Changes in spatial self-consciousness elicit grid cell-like representation in the entorhinal cortex.

Grid cells in the entorhinal cortex (EC) encode an individual's location in space, integrating both environmental and multisensory bodily cues. Notably, body-derived signals are also primary signals for the sense of self. While studies have demonstrated that continuous application of visuo-tactile bodily stimuli can induce perceptual shifts in self-location, it remains unexplored whether these illusory changes suffice to trigger grid cell-like representation (GCLR) within the EC, and how this compares to GCLR during conventional virtual navigation. To address this, we systematically induced illusory drifts in self-location toward controlled directions using visuo-tactile bodily stimulation, while maintaining the subjects' visual viewpoint fixed (absent conventional virtual navigation). Subsequently, we evaluated the corresponding GCLR in the EC through functional MRI analysis. Our results reveal that illusory changes in perceived self-location (independent of changes in environmental navigation cues) can indeed evoke entorhinal GCLR, correlating in strength with the magnitude of perceived self-location, and characterized by similar grid orientation as during conventional virtual navigation in the same virtual room. These data demonstrate that the same grid-like representation is recruited when navigating based on environmental, mainly visual cues, or when experiencing illusory forward drifts in self-location, driven by perceptual multisensory bodily cues.

Experimentally-evidenced personality alterations following meningioma resection: A case report.

Personality changes following neurosurgical procedures remain poorly understood and pose a major concern for patients, rendering a strong need for predictive biomarkers. Here we report a case of a female patient in her 40s who underwent resection of a large sagittal sinus meningioma with bilateral extension, including resection and ligation of the superior sagittal sinus, that resulted in borderline personality disorder. Importantly, we captured clinically-observed personality changes in a series of experiments assessing self-other voice discrimination, one of the experimental markers for self-consciousness. In all experiments, the patient consistently confused self- and other voices - i.e., she misattributed other-voice stimuli to herself and self-voice stimuli to others. Moreover, the electroencephalogram (EEG) microstate, that was in healthy participants observed when hearing their own voice, in this patient occurred for other-voice stimuli. We hypothesize that the patient's personality alterations resulted from a gradual development of a venous collateral hemodynamic network that impacted venous drainage of brain areas associated with self-consciousness. In addition, resection and ligation of the superior sagittal sinus significantly aggravated personality alterations through postoperative decompensation of a direct frontal lobe compression. Experimentally mirroring clinical observations, these findings are of high relevance for developing biomarkers of post-surgical personality alterations.

Image Mapping Accuracy Evaluation Using UAV with Standalone, Differential (RTK), and PPP GNSS Positioning Techniques in an Abandoned Mine Site.

Global navigation satellite systems (GNSSs) provide a common positioning method that utilizes satellite signals to determine the spatial location of a receiver. However, there are several error factors in standalone GNSS positioning due to instrumental, procedural, and environmental factors that arise during the signal transmission process, and the final positioning error can be up to several meters or greater in length. Thus, real-time kinematic (RTK) correction and post-mission precise point positioning (PPP) processing technologies are proposed to improve accuracy and accomplish precise position measurements. To evaluate the geolocation accuracy of mosaicked UAV images of an abandoned mine site, we compared each orthomosaic image and digital elevation model obtained using standalone GNSS positioning, differential (RTK) GNSS positioning, and post-mission PPP processing techniques. In the three types of error evaluation measure (i.e., relative camera location error, ground control points-based absolute image mapping error, and volumetric difference of mine tailings), we found that the RTK GNSS positioning method obtained the best performance in terms of the relative camera location error and the absolute image mapping error evaluations, and the PPP post-processing correction effectively reduced the error (69.5% of the average total relative camera location error and 59.3% of the average total absolute image mapping error) relative to the standalone GNSS positioning method. Although differential (RTK) GNSS positioning is widely used in positioning applications that require very high accuracy, post-mission PPP processing can also be used in various fields in which it is either not feasible to operate expensive equipment to receive RTK GNSS signals or network RTK services are unavailable.

Breathing is coupled with voluntary initiation of mental imagery.

Previous research has suggested that bodily signals from internal organs are associated with diverse cortical and subcortical processes involved in sensory-motor functions, beyond homeostatic reflexes. For instance, a recent study demonstrated that the preparation and execution of voluntary actions, as well as its underlying neural activity, are coupled with the breathing cycle. In the current study, we investigated whether such breathing-action coupling is limited to voluntary motor action or whether it is also present for mental actions not involving any overt bodily movement. To answer this question, we recorded electroencephalography (EEG), electromyography (EMG), and respiratory signals while participants were conducting a voluntary action paradigm including self-initiated motor execution (ME), motor imagery (MI), and visual imagery (VI) tasks. We observed that the voluntary initiation of ME, MI, and VI are similarly coupled with the respiration phase. In addition, EEG analysis revealed the existence of readiness potential (RP) waveforms in all three tasks (i.e., ME, MI, VI), as well as a coupling between the RP amplitude and the respiratory phase. Our findings show that the voluntary initiation of both imagined and overt action is coupled with respiration, and further suggest that the breathing system is involved in preparatory processes of voluntary action by contributing to the temporal decision of when to initiate the action plan, regardless of whether this culminates in overt movements.

Sense of self impacts spatial navigation and hexadirectional coding in human entorhinal cortex.

Grid cells in entorhinal cortex (EC) encode an individual's location in space and rely on environmental cues and self-motion cues derived from the individual's body. Body-derived signals are also primary signals for the sense of self and based on integrated sensorimotor signals (proprioceptive, tactile, visual, motor) that have been shown to enhance self-centered processing. However, it is currently unknown whether such sensorimotor signals that modulate self-centered processing impact grid cells and spatial navigation. Integrating the online manipulation of bodily signals, to modulate self-centered processing, with a spatial navigation task and an fMRI measure to detect grid cell-like representation (GCLR) in humans, we report improved performance in spatial navigation and decreased GCLR in EC. This decrease in entorhinal GCLR was associated with an increase in retrosplenial cortex activity, which was correlated with participants' navigation performance. These data link self-centered processes during spatial navigation to entorhinal and retrosplenial activity and highlight the role of different bodily factors at play when navigating in VR.

Breathing affects self-other voice discrimination in a bodily state associated with somatic passivity.

A growing number of studies have focused on identifying cognitive processes that are modulated by interoceptive signals, particularly in relation to the respiratory or cardiac cycle. Considering the fundamental role of interoception in bodily self-consciousness, we here investigated whether interoceptive signals also impact self-voice perception. We applied an interactive, robotic paradigm associated with somatic passivity (a bodily state characterized by illusory misattribution of self-generated touches to someone else) to investigate whether somatic passivity impacts self-voice perception as a function of concurrent interoceptive signals. Participants' breathing and heartbeat signals were recorded while they performed two self-voice tasks (self-other voice discrimination and loudness perception) and while simultaneously experiencing two robotic conditions (somatic passivity condition; control condition). Our data reveal that respiration, but not cardiac activity, affects self-voice perception: participants were better at discriminating self-voice from another person's voice during the inspiration phase of the respiration cycle. Moreover, breathing effects were prominent in participants experiencing somatic passivity and a different task with the same stimuli (i.e., judging the loudness and not identity of the voices) was unaffected by breathing. Combining interoception and voice perception with self-monitoring framework, these data extend findings on breathing-dependent changes in perception and cognition to self-related processing.

The perspectives of mapping and monitoring of the sense of self in neurosurgical patients.

Surgical treatment of tumors, epileptic foci or of vascular origin, requires a detailed individual pre-surgical workup and intra-operative surveillance of brain functions to minimize the risk of post-surgical neurological deficits and decline of quality of life. Most attention is attributed to language, motor functions, and perception. However, higher cognitive functions such as social cognition, personality, and the sense of self may be affected by brain surgery. To date, the precise localization and the network patterns of brain regions involved in such functions are not yet fully understood, making the assessment of risks of related post-surgical deficits difficult. It is in the interest of neurosurgeons to understand with which neural systems related to selfhood and personality they are interfering during surgery. Recent neuroscience research using virtual reality and clinical observations suggest that the insular cortex, medial prefrontal cortex, and temporo-parietal junction are important components of a neural system dedicated to self-consciousness based on multisensory bodily processing, including exteroceptive and interoceptive cues (bodily self-consciousness (BSC)). Here, we argue that combined extra- and intra-operative approaches using targeted cognitive testing, functional imaging and EEG, virtual reality, combined with multisensory stimulations, may contribute to the assessment of the BSC and related cognitive aspects. Although the usefulness of particular biomarkers, such as cardiac and respiratory signals linked to virtual reality, and of heartbeat evoked potentials as a surrogate marker for intactness of multisensory integration for intra-operative monitoring has to be proved, systemic and automatized testing of BSC in neurosurgical patients will improve future surgical outcome.

Behavioral and neurophysiological evidence for altered interoceptive bodily processing in chronic pain.

Whereas impaired multisensory processing of bodily stimuli and distorted body representation are well-established in various chronic pain disorders, such research has focused on exteroceptive bodily cues and neglected bodily signals from the inside of the body (or interoceptive signals). Extending existing basic and clinical research, we investigated for the first time interoception and its neurophysiological correlates in patients with complex regional pain syndrome (CRPS). In three different experiments, including a total of 36 patients with CRPS and 42 aged-gender matched healthy controls, we measured interoceptive sensitivity (heart beat counting task, HBC) and neural responses to heartbeats (heartbeat evoked potentials, HEPs). As hypothesized, we observed reduced sensitivity in perceiving interoceptive bodily stimuli, i.e. their heartbeat, in two independent samples of CRPS patients (studies 1 and 2). Moreover, the cortical processing of their heartbeat, i.e. the HEP, was reduced compared to controls (study 3) and reduced interoceptive sensitivity and HEPs were related to CRPS patients' motor impairment and pain duration. By providing consistent evidence for impaired processing of interoceptive bodily cues in CRPS, this study shows that the perceptual changes occurring in chronic pain include signals originating from the visceral organs, suggesting changes in the neural body representation, that includes next to exteroceptive, also interoceptive bodily signals. By showing that impaired interoceptive processing is associated with clinical symptoms, our findings also encourage the use of interoceptive-related information in future rehabilitation for chronic pain.

Breathing is coupled with voluntary action and the cortical readiness potential.

Voluntary action is a fundamental element of self-consciousness. The readiness potential (RP), a slow drift of neural activity preceding self-initiated movement, has been suggested to reflect neural processes underlying the preparation of voluntary action; yet more than fifty years after its introduction, interpretation of the RP remains controversial. Based on previous research showing that internal bodily signals affect sensory processing and ongoing neural activity, we here investigated the potential role of interoceptive signals in voluntary action and the RP. We report that (1) participants initiate voluntary actions more frequently during expiration, (2) this respiration-action coupling is absent during externally triggered actions, and (3) the RP amplitude is modulated depending on the respiratory phase. Our findings demonstrate that voluntary action is coupled with the respiratory system and further suggest that the RP is associated with fluctuations of ongoing neural activity that are driven by the involuntary and cyclic motor act of breathing.

Heartbeat-evoked cortical responses: Underlying mechanisms, functional roles, and methodological considerations.

The heart continuously and cyclically communicates with the brain. Beyond homeostatic regulation and sensing, recent neuroscience research has started to shed light on brain-heart interactions in diverse cognitive and emotional processes. In particular, neural responses to heartbeats, as measured with the so-called heartbeat-evoked potential, have been shown to be useful for investigating cortical activity processing cardiac signals. In this review, we first overview and discuss the basic properties of the HEP such as underlying physiological pathways, brain regions, and neural mechanisms. We then provide a systematic review of the mental processes associated with cortical HEP activations, notably heartbeat perception, emotional feelings, perceptual awareness, and self-consciousness, in healthy subjects and clinical populations. Finally, we discuss methodological issues regarding the experimental design and data analysis for separating genuine HEP components from physiological artifacts (e.g., cardiac field artifact, pulse artifact) or other neural activities that are not specifically associated with the heartbeat. Findings from this review suggest that when intrinsic limitations (e.g., artifacts) are carefully controlled, the HEP could provide a reliable neural measure for investigating brain-viscera interactions in diverse mental processes.

Coupling Inner and Outer Body for Self-Consciousness.

Although recent studies on self-consciousness emphasized the importance of bodily processing and multisensory integration, such research has focused solely on bodily signals originating from the outside of the body (i.e., exteroceptive bodily signals) or internal bodily signals from visceral organs (i.e., interoceptive bodily signals) and how each system contributes to self-consciousness, without much interaction between the two approaches. Reviewing the latest evidence on interoceptive bodily processing and the combination of exteroceptive and interoceptive bodily signals for self-consciousness, we propose an integrated neural system reconciling these two largely separated views and delineate how it accounts for fundamental aspects of self-consciousness such as self-identification and self-location, as well as its experienced global unity and temporal continuity.

Increased heartbeat-evoked potential during REM sleep in nightmare disorder.

Nightmares are characterized by the experience of strong negative emotions occurring mainly during REM sleep. Some people suffer from nightmare disorder, which is defined by the repeated occurrence of nightmares and by significant distress in wakefulness. Yet, whether frequent nightmares relate to a general increase in emotional reactivity or arousal during sleep remains unclear. To address this question, we recorded heartbeat-evoked potentials (HEPs) during wakefulness, NREM and REM sleep in patients with nightmare disorder and healthy participants. The HEP represents a cortical (EEG) response to the heartbeat and indexes brain-body interactions, such as interoceptive processing and intrinsic levels of arousal. HEP amplitude is typically increased during states of high emotional arousal and motivation, and is decreased in depression. Here we compared the amplitude of HEPs between nightmare patients and healthy controls separately during AWAKE, NREM, REM periods, and found higher HEP amplitude in nightmare patients compared to healthy controls over a cluster of frontal regions only during REM sleep. This effect was not paralleled by any group difference in cardiac control measures (e.g. heart rate variability, interbeat interval). These findings corroborate the notion that nightmares are essentially a REM pathology and suggest that increased emotional arousal during REM sleep, as measured by HEP, is a physiological condition responsible for frequent nightmares. This result also supports that HEP may be used as a biomarker of increased emotional and sensory processing during REM sleep in these patients.

Sad faces increase the heartbeat-associated interoceptive information flow within the salience network: a MEG study.

The somatic marker hypothesis proposes that the cortical representation of visceral signals is a crucial component of emotional processing. No previous study has investigated the information flow among brain regions that process visceral information during emotional perception. In this magnetoencephalography study of 32 healthy subjects of either sex, heartbeat-evoked responses (HERs), which reflect the cortical processing of heartbeats, were modulated by the perception of a sad face. The modulation effect was localized to the prefrontal cortices, the globus pallidus, and an interoceptive network including the right anterior insula (RAI) and dorsal anterior cingulate cortex (RdACC). Importantly, our Granger causality analysis provides the first evidence for the increased flow of heartbeat information from the RAI to the RdACC during sad face perception. Moreover, using a surrogate R-peak analysis, we have shown that this HER modulation effect was time-locked to heartbeats. These findings advance the understanding of brain-body interactions during emotional processing.

Audio-Tactile and Peripersonal Space Processing Around the Trunk in Human Parietal and Temporal Cortex: An Intracranial EEG Study.

Interactions with the environment happen within one's peripersonal space (PPS)-the space surrounding the body. Studies in monkeys and humans have highlighted a multisensory distributed cortical network representing the PPS. However, knowledge about the temporal dynamics of PPS processing around the trunk is lacking. Here, we recorded intracranial electroencephalography (iEEG) in humans while administering tactile stimulation (T), approaching auditory stimuli (A), and the 2 combined (AT). To map PPS, tactile stimulation was delivered when the sound was far, intermediate, or close to the body. The 19% of the electrodes showed AT multisensory integration. Among those, 30% showed a PPS effect, a modulation of the response as a function of the distance between the sound and body. AT multisensory integration and PPS effects had similar spatiotemporal characteristics, with an early response (~50 ms) in the insular cortex, and later responses (~200 ms) in precentral and postcentral gyri. Superior temporal cortex showed a different response pattern with AT multisensory integration at ~100 ms without a PPS effect. These results, represent the first iEEG delineation of PPS processing in humans and show that PPS and multisensory integration happen at similar neural sites and time periods, suggesting that PPS representation is based on a spatial modulation of multisensory integration.

Audio-visual sensory deprivation degrades visuo-tactile peri-personal space.

Self-perception is scaffolded upon the integration of multisensory cues on the body, the space surrounding the body (i.e., the peri-personal space; PPS), and from within the body. We asked whether reducing information available from external space would change: PPS, interoceptive accuracy, and self-experience. Twenty participants were exposed to 15 min of audio-visual deprivation and performed: (i) a visuo-tactile interaction task measuring their PPS; (ii) a heartbeat perception task measuring interoceptive accuracy; and (iii) a series of questionnaires related to self-perception and mental illness. These tasks were carried out in two conditions: while exposed to a standard sensory environment and under a condition of audio-visual deprivation. Results suggest that while PPS becomes ill defined after audio-visual deprivation, interoceptive accuracy is unaltered at a group-level, with some participants improving and some worsening in interoceptive accuracy. Interestingly, correlational individual differences analyses revealed that changes in PPS after audio-visual deprivation were related to interoceptive accuracy and self-reports of "unusual experiences" on an individual subject basis. Taken together, the findings argue for a relationship between the malleability of PPS, interoceptive accuracy, and an inclination toward aberrant ideation often associated with mental illness.

Bodily self-consciousness and its disorders.

Research in clinical and human neuroscience indicates that important brain mechanisms of self-consciousness are based on the integration of multisensory bodily signals (i.e., bodily self-consciousness: BSC), including signals coming from outside our body (i.e., exteroceptive signals, such as tactile, auditory, and visual information) and the inside of our body (i.e., interoceptive signals). In this chapter, we discuss selected behavioral and neuroimaging studies about how multisensory integration generates and modulates BSC in humans, with particular relevance to parietal mechanisms. We then review the neurology of disorders of BSC after acquired brain damage or dysfunction, ranging from body attentional disorders to delusional and illusory deficits about the patient's own body, associated with a breakdown of the link between the body and the self.

Neural Sources and Underlying Mechanisms of Neural Responses to Heartbeats, and their Role in Bodily Self-consciousness: An Intracranial EEG Study.

Recent research has shown that heartbeat-evoked potentials (HEPs), brain activity in response to heartbeats, are a useful neural measure for investigating the functional role of brain-body interactions in cognitive processes including self-consciousness. In 2 experiments, using intracranial electroencephalography (EEG), we investigated (1) the neural sources of HEPs, (2) the underlying mechanisms for HEP generation, and (3) the functional role of HEPs in bodily self-consciousness. In Experiment-1, we found that shortly after the heartbeat onset, phase distributions across single trials were significantly concentrated in 10% of the recording sites, mainly in the insula and the operculum, but also in other regions including the amygdala and fronto-temporal cortex. Such phase concentration was not accompanied by increased spectral power, and did not correlate with spectral power changes, suggesting that a phase resetting, rather than an additive "evoked potential" mechanism, underlies HEP generation. In Experiment-2, we further aimed to anatomically refine previous scalp EEG data that linked HEPs with bodily self-consciousness. We found that HEP modulations in the insula reflected an experimentally induced altered sense of self-identification. Collectively, these results provide novel and solid electrophysiological evidence on the neural sources and underlying mechanisms of HEPs, and their functional role in self-consciousness.

The neural monitoring of visceral inputs, rather than attention, accounts for first-person perspective in conscious vision.

Why should a scientist whose aim is to unravel the neural mechanisms of perception consider brain-body interactions seriously? Brain-body interactions have traditionally been associated with emotion, effort, or stress, but not with the "cold" processes of perception and attention. Here, we review recent experimental evidence suggesting a different picture: the neural monitoring of bodily state, and in particular the neural monitoring of the heart, affects visual perception. The impact of spontaneous fluctuations of neural responses to heartbeats on visual detection is as large as the impact of explicit manipulations of spatial attention in perceptual tasks. However, we propose that the neural monitoring of visceral inputs plays a specific role in conscious perception, distinct from the role of attention. The neural monitoring of organs such as the heart or the gut would generate a subject-centered reference frame, from which the first-person perspective inherent to conscious perception can develop. In this view, conscious perception results from the integration of visual content with first-person perspective.

Transient Modulations of Neural Responses to Heartbeats Covary with Bodily Self-Consciousness.

UNLABELLED: Recent research has investigated self-consciousness associated with the multisensory processing of bodily signals (e.g., somatosensory, visual, vestibular signals), a notion referred to as bodily self-consciousness, and these studies have shown that the manipulation of bodily inputs induces changes in bodily self-consciousness such as self-identification. Another line of research has highlighted the importance of signals from the inside of the body (e.g., visceral signals) and proposed that neural representations of internal bodily signals underlie self-consciousness, which to date has been based on philosophical inquiry, clinical case studies, and behavioral studies. Here, we investigated the relationship of bodily self-consciousness with the neural processing of internal bodily signals. By combining electrical neuroimaging, analysis of peripheral physiological signals, and virtual reality technology in humans, we show that transient modulations of neural responses to heartbeats in the posterior cingulate cortex covary with changes in bodily self-consciousness induced by the full-body illusion. Additional analyses excluded that measured basic cardiorespiratory parameters or interoceptive sensitivity traits could account for this finding. These neurophysiological data link experimentally the cortical mapping of the internal body to self-consciousness. SIGNIFICANCE STATEMENT: What are the brain mechanisms of self-consciousness? Prominent views propose that the neural processing associated with signals from the internal organs (such as the heart and the lung) plays a critical role in self-consciousness. Although this hypothesis dates back to influential views in philosophy and psychology (e.g., William James), definitive experimental evidence supporting this idea is lacking despite its recent impact in neuroscience. In the present study, we show that posterior cingulate activities responding to heartbeat signals covary with changes in participants' conscious self-identification with a body that were manipulated experimentally using virtual reality technology. Our finding provides important neural evidence about the long-standing proposal that self-consciousness is linked to the cortical processing of internal bodily signals.