Early visual areas contain specific information about visual items maintained in working memory, suggesting a role for early visual cortex in more complex cognitive functions [1-4]. It is an open question, however, whether these areas also underlie the ability to internally generate images de novo (i.e., mental imagery). Research on mental imagery has to this point focused mostly on whether mental images activate early sensory areas, with mixed results [5-7]. Recent studies suggest that multivariate pattern analysis of neural activity patterns in visual regions can reveal content-specific representations during cognitive processes, even though overall activation levels are low [1-4]. Here, we used this approach [8, 9] to study item-specific activity patterns in early visual areas (V1-V3) when these items are internally generated. We could reliably decode stimulus identity from neural activity patterns in early visual cortex during both working memory and mental imagery. Crucially, these activity patterns resembled those evoked by bottom-up visual stimulation, suggesting that mental images are indeed "perception-like" in nature. These findings suggest that the visual cortex serves as a dynamic "blackboard" [10, 11] that is used during both bottom-up stimulus processing and top-down internal generation of mental content.

A relatively new analysis technique, known as neural decoding or multivariate pattern analysis (MVPA), has become increasingly popular for cognitive neuroimaging studies over recent years. These techniques promise to uncover the representational contents of neural signals, as well as the underlying code and the dynamic profile thereof. A field in which these techniques have led to novel insights in particular is that of visual working memory (VWM). In the present study, we subjected human volunteers to a combined VWM/imagery task while recording their neural signals using magnetoencephalography (MEG). We applied multivariate decoding analyses to uncover the temporal profile underlying the neural representations of the memorized item. Analysis of gaze position however revealed that our results were contaminated by systematic eye movements, suggesting that the MEG decoding results from our originally planned analyses were confounded. In addition to the eye movement analyses, we also present the original analyses to highlight how these might have readily led to invalid conclusions. Finally, we demonstrate a potential remedy, whereby we train the decoders on a functional localizer that was specifically designed to target bottom-up sensory signals and as such avoids eye movements. We conclude by arguing for more awareness of the potentially pervasive and ubiquitous effects of eye movement-related confounds.

The study of visual working memory (VWM) has recently seen revitalization with the emergence of new insights and theories regarding its neural underpinnings. One crucial ingredient responsible for this progress is the rise of neural decoding techniques. These techniques promise to uncover the representational contents of neural signals, as well as the underlying code and the dynamic profile thereof. Here, we aimed to contribute to the field by subjecting human volunteers to a combined VWM/imagery task, while recording and decoding their neural signals as measured by MEG. At first sight, the results seem to provide evidence for a persistent, stable representation of the memorandum throughout the delay period. However, control analyses revealed that these findings can be explained by subtle, VWM-specific eye movements. As a potential remedy, we demonstrate the use of a functional localizer, which was specifically designed to target bottom-up sensory signals and as such avoids eye movements, to train the neural decoders. This analysis revealed a sustained representation for approximately 1 second, but no longer throughout the entire delay period. We conclude by arguing for more awareness of the potentially pervasive and ubiquitous effects of eye movement-related confounds.Significance statementVisual working memory is an important aspect of higher cognition and has been subject of much investigation within the field of cognitive neuroscience. Over recent years, these studies have increasingly relied on the use of neural decoding techniques. Here, we show that neural decoding may be susceptible to confounds induced by stimulus-specific eye movements. Such eye movements during working memory have been reported before, and may in fact be a common phenomenon. Given the widespread use of neural decoding and the potentially contaminating effects of eye movements, we therefore believe that our results are of significant relevance for the field.

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For the development of minimally invasive brain-computer interfaces (BCIs), it is important to accurately localize the area of implantation. Using fMRI, we investigated which brain areas are involved in motor imagery. Twelve healthy subjects performed a motor execution and imagery task during separate fMRI and EEG measurements. fMRI results showed that during imagery, premotor and parietal areas were most robustly activated in individual subjects, but surprisingly, no activation was found in the primary motor cortex. EEG results showed that spectral power decreases in contralateral sensorimotor rhythms (8-24 Hz) during both movement and imagery. To further verify the involvement of the motor imagery areas found with fMRI, one epilepsy patient performed the same task during both fMRI and ECoG recordings. Significant ECoG low (8-24 Hz) and high (65-95 Hz) frequency power changes were observed selectively on premotor cortex and these co-localized with fMRI. During a subsequent BCI task, excellent performance (91%) was obtained based on ECoG power changes from the localized premotor area. These results indicate that other areas than the primary motor area may be more reliably activated during motor imagery. Specifically, the premotor cortex may be a better area to implant an invasive BCI.

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