The Coding Logic of Interoception.

Interoception, the ability to precisely and timely sense internal body signals, is critical for life. The interoceptive system monitors a large variety of mechanical, chemical, hormonal, and pathological cues using specialized organ cells, organ innervating neurons, and brain sensory neurons. It is important for maintaining body homeostasis, providing motivational drives, and regulating autonomic, cognitive, and behavioral functions. However, compared to external sensory systems, our knowledge about how diverse body signals are coded at a system level is quite limited. In this review, we focus on the unique features of interoceptive signals and the organization of the interoceptive system, with the goal of better understanding the coding logic of interoception.

Interoceptive signals shape the earliest markers and neural pathway to awareness at the visual threshold.

Variations in interoceptive signals from the baroreceptors (BRs) across the cardiac and respiratory cycle can modulate cortical excitability and so affect awareness. It remains debated at what stages of processing they affect awareness-related event-related potentials (ERPs) in different sensory modalities. We investigated the influence of the cardiac (systole/diastole) and the respiratory (inhalation/exhalation) phase on awareness-related ERPs. Subjects discriminated visual threshold stimuli while their electroencephalogram, electrocardiogram, and respiration were simultaneously recorded. We compared ERPs and their intracranial generators for stimuli classified correctly with and without awareness as a function of the cardiac and respiratory phase. Cyclic variations of interoceptive signals from the BRs modulated both the earliest electrophysiological markers and the trajectory of brain activity when subjects became aware of the stimuli: an early sensory component (P1) was the earliest marker of awareness for low (diastole/inhalation) and a perceptual component (visual awareness negativity) for high (systole/exhalation) BR activity, indicating that BR signals interfere with the sensory processing of the visual input. Likewise, activity spread from the primary visceral cortex (posterior insula) to posterior parietal cortices during high and from associative interoceptive centers (anterior insula) to the prefrontal cortex during low BR activity. Consciousness is thereby resolved in cognitive/associative regions when BR is low and in perceptual centers when it is high. Our results suggest that cyclic fluctuations of BR signaling affect both the earliest markers of awareness and the brain processes underlying conscious awareness.

When the Heart Meets the Mind: Exploring the Brain-Heart Interaction during Time Perception.

Recent studies suggest that time estimation relies on bodily rhythms and interoceptive signals. We provide the first direct electrophysiological evidence suggesting an association between the brain's processing of heartbeat and duration judgment. We examined heartbeat-evoked potential (HEP) and contingent negative variation (CNV) during an auditory duration-reproduction task and a control reaction-time task spanning 4, 8, and 12 s intervals, in both male and female participants. Interoceptive awareness was assessed with the Self-Awareness Questionnaire (SAQ) and interoceptive accuracy through the heartbeat-counting task (HCT). Results revealed that SAQ scores, but not the HCT, correlated with mean reproduced durations with higher SAQ scores associating with longer and more accurate duration reproductions. Notably, the HEP amplitude changes during the encoding phase of the timing task, particularly within 130-270 ms (HEP1) and 470-520 ms (HEP2) after the R-peak, demonstrated interval-specific modulations that did not emerge in the control task. A significant ramp-like increase in HEP2 amplitudes occurred during the duration-encoding phase of the timing but not during the control task. This increase within the reproduction phase of the timing task correlated significantly with the reproduced durations for the 8 s and the 4 s intervals. The larger the increase in HEP2, the greater the under-reproduction of the estimated duration. CNV components during the encoding phase of the timing task were more negative than those in the reaction-time task, suggesting greater executive resources orientation toward time. We conclude that interoceptive awareness (SAQ) and cortical responses to heartbeats (HEP) predict duration reproductions, emphasizing the embodied nature of time.

Blind individuals' enhanced ability to sense their own heartbeat is related to the thickness of their occipital cortex.

Blindness is associated with heightened sensory abilities, such as improved hearing and tactile acuity. Moreover, recent evidence suggests that blind individuals are better than sighted individuals at perceiving their own heartbeat, suggesting enhanced interoceptive accuracy. Structural changes in the occipital cortex have been hypothesized as the basis of these behavioral enhancements. Indeed, several studies have shown that congenitally blind individuals have increased cortical thickness within occipital areas compared to sighted individuals, but how these structural differences relate to behavioral enhancements is unclear. This study investigated the relationship between cardiac interoceptive accuracy and cortical thickness in 23 congenitally blind individuals and 23 matched sighted controls. Our results show a significant positive correlation between performance in a heartbeat counting task and cortical thickness only in the blind group, indicating a connection between structural changes in occipital areas and blind individuals' enhanced ability to perceive heartbeats.

PIEZO ion channels: force sensors of the interoceptive nervous system.

Many organs are designed to move: the heart pumps each second, the gastrointestinal tract squeezes and churns to digest food, and we contract and relax skeletal muscles to move our bodies. Sensory neurons of the peripheral nervous system detect signals from bodily tissues, including the forces generated by these movements, to control physiology. The processing of these internal signals is called interoception, but this is a broad term that includes a wide variety of both chemical and mechanical sensory processes. Mechanical senses are understudied, but rapid progress has been made in the last decade, thanks in part to the discovery of the mechanosensory PIEZO ion channels (Coste et al., 2010). The role of these mechanosensors within the interoceptive nervous system is the focus of this review. In defining the transduction molecules that govern mechanical interoception, we will have a better grasp of how these signals drive physiology.

Homeostatic Feelings and the Emergence of Consciousness.

In this article, we summarize our views on the problem of consciousness and outline the current version of a novel hypothesis for how conscious minds can be generated in mammalian organisms. We propose that a mind can be considered conscious when three processes are in place: the first is a continuous generation of interoceptive feelings, which results in experiencing of the organism's internal operations; the second is the equally continuous production of images, generated according to the organism's sensory perspective relative to its surround; the third combines feeling/experience and perspective resulting in a process of subjectivity relative to the image contents. We also propose a biological basis for these three components: the peripheral and central physiology of interoception and exteroception help explain the implementation of the first two components, whereas the third depends on central nervous system integration, at multiple levels, from spinal cord, brainstem, and diencephalic nuclei, to selected regions of the mesial cerebral cortices.

Listen to your heart: Trade-off between cardiac interoceptive processing and visual exteroceptive processing.

Internal bodily signals, such as heartbeats, can influence conscious perception of external sensory information. Spontaneous shifts of attention between interoception and exteroception have been proposed as the underlying mechanism, but direct evidence is lacking. Here, we used steady-state visual evoked potential (SSVEP) frequency tagging to independently measure the neural processing of visual stimuli that were concurrently presented but varied in heartbeat coupling in healthy participants. Although heartbeat coupling was irrelevant to participants' task of detecting brief color changes, we found decreased SSVEPs for systole-coupled stimuli and increased SSVEPs for diastole-coupled stimuli, compared to non-coupled stimuli. These results suggest that attentional and representational resources allocated to visual stimuli vary according to fluctuations in cardiac-related signals across the cardiac cycle, reflecting spontaneous and immediate competition between cardiac-related signals and visual events. Furthermore, frequent coupling of visual stimuli with stronger cardiac-related signals not only led to a larger heartbeat evoked potential (HEP) but also resulted in a smaller color change evoked N2 component, with the increase in HEP amplitude associated with a decrease in N2 amplitude. These findings indicate an overall or longer-term increase in brain resources allocated to the internal domain at the expense of reduced resources available for the external domain. Our study highlights the dynamic reallocation of limited processing resources across the internal-external axis and supports the trade-off between interoception and exteroception.

Heartbeat-related spectral perturbation of electroencephalogram reflects dynamic interoceptive attention states in the trial-by-trial classification analysis.

Attending to heartbeats for interoceptive awareness initiates distinct electrophysiological responses synchronized with the R-peaks of an electrocardiogram (ECG), such as the heartbeat-evoked potential (HEP). Beyond HEP, this study proposes heartbeat-related spectral perturbation (HRSP), a time-frequency map of the R-peak locked electroencephalogram (EEG), and explores its characteristics in identifying interoceptive attention states using a classification approach. HRSPs of EEG brain components specified by independent component analysis (ICA) were used for the offline and online classification of interoceptive states. A convolutional neural network (CNN) designed specifically for HRSP was applied to publicly available data from a binary-state experiment (attending to self-heartbeats and white noise) and data from our four-state classification experiment (attending to self-heartbeats, white noise, time passage, and toe) with diverse input feature conditions of HRSP. From the dynamic state perspective, we evaluated the primary frequency bands of HRSP and the minimal number of averaging epochs required to reflect changing interoceptive attention states without compromising accuracy. We also assessed the utility of group ICA and models for classifying HRSP in new participants. The CNN for trial-by-trial HRSP with actual R-peaks demonstrated significantly higher classification accuracy than HRSP with sham, i.e., randomly positioned, R-peaks. Gradient-weighted class activation mapping highlighted the prominent role of theta and alpha bands between 200-600 ms post-R-peak-features absent in classifications using sham HRSPs. Online classification benefits from employing a group ICA and classification model, ensuring reliable accuracy without individual EEG precollection. These results suggest HRSP's potential to reflect interoceptive attention states, proposing transformative implications for clinical applications.

The anterior cingulate cortex is involved in intero-exteroceptive integration for spatial image transformation of the self-body.

Spatial image transformation of the self-body is a fundamental function of visual perspective-taking. Recent research underscores the significance of intero-exteroceptive information integration to construct representations of our embodied self. This raises the intriguing hypothesis that interoceptive processing might be involved in the spatial image transformation of the self-body. To test this hypothesis, the present study used functional magnetic resonance imaging to measure brain activity during an arm laterality judgment (ALJ) task. In this task, participants were tasked with discerning whether the outstretched arm of a human figure, viewed from the front or back, was the right or left hand. The reaction times for the ALJ task proved longer when the stimulus presented orientations of 0°, 90°, and 270° relative to the upright orientation, and when the front view was presented rather than the back view. Reflecting the increased reaction time, increased brain activity was manifested in a cluster centered on the dorsal anterior cingulate cortex (ACC), suggesting that the activation reflects the involvement of an embodied simulation in ALJ. Furthermore, this cluster of brain activity exhibited overlap with regions where the difference in activation between the front and back views positively correlated with the participants' interoceptive sensitivity, as assessed through the heartbeat discrimination task, within the pregenual ACC. These results suggest that the ACC plays an important role in integrating intero-exteroceptive cues to spatially transform the image of our self-body.

Where alcohol use disorder meets interoception: A meta-analytic view on structural and functional neuroimaging data.

Alcohol use disorder (AUD) has been associated with changes in the processing of internal body signals, known as interoception. Changes in brain structure, particularly in the insula, are thought to underlie impaired interoception. As studies specifically investigating this association are largely lacking, this analysis takes an approach that compares meta-analytic results on interoception with recently published meta-analytic results on gray matter reduction in AUD. A systematic literature search identified 25 eligible interoception studies. Activation likelihood estimation (ALE) was used to test for spatial convergence of study results. Overlap between interoception and AUD clusters was tested using conjunction analysis. Meta-analytic connectivity modeling (MACM) and resting-state functional connectivity were used to identify the functional network of interoception and to test where this network overlapped with AUD meta-analytic clusters. The results were characterized using behavioral domain analysis. The interoception ALE identified a cluster in the left middle insula. There was no overlap with clusters of reduced gray matter in AUD. MACM analysis of the interoception cluster revealed a large network located in the insulae, thalami, basal nuclei, cingulate and medial frontal cortices, and pre- and postcentral gyri. Resting state analysis confirmed this result, showing the strongest connections to nodes of the salience- and somatomotor network. Five of the eight clusters that showed a structural reduction in AUD were located within these networks. The behavioral profiles of these clusters were suggestive of higher-level processes such as salience control, somatomotor functions, and skin sensations. The results suggest an altered salience mapping of interoceptive signals in AUD, consistent with current models. Connections to the somatomotor network may be related to action control and integration of skin sensations. Mindfulness-based interventions, pleasurable touch, and (deep) transcranial magnetic stimulation may be targeted interventions that reduce interoceptive deficits in AUD and thus contribute to drug use reduction and relapse prevention.

When the interoceptive and conceptual clash: The case of oppositional phenomenal self-modelling in Tourette syndrome.

Tourette syndrome (TS) has been associated with a rich set of symptoms that are said to be uncomfortable, unwilled, and effortful to manage. Furthermore, tics, the canonical characteristic of TS, are multifaceted, and their onset and maintenance is complex. A formal account that integrates these features of TS symptomatology within a plausible theoretical framework is currently absent from the field. In this paper, we assess the explanatory power of hierarchical generative modelling in accounting for TS symptomatology from the perspective of active inference. We propose a fourfold analysis of sensory, motor, and cognitive phenomena associated with TS. In Section 1, we characterise tics as a form of action aimed at sensory attenuation. In Section 2, we introduce the notion of epistemic ticcing and describe such behaviour as the search for evidence that there is an agent (i.e., self) at the heart of the generative hierarchy. In Section 3, we characterise both epistemic (sensation-free) and nonepistemic (sensational) tics as habitual behaviour. Finally, in Section 4, we propose that ticcing behaviour involves an inevitable conflict between distinguishable aspects of selfhood; namely, between the minimal phenomenal sense of self-which is putatively underwritten by interoceptive inference-and the explicit preferences that constitute the individual's conceptual sense of self. In sum, we aim to provide an empirically informed analysis of TS symptomatology under active inference, revealing a continuity between covert and overt features of the condition.

The neurobiology of aesthetic chills: How bodily sensations shape emotional experiences.

The phenomenon of aesthetic chills-shivers and goosebumps associated with either rewarding or threatening stimuli-offers a unique window into the brain basis of conscious reward because of their universal nature and simultaneous subjective and physical counterparts. Elucidating the neural mechanisms underlying aesthetic chills can reveal fundamental insights about emotion, consciousness, and the embodied mind. What is the precise timing and mechanism of bodily feedback in emotional experience? How are conscious feelings and motivations generated from interoceptive predictions? What is the role of uncertainty and precision signaling in shaping emotions? How does the brain distinguish and balance processing of rewards versus threats? We review neuroimaging evidence and highlight key questions for understanding how bodily sensations shape conscious feelings. This research stands to advance models of brain-body interactions shaping affect and may lead to novel nonpharmacological interventions for disorders of motivation and pleasure.

The Adaptiveness of Emotion Regulation Variability and Interoceptive Attention in Daily Life.

OBJECTIVE: In daily life, we must dynamically and flexibly deploy strategies to regulate our emotions, which depends on awareness of emotions and internal bodily signals. Variability in emotion-regulation strategy use may predict fewer negative emotions, especially when people pay more attention to their bodily states-or have greater "interoceptive attention" (IA). Using experience sampling, this study aimed to test whether IA predicts variability in strategy use and whether this variability and IA together predict negative affect. METHODS: University student participants ( n = 203; 165 females; Mage = 20.68, SD age = 1.84) completed trait questionnaires and reported state levels of IA, emotional awareness, negative affect, and emotion-regulation strategies, seven times daily for 1 week. RESULTS: State IA significantly predicted between-strategy variability, which was mediated by emotional awareness (indirect effect = 0.002, 95% confidence interval = <0.001-0.003). Between-strategy variability was associated with lower negative affect, particularly when individuals had higher state IA (simple slope = -0.83, t = -5.87, p < .001) versus lower IA (simple slope = -0.31, t = -2.62, p = .009). CONCLUSIONS: IA appears to facilitate adaptative emotion regulation and help alleviate negative affect. Findings underscore the key roles of IA and emotion-regulation flexibility in mental health.

Testing the monitor and acceptance theory: the role of training-induced changes in monitoring- and acceptance-related capacities after attention-based, socio-emotional, or socio-cognitive mental training in reducing cortisol stress reactivity.

Mindfulness-based interventions have become a popular means to reduce stress. However, the specific mechanisms driving observed stress reduction remain understudied. The Monitor and Acceptance Theory suggests that the cultivation of monitoring and acceptance skills are necessary moderators of practice-induced stress reduction. In the context of the ReSource Project, a large healthy adult sample underwent three 3-month mental training modules targeting either attentional (Presence module), socio-affective (Affect module) or socio-cognitive skills (Perspective module). In the current study, the development of a range of inter-individual differences in mindfulness-, interoception- and compassion-related traits - which mapped to either monitoring or acceptance categories - was tracked. The relationship of these training-induced changes with cortisol stress reactivity after the three distinct 3-month training modules was explored. We found that stress sensitivity was particularly modulated by a differential adaptivity of one cultivated attentional capacity - Attention regulation - which predicted higher cortisol reactivity after mere attention training (Presence) but was associated with lower stress-induced cortisol release after additional socio-affective and socio-cognitive practice (Affect and Perspective). However, this effect did not survive multiple comparisons correction, and analyses were limited by the sample size available. We conclude that our study provides preliminary support of the Monitor and Acceptance Theory, lending weight to the advantage of primary attentional increases in order to fully harness the beneficial effects of socio-affective training, ultimately leading to stress reduction. Although training-induced increases in acceptance were not directly shown to contribute to lowering cortisol stress reactivity, the data suggest an additional benefit of socio-affective and socio-cognitive training that is not directly captured within the current analyses. Our study corroborates the importance of going beyond the training of attention monitoring to foster stress resilience, and highlights that mental training relies on the co-development of several interacting processes to successfully attenuate stress. Further exploring the overarching concept of acceptance in future research may prove beneficial to the theoretical framework of MAT, and in understanding the processes by which stress reduction occurs.

Generalization of a positive-feature interoceptive morphine occasion setter across the rat estrous cycle.

INTRODUCTION: Interoceptive stimuli elicited by drug administration acquire conditioned modulatory properties of the induction of conditioned appetitive behaviours by exteroceptive cues. This effect may be modeled using a drug discrimination task in which the drug stimulus is trained as a positive-feature (FP) occasion setter (OS) that disambiguates the relation between an exteroceptive light conditioned stimulus (CS) and a sucrose unconditioned stimulus (US). We previously reported that females are less sensitive to generalization of a FP morphine OS than males, so we investigated the role of endogenous ovarian hormones in this difference. METHODS: Male and female rats received intermixed injections of 3.2 mg/kg morphine or saline before each daily training session. Training consisted of 8 presentations of the CS, each followed by access to sucrose on morphine, but not saline sessions. Following acquisiton, rats were tested for generalization of the morphine stimulus to 0, 1.0, 3.2, and 5.4 mg/kg morphine. Female rats were monitored for estrous cyclicity using vaginal cytology throughout the study. RESULTS: Both sexes acquired stable drug discrimination. A gradient of generalization was measured across morphine doses and this behaviour did not differ by sex, nor did it differ across the estrous cycle in females. CONCLUSIONS: Morphine generalization is independent of fluctuations in levels of sex and endogenous gonadal hormones in females under these experimental conditions.

Individuals lack the ability to accurately detect emotional piloerection.

Piloerection (e.g., goosebumps) is an essential thermoregulatory and social signaling mechanism in non-human animals. Although humans also experience piloerection-often being perceived as an indicator of profound emotional experiences-its comparatively less effective role in thermoregulation and communication might influence our capacity to monitor its occurrence. We present three studies (total N = 617) demonstrating participants' general inability to detect their own piloerection events and their lack of awareness that piloerection occurs with a similar frequency on multiple anatomical locations. Self-reported goosebumps were more frequent than observed piloerection. However, only 31.8% of self-reports coincided with observable piloerection, a bias unrelated to piloerection intensity, anatomical location, heart-rate variability, or interoceptive awareness. We also discovered a self-report bias for the forearm, contradicting the observation that piloerection occurs with equal frequency on multiple anatomical locations. Finally, there was low correspondence between self-reports of being "emotionally moved" and observed piloerection. These counterintuitive findings not only highlight a disconnect between an obvious physiological response and our capacity for self-monitoring, but they underscore a fascinating divergence between human and non-human species. Although piloerection is vital in non-human organisms, the connection between piloerection and psychological experience in humans may be less significant than previously assumed, possibly due to its diminished evolutionary relevance.

The duration discrimination respiratory task: A new test to measure respiratory interoceptive accuracy.

Interoception, which refers to the perception of body's internal state, is implicated in emotional processes and psychopathological disorders. Over the last decades, different tools have been developed to measure interoceptive accuracy, or the ability to accurately perceive physiological signals. Most of these tools have focused on cardiac interoception, but respiratory interoception has been less investigated due to the more complex and less portable equipment required. In this study, we suggest a new duration discrimination respiratory (DDR) task that does not require complex equipment. Using an adaptive staircase procedure, this task aims to determine an individual's ability to detect exhalation longer than their resting reference duration. One hundred and twenty-three healthy subjects completed the DDR task, an interoceptive task of heart rate discrimination, and filled out questionnaires on interoceptive awareness (Multidimensional Assessment of Interoceptive Awareness), alexithymia (Toronto Alexithymia Scale [TAS]), affects (Positive and Negative Affect Scale [PANAS]), and anamnestic. Results demonstrated a good internal consistency (Cronbach's alpha = .93) of the DDR task. On average, subjects needed 99.22% (SD = 36.38) of their reference exhalation time in addition to reference exhalation to detect a prolonged exhalation. Higher self-reported fitness levels, not counting during the DDR task and lower difficulty in describing feelings (TAS subscale), predicted higher respiratory discrimination duration. In conclusion, this study demonstrates the utility of the DDR task as a valid measure of interoception.

Interoceptive awareness mediated the effects of a 15-minute diaphragmatic breathing on empathy for pain: A randomized controlled trial.

Although empathy for pain plays an important role in positive interpersonal relationships and encourages engagement in prosocial behavior, it remains largely unknown whether empathy for pain could be effectively altered by psychophysiological techniques. This study aimed to investigate the impact of a single session of diaphragmatic breathing practice on empathy for pain and examine the potential mechanism involving interoceptive awareness. A total of 66 healthy participants were randomly assigned to the intervention group or the control group. The intervention group received a 15-minute diaphragmatic breathing (DB) practice with real-time biofeedback, while the control group was to gaze at a black screen at rest and not engaged in any other activities. Before and after the invention, all participants were instructed to evaluate the intensity and unpleasantness of empathy for pain while watching different pictures with pain or non-pain conditions. The Multidimensional Assessment of Interoceptive Awareness (MAIA) was then administered to measure interoceptive awareness. The results indicated a significant interaction between group and time with regard to empathy for pain and MAIA. The DB group showed a statistically significant decrease in both pain intensity and unpleasantness during the pain picture condition, as well as a noteworthy increase in MAIA scores. The control group did not demonstrate any substantial changes. More importantly, the regulation of attention, a dimension of MAIA, had a significant mediating effect on the impact of diaphragmatic breathing on reported unpleasantness. Diaphragmatic breathing could serve as a simple, convenient, and practical strategy to optimize human empathy for pain that warrants further investigation, which has important implications not only for individuals with impaired empathy for pain but also for the improvement of interoceptive awareness.

Interoceptive training with real-time haptic versus visual heartbeat feedback.

The perception of signals from within the body, known as interoception, is increasingly recognized as a prerequisite for physical and mental health. This study is dedicated to the development of effective technological approaches for enhancing interoceptive abilities. We provide evidence of the effectiveness and practical feasibility of a novel real-time haptic heartbeat supplementation technology combining principles of biofeedback and sensory augmentation. In a randomized controlled study, we applied the developed naturalistic haptic feedback on a group of 30 adults, while another group of 30 adults received more traditional real-time visual heartbeat feedback. A single session of haptic, but not visual heartbeat feedback resulted in increased interoceptive accuracy and confidence, as measured by the heart rate discrimination task, and in a shift of attention toward the body. Participants rated the developed technology as more helpful and pleasant than the visual feedback, thus indicating high user satisfaction. The study highlights the importance of matching sensory characteristics of the feedback provided to the natural bodily prototype. Our work suggests that real-time haptic feedback might be a superior approach for strengthening the mind-body connection in interventions for physical and mental health.

Insights into interoceptive and emotional processing: Lessons from studies on insular HD-tDCS.

Interoception, the processing of internal bodily signals, is proposed as the fundamental mechanism underlying emotional experiences. Interoceptive and emotional processing appear distorted in psychiatric disorders. However, our understanding of the neural structures involved in both processes remains limited. To explore the feasibility of enhancing interoception and emotion, we conducted two studies using high-definition transcranial direct current stimulation (HD-tDCS) applied to the right anterior insula. In study one, we compared the effects of anodal HD-tDCS and sham tDCS on interoceptive abilities (sensibility, confidence, accuracy, emotional evaluation) in 52 healthy subjects. Study two additionally included physical activation through ergometer cycling at the beginning of HD-tDCS and examined changes in interoceptive and emotional processing in 39 healthy adults. In both studies, HD-tDCS was applied in a single-blind cross-over online design with two separate sessions. Study one yielded no significant effects of HD-tDCS on interoceptive dimensions. In study two, significant improvements in interoceptive sensibility and confidence were observed over time with physical preactivation, while no differential effects were found between sham and insula stimulation. The expected enhancement of interoceptive and emotional processing following insula stimulation was not observed. We conclude that HD-tDCS targeting the insula does not consistently increase interoceptive or emotional variables. The observed increase in interoceptive sensibility may be attributed to the activation of the interoceptive network through physical activity or training effects. Future research on HD-tDCS involving interoceptive network structures could benefit from protocols targeting larger regions within the network, rather than focusing solely on insula stimulation.