Within-Category Decoding of Information in Different Attentional States in Short-Term Memory.

A long-standing assumption of cognitive neuroscience has been that working memory (WM) is accomplished by sustained, elevated neural activity. More recently, theories of WM have expanded this view by describing different attentional states in WM with differing activation levels. Several studies have used multivariate pattern analysis (MVPA) of functional magnetic resonance imaging (fMRI) and electroencephalography (EEG) data to study neural activity corresponding to these WM states. Intriguingly, no evidence was found for active neural representations for information held in WM outside the focus of attention ("unattended memory items," UMIs), suggesting that only attended memory items (AMIs) are accompanied by an active trace. However, these results depended on category-level decoding, which lacks sensitivity to neural representations of individual items. Therefore, we employed a WM task in which subjects remembered the directions of motion of two dot arrays, with a retrocue indicating which was relevant for an imminent memory probe (the AMI). This design allowed MVPA decoding of delay-period fMRI signal at the stimulus-item level, affording a more sensitive test of the neural representation of UMIs. Whereas evidence for the AMI was reliably high, evidence for the UMI dropped to baseline, consistent with the notion that different WM attentional states may have qualitatively different mechanisms of retention.

Reactivation of latent working memories with transcranial magnetic stimulation.

The ability to hold information in working memory is fundamental for cognition. Contrary to the long-standing view that working memory depends on sustained, elevated activity, we present evidence suggesting that humans can hold information in working memory via "activity-silent" synaptic mechanisms. Using multivariate pattern analyses to decode brain activity patterns, we found that the active representation of an item in working memory drops to baseline when attention shifts away. A targeted pulse of transcranial magnetic stimulation produced a brief reemergence of the item in concurrently measured brain activity. This reactivation effect occurred and influenced memory performance only when the item was potentially relevant later in the trial, which suggests that the representation is dynamic and modifiable via cognitive control. The results support a synaptic theory of working memory.

Distributed patterns of activity in sensory cortex reflect the precision of multiple items maintained in visual short-term memory.

Traditionally, load sensitivity of sustained, elevated activity has been taken as an index of storage for a limited number of items in visual short-term memory (VSTM). Recently, studies have demonstrated that the contents of a single item held in VSTM can be decoded from early visual cortex, despite the fact that these areas do not exhibit elevated, sustained activity. It is unknown, however, whether the patterns of neural activity decoded from sensory cortex change as a function of load, as one would expect from a region storing multiple representations. Here, we use multivoxel pattern analysis to examine the neural representations of VSTM in humans across multiple memory loads. In an important extension of previous findings, our results demonstrate that the contents of VSTM can be decoded from areas that exhibit a transient response to visual stimuli, but not from regions that exhibit elevated, sustained load-sensitive delay-period activity. Moreover, the neural information present in these transiently activated areas decreases significantly with increasing load, indicating load sensitivity of the patterns of activity that support VSTM maintenance. Importantly, the decrease in classification performance as a function of load is correlated with within-subject changes in mnemonic resolution. These findings indicate that distributed patterns of neural activity in putatively sensory visual cortex support the representation and precision of information in VSTM.

Prioritized maps of space in human frontoparietal cortex.

Priority maps are theorized to be composed of large populations of neurons organized topographically into a map of gaze-centered space whose activity spatially tags salient and behaviorally relevant information. Here, we identified four priority map candidates along human posterior intraparietal sulcus (IPS0-IPS3) and two along the precentral sulcus (PCS) that contained reliable retinotopically organized maps of contralateral visual space. Persistent activity increased from posterior-to-anterior IPS areas and from inferior-to-superior PCS areas during the maintenance of a working memory representation, the maintenance of covert attention, and the maintenance of a saccade plan. Moreover, decoders trained to predict the locations on one task (e.g., working memory) cross-predicted the locations on other tasks (e.g., attention) in superior PCS and IPS2, suggesting that these patterns of maintenance activity may be interchangeable across the tasks. Such properties make these two areas in frontal and parietal cortex viable priority map candidates.

The relationship between working memory storage and elevated activity as measured with functional magnetic resonance imaging.

Does the sustained, elevated neural activity observed during working memory tasks reflect the short-term retention of information? Functional magnetic resonance imaging (fMRI) data of delayed recognition of visual motion in human participants were analyzed with two methods: a general linear model (GLM) and multivoxel pattern analysis. Although the GLM identified sustained, elevated delay-period activity in superior and lateral frontal cortex and in intraparietal sulcus, pattern classifiers were unable to recover trial-specific stimulus information from these delay-active regions. The converse-no sustained, elevated delay-period activity but successful classification of trial-specific stimulus information-was true of posterior visual regions, including area MT+ (which contains both middle temporal area and medial superior temporal area) and calcarine and pericalcarine cortex. In contrast to stimulus information, pattern classifiers were able to extract trial-specific task instruction-related information from frontal and parietal areas showing elevated delay-period activity. Thus, the elevated delay-period activity that is measured with fMRI may reflect processes other than the storage, per se, of trial-specific stimulus information. It may be that the short-term storage of stimulus information is represented in patterns of (statistically) "subthreshold" activity distributed across regions of low-level sensory cortex that univariate methods cannot detect.

Cortical projection topography of the human splenium: hemispheric asymmetry and individual differences.

The corpus callosum is the largest white matter pathway in the human brain. The most posterior portion, known as the splenium, is critical for interhemispheric communication between visual areas. The current study employed diffusion tensor imaging to delineate the complete cortical projection topography of the human splenium. Homotopic and heterotopic connections were revealed between the splenium and the posterior visual areas, including the occipital and the posterior parietal cortices. In nearly one third of participants, there were homotopic connections between the primary visual cortices, suggesting interindividual differences in splenial connectivity. There were also more instances of connections with the right hemisphere, indicating a hemispheric asymmetry in interhemispheric connectivity within the splenium. Combined, these findings demonstrate unique aspects of human interhemispheric connectivity and provide anatomical bases for hemispheric asymmetries in visual processing and a long-described hemispheric asymmetry in speed of interhemispheric communication for visual information.