The relationship between alpha power and heart rate variability commonly seen in various mental states.

The extensive exploration of the correlation between electroencephalogram (EEG) and heart rate variability (HRV) has yielded inconsistent outcomes, largely attributable to variations in the tasks employed in the studies. The direct relationship between EEG and HRV is further complicated by alpha power, which is susceptible to influences such as mental fatigue and sleepiness. This research endeavors to examine the brain-heart interplay typically observed during periods of music listening and rest. In an effort to mitigate the indirect effects of mental states on alpha power, subjective fatigue and sleepiness were measured during rest, while emotional valence and arousal were evaluated during music listening. Partial correlation analyses unveiled positive associations between occipital alpha2 power (10-12 Hz) and nHF, an indicator of parasympathetic activity, under both music and rest conditions. These findings underscore brain-heart interactions that persist even after the effects of other variables have been accounted for.

Theoretical and Technical Issues Concerning the Measurement of Alpha Frequency and the Application of Signal Detection Theory: Comment on Buergers and Noppeney (2022).

Classical and recent evidence has suggested that alpha oscillations play a critical role in temporally discriminating or binding successively presented items. Challenging this view, Buergers and Noppeney [Buergers, S., & Noppeney, U. The role of alpha oscillations in temporal binding within and across the senses. Nature Human Behaviour, 6, 732-742, 2022] found that by combining EEG, psychophysics, and signal detection theory, neither prestimulus nor resting-state alpha frequency influences perceptual sensitivity and bias in the temporal binding task. We propose the following four points that should be considered when interpreting the role of alpha oscillations, and especially their frequency, on perceptual temporal binding: (1) Multiple alpha components can be contaminated in conventional EEG analysis; (2) the effect of alpha frequency on perception will interact with alpha power; (3) prestimulus and resting-state alpha frequency can be different from poststimulus alpha frequency, which is the frequency during temporal binding and should be more directly related to temporal binding; and (4) when applying signal detection theory under the assumption of equal variance, the assumption is often incomplete and can be problematic (e.g., the magnitude relationships between individuals in parametric sensitivity may change when converted into nonparametric sensitivity). Future directions, including solutions to each of the issues, are discussed.

Macromolecular tissue volume mapping of lateral geniculate nucleus subdivisions in living human brains.

The lateral geniculate nucleus (LGN) is a key thalamic nucleus in the visual system, which has an important function in relaying retinal visual input to the visual cortex. The human LGN is composed mainly of magnocellular (M) and parvocellular (P) subdivisions, each of which has different stimulus selectivity in neural response properties. Previous studies have discussed the potential relationship between LGN subdivisions and visual disorders based on psychophysical data on specific types of visual stimuli. However, these relationships remain speculative because non-invasive measurements of these subdivisions are difficult due to the small size of the LGN. Here we propose a method to identify these subdivisions by combining two structural MR measures: high-resolution proton-density weighted images and macromolecular tissue volume (MTV) maps. We defined the M and P subdivisions based on MTV fraction data and tested the validity of the definition by (1) comparing the data with that from human histological studies, (2) comparing the data with functional magnetic resonance imaging measurements on stimulus selectivity, and (3) analyzing the test-retest reliability. The findings demonstrated that the spatial organization of the M and P subdivisions was consistent across subjects and in line with LGN subdivisions observed in human histological data. Moreover, the difference in stimulus selectivity between the subdivisions identified using MTV was consistent with previous physiology literature. The definition of the subdivisions based on MTV was shown to be robust over measurements taken on different days. These results suggest that MTV mapping is a promising approach for evaluating the tissue properties of LGN subdivisions in living humans. This method potentially will enable neuroscientific and clinical hypotheses about the human LGN subdivisions to be tested.

Neural correlates of impulsive compulsive behaviors in Parkinson's disease: A Japanese retrospective study.

BACKGROUND: Impulsive compulsive behaviors (ICBs) often disturb patients with Parkinson's Disease (PD), of which impulse control disorder (ICD) and dopamine dysregulation syndrome (DDS) are two major subsets. The nucleus accumbens (NAcc) is involved in ICB; however, it remains unclear how the NAcc affects cortical function and defines the different behavioral characteristics of ICD and DDS. OBJECTIVES: To identify the cortico-striatal network primarily involved in ICB and the differences in these networks between patients with ICD and DDS using structural and resting-state functional magnetic resonance imaging. METHODS: Patients with PD were recruited using data from a previous cohort study and divided into those with ICB (ICB group) and without ICB (non-ICB group) using the Japanese version of the Questionnaire for Impulsive Compulsive Disorders in Parkinson's Disease (J-QUIP). From these two groups, we extracted 37 pairs matched for age, sex, disease duration, and levodopa equivalent daily dose of dopamine agonists. Patients with ICB were further classified as having ICD or DDS based on the J-QUIP subscore. General linear models were used to compare gray matter volume and functional connectivity (FC) of the NAcc, caudate, and putamen between the ICB and non-ICB groups and between patients with ICD and those with DDS. RESULTS: We found no significant differences in gray matter volumebetween the ICB and non-ICB groups or between patients with ICD and those with DDS. Compared with the non-ICB group, the FC of the right NAcc in the ICB group was lower in the bilateral ventromedial prefrontal cortex and higher in the left middle occipital gyrus. Furthermore, patients with DDS showed higher FC between the right putamen and left superior temporal gyrus and higher FC between the left caudate and bilateral middle occipital gyrus than patients with ICD. In contrast, patients with ICD exhibited higher FC between the left NAcc and the right posterior cingulate cortex than patients with DDS. CONCLUSIONS: The functionally altered network between the right NAcc and ventromedial prefrontal cortex was associated with ICB in PD. In addition, the surrounding cortico-striatal networks may differentiate the behavioral characteristics of patients with ICD and those with DDS.

Can enhancement and suppression concurrently guide attention? An assessment at the individual level.

BACKGROUND: Although people can pay attention to targets while ignoring distractors, previous research suggests that target enhancement and distractor suppression work separately and independently. Here, we sought to replicate previous findings and re-establish their independence. METHODS: We employed an internet-based psychological experiment. We presented participants with a visual search task in which they searched for a specified shape with or without a singleton. We replicated the singleton-presence benefit in search performance, but this effect was limited to cases where the target color was fixed across all trials. In a randomly intermixed probe task (30% of all trials), the participants searched for a letter among colored probes; we used this task to assess how far attention was separately allocated toward the target or distractor dimensions. RESULTS: We found a negative correlation between target enhancement and distractor suppression, indicating that the participants who paid closer attention to target features ignored distractor features less effectively and vice versa. Averaged data showed no benefit from target color or cost from distractor color, possibly because of the substantial differences in strategy across participants. CONCLUSIONS: These results suggest that target enhancement and distractor suppression guide attention in mutually dependent ways and that the relative contribution of these components depends on the participants' search strategy.

Building a decoder of perceptual decisions from microsaccades and pupil size.

Many studies have reported neural correlates of visual awareness across several brain regions, including the sensory, parietal, and frontal areas. In most of these studies, participants were instructed to explicitly report their perceptual experience through a button press or verbal report. It is conceivable, however, that explicit reporting itself may trigger specific neural responses that can confound the direct examination of the neural correlates of visual awareness. This suggests the need to assess visual awareness without explicit reporting. One way to achieve this is to develop a technique to predict the visual awareness of participants based on their peripheral responses. Here, we used eye movements and pupil sizes to decode trial-by-trial changes in the awareness of a stimulus whose visibility was deteriorated due to adaptation-induced blindness (AIB). In the experiment, participants judged whether they perceived a target stimulus and rated the confidence they had in their perceptual judgment, while their eye movements and pupil sizes were recorded. We found that not only perceptual decision but also perceptual confidence can be separately decoded from the eye movement and pupil size. We discuss the potential of this technique with regard to assessing visual awareness in future neuroimaging experiments.

Frequency- and phase-dependent effects of auditory entrainment on attentional blink.

Attentional blink (AB) is the impaired detection of a second target (T2) after a first target has been identified. In this paper, we investigated the functional roles of alpha and theta oscillations on AB by determining how much preceding rhythmic auditory stimulation affected the performance of AB. Healthy young adults participated in the experiment online. We found that when two targets were embedded in rapid serial visual presentation (RSVP) of distractors at 10 Hz (i.e., alpha frequency), the magnitude of AB increased with auditory stimuli. The increase was limited to the case when the frequency and phase of auditory stimuli matched the following RSVP stream. On the contrary, when only two targets were presented without distractors, auditory stimuli at theta, not alpha, increased the AB magnitude. These results indicate that neural oscillations at two different frequencies, namely, alpha and theta, are involved in attentional blink.

Microstructural Properties of Human Brain Revealed by Fractional Anisotropy Can Predict the After-Effect of Intermittent Theta Burst Stimulation.

Intermittent theta burst stimulation (iTBS) delivered by transcranial magnetic stimulation (TMS) produces a long-term potentiation-like after-effect useful for investigations of cortical function and of potential therapeutic value. However, the iTBS after-effect over the primary motor cortex (M1) as measured by changes in motor evoked potential (MEP) amplitude exhibits a largely unexplained variability across individuals. Here, we present evidence that individual differences in white matter (WM) and gray matter (GM) microstructural properties revealed by fractional anisotropy (FA) predict the magnitude of the iTBS-induced after-effect over M1. The MEP amplitude change in the early phase (5-10 min post-iTBS) was associated with FA values in WM tracts such as right superior longitudinal fasciculus and corpus callosum. By contrast, the MEP amplitude change in the late phase (15-30 min post-iTBS) was associated with FA in GM, primarily in right frontal cortex. These results suggest that the microstructural properties of regions connected directly or indirectly to the target region (M1) are crucial determinants of the iTBS after-effect. FA values indicative of these microstructural differences can predict the potential effectiveness of repetitive TMS for both investigational use and clinical application.

The DecNef collection, fMRI data from closed-loop decoded neurofeedback experiments.

Decoded neurofeedback (DecNef) is a form of closed-loop functional magnetic resonance imaging (fMRI) combined with machine learning approaches, which holds some promises for clinical applications. Yet, currently only a few research groups have had the opportunity to run such experiments; furthermore, there is no existing public dataset for scientists to analyse and investigate some of the factors enabling the manipulation of brain dynamics. We release here the data from published DecNef studies, consisting of 5 separate fMRI datasets, each with multiple sessions recorded per participant. For each participant the data consists of a session that was used in the main experiment to train the machine learning decoder, and several (from 3 to 10) closed-loop fMRI neural reinforcement sessions. The large dataset, currently comprising more than 60 participants, will be useful to the fMRI community at large and to researchers trying to understand the mechanisms underlying non-invasive modulation of brain dynamics. Finally, the data collection size will increase over time as data from newly run DecNef studies will be added.

V1 Projection Zone Signals in Human Macular Degeneration Depend on Task Despite Absence of Visual Stimulus.

Identifying the plastic and stable components of the visual cortex after retinal loss is an important topic in visual neuroscience and neuro-ophthalmology.1-5 Humans with juvenile macular degeneration (JMD) show significant blood-oxygen-level-dependent (BOLD) responses in the primary visual area (V1) lesion projection zone (LPZ),6 despite the absence of the feedforward signals from the degenerated retina. Our previous study7 reported that V1 LPZ responds to full-field visual stimuli during the one-back task (OBT), not during passive viewing, suggesting the involvement of task-related feedback signals. Aiming to clarify whether visual inputs to the intact retina are necessary for the LPZ responses, here, we measured BOLD responses to tactile and auditory stimuli for both JMD patients and control participants with and without OBT. Participants were instructed to close their eyes during the experiment for the purpose of eliminating retinal inputs. Without OBT, no V1 responses were detected in both groups of participants. With OBT, to the contrary, both stimuli caused substantial V1 responses in JMD patients, but not controls. Furthermore, we also found that the task-dependent activity in V1 LPZ became less pronounced when JMD patients opened their eyes, suggesting that task-related feedback signals can be partially suppressed by residual feedforward signals. Modality-independent V1 LPZ responses only in the task condition suggest that V1 LPZ responses reflect task-related feedback signals rather than reorganized feedforward visual inputs.

Atypical spatial frequency dependence of visual metacognition among schizophrenia patients.

Although altered early stages of visual processing have been reported among schizophrenia patients, how such atypical visual processing may affect higher-level cognition remains largely unknown. Here we tested the hypothesis that metacognitive performance may be atypically modulated by spatial frequency (SF) of visual stimuli among individuals with schizophrenia, given their altered magnocellular function. To study the effect of SF on metacognitive performance, we asked patients and controls to perform a visual detection task on gratings with different SFs and report confidence, and analyzed the data using the signal detection theoretic measure meta-d'. Control subjects showed better metacognitive performance after yes- (stimulus presence) than after no- (stimulus absence) responses ('yes-response advantage') for high SF (HSF) stimuli but not for low SF (LSF) stimuli. The patients, to the contrary, showed a 'yes-response advantage' not only for HSF but also for LSF stimuli, indicating atypical SF dependency of metacognition. An fMRI experiment using the same task revealed that the dorsolateral prefrontal cortex (DLPFC), known to be crucial for metacognition, shows activity mirroring the behavioral results: decoding accuracy of perceptual confidence in DLPFC was significantly higher for HSF than for LSF stimuli in controls, whereas this decoding accuracy was independent of SF in patients. Additionally, the functional connectivity of DLPFC with parietal and visual areas was modulated by SF and response type (yes/no) in a different manner between controls and patients. While individuals without schizophrenia may flexibly adapt metacognitive computations across SF ranges, patients may employ a different mechanism that is independent of SF. Because visual stimuli of low SF have been linked to predictive top-down processing, this may reflect atypical functioning in these processes in schizophrenia.

Predicting Neural Response Latency of the Human Early Visual Cortex from MRI-Based Tissue Measurements of the Optic Radiation.

Although the non-invasive measurement of visually evoked responses has been extensively studied, the structural basis of variabilities in latency in healthy humans is not well understood. We investigated how tissue properties of optic radiation could predict interindividual variability in the latency of the initial visually evoked component (C1), which may originate from the primary visual cortex (V1). We collected C1 peak latency data using magnetoencephalography (MEG) and checkerboard stimuli, and multiple structural magnetic resonance imaging (MRI) data from 20 healthy subjects. While we varied the contrast and position of the stimuli, the C1 measurement was most reliable when high-contrast stimuli were presented to the lower visual field (LVF). We then attempted to predict interindividual variability in C1 peak latency in this stimulus condition with a multiple regression model using MRI parameters along the optic radiation. We found that this model could predict >20% of variance in C1 latency, when the data were averaged across the hemispheres. The model using the corticospinal tract did not predict variability in C1 latency, suggesting that there is no evidence for generalization to a non-visual tract. In conclusion, our results suggest that the variability in neural latencies in the early visual cortex in healthy subjects can be partly explained by tissue properties along the optic radiation. We discuss the challenges of predicting neural latency using current structural neuroimaging methods and other factors that may explain interindividual variance in neural latency.

Inter-individual Differences in Occipital Alpha Oscillations Correlate with White Matter Tissue Properties of the Optic Radiation.

Neural oscillations at ∼10 Hz, called alpha oscillations, are one of the most prominent components of neural oscillations in the human brain. In recent years, characteristics (power/frequency/phase) of occipital alpha oscillations have been correlated with various perceptual phenomena. However, the relationship between inter-individual differences in alpha oscillatory characteristics and the properties of the underlying brain structures, such as white matter pathways, is unclear. A possibility is that intrinsic occipital alpha oscillations are mediated by thalamocortical interaction; we hypothesized that the most promising candidate for characterizing the intrinsic alpha oscillation is optic radiation (OR), which is the geniculo-cortical pathway carrying signals between the lateral geniculate nucleus (LGN) and primary visual cortex (V1). We used resting-state magnetoencephalography (MEG) and diffusion-weighted/quantitative magnetic resonance imaging (MRI) (dMRI/qMRI) to correlate the frequency and power of occipital alpha oscillations with the tissue properties of the OR by focusing on the different characteristics across individuals. We found that the peak alpha frequency (PAF) negatively correlated with intracellular volume fraction (ICVF), reflecting diffusion properties in intracellular (axonal) space, whereas the peak alpha power was not correlated with any tissue properties measurements. No significant correlation was found between OR and beta frequency/amplitude or between other white matter tract connecting parietal and inferotemporal cortex and alpha frequency/amplitude. These results support the hypothesis that an interaction between thalamic nuclei and early visual areas is essential for the occipital alpha oscillatory rhythm.

Threat Anticipation in Pulvinar and in Superficial Layers of Primary Visual Cortex (V1). Evidence from Layer-Specific Ultra-High Field 7T fMRI.

The perceptual system gives priority to threat-relevant signals with survival value. In addition to the processing initiated by sensory inputs of threat signals, prioritization of threat signals may also include processes related to threat anticipation. These neural mechanisms remain largely unknown. Using ultra-high-field 7 tesla (7T) fMRI, we show that anticipatory processing takes place in the early stages of visual processing, specifically in the pulvinar and V1. When anticipation of a threat-relevant fearful face target triggered false perception of not-presented target, there was enhanced activity in the pulvinar as well as in the V1 superficial-cortical-depth (layers 1-3). The anticipatory activity was absent in the LGN or higher visual cortical areas (V2-V4). The effect in V1 was specific to the perception of fearful face targets and did not generalize to happy face targets. A preliminary analysis showed that the connectivity between the pulvinar and V1 superficial-cortical-depth was enhanced during false perception of threat, indicating that the pulvinar and V1 may interact in preparation of anticipated threat. The anticipatory processing supported by the pulvinar and V1 may play an important role in non-sensory-input-driven anxiety states.

Microstructural properties of the vertical occipital fasciculus explain the variability in human stereoacuity.

Stereopsis is a fundamental visual function that has been studied extensively. However, it is not clear why depth discrimination (stereoacuity) varies more significantly among people than other modalities. Previous studies have reported the involvement of both dorsal and ventral visual areas in stereopsis, implying that not only neural computations in cortical areas but also the anatomical properties of white matter tracts connecting those areas can impact stereopsis. Here, we studied how human stereoacuity relates to white matter properties by combining psychophysics, diffusion MRI (dMRI), and quantitative MRI (qMRI). We performed a psychophysical experiment to measure stereoacuity and, in the same participants, we analyzed the microstructural properties of visual white matter tracts on the basis of two independent measurements, dMRI (fractional anisotropy, FA) and qMRI (macromolecular tissue volume; MTV). Microstructural properties along the right vertical occipital fasciculus (VOF), a major tract connecting dorsal and ventral visual areas, were highly correlated with measures of stereoacuity. This result was consistent for both FA and MTV, suggesting that the behavioral-structural relationship reflects differences in neural tissue density, rather than differences in the morphological configuration of fibers. fMRI confirmed that binocular disparity stimuli activated the dorsal and ventral visual regions near VOF endpoints. No other occipital tracts explained the variance in stereoacuity. In addition, the VOF properties were not associated with differences in performance on a different psychophysical task (contrast detection). These series of experiments suggest that stereoscopic depth discrimination performance is, at least in part, constrained by dorso-ventral communication through the VOF.

A non-invasive, quantitative study of broadband spectral responses in human visual cortex.

Currently, non-invasive methods for studying the human brain do not routinely and reliably measure spike-rate-dependent signals, independent of responses such as hemodynamic coupling (fMRI) and subthreshold neuronal synchrony (oscillations and event-related potentials). In contrast, invasive methods-microelectrode recordings and electrocorticography (ECoG)-have recently measured broadband power elevation in field potentials (~50-200 Hz) as a proxy for locally averaged spike rates. Here, we sought to detect and quantify stimulus-related broadband responses using magnetoencephalography (MEG). Extracranial measurements like MEG and EEG have multiple global noise sources and relatively low signal-to-noise ratios; moreover high frequency artifacts from eye movements can be confounded with stimulus design and mistaken for signals originating from brain activity. For these reasons, we developed an automated denoising technique that helps reveal the broadband signal of interest. Subjects viewed 12-Hz contrast-reversing patterns in the left, right, or bilateral visual field. Sensor time series were separated into evoked (12-Hz amplitude) and broadband components (60-150 Hz). In all subjects, denoised broadband responses were reliably measured in sensors over occipital cortex, even in trials without microsaccades. The broadband pattern was stimulus-dependent, with greater power contralateral to the stimulus. Because we obtain reliable broadband estimates with short experiments (~20 minutes), and with sufficient signal-to-noise to distinguish responses to different stimuli, we conclude that MEG broadband signals, denoised with our method, offer a practical, non-invasive means for characterizing spike-rate-dependent neural activity for addressing scientific questions about human brain function.

[Modulation of Metacognition with Decoded Neurofeedback].

Humans often assess their confidence in their own perception, e.g., feeling "confident" or "certain" of having seen a friend, or feeling "uncertain" about whether the phone rang. The neural mechanism underlying the metacognitive function that reflects subjective perception still remains under debate. We have previously used decoded neurofeedback (DecNef) to demonstrate that manipulating the multivoxel activation patterns in the frontoparietal network modulates perceptual confidence without affecting perceptual performance. The results provided clear evidence for a dissociation between perceptual confidence and performance and suggested a distinct role of the frontoparietal network in metacognition.

Illusory Jitter Perceived at the Frequency of Alpha Oscillations.

Neural oscillations, such as alpha (8-13 Hz), beta (13-30 Hz), and gamma (30-100 Hz), are widespread across cortical areas, and their possible functional roles include feature binding [1], neuronal communication [2, 3], and memory [1, 4]. The most prominent signal among these neural oscillations is the alpha oscillation. Although accumulating evidence suggests that alpha oscillations correlate with various aspects of visual processing [5-18], the number of studies proving their causal contribution in visual perception is limited [11, 16-18]. Here we report that illusory visual vibrations are consciously experienced at the frequency of intrinsic alpha oscillations. We employed an illusory jitter perception termed the motion-induced spatial conflict [19] that originates from the cyclic interaction between motion and shape processing. Comparison between the perceived frequency of illusory jitter and the peak alpha frequency (PAF) measured using magnetoencephalography (MEG) revealed that the inter- and intra-participant variations of the PAF are mirrored by an illusory jitter perception. More crucially, psychophysical and MEG measurements during amplitude-modulated current stimulation [20] showed that the PAF can be artificially manipulated, which results in a corresponding change in the perceived jitter frequency. These results suggest the causal contribution of neural oscillations at the alpha frequency in creating temporal characteristics of visual perception. Our results suggest that cortical areas, dorsal and ventral visual areas in this case, are interacting at the frequency of alpha oscillations [2, 3, 21-27].

[Associative Learning between Orientation and Color in Early Visual Areas].

Associative learning is an essential neural phenomenon where the contingency of different items increases after training. Although associative learning has been found to occur in many brain regions, there is no clear evidence that associative learning of visual features occurs in early visual areas. Here, we developed an associative decoded functional magnetic resonance imaging (fMRI) neurofeedback (A-DecNef) to determine whether associative learning of color and orientation can be induced in early visual areas. During the three days' training, A-DecNef induced fMRI signal patterns that corresponded to a specific target color (red) mostly in early visual areas while a vertical achromatic grating was simultaneously, physically presented to participants. Consequently, participants' perception of "red" was significantly more frequently than that of "green" in an achromatic vertical grating. This effect was also observed 3 to 5 months after training. These results suggest that long-term associative learning of two different visual features such as color and orientation, was induced most likely in early visual areas. This newly extended technique that induces associative learning may be used as an important tool for understanding and modifying brain function, since associations are fundamental and ubiquitous with respect to brain function.