Spontaneous variability predicts compensative motor response in vocal pitch control.

Our motor system uses sensory feedback to keep desired performance. From this view, motor fluctuation is not simply 'noise' inevitably caused in the nervous system but would play a role in generating variations to explore better outcomes via sensory feedback. Vocalization system offers a good model for studying such sensory-motor interactions since we regulate vocalization by hearing our own voice. This behavior is typically observed as compensatory responses in vocalized pitch, or fundamental frequency (fo), when artificial fo shifts were induced in the auditory feedback. However, the relationship between adaptive regulation and motor exploration in vocalization has remained unclear. Here we investigated behavioral variability in spontaneous vocal fo and compensatory responses against fo shifts in the feedback, and demonstrated that larger spontaneous fluctuation correlates with greater compensation in vocal fo. This correlation was found in slow components (≤ 5 Hz) of the spontaneous fluctuation but not in fast components (between 6 and 30 Hz), and the slow one was amplified during the compensatory responses. Furthermore, the compensatory ratio was reduced when large fo shifts were applied to the auditory feedback, as if reflecting the range of motor exploration. All these findings consistently suggest the functional role of motor variability in the exploration of better vocal outcomes.

Individual variability in the nonlinear development of the corpus callosum during infancy and toddlerhood: a longitudinal MRI analysis.

The human brain spends several years bootstrapping itself through intrinsic and extrinsic modulation, thus gradually developing both spatial organization and functions. Based on previous studies on developmental patterns and inter-individual variability of the corpus callosum (CC), we hypothesized that inherent variations of CC shape among infants emerge, depending on the position within the CC, along the developmental timeline. Here we used longitudinal magnetic resonance imaging data from infancy to toddlerhood and investigated the area, thickness, and shape of the midsagittal plane of the CC by applying multilevel modeling. The shape characteristics were extracted using the Procrustes method. We found nonlinearity, region-dependency, and inter-individual variability, as well as intra-individual consistencies, in CC development. Overall, the growth rate is faster in the first year than in the second year, and the trajectory differs between infants; the direction of CC formation in individual infants was determined within six months and maintained to two years. The anterior and posterior subregions increase in area and thickness faster than other subregions. Moreover, we clarified that the growth rate of the middle part of the CC is faster in the second year than in the first year in some individuals. Since the division of regions exhibiting different tendencies coincides with previously reported divisions based on the diameter of axons that make up the region, our results suggest that subregion-dependent individual variability occurs due to the increase in the diameter of the axon caliber, myelination partly due to experience and axon elimination during the early developmental period.

Oscillator decomposition of infant fNIRS data.

The functional near-infrared spectroscopy (fNIRS) can detect hemodynamic responses in the brain and the data consist of bivariate time series of oxygenated hemoglobin (oxy-Hb) and deoxygenated hemoglobin (deoxy-Hb) on each channel. In this study, we investigate oscillatory changes in infant fNIRS signals by using the oscillator decompisition method (OSC-DECOMP), which is a statistical method for extracting oscillators from time series data based on Gaussian linear state space models. OSC-DECOMP provides a natural decomposition of fNIRS data into oscillation components in a data-driven manner and does not require the arbitrary selection of band-pass filters. We analyzed 18-ch fNIRS data (3 minutes) acquired from 21 sleeping 3-month-old infants. Five to seven oscillators were extracted on most channels, and their frequency distribution had three peaks in the vicinity of 0.01-0.1 Hz, 1.6-2.4 Hz and 3.6-4.4 Hz. The first peak was considered to reflect hemodynamic changes in response to the brain activity, and the phase difference between oxy-Hb and deoxy-Hb for the associated oscillators was at approximately 230 degrees. The second peak was attributed to cardiac pulse waves and mirroring noise. Although these oscillators have close frequencies, OSC-DECOMP can separate them through estimating their different projection patterns on oxy-Hb and deoxy-Hb. The third peak was regarded as the harmonic of the second peak. By comparing the Akaike Information Criterion (AIC) of two state space models, we determined that the time series of oxy-Hb and deoxy-Hb on each channel originate from common oscillatory activity. We also utilized the result of OSC-DECOMP to investigate the frequency-specific functional connectivity. Whereas the brain oscillator exhibited functional connectivity, the pulse waves and mirroring noise oscillators showed spatially homogeneous and independent changes. OSC-DECOMP is a promising tool for data-driven extraction of oscillation components from biological time series data.

Errata: Best practices for fNIRS publications.

[This corrects the article DOI: 10.1117/1.NPh.8.1.012101.].

Best practices for fNIRS publications.

The application of functional near-infrared spectroscopy (fNIRS) in the neurosciences has been expanding over the last 40 years. Today, it is addressing a wide range of applications within different populations and utilizes a great variety of experimental paradigms. With the rapid growth and the diversification of research methods, some inconsistencies are appearing in the way in which methods are presented, which can make the interpretation and replication of studies unnecessarily challenging. The Society for Functional Near-Infrared Spectroscopy has thus been motivated to organize a representative (but not exhaustive) group of leaders in the field to build a consensus on the best practices for describing the methods utilized in fNIRS studies. Our paper has been designed to provide guidelines to help enhance the reliability, repeatability, and traceability of reported fNIRS studies and encourage best practices throughout the community. A checklist is provided to guide authors in the preparation of their manuscripts and to assist reviewers when evaluating fNIRS papers.

Unconscious and Distinctive Control of Vocal Pitch and Timbre During Altered Auditory Feedback.

Vocal control plays a critical role in smooth social communication. Speakers constantly monitor auditory feedback (AF) and make adjustments when their voices deviate from their intentions. Previous studies have shown that when certain acoustic features of the AF are artificially altered, speakers compensate for this alteration in the opposite direction. However, little is known about how the vocal control system implements compensations for alterations of different acoustic features, and associates them with subjective consciousness. The present study investigated whether compensations for the fundamental frequency (F0), which corresponds to perceived pitch, and formants, which contribute to perceived timbre, can be performed unconsciously and independently. Forty native Japanese speakers received two types of altered AF during vowel production that involved shifts of either only the formant frequencies (formant modification; Fm) or both the pitch and formant frequencies (pitch + formant modification; PFm). For each type, three levels of shift (slight, medium, and severe) in both directions (increase or decrease) were used. After the experiment, participants were tested for whether they had perceived a change in the F0 and/or formants. The results showed that (i) only formants were compensated for in the Fm condition, while both the F0 and formants were compensated for in the PFm condition; (ii) the F0 compensation exhibited greater precision than the formant compensation in PFm; and (iii) compensation occurred even when participants misperceived or could not explicitly perceive the alteration in AF. These findings indicate that non-experts can compensate for both formant and F0 modifications in the AF during vocal production, even when the modifications are not explicitly or correctly perceived, which provides further evidence for a dissociation between conscious perception and action in vocal control. We propose that such unconscious control of voice production may enhance rapid adaptation to changing speech environments and facilitate mutual communication.

Developmental changes in cortical sensory processing during wakefulness and sleep.

Infants are exposed to auditory and visual information during sleep as well as wakefulness. Little is known, however, about the differences in cortical processing of sensory input between these different behavioral states. In the present study, cortical hemodynamic responses to auditory and visual stimuli during wakefulness and sleep were measured in infants aged 2-10 months using functional near-infrared spectroscopy (fNIRS). While asynchronously presented auditory and visual stimuli during wakefulness induced focal responses in the corresponding sensory regions of the occipital and temporal cortices, the responses to the same stimuli during sleep were dramatically different. Auditory stimuli during sleep induced global responses over the frontal, temporal, and occipital regions, and the response pattern did not change between 2 and 10 months of age. In contrast, visual stimuli during sleep induced responses in the occipital cortex, and the response pattern exhibited developmental changes from a pattern of activation to one of deactivation around a half year of age. The functional connectivity among the cortical regions was generally higher during sleep than during wakefulness. The hemoglobin phase of oxygenation and deoxygenation (hPod) and the phase locking index of hPod (hPodL) showed general developmental changes and behavioral state dependent differences but no significant differences were seen between the stimulus types. The results suggest that the behavioral states have a fundamental impact on cortical sensory processing; (1) sensory processing during wakefulness is performed in more localized regions, (2) auditory processing is active during both wakefulness and sleep, (3) visual processing undergoes development of inhibitory mechanisms during sleep, and (4) these phenomena primarily reflect neural development rather than vascular and metabolic development.

Explicit Performance in Girls and Implicit Processing in Boys: A Simultaneous fNIRS-ERP Study on Second Language Syntactic Learning in Young Adolescents.

Learning a second language (L2) proceeds with individual approaches to proficiency in the language. Individual differences including sex, as well as working memory (WM) function appear to have strong effects on behavioral performance and cortical responses in L2 processing. Thus, by considering sex and WM capacity, we examined neural responses during L2 sentence processing as a function of L2 proficiency in young adolescents. In behavioral tests, girls significantly outperformed boys in L2 tests assessing proficiency and grammatical knowledge, and in a reading span test (RST) assessing WM capacity. Girls, but not boys, showed significant correlations between L2 tests and RST scores. Using functional near-infrared spectroscopy (fNIRS) and event-related potential (ERP) simultaneously, we measured cortical responses while participants listened to syntactically correct and incorrect sentences. ERP data revealed a grammaticality effect only in boys in the early time window (100-300 ms), implicated in phrase structure processing. In fNIRS data, while boys had significantly increased activation in the left prefrontal region implicated in syntactic processing, girls had increased activation in the posterior language-related region involved in phonology, semantics, and sentence processing with proficiency. Presumably, boys implicitly focused on rule-based syntactic processing, whereas girls made full use of linguistic knowledge and WM function. The present results provide important fundamental data for learning and teaching in L2 education.

Spatial variation in the hemoglobin phase of oxygenation and deoxygenation in the developing cortex of infants.

Spontaneous low-frequency oscillatory changes in oxygenated hemoglobin (oxy-Hb) and deoxygenated hemoglobin (deoxy-Hb) are observed using functional near-infrared spectroscopy (fNIRS). A previous study showed that the time-averaged phase difference between oxy-Hb and deoxy-Hb changes, referred to as hemoglobin phase of oxygenation and deoxygenation (hPod), is sensitive to the development of the cortex. We examined phase-locking index of hPod, referred to as [Formula: see text], in addition to hPod, in neonates and 3- and 6-month-old infants using the 94-channel fNIRS data, which covered large lateral regions of the cortex. The results showed that (1) developmental changes in hPod exhibited spatial dependency; (2) [Formula: see text] increased between the neonate group and 3-month-old infant group over the posterior, but not anterior, regions of the cortex; and (3) the cortical regions of each age group were clustered in several domains with specific characteristics of hPod and [Formula: see text]. This study indicates that the neonatal cortex is composed of regions with specific characteristics of hPod and [Formula: see text], and drastic changes occur between the neonatal period and 3 months of age. This study suggests that hPod and [Formula: see text] are sensitive to the cortical region-specific development of the circulatory, blood flow, metabolic, and neurovascular functions in young infants.

Macroanatomical Landmarks Featuring Junctions of Major Sulci and Fissures and Scalp Landmarks Based on the International 10-10 System for Analyzing Lateral Cortical Development of Infants.

The topographic relationships between the macroanatomical structure of the lateral cortex, including sulci and fissures, and anatomical landmarks on the external surface of the head are known to be consistent. This allows the coregistration of EEG electrodes or functional near-infrared spectroscopy over the scalp with underlying cortical regions. However, limited information is available as to whether the topographic relationships are maintained in rapidly developing infants, whose brains and heads exhibit drastic growth. We used MRIs of infants ranging in age from 3 to 22 months old, and identified 20 macroanatomical landmarks, featuring the junctions of major sulci and fissures, as well as cranial landmarks and virtually determined positions of the international 10-20 and 10-10 systems. A Procrustes analysis revealed developmental trends in changes of shape in both the cortex and head. An analysis of Euclidian distances between selected pairs of cortical landmarks at standard stereotactic coordinates showed anterior shifts of the relative positions of the premotor and parietal cortices with age. Finally, cortical landmark positions and their spatial variability were compared with 10-10 landmark positions. The results indicate that variability in the distribution of each macroanatomical landmark was much smaller than the pitch of the 10-10 landmarks. This study demonstrates that the scalp-based 10-10 system serves as a good frame of reference in infants not only for assessing the development of the macroanatomy of the lateral cortical structure, but also for functional studies of cortical development using transcranial modalities such as EEG and fNIRS.

Development of a neurofeedback protocol targeting the frontal pole using near-infrared spectroscopy.

AIM: Neurofeedback has been studied with the aim of controlling cerebral activity. Near-infrared spectroscopy is a non-invasive neuroimaging technique used for measuring hemoglobin concentration changes in cortical surface areas with high temporal resolution. Thus, near-infrared spectroscopy may be useful for neurofeedback, which requires real-time feedback of repeated brain activation measurements. However, no study has specifically targeted neurofeedback, using near-infrared spectroscopy, in the frontal pole cortex. METHODS: We developed an original near-infrared spectroscopy neurofeedback system targeting the frontal pole cortex. Over a single day of testing, each healthy participant (n = 24) received either correct or incorrect (Sham) feedback from near-infrared spectroscopy signals, based on a crossover design. RESULTS: Under correct feedback conditions, significant activation was observed in the frontal pole cortex (P = 0.000073). Additionally, self-evaluation of control and metacognitive beliefs were associated with near-infrared spectroscopy signals (P = 0.006). CONCLUSION: The neurofeedback system developed in this study might be useful for developing control of frontal pole cortex activation.

Detection of resting state functional connectivity using partial correlation analysis: A study using multi-distance and whole-head probe near-infrared spectroscopy.

Multichannel near-infrared spectroscopy (NIRS) is a functional neuroimaging modality that enables easy-to-use and noninvasive measurement of changes in blood oxygenation levels. We developed a clinically-applicable method for estimating resting state functional connectivity (RSFC) with NIRS using a partial correlation analysis to reduce the influence of extraneural components. Using a multi-distance probe arrangement NIRS, we measured resting state brain activity for 8min in 17 healthy participants. Independent component analysis was used to extract shallow and deep signals from the original NIRS data. Pearson's correlation calculated from original signals was significantly higher than that calculated from deep signals, while partial correlation calculated from original signals was comparable to that calculated from deep (cerebral-tissue) signals alone. To further test the validity of our method, we also measured 8min of resting state brain activity using a whole-head NIRS arrangement consisting of 17 cortical regions in 80 healthy participants. Significant RSFC between neighboring, interhemispheric homologous, and some distant ipsilateral brain region pairs was revealed. Additionally, females exhibited higher RSFC between interhemispheric occipital region-pairs, in addition to higher connectivity between some ipsilateral pairs in the left hemisphere, when compared to males. The combined results of the two component experiments indicate that partial correlation analysis is effective in reducing the influence of extracerebral signals, and that NIRS is able to detect well-described resting state networks and sex-related differences in RSFC.

Referential framework for transcranial anatomical correspondence for fNIRS based on manually traced sulci and gyri of an infant brain.

Functional near infrared spectroscopy (fNIRS), which is compact, portable, and tolerant of body movement, is suitable for monitoring infant brain functions. Nevertheless, fNIRS also poses a technical problem in that it cannot provide structural information. Supplementation with structural magnetic resonance images (MRI) is not always feasible for infants who undergo fNIRS measurement. Probabilistic registration methods using an MRI database instead of subjects' own MRIs are optimized for adult studies and offer only limited resources for infant studies. To overcome this, we used high-quality infant MRI data for a 12-month-old infant and manually delineated segmented gyri from among the highly visible macroanatomies on the lateral cortical surface. These macroanatomical regions are primarily linked to the spherical coordinate system based on external cranial landmarks, and further to traditional 10-20-based head-surface positioning systems. While macroanatomical structures were generally comparable between adult and infant atlases, differences were found in the parietal lobe, which was positioned posteriorly at the vertex in the infant brain. The present study provides a referential framework for macroanatomical analyses in infant fNIRS studies. With this resource, multichannel fNIRS functional data could be analyzed in reference to macroanatomical structures through virtual and probabilistic registrations without acquiring subject-specific MRIs.

A brain of two halves: insights into interhemispheric organization provided by near-infrared spectroscopy.

The discovery of functional lateralization and localization of the brain marked the beginning of a new era in neuroscience. While the past 150 years of research have provided a great deal of knowledge of hemispheric differences and functional relationships, the precise organization of functional laterality remains a topic of intense debate. Here I will shed light on the functional organization of the two hemispheres by reviewing some of the most recent functional near-infrared spectroscopy (NIRS) studies that have reported hemispheric differences in activation patterns. Most NIRS studies using visual stimuli, which revealed functional differentiation between the hemispheres, have reported unilateral activation, i.e., significant levels of activation in only one hemisphere. Auditory stimuli, including speech sounds, elicited bilateral activation, while the limited number of studies on young infants revealed primarily unilateral activation. The stimulus modality and the age of the participants therefore determine whether the resulting cortical activation is unilateral or bilateral. By combining a review of the existing literature with NIRS results regarding homologous connectivity across hemispheres, I hypothesized that the origin of functional lateralization changes from the independence of each hemispheric region, to mutual inhibition between homologous regions during development. Future studies applying multi-modal measurements along with NIRS and spatiotemporal analyses will further deepen our understanding of the interhemispheric organization of brain function.

Frequency-specific network topologies in the resting human brain.

A community is a set of nodes with dense inter-connections, while there are sparse connections between different communities. A hub is a highly connected node with high centrality. It has been shown that both "communities" and "hubs" exist simultaneously in the brain's functional connectivity network (FCN), as estimated by correlations among low-frequency spontaneous fluctuations in functional magnetic resonance imaging (fMRI) signal changes (0.01-0.10 Hz). This indicates that the brain has a spatial organization that promotes both segregation and integration of information. Here, we demonstrate that frequency-specific network topologies that characterize segregation and integration also exist within this frequency range. In investigating the coherence spectrum among 87 brain regions, we found that two frequency bands, 0.01-0.03 Hz (very low frequency [VLF] band) and 0.07-0.09 Hz (low frequency [LF] band), mainly contributed to functional connectivity. Comparing graph theoretical indices for the VLF and LF bands revealed that the network in the former had a higher capacity for information segregation between identified communities than the latter. Hubs in the VLF band were mainly located within the anterior cingulate cortices, whereas those in the LF band were located in the posterior cingulate cortices and thalamus. Thus, depending on the timescale of brain activity, at least two distinct network topologies contributed to information segregation and integration. This suggests that the brain intrinsically has timescale-dependent functional organizations.

Greater contribution of cerebral than extracerebral hemodynamics to near-infrared spectroscopy signals for functional activation and resting-state connectivity in infants.

While near-infrared spectroscopy (NIRS) has been increasingly applied to neuroimaging and functional connectivity studies in infants, it has not been quantitatively examined as to what extent the deep tissue (such as cerebral tissue) as opposed to shallow tissue (such as scalp), contributes to NIRS signals measured in infants. A method for separating the effects of deep- and shallow-tissue layers was applied to data of nine sleeping three-month-old infants who had been exposed to 3-s speech sounds or silence (i.e., resting state) and whose hemodynamic changes over their bilateral temporal cortices had been measured by using an NIRS system with multiple source-detector (S-D) distances. The deep-layer contribution was found to be large during resting [67% at S-D 20 mm, 78% at S-D 30 mm for oxygenated hemoglobin (oxy-Hb)] as well as during the speech condition (72% at S-D 20 mm, 82% at S-D 30 mm for oxy-Hb). A left-right connectivity analysis showed that correlation coefficients between left and right channels did not differ between original- and deep-layer signals under no-stimulus conditions and that of original- and deep-layer signals were larger than those of the shallow layer. These results suggest that NIRS signals obtained in infants with appropriate S-D distances largely reflected cerebral hemodynamic changes.

Acoustic cues for the recognition of self-voice and other-voice.

Self-recognition, being indispensable for successful social communication, has become a major focus in current social neuroscience. The physical aspects of the self are most typically manifested in the face and voice. Compared with the wealth of studies on self-face recognition, self-voice recognition (SVR) has not gained much attention. Converging evidence has suggested that the fundamental frequency (F0) and formant structures serve as the key acoustic cues for other-voice recognition (OVR). However, little is known about which, and how, acoustic cues are utilized for SVR as opposed to OVR. To address this question, we independently manipulated the F0 and formant information of recorded voices and investigated their contributions to SVR and OVR. Japanese participants were presented with recorded vocal stimuli and were asked to identify the speaker-either themselves or one of their peers. Six groups of 5 peers of the same sex participated in the study. Under conditions where the formant information was fully preserved and where only the frequencies lower than the third formant (F3) were retained, accuracies of SVR deteriorated significantly with the modulation of the F0, and the results were comparable for OVR. By contrast, under a condition where only the frequencies higher than F3 were retained, the accuracy of SVR was significantly higher than that of OVR throughout the range of F0 modulations, and the F0 scarcely affected the accuracies of SVR and OVR. Our results indicate that while both F0 and formant information are involved in SVR, as well as in OVR, the advantage of SVR is manifested only when major formant information for speech intelligibility is absent. These findings imply the robustness of self-voice representation, possibly by virtue of auditory familiarity and other factors such as its association with motor/articulatory representation.

Semantic categories and contexts of written words affect the early ERP component.

We investigated the spatiotemporal patterns of event-related potentials to examine how semantic categories affect early stages of written word processing in the brain. We used a semantic priming paradigm in which the categories of prime and target words were the same (animate or inanimate). Event-related potentials were recorded while native Japanese participants made semantic-relatedness judgments of word pairs. Semantic category effects were observed approximately 150 ms after the target item was presented. Inanimate words elicited greater positive deflection than animate words, but no significant difference was observed for prime item processing. Source modeling estimated that the observed effects were induced by neural mechanisms occurring in the anterior temporal region, the posterior portion of the inferior temporal region, and the inferior parietal regions in both the hemispheres. Our results suggest that semantic category effects in the early stages of word processing are evoked by multiple cortical regions related to top-down processing of the preceding semantic information.

Effect of auditory input on activations in infant diverse cortical regions during audiovisual processing.

A fundamental question with regard to perceptual development is how multisensory information is processed in the brain during the early stages of development. Although a growing body of evidence has shown the early emergence of modality-specific functional differentiation of the cortical regions, the interplay between sensory inputs from different modalities in the developing brain is not well understood. To study the effects of auditory input during audio-visual processing in 3-month-old infants, we evaluated the spatiotemporal cortical hemodynamic responses of 50 infants while they perceived visual objects with or without accompanying sounds. The responses were measured using 94-channel near-infrared spectroscopy over the occipital, temporal, and frontal cortices. The effects of sound manipulation were pervasive throughout the diverse cortical regions and were specific to each cortical region. Visual stimuli co-occurring with sound induced the early-onset activation of the early auditory region, followed by activation of the other regions. Removal of the sound stimulus resulted in focal deactivation in the auditory regions and reduced activation in the early visual region, the association region of the temporal and parietal cortices, and the anterior prefrontal regions, suggesting multisensory interplay. In contrast, equivalent activations were observed in the lateral occipital and lateral prefrontal regions, regardless of sound manipulation. Our findings indicate that auditory input did not generally enhance overall activation in relation to visual perception, but rather induced specific changes in each cortical region. The present study implies that 3-month-old infants may perceive audio-visual multisensory inputs by using the global network of functionally differentiated cortical regions.

A NIRS-fMRI study of resting state network.

Resting state functional connectivity, which is defined as temporal correlation of spontaneous activity between diverse brain regions, has been reported to form resting state networks (RSNs), consisting of a specific set of brain regions, based on functional magnetic resonance imaging (fMRI). Recently, studies using near-infrared spectroscopy (NIRS) reported that NIRS signals also show temporal correlation between different brain regions. The local relationship between NIRS and fMRI signals has been examined by simultaneously recording these signals when participants perform tasks or respond to stimuli. However, the NIRS-fMRI signal relationship during the resting state has been reported only between NIRS signals obtained within limited regions and whole brain fMRI signals. Therefore, it remains unclear whether NIRS signals obtained at diverse regions correlate with regional fMRI signals close to the NIRS measurement channels, especially in relation to the RSNs. In this study, we tested whether the signals measured by these different modalities during the resting state have the consistent characteristics of the RSNs. Specifically, NIRS signals during the resting state were acquired over the frontal, temporal, and occipital cortices while whole brain fMRI data was simultaneously recorded. First, by projecting the NIRS channel positions over the cerebral cortical surface, we identified the most likely anatomical locations of all NIRS channels used in the study. Next, to investigate the regional signal relationship between NIRS and fMRI, we calculated the cross-correlation between NIRS signals and fMRI signals in the brain regions adjacent to each NIRS channel. For each NIRS channel, we observed the local maxima of correlation coefficients between NIRS and fMRI signals within a radius of 2 voxels from the projection point. Furthermore, we also found that highly correlated voxels with the NIRS signal were mainly localized within brain tissues for all NIRS channels, with the exception of 2 frontal channels. Finally, by calculating the correlation between NIRS signals at a channel and whole brain fMRI signals, we observed that NIRS signals correlate with fMRI signals not only within brain regions adjacent to NIRS channels but also within distant brain regions constituting RSNs, such as the dorsal attention, fronto-parietal control, and default mode networks. These results support the idea that NIRS signals obtained at several cortical regions during the resting state mainly reflect regional spontaneous hemodynamic fluctuations that originate from spontaneous cortical activity, and include information that characterizes the RSNs. Because NIRS is relatively easy to use and a less physically demanding neuroimaging technique, our findings should facilitate a broad application of this technique to examine RSNs, especially for clinical populations and conditions unsuitable for fMRI.