the cued item when the cue was a spatial pre-cue (Experiment 1) and 5-month-old infants exhibited enhanced memory for the cued item when the cue was relative motion (Experiment 2). These results demonstrate for the first time that infants younger than 6 months can encode information in VSTM about individual items in multiple-object arrays, and that attention-directing cues influence both perceptual and VSTM encoding of stimuli in infants as in adults.

Distinguishing between lateralized and nonlateralized brain activity associated with visual short-term memory: fMRI, MEG, and EEG evidence from the same observers

Previous functional neuroimaging studies have shown that maintenance of centrally presented objects in visual short-term memory (VSTM) leads to bilateral increases of BOLD activations in IPS/IOS cortex, while prior electrophysiological work suggests that maintaining stimuli encoded from a single hemifield leads to a sustained posterior contralateral negativity (SPCN) in electrophysiology and magnetoencephalography. These two findings have never been investigated using the same physiological measures. We recorded the BOLD response using fMRI, magnetoencephalography (MEG), and electrophysiology (EEG), while subjects encoded visual stimuli from a single hemifield of a balanced display. The EEG showed an SPCN. However, no SPCN-like activation was observed in the BOLD signals. The BOLD response in parietal cortex remained bilateral, even after unilateral encoding of the stimuli, but MEG showed both bilateral and contralateral activations, each likely reflecting a sub portion of the neuronal populations participating in the maintenance of information in VSTM. Contrary to the assumption that BOLD, EEG, and MEG responses – that were each linked to the maintenance of information in VSTM – are markers of the same neuronal processes, our findings suggest that each technique reveals a somewhat distinct but overlapping neural signature of the mechanisms supporting visual short-term memory.

The decay characteristics of size, color, and shape information in visual short-term memory

The decay characteristics of size, color, and shape information in visual short-term memory

Transcranial magnetic stimulation reveals the content of visual short-term memory in the visual cortex

Cortical areas involved in sensory analysis are also believed to be involved in short-term storage of that sensory information. Here we investigated whether transcranial magnetic stimulation (TMS) can reveal the content of visual short-term memory (VSTM) by bringing this information to visual awareness. Subjects were presented with two random-dot displays (moving either to the left or to the right) and they were required to maintain one of these in VSTM. In Experiment 1, TMS was applied over the motion-selective area V5/MT+ above phosphene threshold during the maintenance phase. The reported phosphene contained motion features of the memory item, when the phosphene spatially overlapped with memory item. Specifically, phosphene motion was enhanced when the memory item moved in the same direction as the subjects' V5/MT+ baseline phosphene, whereas it was reduced when the motion direction of the memory item was incongruent with that of the baseline V5/MT+ phosphene. There was no effect on phosphene reports when there was no spatial overlap between the phosphene and the memory item. In Experiment 2, VSTM maintenance did not influence the appearance of phosphenes induced from the lateral occipital region. These interactions between VSTM maintenance and phosphene appearance demonstrate that activity in V5/MT+ reflects the motion qualities of items maintained in VSTM. Furthermore, these results also demonstrate that information in VSTM can modulate the pattern of visual activation reaching awareness, providing evidence for the view that overlapping neuronal populations are involved in conscious visual perception and VSTM.

Retention Interval Affects Visual Short-Term Memory Encoding

Humans can efficiently store fine-detailed facial emotional information in visual short-term memory for several seconds. However, an unresolved question is whether the same neural mechanisms underlie high-fidelity short-term memory for emotional expressions at different retention intervals. Here we show that retention interval affects the neural processes of short-term memory encoding using a delayed facial emotion discrimination task. The early sensory P100 component of the event-related potentials (ERP) was larger in the 1-s interstimulus interval (ISI) condition than in the 6-s ISI condition, whereas the face-specific N170 component was larger in the longer ISI condition. Furthermore, the memory-related late P3b component of the ERP responses was also modulated by retention interval: it was reduced in the 1-s ISI as compared with the 6-s condition. The present findings cannot be explained based on differences in sensory processing demands or overall task difficulty because there was no difference in the stimulus information and subjects' performance between the two different ISI conditions. These results reveal that encoding processes underlying high-precision short-term memory for facial emotional expressions are modulated depending on whether information has to be stored for one or for several seconds.

Posterior parietal cortex mediates encoding and maintenance processes in change blindness

It is commonly accepted that right posterior parietal cortex (PPC) plays an important role in updating spatial representations, directing visuospatial attention, and planning actions. However, recent studies suggest that right PPC may also be involved in processes that are more closely associated with our visual awareness as its activation level positively correlates with successful conscious change detection (Beck, D.M., Rees, G., Frith, C.D., & Lavie, N. (2001). Neural correlates of change detection and change blindness. Nature Neuroscience, 4, 645–650.). Furthermore, disruption of its activity increases the occurrences of change blindness, thus suggesting a causal role for right PPC in change detection (Beck, D.M., Muggleton, N., Walsh, V., & Lavie, N. (2006). Right parietal cortex plays a critical role in change blindness. Cerebral Cortex, 16, 712–717.). In the context of a 1-shot change detection paradigm, we applied transcranial magnetic stimulation (TMS) during different time intervals to elucidate the temporally precise involvement of PPC in change detection. While subjects attempted to detect changes between two image sets separated by a brief time interval, TMS was applied either during the presentation of picture 1 when subjects were encoding and maintaining information into visual short-term memory, or picture 2 when subjects were retrieving information relating to picture 1 and comparing it to picture 2. Our results show that change blindness occurred more often when TMS was applied during the viewing of picture 1, which implies that right PPC plays a crucial role in the processes of encoding and maintaining information in visual short-term memory. In addition, since our stimuli did not involve changes in spatial locations, our findings also support previous studies suggesting that PPC may be involved in the processes of encoding non-spatial visual information (Todd, J.J. & Marois, R. (2004). Capacity limit of visual short-term memory in human posterior parietal cortex. Nature, 428, 751–754.).

Bilateral parietal and contralateral responses during maintenance of unilaterally encoded objects in visual short‐term memory: Evidence from magnetoencephalography

A component of the event-related magnetic field (ERMF) response was observed in magnetoencephalographic signals recorded during the maintenance of information in visual short-term memory (VSTM). This sustained posterior contralateral magnetic (SPCM) field is likely the magnetic equivalent of the sustained posterior contralateral negativity (SPCN) found in electrophysiology. Magnetoencephalography data showed, at the sensor level, a bilateral activation over the parietal cortex that increased in amplitude for higher memory load. Others sensors, also over the parietal cortex, showed an activation pattern similar to the SPCN with higher activation for the hemisphere contralateral to the visual field from which visual information was encoded. These two activation patterns suggest that the SPCN and SPCM are generated by a network of cortical sources that includes bilateral parietal loci, likely intra-parietal/intra-occipital cortex, and contralateral parietal sources.

Updating objects in visual short-term memory is feature selective

The purpose of this study was to examine whether the process of updating information in visual short-term memory (VSTM) is object based. We investigated whether modifying the memory of one feature of an object would automatically promote refreshing the memory of all of its other features. The results showed that the facilitative effect of updating was specific to the updated feature of an object and did not spread to its nonupdated features. This feature-selective effect suggests that updating VSTM is not object based (Experiment 1), even though storage was object based (Experiment 2). Control experiments ruled out strategy-based (Experiment 3) and stimulus-related (Experiments 4-6) accounts. Feature-selective updating may indicate that the mechanism used to modify the contents of memory may have a different basis than that used to encode or store information in memory.

Neuropsychological evidence for a spatial bias in visual short-term memory after left posterior ventral damage

For the first time, we report a spatial bias in visual short-term memory (VSTM) after left medial and inferior occipito-temporal damage. Our patient D.M. showed a spatial bias in report from VSTM, being more accurate at reporting stimuli presented in her left than her right visual field (Experiment 1). This spatial bias could not be attributed to a visual field deficit (Experiment 2) and was based on the relative rather than the absolute locations of the stimuli (Experiment 3). It was reduced when the transfer of items to VSTM was facilitated—for example, by grouping stimuli (Experiment 4) or by reducing the number of items to be remembered (Experiment 5). The spatial bias was attenuated when items moved from right to centre or left to centre, and D.M. was cued to report the item that would have been on the right or left, had the movement continued (Experiment 6). We conclude that posterior ventral damage can impair both the consolidation of new information in VSTM and the explicit report of consolidated information from VSTM.

Dissociation of the N2pc and sustained posterior contralateral negativity in a choice response task

The N2pc, a greater negativity at posterior electrodes on the side contralateral to an attended visual stimulus, usually between 180 and 280 ms, is thought to reflect the moment-to-moment deployment of visual–spatial attention. In tasks that require the retention of information in visual short-term memory, there is also a sustained posterior contralateral negativity (SPCN) that often begins about 300–400 ms after stimulus onset and that persists for the duration of the retention interval. A positive-going deflection at around 300 ms often separates the N2pc and the SPCN. An SPCN is also observed in tasks that are not defined as memory tasks, but that presumably engage visual short-term memory as an intermediate processing buffer (e.g., in order to make a choice response to a briefly-presented visual stimulus). The SPCN in memory tasks has been shown to increase in amplitude as the memory load is increased. We used this property of the SPCN to verify that the SPCN observed during the performance of a choice task with a response following each stimulus display is related to the SPCN observed in tasks that are structured as memory tasks. Using identical physical stimuli, we manipulated the hypothesized memory load across trial blocks by instructions either to encode only one stimulus or two stimuli. We observed an increase of the amplitude of the SPCN as memory load increased, with no concomitant increase in the amplitude of the N2pc that immediately preceded it. The results provide a clear dissociation between the N2pc (spatial attention, not affected by memory load) and the SPCN (visual short-term memory, sharply sensitive to memory load).

Medial Temporal Lobe Activity Predicts Successful Relational Memory Binding

Previous neuropsychological findings have implicated medial temporal lobe (MTL) structures in retaining object-location relations over the course of short delays, but MTL effects have not always been reported in neuroimaging investigations with similar short-term memory requirements. Here, we used event-related functional magnetic resonance imaging to test the hypothesis that the hippocampus and related MTL structures support accurate retention of relational memory representations, even across short delays. On every trial, four objects were presented, each in one of nine possible locations of a three-dimensional grid. Participants were to mentally rotate the grid and then maintain the rotated representation in anticipation of a test stimulus: a rendering of the grid, rotated 90 degrees from the original viewpoint. The test stimulus was either a "match" display, in which object-location relations were intact, or a "mismatch" display, in which one object occupied a new, previously unfilled location (mismatch position), or two objects had swapped locations (mismatch swap). Encoding phase activation in anterior and posterior regions of the left hippocampus, and in bilateral perirhinal cortex, predicted subsequent accuracy on the short-term memory decision, as did bilateral posterior hippocampal activity after the test stimulus. Notably, activation in these posterior hippocampal regions was also sensitive to the degree to which object-location bindings were preserved in the test stimulus; activation was greatest for match displays, followed by mismatch-position displays, and finally mismatch-swap displays. These results indicate that the hippocampus and related MTL structures contribute to successful encoding and retrieval of relational information in visual short-term memory.