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In a dynamic environment, the already limited information that human working memory can maintain needs to be constantly updated to optimally guide behaviour. Indeed, previous studies showed that working memory representations are continuously being transformed during delay periods leading up to a response. This goes hand-in-hand with the removal of task-irrelevant items. However, does such removal also include veridical, original stimuli, as they were prior to transformation? Here we aimed to assess the neural representation of task-relevant transformed representations, compared to the no-longer-relevant veridical representations they originated from. We applied multivariate pattern analysis to electroencephalographic data during maintenance of orientation gratings with and without mental rotation. During maintenance, we perturbed the representational network by means of a visual impulse stimulus, and were thus able to successfully decode veridical as well as imaginary, transformed orientation gratings from impulse-driven activity. On the one hand, the impulse response reflected only task-relevant (cued), but not task-irrelevant (uncued) items, suggesting that the latter were quickly discarded from working memory. By contrast, even though the original cued orientation gratings were also no longer task-relevant after mental rotation, these items continued to be represented next to the rotated ones, in different representational formats. This seemingly inefficient use of scarce working memory capacity was associated with reduced probe response times and may thus serve to increase precision and flexibility in guiding behaviour in dynamic environments.
The activity-silent framework of working memory (WM) posits that the neural activity during object perception and encoding leaves behind patterned, “activity-silent” neural traces that enable WM maintenance without the need for continuous, memory-specific neural activity. The presence of such traces in the memory network subsequently patterns its responses to external stimulation, which can be used to readout the contents of WM using an impulse perturbation or “pinging” approach. The extent to which the neural impulse response is patterned by the WM network should be modulated by the physical overlap between the initial memory item and the subsequent external perturbation stimulus, with higher overlap increasing WM readout. Here we tested this prediction in a delayed orientation match-to-sample task, by either matching or mismatching task-irrelevant spatial frequencies between memory items and impulse stimuli, and between memory items and probes. Matching frequencies resulted in faster behavioral response times, and increased the WM-specificity of the neural impulse response as measured from the EEG signal. We found no evidence that matching spatial frequencies resulted in globally stronger or different neural responses, but rather in distinct neural activation patterns. The beneficial effects of feature matching in our task support the tenets of the activity-silent framework of WM, and confirm that impulse perturbation interacts directly with the representations that are held in memory.
The activity-silent framework of working memory (WM) posits that the neural activity during object perception and encoding leaves behind patterned, “activity-silent” neural traces that enable WM maintenance without the need for continuous, memory-specific neural activity. The presence of such traces in the memory network subsequently patterns its responses to external stimulation, which can be used to readout the contents of WM using an impulse perturbation or “pinging” approach. The extent to which the neural impulse response is patterned by the WM network should be modulated by the physical overlap between the initial memory item and the subsequent external perturbation stimulus, with higher overlap increasing WM readout. Here we tested this prediction in a delayed orientation match-to-sample task, by either matching or mismatching task-irrelevant spatial frequencies between memory items and impulse stimuli, and between memory items and probes. Matching frequencies resulted in faster behavioral response times, and increased the WM-specificity of the neural impulse response as measured from the EEG signal. We found no evidence that matching spatial frequencies resulted in globally stronger or different neural responses, but rather in distinct neural activation patterns. The beneficial effects of feature matching in our task support the tenets of the activity-silent framework of WM, and confirm that impulse perturbation interacts directly with the representations that are held in memory.
Behaviorally irrelevant feature matching increases neural and behavioral working memory readout
(2024)
There is an ongoing debate about whether working memory (WM) maintenance relies on persistent activity and/or short-term synaptic plasticity. This is a challenging question, because neuroimaging techniques in cognitive neuroscience measure activity only. Recently, neural perturbation techniques have been developed to tackle this issue, such as visual impulse perturbation or “pinging”, which reveals (un)attended WM content during maintenance. There are contrasting explanations of how pinging reveals WM content, which is central to the debate. Pinging could reveal mnemonic representations by perturbing content-specific networks or by increasing the neural signal-to-noise ratio of active neural states. Here we tested the extent to which the neural impulse response is patterned by the WM network, by presenting two different impulse stimuli. If the impulse interacts with WM networks, the WM-specific impulse response should be enhanced by physical overlap between the initial memory item and the subsequent external perturbation stimulus. This prediction was tested in a delayed orientation match-to-sample task by matching or mismatching task-irrelevant spatial frequencies between memory items and impulse stimuli, as well as probes. Matching probe spatial frequency with memory items resulted in faster behavioral response times and matching impulse spatial frequency with memory items increased the specificity of the neural impulse response as measured from EEG. Matching spatial frequencies did neither result in globally stronger neural responses nor in a larger decrease in trial-to-trial variability compared to mismatching spatial frequencies. The improved neural and behavioural readout of irrelevant feature matching provide evidence that impulse perturbation interacts directly with the memory representations.
