Biology subjects

De Lucia, M.

Publications and source records attributed to De Lucia, M..

9 recordsLinked to original sources

bioRxiv2026

Coordinated Brain-Heart Dynamics During Human NREM Sleep

Communication between the brain and cardiovascular system is increasingly recognized as a fundamental determinant of health, yet the mechanisms coordinating neural and cardiac rhythms remain poorly understood. Sleep provides a unique physiological state for investigating neurovisceral integration because cortical slow oscillations (SOs), thalamocortical sleep spindles, and autonomic activity undergo profound reorganization during non-rapid eye movement (NREM) sleep. We analyzed overnight high-density electroencephalography and electrocardiography recordings from 30 healthy adults and quantified interactions among SOs, sleep spindles, cardiac R-peaks, heart rate variability (HRV), spectral slope, and Lempel-Ziv complexity. To assess integrated brain-heart coordination, we developed a Joint Phase-Amplitude Coupling (JPAC) metric that quantifies the corelation between SO phase-dependent modulation of spindle amplitude and cardiac timing. SO-spindle coupling showed the expected fronto-central distribution but was not significantly related to autonomic or cortical state markers. In contrast, SO-R-peak coupling and JPAC systematically tracked physiological state. Stronger JPAC was associated with higher high-frequency HRV, steeper spectral slopes, and lower neural complexity, indicating enhanced coupling during synchronized, parasympathetically dominated NREM sleep. These findings demonstrate coordinated organization of cortical, thalamocortical, and cardiac rhythms within a common oscillatory framework and identify NREM sleep as a state and potential treatment target of large-scale neurovisceral integration.

neuroscience↗

bioRxiv2026

Heartbeat-locked auditory deviations slow down cardiac activity

Our responses to environmental inputs depend on the variations of our own physiological activity. However, the mechanisms by which the integration of sensory information with interoceptive signals shape bodily responses to external events remain debated. In this pre-registered study, we hypothesized three possible mechanisms underlying such exteroceptive-interoceptive integration: cardiac surprise, active inference, and dynamic coupling. To test them, we implemented a closed-loop stimulation procedure to play auditory deviations from sequences either synchronized or not with heartbeats which varied in type (omissions or rare tones) and predictability (random or regular intervals). First, we replicated previous findings that cardiac activity slows down in response to sound omissions only when sounds are synchronized with heartbeats. Second, we showed that this effect extends to rare tones, excluding the dynamic coupling hypothesis. Third, we demonstrated that these responses do not depend on the predictability of auditory deviations, excluding both cardiac surprise and active inference hypotheses. In a control experiment, we further observed that behavioral responses depend on the type and predictability of auditory deviants: participants can discriminate subjectively which sounds were synchronized with their own heartbeats without evidence of a relationship to interoception nor cardiac responses. Overall, these results demonstrate that auditory deviations slow down cardiac responses when locked to heartbeats but independently from their type and regularity, calling for novel hypotheses to account for the interoceptive-exteroceptive integration of sensory signals into cardiac activity. Impact statementUsing a cardio-audio synchrony task, we show that cardiac responses slow down upon heartbeat-locked auditory deviations independently from their type or regularity, suggesting a simple, fundamental mechanism of integration of internal and external signals into bodily responses to the environment. The heart may provide a straightforward way to study basic self-related processes, without depending on behavior or self-report, which is especially valuable for individuals who are unable to respond or communicate.

physiology↗

bioRxiv2026

Sensory processing reallocation from external to internal signals in REM sleep

The brain continuously integrates information from the external environment (exteroception) and the internal bodily milieu (interoception). How the balance between these two processing streams shifts across vigilance states with differing levels of environmental responsiveness, however, remains poorly understood. Here, we examined neural responses to external auditory and internal cardiac signals across wakefulness and REM sleep microstates - tonic and phasic REM - which are characterized by progressively reduced responsiveness to external stimulation. High-density EEG was recorded in healthy participants (n=25). Auditory evoked potentials (AEPs) and heartbeat evoked potentials (HEPs) served as indices of exteroception and interoception, respectively, and were compared across vigilance states. AEPs progressively decreased from wakefulness to tonic REM and were most attenuated during phasic REM. In contrast, HEPs were preserved across REM microstates and were enhanced relative to wakefulness, indicating sustained - and even amplified - processing of cardiac signals during REM sleep. To quantify the relative weighting of external and internal signals, we introduce an exteroceptive-interoceptive index, defined as the ratio of auditory to cardiac neural responses. This index decreased systematically across vigilance states, revealing a graded shift from externally oriented processing during wakefulness to internally oriented processing during phasic REM, with tonic REM occupying an intermediate position. Together, these findings demonstrate that while responsiveness to external stimuli diminishes during phasic REM, the brain continues to prioritize physiologically relevant internal signals. The exteroceptive-interoceptive balance may thus provide a novel, mechanistically grounded marker of altered consciousness, particularly informative in contexts where behavioural responsiveness cannot be assessed. Graphical abstract O_FIG O_LINKSMALLFIG WIDTH=200 HEIGHT=141 SRC="FIGDIR/small/712081v1_ufig1.gif" ALT="Figure 1"> View larger version (19K): org.highwire.dtl.DTLVardef@1e46a9borg.highwire.dtl.DTLVardef@112f050org.highwire.dtl.DTLVardef@5f5249org.highwire.dtl.DTLVardef@135cc4_HPS_FORMAT_FIGEXP M_FIG C_FIG

neuroscience↗

bioRxiv2025

Impact of the pulse artifact on evoked activity in human wakefulness and sleep

The investigation of the neural evoked response to the heartbeat quantified using electroencephalography (i.e., heartbeat evoked potentials or HEPs), has gained recent attention in neuroscience, notably as a measure of interoception, the sensory system responding to internal bodily states. One main challenge in measuring HEPs is their susceptibility to cardiac artifacts contamination, including the cardiac field artifact and the pulse artifact (PA), the latter possibly caused by the mechanical pressure of pulsating vessels. Here, we aimed at assessing the impact of PAs on HEPs and auditory evoked potentials (AEPs, a proxy of the neural responses to exteroceptive sensory stimuli). To this aim, we compared two pre-processing pipelines using independent component analysis in healthy volunteers (N=30). The first, standard, pipeline excluded ocular, muscle, sweating-related activity and cardiac-related activity stemming from the cardiac field artifact. The second, pairwise phase consistency (PPC) pipeline, in addition to the removal of the aforementioned components, used the quantitative metric of PPC between independent components and the ECG to remove the cardiac-related PA. We tested how these two pre-processing approaches influenced HEPs and AEPs recorded under diverse neurophysiological conditions (wakefulness, N2, N3, and REM sleep). Comparing the HEPs from the standard and the PPC approaches (cluster-based permutation statistics, p<0.05, two-tailed) revealed a significant effect of PAs, particularly in wakefulness, followed by REM and N2 sleep, with Cohens d effect sizes of 1.92, 0.95 and 0.88, respectively. By contrast, PA correction had a negligible effect (p>0.05, two-tailed) on the HEPs in N3 sleep and on the AEPs in all vigilance states. Our results emphasize the need to account for the PA as a significant confounding factor when comparing HEPs across groups with varying vascular or cardiac conditions.

neuroscience↗

bioRxiv2025

Cardiac-locked auditory stimulation modulates pupil and neural dynamics

The human brain is sensitive to temporal regularities across bodily and environmental signals. Here, we investigated the pupil and neural correlates of regularity encoding established across cardiac and auditory stimuli. Auditory sequences were presented in synchrony with the heartbeat (synchronous), at a fixed pace, or without temporal regularity while recording pupillometry, electroencephalography, and electrocardiography in healthy participants. Sounds evoked typical pupil dilation in all conditions. However, only in the synchronous condition, pupil dilation progressively decreased over the course of the sequence, possibly reflecting adaptation to the repeated cardio-auditory alignment. A concurrent increase in global EEG activity suggested enhanced cortical processing in response to the synchronous sequence. Pupil constriction was driven by participants with higher heart rate, indicating that pupil adaptation mostly occurs in response to fast auditory sequences. Cardio-audio regularity encoding manifests as a pupil adaptation and an amplification of global EEG activity, likely reflecting improved temporal prediction precision.

neuroscience↗

bioRxiv2024

Multisensory integration in Peripersonal Space indexes consciousness states in sleep and disorders of consciousness

Conscious experience encompasses not only the awareness of external objects, but also a phenomenal representation of the embodied subject of the experience. The latter is mediated by the integration of multisensory stimuli between the body and the environment, a process mediated by the Peripersonal Space (PPS) system. Here we thus tested the hypothesis that a neural marker of PPS representation may index the presence of conscious experience. Using high-density EEG in awake participants, we identified a "PPS index", characterized by high-beta oscillations in centroparietal regions during the integration of audiotactile stimuli presented near versus far from the body. We then examined this marker across two models of altered consciousness, i.e., sleep and disorders of consciousness. The PPS index persisted during dreaming and waking conscious states but was absent during dreamless, unconscious states. Moreover, the same index predicted behavioural measures of consciousness and clinical outcome in patients recovering from disorders of consciousness. These results suggest that multisensory integration within the PPS is tightly linked to the presence of conscious experience.

neuroscience↗

bioRxiv2022

Single-subject EEG measurement of interhemispheric transfer-time for the in-vivo estimation of axonal morphology

Assessing axonal morphology in-vivo opens new avenues for the combined study of brain structure and function. A novel approach has recently been introduced to estimate the morphology of axonal fibers from the combination of MRI data and EEG measures of the interhemispheric transfer time (IHTT). In the original study, the IHTT measures were computed from EEG data averaged across a group, leading to bias of the axonal morphology estimates. Here, we seek to estimate axonal morphology from individual measures of IHTT, obtained from EEG data acquired in a visual evoked potential experiment. Subject-specific IHTTs are computed in a data-driven framework with minimal a priori constraints, based on the maximal peak of neural responses to visual stimuli within periods of statistically significant evoked activity in the inverse solution space. The subject-specific IHTT estimates ranged from 8 to 29 ms except for one participant and the between-session variability was comparable to the differences in IHTT between subjects. The scale parameter of the axonal radius distribution, computed from the IHTT estimates and the MRI data, ranged from 0 to 0.79 m. The change in axonal g-ratio with axonal radius ranged from 0.62 to 0.81 m-. The single-subject measurement of the IHTT yields estimates of axonal morphology that are consistent with histological values. However, improvement of the repeatability of the IHTT estimates is required to improve the specificity of the single-subject axonal morphology estimates.

neuroscience↗

bioRxiv2022

Cardio-audio synchronization elicits prediction in auditory sequences during human wakefulness and sleep

The human brain can infer temporal regularities in auditory sequences with fixed sound-to-sound intervals and in pseudo-regular sequences where sound onsets are locked to cardiac inputs. Here, we investigated auditory and cardio-audio regularity encoding during sleep, when reduced vigilance may result in altered bodily and environmental stimulus processing. Using electroencephalography and electrocardiography in healthy volunteers (N=26) during wakefulness and sleep, we measured the response to unexpected sound omissions within three auditory regularity conditions: synchronous, where sound and heartbeat are temporally locked, isochronous, with fixed sound-to-sound intervals, and a control condition without specific regularity. During wakefulness and all sleep stages, the cardiac signal following sound omissions exhibited a deceleration over time in the synchronous condition only. At the neural level, both the synchronous and isochronous sequences gave rise to a neural omission response in wakefulness and N2 sleep. Regularity encoding in the synchronous and isochronous conditions was characterized by a modulation of the background oscillatory activity in N2 sleep, outlining a possible mechanism through which the brain aligns periods of high neuronal excitability to the expected sound onset. The violation of auditory and cardio-audio regularity elicits cardiac and neural surprise responses across vigilance stages. Significance StatementAcross vigilance states, the human brain can generate predictions about the future based on past sensory regularities. While this is evident for environmental stimuli, the role of bodily signals in forming sensory prediction remains unknown. Here, we show that the human brain utilizes the temporal relationship between cardiac and auditory inputs in order to anticipate upcoming sounds during wakefulness and sleep. After presenting sounds in synchrony with the ongoing heartbeat, a sound omission elicited both a heartbeat deceleration and a prediction error signal as measured by the electroencephalographic response. Heartbeat signals support auditory regularity encoding during sleep and wakefulness, highlighting one mechanism for optimizing the detection of unexpected stimuli by taking advantage of the continuously monitored cardiac signals.

neuroscience↗

bioRxiv2022

In-vivo estimation of axonal morphology from MRI and EEG data

PurposeWe present a novel approach that allows the estimation of morphological features of axonal fibers from data acquired in-vivo in humans. This approach allows the assessment of white matter microscopic properties non-invasively with improved specificity. TheoryThe proposed approach is based on a biophysical model of Magnetic Resonance Imaging (MRI) data and of axonal conduction velocity estimates obtained with Electroencephalography (EEG). In a white matter tract of interest, these data depend on 1) the distribution of axonal radius - P(r)- and 2) the g-ratio of the individual axons that compose this tract - g(r). P(r)is assumed to follow a Gamma distribution with mode and scale parameters, M and{theta} , and g(r) is described by a power-law with parameters and {beta}. MethodsMRI and EEG data were recorded from 14 healthy volunteers. MRI data were collected with a 3T scanner. MRI g-ratio maps were computed and sampled along the visual transcallosal tract. EEG data were recorded using a 128-lead system with a visual Poffenberg paradigm. The interhemispheric transfer time and axonal conduction velocity were computed from the EEG current density at the group level. Using the MRI and EEG measures and the proposed model, we estimated morphological properties of axons in the visual transcallosal tract. ResultsThe estimated interhemispheric transfer time was 11.72{+/-}2.87 ms, leading to an average conduction velocity across subjects of 13.22{+/-}1.18 m/s. Out of the 4 free parameters of the proposed model, we estimated{theta} - the width of the right tail of the axonal radius distribution and {beta} - the scaling factor of the axonal g-ratio, a measure of fiber myelination. Across subjects, the parameter{theta} was 0.40{+/-}0.07 {micro}m and the parameter {beta} was 0.67{+/-}0.02 {micro}m-. ConclusionsThe estimates of axonal radius and myelination are consistent with histological findings, illustrating the feasibility of this approach. The proposed method allows the measurement of the distribution of axonal radius and myelination within a white matter tract, opening new avenues for the combined study of brain structure and function, and for in-vivo histological studies of the human brain.

neuroscience↗