Neurofeedback training of executive function in autism spectrum disorder: distinct effects on brain activity levels and compensatory connectivity changes.

BACKGROUND: Deficits in executive function (EF) are consistently reported in autism spectrum disorders (ASD). Tailored cognitive training tools, such as neurofeedback, focused on executive function enhancement might have a significant impact on the daily life functioning of individuals with ASD. We report the first real-time fMRI neurofeedback (rt-fMRI NF) study targeting the left dorsolateral prefrontal cortex (DLPFC) in ASD. METHODS: Thirteen individuals with autism without intellectual disability and seventeen neurotypical individuals completed a rt-fMRI working memory NF paradigm, consisting of subvocal backward recitation of self-generated numeric sequences. We performed a region-of-interest analysis of the DLPFC, whole-brain comparisons between groups and, DLPFC-based functional connectivity. RESULTS: The ASD and control groups were able to modulate DLPFC activity in 84% and 98% of the runs. Activity in the target region was persistently lower in the ASD group, particularly in runs without neurofeedback. Moreover, the ASD group showed lower activity in premotor/motor areas during pre-neurofeedback run than controls, but not in transfer runs, where it was seemingly balanced by higher connectivity between the DLPFC and the motor cortex. Group comparison in the transfer run also showed significant differences in DLPFC-based connectivity between groups, including higher connectivity with areas integrated into the multidemand network (MDN) and the visual cortex. CONCLUSIONS: Neurofeedback seems to induce a higher between-group similarity of the whole-brain activity levels (including the target ROI) which might be promoted by changes in connectivity between the DLPFC and both high and low-level areas, including motor, visual and MDN regions.

Multi-scale analysis of acupuncture mechanisms for motor and sensory cortex activity based on SEEG data.

Acupuncture, a traditional Chinese therapy, is gaining attention for its impact on the brain. While existing electroencephalogram and functional magnetic resonance image research has made significant contributions, this paper utilizes stereo-electroencephalography data for a comprehensive exploration of neurophysiological effects. Employing a multi-scale approach, channel-level analysis reveals notable $\delta $-band activity changes during acupuncture. At the brain region level, acupuncture modulated connectivity between the paracentral lobule and the precentral gyrus. Whole-brain analysis indicates acupuncture's influence on network organization, and enhancing $E_{glob}$ and increased interaction between the motor and sensory cortex. Brain functional reorganization is an important basis for functional recovery or compensation after central nervous system injury. The use of acupuncture to stimulate peripheral nerve trunks, muscle motor points, acupoints, etc., in clinical practice may contribute to the reorganization of brain function. This multi-scale perspective provides diverse insights into acupuncture's effects. Remarkably, this paper pioneers the introduction of stereo-electroencephalography data, advancing our understanding of acupuncture's mechanisms and potential therapeutic benefits in clinical settings.

Brain wave oscillations as an objective neurophysiological biomarker of homeopathic subjective well-being [Pré-print]

Homeopathy uses the "similitude principle" to arouse a therapeutic reaction in the body against its own disorders. For this to occur optimally, the medicinal pathogenetic effects must present similarity with the totality of the individual's symptoms. To assess if this similarity has been successfully achieved, Hahnemann states that "improvement in the disposition and mind"­i.e., subjective well-being­is the most important parameter to consider. Aim Our aim was to perform a narrative review of the literature, exploring what is known about subjective well-being as a marker of therapeutic action, and to formulate ways in which subjective well-being might be quantifiable and applied in future homeopathy research. The concept of subjective well-being has been extensively studied in the complementary and conventional medical literature. Improved well-being has been observed in clinical trials, including those in the fields of positive psychology and meditation. Positive subjective outcomes of this nature are supported by objective evidence through associated changes in brain oscillatory activity using electroencephalography and/or "brain mapping" by functional magnetic resonance imaging. Neurophysiological responses in the brain have been identified in subjects after they ingested a homeopathic medicine. The concept of subjective well-being is supported by a body of literature and is a measurable entity. When viewed from the perspective of electrophysiological changes, brain activity is an objective neurophysiological biomarker with a potential to quantify individual well-being in the context of homeopathy research.

Psilocybin enhances insightfulness in meditation: a perspective on the global topology of brain imaging during meditation.

In this study, for the first time, we explored a dataset of functional magnetic resonance images collected during focused attention and open monitoring meditation before and after a five-day psilocybin-assisted meditation retreat using a recently established approach, based on the Mapper algorithm from topological data analysis. After generating subject-specific maps for two groups (psilocybin vs. placebo, 18 subjects/group) of experienced meditators, organizational principles were uncovered using graph topological tools, including the optimal transport (OT) distance, a geometrically rich measure of similarity between brain activity patterns. This revealed characteristics of the topology (i.e. shape) in space (i.e. abstract space of voxels) and time dimension of whole-brain activity patterns during different styles of meditation and psilocybin-induced alterations. Most interestingly, we found that (psilocybin-induced) positive derealization, which fosters insightfulness specifically when accompanied by enhanced open-monitoring meditation, was linked to the OT distance between open-monitoring and resting state. Our findings suggest that enhanced meta-awareness through meditation practice in experienced meditators combined with potential psilocybin-induced positive alterations in perception mediate insightfulness. Together, these findings provide a novel perspective on meditation and psychedelics that may reveal potential novel brain markers for positive synergistic effects between mindfulness practices and psilocybin.

Speech-induced suppression during natural dialogues.

When engaged in a conversation, one receives auditory information from the other's speech but also from their own speech. However, this information is processed differently by an effect called Speech-Induced Suppression. Here, we studied brain representation of acoustic properties of speech in natural unscripted dialogues, using electroencephalography (EEG) and high-quality speech recordings from both participants. Using encoding techniques, we were able to reproduce a broad range of previous findings on listening to another's speech, and achieving even better performances when predicting EEG signal in this complex scenario. Furthermore, we found no response when listening to oneself, using different acoustic features (spectrogram, envelope, etc.) and frequency bands, evidencing a strong effect of SIS. The present work shows that this mechanism is present, and even stronger, during natural dialogues. Moreover, the methodology presented here opens the possibility of a deeper understanding of the related mechanisms in a wider range of contexts.

No replication of direct neuronal activity-related (DIANA) fMRI in anesthetized mice.

Direct imaging of neuronal activity (DIANA) by functional magnetic resonance imaging (fMRI) could be a revolutionary approach for advancing systems neuroscience research. To independently replicate this observation, we performed fMRI experiments in anesthetized mice. The blood oxygenation level-dependent (BOLD) response to whisker stimulation was reliably detected in the primary barrel cortex before and after DIANA experiments; however, no DIANA-like fMRI peak was observed in individual animals' data with the 50 to 300 trials. Extensively averaged data involving 1050 trials in six mice showed a flat baseline and no detectable neuronal activity-like fMRI peak. However, spurious, nonreplicable peaks were found when using a small number of trials, and artifactual peaks were detected when some outlier-like trials were excluded. Further, no detectable DIANA peak was observed in the BOLD-responding thalamus from the selected trials with the neuronal activity-like reference function in the barrel cortex. Thus, we were unable to replicate the previously reported results without data preselection.

Validation of real-time fMRI neurofeedback procedure for cognitive training using counterbalanced active-sham study design.

Investigation of neural mechanisms of real-time functional MRI neurofeedback (rtfMRI-nf) training requires an efficient study control approach. A common rtfMRI-nf study design involves an experimental group, receiving active rtfMRI-nf, and a control group, provided with sham rtfMRI-nf. We report the first study in which rtfMRI-nf procedure is controlled through counterbalancing training runs with active and sham rtfMRI-nf for each participant. Healthy volunteers (n = 18) used rtfMRI-nf to upregulate fMRI activity of an individually defined target region in the left dorsolateral prefrontal cortex (DLPFC) while performing tasks that involved mental generation of a random numerical sequence and serial summation of numbers in the sequence. Sham rtfMRI-nf was provided based on fMRI activity of a different brain region, not involved in these tasks. The experimental procedure included two training runs with the active rtfMRI-nf and two runs with the sham rtfMRI-nf, in a randomized order. The participants achieved significantly higher fMRI activation of the left DLPFC target region during the active rtfMRI-nf conditions compared to the sham rtfMRI-nf conditions. fMRI functional connectivity of the left DLPFC target region with the nodes of the central executive network was significantly enhanced during the active rtfMRI-nf conditions relative to the sham conditions. fMRI connectivity of the target region with the nodes of the default mode network was similarly enhanced. fMRI connectivity changes between the active and sham conditions exhibited meaningful associations with individual performance measures on the Working Memory Multimodal Attention Task, the Approach-Avoidance Task, and the Trail Making Test. Our results demonstrate that the counterbalanced active-sham study design can be efficiently used to investigate mechanisms of active rtfMRI-nf in direct comparison to those of sham rtfMRI-nf. Further studies with larger group sizes are needed to confirm the reported findings and evaluate clinical utility of this study control approach.

Increased Thalamic Connectivity in Juvenile Myoclonic Epilepsy Based on Electroencephalography Source-Level Analysis.

Background: This study investigated alterations in the intrinsic thalamic network of patients with juvenile myoclonic epilepsy (JME) based on an electroencephalography (EEG) source-level analysis. Materials and Methods: We enrolled patients newly diagnosed with JME as well as healthy controls. The assessments were conducted in the resting state. We computed sources based on the scalp electrical potentials using a minimum-norm imaging method and a standardized, low-resolution, brain electromagnetic tomography approach. To create a functional connectivity matrix, we used the Talairach atlas to define thalamic nodes and applied the coherence method to measure brain synchronization as edges. We then calculated the intrinsic thalamic network using graph theory. We compared the intrinsic thalamic network of patients with JME with those of healthy controls. Results: This study included 67 patients with JME and 66 healthy controls. EEG source-level analysis revealed significant differences in the intrinsic thalamic networks between patients with JME and healthy controls. The measures of functional connectivity (radius, diameter, and characteristic path length) were significantly lower in patients with JME than in healthy controls (radius: 2.769 vs. 3.544, p = 0.015; diameter: 4.464 vs. 5.443, p = 0.024; and characteristic path length: 2.248 vs. 2.616, p = 0.046). Conclusions: We demonstrated alterations in the intrinsic thalamic network in patients with JME compared with those in healthy controls based on the EEG source-level analysis. These findings indicated increased thalamic connectivity in the JME group. These intrinsic thalamic network changes may be related to the pathophysiology of JME.

Beyond boundaries: investigating shared and divergent connectivity in the pre-/postcentral gyri and supplementary motor area.

OBJECTIVE: This study aimed to comprehensively investigate the functional connectivity of key brain regions involved in motor and sensory functions, namely the precentral gyrus, postcentral gyrus and supplementary motor area (SMA). Using advanced MRI, the objective was to understand the neurophysiological integrative characterizations of these regions by examining their connectivity with eight distinct functional brain networks. The goal was to uncover their roles beyond conventional motor and sensory functions, contributing to a more holistic understanding of brain functioning. METHODS: The study involved 198 healthy volunteers, with the primary methodology being functional connectivity analysis using advanced MRI techniques. The bilateral precentral gyrus, postcentral gyrus and SMA served as seed regions, and their connectivity with eight distinct brain regional functional networks was investigated. This approach allowed for the exploration of synchronized activity between these critical brain areas, shedding light on their integrated functioning and relationships with other brain networks. RESULTS: The study revealed a nuanced landscape of functional connectivity for the precentral gyrus, postcentral gyrus and SMA with the main functional brain networks. Despite their high functional connectedness, these regions displayed diverse functional integrations with other networks, particularly in the salience, visual, cerebellar and language networks. Specific data and statistical significance were not provided in the abstract, but the results suggested unique and distinct roles for each brain area in sophisticated cognitive tasks beyond their conventional motor and sensory functions. CONCLUSION: The study emphasized the multifaceted roles of the precentral gyrus, postcentral gyrus and SMA. Beyond their crucial involvement in motor and sensory functions, these regions exhibited varied functional integrations with different brain networks. The observed disparities, especially in the salience, visual, cerebellar and language networks, indicated a nuanced and specialized involvement of these regions in diverse cognitive functions. The study underscores the importance of considering the broader neurophysiological landscape to comprehend the intricate roles of these brain areas, contributing to ongoing efforts in unraveling the complexities of brain function.

Mapping Subcortico-Cortical Coupling-A Comparison of Thalamic and Subthalamic Oscillations.

BACKGROUND: The ventral intermediate nucleus of the thalamus (VIM) is an effective target for deep brain stimulation in tremor patients. Despite its therapeutic importance, its oscillatory coupling to cortical areas has rarely been investigated in humans. OBJECTIVES: The objective of this study was to identify the cortical areas coupled to the VIM in patients with essential tremor. METHODS: We combined resting-state magnetoencephalography with local field potential recordings from the VIM of 19 essential tremor patients. Whole-brain maps of VIM-cortex coherence in several frequency bands were constructed using beamforming and compared with corresponding maps of subthalamic nucleus (STN) coherence based on data from 19 patients with Parkinson's disease. In addition, we computed spectral Granger causality. RESULTS: The topographies of VIM-cortex and STN-cortex coherence were very similar overall but differed quantitatively. Both nuclei were coupled to the ipsilateral sensorimotor cortex in the high-beta band; to the sensorimotor cortex, brainstem, and cerebellum in the low-beta band; and to the temporal cortex, brainstem, and cerebellum in the alpha band. High-beta coherence to sensorimotor cortex was stronger for the STN (P = 0.014), whereas low-beta coherence to the brainstem was stronger for the VIM (P = 0.017). Although the STN was driven by cortical activity in the high-beta band, the VIM led the sensorimotor cortex in the alpha band. CONCLUSIONS: Thalamo-cortical coupling is spatially and spectrally organized. The overall similar topographies of VIM-cortex and STN-cortex coherence suggest that functional connections are not necessarily unique to one subcortical structure but might reflect larger frequency-specific networks involving VIM and STN to a different degree. © 2024 The Authors. Movement Disorders published by Wiley Periodicals LLC on behalf of International Parkinson and Movement Disorder Society.

Processing of auditory novelty in human cortex during a semantic categorization task.

Auditory semantic novelty - a new meaningful sound in the context of a predictable acoustical environment - can probe neural circuits involved in language processing. Aberrant novelty detection is a feature of many neuropsychiatric disorders. This large-scale human intracranial electrophysiology study examined the spatial distribution of gamma and alpha power and auditory evoked potentials (AEP) associated with responses to unexpected words during performance of semantic categorization tasks. Participants were neurosurgical patients undergoing monitoring for medically intractable epilepsy. Each task included repeatedly presented monosyllabic words from different talkers ("common") and ten words presented only once ("novel"). Targets were words belonging to a specific semantic category. Novelty effects were defined as differences between neural responses to novel and common words. Novelty increased task difficulty and was associated with augmented gamma, suppressed alpha power, and AEP differences broadly distributed across the cortex. Gamma novelty effect had the highest prevalence in planum temporale, posterior superior temporal gyrus (STG) and pars triangularis of the inferior frontal gyrus; alpha in anterolateral Heschl's gyrus (HG), anterior STG and middle anterior cingulate cortex; AEP in posteromedial HG, lower bank of the superior temporal sulcus, and planum polare. Gamma novelty effect had a higher prevalence in dorsal than ventral auditory-related areas. Novelty effects were more pronounced in the left hemisphere. Better novel target detection was associated with reduced gamma novelty effect within auditory cortex and enhanced gamma effect within prefrontal and sensorimotor cortex. Alpha and AEP novelty effects were generally more prevalent in better performing participants. Multiple areas, including auditory cortex on the superior temporal plane, featured AEP novelty effect within the time frame of P3a and N400 scalp-recorded novelty-related potentials. This work provides a detailed account of auditory novelty in a paradigm that directly examined brain regions associated with semantic processing. Future studies may aid in the development of objective measures to assess the integrity of semantic novelty processing in clinical populations.

Penalized canonical correlation analysis reveals a relationship between temperament traits and brain oscillations during mind wandering.

INTRODUCTION: There has been a growing interest in studying brain activity under naturalistic conditions. However, the relationship between individual differences in ongoing brain activity and psychological characteristics is not well understood. We investigated this connection, focusing on the association between oscillatory activity in the brain and individually characteristic dispositional traits. Given the variability of unconstrained resting states among individuals, we devised a paradigm that could harmonize the state of mind across all participants. METHODS: We constructed task contrasts that included focused attention (FA), self-centered future planning, and rumination on anxious thoughts triggered by visual imagery. Magnetoencephalography was recorded from 28 participants under these 3 conditions for a duration of 16 min. The oscillatory power in the alpha and beta bands was converted into spatial contrast maps, representing the difference in brain oscillation power between the two conditions. We performed permutation cluster tests on these spatial contrast maps. Additionally, we applied penalized canonical correlation analysis (CCA) to study the relationship between brain oscillation patterns and behavioral traits. RESULTS: The data revealed that the FA condition, as compared to the other conditions, was associated with higher alpha and beta power in the temporal areas of the left hemisphere and lower alpha and beta power in the parietal areas of the right hemisphere. Interestingly, the penalized CCA indicated that behavioral inhibition was positively correlated, whereas anxiety was negatively correlated, with a pattern of high oscillatory power in the bilateral precuneus and low power in the bilateral temporal regions. This unique association was found in the anxious-thoughts condition when contrasted with the focused-attention condition. CONCLUSION: Our findings suggest individual temperament traits significantly affect brain engagement in naturalistic conditions. This research underscores the importance of considering individual traits in neuroscience and offers an effective method for analyzing brain activity and psychological differences.

High-resolution EEG source localization in personalized segmentation-free head model with multi-dipole fitting.

Objective. Electroencephalograms (EEGs) are often used to monitor brain activity. Several source localization methods have been proposed to estimate the location of brain activity corresponding to EEG readings. However, only a few studies evaluated source localization accuracy from measured EEG using personalized head models in a millimeter resolution. In this study, based on a volume conductor analysis of a high-resolution personalized human head model constructed from magnetic resonance images, a finite difference method was used to solve the forward problem and to reconstruct the field distribution.Approach. We used a personalized segmentation-free head model developed using machine learning techniques, in which the abrupt change of electrical conductivity occurred at the tissue interface is suppressed. Using this model, a smooth field distribution was obtained to address the forward problem. Next, multi-dipole fitting was conducted using EEG measurements for each subject (N= 10 male subjects, age: 22.5 ± 0.5), and the source location and electric field distribution were estimated.Main results.For measured somatosensory evoked potential for electrostimulation to the wrist, a multi-dipole model with lead field matrix computed with the volume conductor model was found to be superior than a single dipole model when using personalized segmentation-free models (6/10). The correlation coefficient between measured and estimated scalp potentials was 0.89 for segmentation-free head models and 0.71 for conventional segmented models. The proposed method is straightforward model development and comparable localization difference of the maximum electric field from the target wrist reported using fMR (i.e. 16.4 ± 5.2 mm) in previous study. For comparison, DUNEuro based on sLORETA was (EEG: 17.0 ± 4.0 mm). In addition, somatosensory evoked magnetic fields obtained by Magnetoencephalography was 25.3 ± 8.5 mm using three-layer sphere and sLORETA.Significance. For measured EEG signals, our procedures using personalized head models demonstrated that effective localization of the somatosensory cortex, which is located in a non-shallower cortex region. This method may be potentially applied for imaging brain activity located in other non-shallow regions.

Extensive functional connectivity between brain areas implicated in mental imagery production and phobic fear during both emotional and neutral mental imagery.

Mental imagery is used by most people in their day-to-day cognition, for example, in planning, daydreaming, or remembering. Importantly, mental imagery has a powerful influence on emotion and is critically involved in many mental disorders. Thus, understanding the link between mental imagery and emotion is of clinical interest. For example, exposure therapy can be successfully conducted using mental imagery of fear-provoking stimuli, i.e., imaginal exposure. In this vein, accumulating evidence shows that mental imagery of a fearful stimulus produces a similar physiological and neural response as actual perception of the stimulus. Alas, knowledge of the neural processes underlying the link between mental imagery and emotion is limited. Functional magnetic resonance imaging data from a previous study on imaginal exposure (N = 30) was used to examine the functional connectivity during the production of phobic and neutral mental imagery. Regions of interest were selected from meta-analyses on brain regions consistently recruited during mental imagery production and phobic fear, respectively. Results showed that these regions were positively correlated during both phobic and neutral mental imagery production. Very few differences in functional connectivity between phobic and neutral imagery were found. Specifically, weaker functional connectivity between the supplemental motor area and a region including parts of the left insula and inferior frontal gyrus was observed during phobic (vs neutral) imagery. In conclusion, our findings suggest that brain regions previously implicated in mental imagery production and phobic fear are highly interconnected during the production of both phobic and neutral imagery.

Older adults preserve audiovisual integration through enhanced cortical activations, not by recruiting new regions.

Effective interactions with the environment rely on the integration of multisensory signals: Our brains must efficiently combine signals that share a common source, and segregate those that do not. Healthy ageing can change or impair this process. This functional magnetic resonance imaging study assessed the neural mechanisms underlying age differences in the integration of auditory and visual spatial cues. Participants were presented with synchronous audiovisual signals at various degrees of spatial disparity and indicated their perceived sound location. Behaviourally, older adults were able to maintain localisation accuracy. At the neural level, they integrated auditory and visual cues into spatial representations along dorsal auditory and visual processing pathways similarly to their younger counterparts but showed greater activations in a widespread system of frontal, temporal, and parietal areas. According to multivariate Bayesian decoding, these areas encoded critical stimulus information beyond that which was encoded in the brain areas commonly activated by both groups. Surprisingly, however, the boost in information provided by these areas with age-related activation increases was comparable across the 2 age groups. This dissociation-between comparable information encoded in brain activation patterns across the 2 age groups, but age-related increases in regional blood-oxygen-level-dependent responses-contradicts the widespread notion that older adults recruit new regions as a compensatory mechanism to encode task-relevant information. Instead, our findings suggest that activation increases in older adults reflect nonspecific or modulatory mechanisms related to less efficient or slower processing, or greater demands on attentional resources.

Cortical depth profiles in primary visual cortex for illusory and imaginary experiences.

Visual illusions and mental imagery are non-physical sensory experiences that involve cortical feedback processing in the primary visual cortex. Using laminar functional magnetic resonance imaging (fMRI) in two studies, we investigate if information about these internal experiences is visible in the activation patterns of different layers of primary visual cortex (V1). We find that imagery content is decodable mainly from deep layers of V1, whereas seemingly 'real' illusory content is decodable mainly from superficial layers. Furthermore, illusory content shares information with perceptual content, whilst imagery content does not generalise to illusory or perceptual information. Together, our results suggest that illusions and imagery, which differ immensely in their subjective experiences, also involve partially distinct early visual microcircuits. However, overlapping microcircuit recruitment might emerge based on the nuanced nature of subjective conscious experience.

Should I stay or should I go? The cerebral bases of street-crossing decision.

An observer willing to cross a street must first estimate if the approaching cars offer enough time to safely complete the task. The brain areas supporting this perception, known as Time-To-Contact (TTC) perception, have been mainly studied through noninvasive correlational approaches. We carried out an experiment in which patients were tested during an awake brain surgery electrostimulation mapping to examine the causal implication of various brain areas in the street-crossing decision process. Forty patients were tested in a gap acceptance task before their surgery to establish a baseline performance. The task was individually adapted upon this baseline level and carried out during their surgery. We acquired and normalized to MNI space the coordinates of the functional areas that influenced task performance. A total of 103 stimulation sites were tested, allowing to establish a large map of the areas involved in the street-crossing decision. Multiple sites were found to impact the gap acceptance decision. A direct implication was however found mostly for sites within the right parietal lobe, while indirect implication was found for sites within the language, motor, or attentional networks. The right parietal lobe can be considered as causally influencing the gap acceptance decision. Other positive sites were all accompanied with dysfunction in other cognitive functions, and therefore should probably not be considered as the site of TTC estimation.

Neural response properties predict perceived contents and locations elicited by intracranial electrical stimulation of human auditory cortex.

Intracranial electrical stimulation (iES) of auditory cortex can elicit sound experiences with a variety of perceived contents (hallucination or illusion) and locations (contralateral or bilateral side), independent of actual acoustic inputs. However, the neural mechanisms underlying this elicitation heterogeneity remain undiscovered. Here, we collected subjective reports following iES at 3062 intracranial sites in 28 patients (both sexes) and identified 113 auditory cortical sites with iES-elicited sound experiences. We then decomposed the sound-induced intracranial electroencephalogram (iEEG) signals recorded from all 113 sites into time-frequency features. We found that the iES-elicited perceived contents can be predicted by the early high-γ features extracted from sound-induced iEEG. In contrast, the perceived locations elicited by stimulating hallucination sites and illusion sites are determined by the late high-γ and long-lasting α features, respectively. Our study unveils the crucial neural signatures of iES-elicited sound experiences in human and presents a new strategy to hearing restoration for individuals suffering from deafness.

Visual mental imagery: Evidence for a heterarchical neural architecture.

Theories of Visual Mental Imagery (VMI) emphasize the processes of retrieval, modification, and recombination of sensory information from long-term memory. Yet, only few studies have focused on the behavioral mechanisms and neural correlates supporting VMI of stimuli from different semantic domains. Therefore, we currently have a limited understanding of how the brain generates and maintains mental representations of colors, faces, shapes - to name a few. Such an undetermined scenario renders unclear the organizational structure of neural circuits supporting VMI, including the role of the early visual cortex. We aimed to fill this gap by reviewing the scientific literature of five semantic domains: visuospatial, face, colors, shapes, and letters imagery. Linking theory to evidence from over 60 different experimental designs, this review highlights three main points. First, there is no consistent activity in the early visual cortex across all VMI domains, contrary to the prediction of the dominant model. Second, there is consistent activity of the frontoparietal networks and the left hemisphere's fusiform gyrus during voluntary VMI irrespective of the semantic domain investigated. We propose that these structures are part of a domain-general VMI sub-network. Third, domain-specific information engages specific regions of the ventral and dorsal cortical visual pathways. These regions partly overlap with those found in visual perception studies (e.g., fusiform face area for faces imagery; lingual gyrus for color imagery). Altogether, the reviewed evidence suggests the existence of domain-general and domain-specific mechanisms of VMI selectively engaged by stimulus-specific properties (e.g., colors or faces). These mechanisms would be supported by an organizational structure mixing vertical and horizontal connections (heterarchy) between sub-networks for specific stimulus domains. Such a heterarchical organization of VMI makes different predictions from current models of VMI as reversed perception. Our conclusions set the stage for future research, which should aim to characterize the spatiotemporal dynamics and interactions among key regions of this architecture giving rise to visual mental images.