Yoni Pertzov

Rapid forgetting prevented by retrospective attention cues.

Recent studies have demonstrated that memory performance can be enhanced by a cue which indicates the item most likely to be subsequently probed, even when that cue is delivered seconds after a stimulus array is extinguished. Although such retro-cuing has attracted considerable interest, the mechanisms underlying it remain unclear. Here, we tested the hypothesis that retro-cues might protect an item from degradation over time. We employed two techniques that previously have not been deployed in retro-cuing tasks. First, we used a sensitive, continuous scale for reporting the orientation of a memorized item, rather than binary measures (change or no change) typically used in previous studies. Second, to investigate the stability of memory across time, we also systematically varied the duration between the retro-cue and report. Although accuracy of reporting uncued objects rapidly declined over short intervals, retro-cued items were significantly more stable, showing negligible decline in accuracy across time and protection from forgetting. Retro-cuing an object’s color was just as advantageous as spatial retro-cues. These findings demonstrate that during maintenance, even when items are no longer visible, attention resources can be selectively redeployed to protect the accuracy with which a cued item can be recalled over time, but with a corresponding cost in recall for uncued items.

Binding deficits in memory following medial temporal lobe damage in patients with voltage-gated potassium channel complex antibody-associated limbic encephalitis

Some prominent studies have claimed that the medial temporal lobe is not involved in retention of information over brief intervals of just a few seconds. However, in the last decade several investigations have reported that patients with medial temporal lobe damage exhibit an abnormally large number of errors when required to remember visual information over brief intervals. But the nature of the deficit and the type of error associated with medial temporal lobe lesions remains to be fully established. Voltage-gated potassium channel complex antibody-associated limbic encephalitis has recently been recognized as a form of treatable autoimmune encephalitis, frequently associated with imaging changes in the medial temporal lobe. Here, we tested a group of these patients using two newly developed visual short-term memory tasks with a sensitive, continuous measure of report. These tests enabled us to study the nature of reporting errors, rather than only their frequency. On both paradigms, voltage-gated potassium channel complex antibody patients exhibited larger errors specifically when several items had to be remembered, but not for a single item. Crucially, their errors were strongly associated with an increased tendency to report the property of the wrong item stored in memory, rather than simple degradation of memory precision. Thus, memory for isolated aspects of items was normal, but patients were impaired at binding together the different properties belonging to an item, e.g. spatial location and object identity, or colour and orientation. This occurred regardless of whether objects were shown simultaneously or sequentially. Binding errors support the view that the medial temporal lobe is involved in linking together different types of information, potentially represented in different parts of the brain, regardless of memory duration. Our novel behavioural measures also have the potential to assist in monitoring response to treatment in patients with memory disorders, such as those with voltage-gated potassium channel complex antibody limbic encephalitis.

Multiple Reference Frames for Saccadic Planning in the Human Parietal Cortex

We apply functional magnetic resonance imaging and multivariate analysis methods to study the coordinate frame in which saccades are represented in the human cortex. Subjects performed a memory-guided saccade task in which equal-amplitude eye movements were executed from several starting points to various directions. Response patterns during the memory period for same-vector saccades were correlated in the frontal eye fields and the intraparietal sulcus (IPS), indicating a retinotopic representation. Interestingly, response patterns in the middle aspect of the IPS were also correlated for saccades made to the same destination point, even when their movement vector was different. Thus, this region also contains information about saccade destination in (at least) a head-centered coordinate frame. This finding may explain behavioral and neuropsychological studies demonstrating that eye movements are also anchored to an egocentric or an allocentric representation of space rather than strictly to the retinal visual input and that parietal cortex is involved in maintaining these representations of space.

Rapid Formation of Spatiotopic Representations As Revealed by Inhibition of Return

Inhibition of return (IOR), a performance decrement for stimuli appearing at recently cued locations, occurs when the target and cue share the same screen position. This is in contrast to cue-based attention facilitation effects that were recently suggested to be mapped in a retinotopic reference frame, the prevailing representation throughout early visual processing stages. Here, we investigate the dynamics of IOR in both reference frames, using a modified cued-location saccadic reaction time task with an intervening saccade between cue and target presentation. Thus, on different trials, the target was present either at the same retinotopic location as the cue, or at the same screen position (e.g., spatiotopic location). IOR was primarily found for targets appearing at the same spatiotopic position as the initial cue, when the cue and target were presented at the same hemifield. This suggests that there is restricted information transfer of cue position across the two hemispheres. Moreover, the effect was maximal when the target was presented 10 ms after the intervening saccade ended and was attenuated in longer delays. In our case, therefore, the representation of previously attended locations (as revealed by IOR) is not remapped slowly after the execution of a saccade. Rather, either a retinotopic representation is remapped rapidly, adjacent to the end of the saccade (using a prospective motor command), or the positions of the cue and target are encoded in a spatiotopic reference frame, regardless of eye position. Spatial attention can therefore be allocated to target positions defined in extraretinal coordinates.

Accumulation of visual information across multiple fixations

Humans often redirect their gaze to the same objects within a scene, even without being consciously aware of it. Here, we investigated what type of visual information is accumulated across recurrent fixations on the same object. On each trial, subjects viewed an array comprised of several objects and were subsequently asked to report on various visual aspects of a randomly chosen target object from that array. Memory performance decreased as more fixations were directed to other objects, following the last fixation on the target object (i.e. post-target fixations). In contrast, performance was enhanced with increasing number of fixations on the target object. However, since the number of post-target fixations and the number of target fixations are usually anti-correlated, memory gain may simply reflect fewer post-target fixations, rather than true accumulation of information. To rule this out, we conducted a second experiment, in which the stimulus disappeared immediately after performing a predefined number of target fixations. Additional fixations on the target object resulted in improved memory performance even under these strict conditions. We conclude that, under the present conditions, various aspects of memory monotonically improve with repeated sampling of the same object.

Forgetting what was where: the fragility of object-location binding.

Although we frequently take advantage of memory for objects locations in everyday life, understanding how an object’s identity is bound correctly to its location remains unclear. Here we examine how information about object identity, location and crucially object-location associations are differentially susceptible to forgetting, over variable retention intervals and memory load. In our task, participants relocated objects to their remembered locations using a touchscreen. When participants mislocalized objects, their reports were clustered around the locations of other objects in the array, rather than occurring randomly. These ‘swap’ errors could not be attributed to simple failure to remember either the identity or location of the objects, but rather appeared to arise from failure to bind object identity and location in memory. Moreover, such binding failures significantly contributed to decline in localization performance over retention time. We conclude that when objects are forgotten they do not disappear completely from memory, but rather it is the links between identity and location that are prone to be broken over time.

The privileged role of location in visual working memory

Reports have conflicted about the possible special role of location in visual working memory (WM). One important question is: Do we maintain the locations of objects in WM even when they are irrelevant to the task at hand? Here we used a continuous response scale to study the types of reporting errors that participants make when objects are presented at the same or at different locations in space. When several objects successively shared the same location, participants exhibited a higher tendency to report features of the wrong object in memory; that is, they responded with features that belonged to objects retained in memory but not probed at retrieval. On the other hand, a similar effect was not observed when objects shared a nonspatial feature, such as color. Furthermore, the effect of location on reporting errors was present even when its manipulation was orthogonal to the task at hand. These findings are consistent with the view that binding together different nonspatial features of an object in memory might be mediated through an object’s location. Hence, spatial location may have a privileged role in WM. The relevance of these findings to conceptual models, as well as to neural accounts of visual WM, is discussed.

Effects of Healthy Ageing on Precision and Binding of Object Location in Visual Short Term Memory

Visual short term memory (STM) declines as people get older, but the nature of this deterioration is not well understood. We tested 139 healthy subjects (19–83 years) who were first required to identify a previously seen object and then report its location using a touchscreen. Results demonstrated an age-related decline in both object identification and localization. Deterioration in localization performance was apparent even when only 1 item had to be remembered, worsening disproportionately with increasing memory load. Thus, age-dependent memory degradation cannot be explained simply by a decrease in the number of items that can be held in visual STM but rather by the precision with which they are recalled. More important, there was no evidence for a significant decrease in object-location binding with increasing age. Thus, although precision for object identity and location declines with age, the ability to associate object identity to its location seems to remain unimpaired. As it has been reported that binding deficits in STM might be the first cognitive signs of early Alzheimer’s disease (AD), the finding that object-location binding processes are relatively intact with normal aging supports the possible suitability of using misbinding as an index measures for probing early diagnosis of AD.

Working memory retrieval as a decision process.

Working memory (WM) is a core cognitive process fundamental to human behavior, yet the mechanisms underlying it remain highly controversial. Here we provide a new framework for understanding retrieval of information from WM, conceptualizing it as a decision based on the quality of internal evidence. Recent findings have demonstrated that precision of WM decreases with memory load. If WM retrieval uses a decision process that depends on memory quality, systematic changes in response time distribution should occur as a function of WM precision. We asked participants to view sample arrays and, after a delay, report the direction of change in location or orientation of a probe. As WM precision deteriorated with increasing memory load, retrieval time increased systematically. Crucially, the shape of reaction time distributions was consistent with a linear accumulator decision process. Varying either task relevance of items or maintenance duration influenced memory precision, with corresponding shifts in retrieval time. These results provide strong support for a decision-making account of WM retrieval based on noisy storage of items. Furthermore, they show that encoding, maintenance, and retrieval in WM need not be considered as separate processes, but may instead be conceptually unified as operations on the same noise-limited, neural representation.

Attention and memory protection: Interactions between retrospective attention cueing and interference

Visual working memory (VWM) and attention have a number of features in common, but despite extensive research it is still unclear how the two interact. Can focused attention improve VWM precision? Can it protect VWM from interference? Here we used a partial-report, continuous-response orientation memory task to examine how attention and interference affect different aspects of VWM and how they interact with one another. Both attention and interference were orthogonally manipulated during the retention interval. Attention was manipulated by presenting informative retro-cues, whereas interference was manipulated by introducing a secondary interfering task. Mixture-model analyses revealed that retro-cues, compared to uninformative cues, improved all aspects of performance: Attention increased recall precision and decreased guessing rate and swap-errors (reporting a wrong item in memory). Similarly, performing a secondary task impaired all aspects of the VWM task. In particular, an interaction between retro-cue and secondary task interference was found primarily for swap-errors. Together these results suggest that both the quantity and quality of VWM representations are sensitive to attention cueing and interference modulations, and they highlight the role of attention in protecting the feature–location associations needed to access the correct items in memory.