Friday 24 April 2015

Research Briefing: organising the contents of working memory

Figure 1. Nicholas Myers
Research Briefing, by Nicholas Myers

Everyone has been in this situation: you are stuck in an endless meeting, and a colleague drones on about a topic of marginal relevance. You begin to zone out and focus on the art hanging in your boss’s office, when suddenly you hear your name mentioned. On high alert, you suddenly shift back to the meeting and scramble to retrieve your colleague’s last sentences. Miraculously, you are able to retrieve a few key words – they must have entered your memory a moment ago, but would have been quickly forgotten if hearing your name had not cued them as potentially vital bits of information.

This phenomenon, while elusive in everyday situations, has been studied experimentally for a number of years now: cues indicating the relevance of a particular item in working memory have a striking benefit to our ability to recall it, even if the cue is presented after the item has already entered memory. See our previous Research Briefing on how retrospective cueing can restore information to the focus of attention in working memory.

In a new article, published in the Journal of Cognitive Neuroscience, we describe a recent experiment that set out to add to our expanding knowledge of how the brain orchestrates these retrospective shifts of attention. We were particularly interested in the potential role of neural synchronization of 10 Hz (or alpha-band) oscillations, because they are important in similar prospective shifts of attention.

Figure 2. Experimental Task Design. [from Myers et al, 2014]
We wanted to examine the similarity of alpha-band responses (and other neural signatures of the engagement of attention) both to retrospective and prospective attention shifts. We needed to come up with a new task that allowed for this comparison. On each trial in our task, experiment volunteers first memorized two visual stimuli. Two seconds later, a second set of two stimuli appeared, so that a total of four stimuli was kept in mind. After a further delay, participants recalled one of the four items.  

In between the presentation of the first and the second set of two items, we sometimes presented a cue: this cue indicated which of the four items would likely be tested at the end of the trial. Crucially, this cue could have either a prospective or a retrospective function, depending on whether it pointed to location where an item had already been presented (a retrospective cue, or retrocue), or to a location where a stimulus was yet to appear (a prospective cue, or precue). This allowed us to examine neural responses to attention-guiding cues that were identical with respect to everything but their forwards- or backwards-looking nature. See Figure 2 for a task schematic.

Figure 3. Results: retro-cueing and pre-cueing
trigger different attention-related ERPs.
[from Myers et al, 2014]
We found marked differences in event-related potential (ERP) profiles between the precue and retrocue conditions. We found evidence that precues primarily generate an anticipatory shift of attention toward the location of an upcoming item: potentials just before the expected appearance of the second set of stimuli reflected the location where volunteers were attending. These included the so-called early directing attention negativity (or 'EDAN') and the late directing attention-related positivity (or 'LDAP'; see Figure 3, middle panel; and see here for a review of attention-related ERPs). Retrocues elicited a different pattern of ERPs that was compatible with an early selection mechanism, but not with stimulus anticipation (i.e., no LDAP, see Figure 3, upper panel). The latter seems plausible, since the cued information was already in memory, and upcoming stimuli were therefore not deserving of attention. In contrast to the distinct ERP patterns, alpha band (8-14 Hz) lateralization was indistinguishable between cue types (reflecting, in both conditions, the location of the cued item; see Figure 4).

Figure 4. Results: retro-cueing and pre-cueing trigger similar patters
of de-synchronisation in low frequency activity (alpha band at ~10Hz).
[from Myers et al, 2014]
What did we learn from this study? Taken together with the ERP results, it seems that alpha-band lateralization can have two distinct roles: after a precue it likely enables anticipatory attention. After a retrocue, however, the alpha-band response may reflect the controlled retrieval of a recently memorized piece of information that has turned out to be more useful than expected, without influencing the brain’s response to upcoming stimuli.

It seems that our senses are capable of storing a limited amount of information on the off chance that it may suddenly become relevant. When this turns out to be the case, top-down control allows us to pick out the relevant information from among all the items quietly rumbling around in sensory brain regions.

Many interesting questions remain that we were not able to address in this study. For example, how do cortical areas responsible for top-down control activate in response to a retrocue, and how do they shuttle cued information into a state that can guide behaviour? 

Key Reference: 

Myers, Walther, Wallis, Stokes & Nobre (2014) Temporal Dynamics of Attention during Encoding versus Maintenance of Working Memory: Complementary Views from Event-related Potentials and Alpha-band Oscillations. Journal of Cognitive Neuroscience (Open Access)

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