Top-down Modulation Research Articles

Brightness illusions drive a neuronal response in the primary visual cortex under top-down modulation

Brightness illusions are a powerful tool in studying vision, yet their neural correlates are poorly understood. Based on a human paradigm, we presented illusory drifting gratings to mice. Primary visual cortex (V1) neurons responded to illusory gratings, matching their direction selectivity for real gratings, and they tracked the spatial phase offset between illusory and real gratings. Illusion responses were delayed compared to real gratings, in line with the theory that processing illusions requires feedback from higher visual areas (HVAs). We provide support for this theory by showing a reduced V1 response to illusions, but not real gratings, following HVAs optogenetic inhibition. Finally, we used the pupil response (PR) as an indirect perceptual report and showed that the mouse PR matches the human PR to perceived luminance changes. Our findings resolve debates over whether V1 neurons are involved in processing illusions and highlight the involvement of feedback from HVAs.

Altered functional connectivity strength of primary visual cortex in subjects with thyroid-associated ophthalmopathy.

Our objective was to explore the disparities in the intrinsic functional connectivity (FC) patterns of primary visual cortex (V1) between patients with thyroid-associated ophthalmopathy (TAO) and healthy controls (HCs) utilizing resting-state functional MRI. Twenty-one patients with TAO (14 males and 7 females; mean age: 54.17 ± 4.83 years) and 21 well-matched HCs (14 males and 7 females; mean age: 55.17 ± 5.37 years) underwent functional MRI scans in the resting-state. We assessed modifications in the intrinsic FC patterns of the V1 in TAO patients using the FC method. Subsequently, the identified alterations in FC regions in the analysis were selected as classification features to distinguish TAO patients from HCs through the support vector machine (SVM) method. The results indicated that, in comparison to HCs, patients with TAO exhibited notably reduced FC values between the left V1 and the bilateral calcarine (CAL), lingual gyrus (LING) and superior occipital gyrus, as well as between the right V1 and the bilateral CAL/LING and the right cerebellum. Furthermore, the SVM classification model based on FC maps demonstrated effective performance in distinguishing TAO patients from HCs, achieving an accuracy of 61.9% using the FC of the left V1 and 64.29% using the FC of the right V1. Our study revealed that patients with TAO manifested disruptions in FC between the V1 and higher visual regions during rest. This might indicate that TAO patients could present with impaired top-down modulations, visual imagery and vision-motor function. These insights could be valuable in understanding the underlying neurobiological mechanisms of vision impairment in individuals with TAO.

Top–down modulation in canonical cortical circuits with short-term plasticity

Cortical dynamics and computations are strongly influenced by diverse GABAergic interneurons, including those expressing parvalbumin (PV), somatostatin (SST), and vasoactive intestinal peptide (VIP). Together with excitatory (E) neurons, they form a canonical microcircuit and exhibit counterintuitive nonlinear phenomena. One instance of such phenomena is response reversal, whereby SST neurons show opposite responses to top-down modulation via VIP depending on the presence of bottom-up sensory input, indicating that the network may function in different regimes under different stimulation conditions. Combining analytical and computational approaches, we demonstrate that model networks with multiple interneuron subtypes and experimentally identified short-term plasticity mechanisms can implement response reversal. Surprisingly, despite not directly affecting SST and VIP activity, PV-to-E short-term depression has a decisive impact on SST response reversal. We show how response reversal relates to inhibition stabilization and the paradoxical effect in the presence of several short-term plasticity mechanisms demonstrating that response reversal coincides with a change in the indispensability of SST for network stabilization. In summary, our work suggests a role of short-term plasticity mechanisms in generating nonlinear phenomena in networks with multiple interneuron subtypes and makes several experimentally testable predictions.

Top-down attention does not modulate mechanical hypersensitivity consecutive to central sensitization: insights from an experimental analysis.

According to the neurocognitive model of attention to pain, when the attentional resources invested in a task unrelated to pain are high, limited cognitive resources can be directed toward the pain. This is supported by experimental studies showing that diverting people's attention away from acute pain leads to experiencing less pain. Theoretical work has suggested that this phenomenon may present a top-down modulatory mechanism for persistent pain as well. However, conclusive empirical evidence is lacking. To fill this gap, we used a preregistered, double-blind, between-subject study design to investigate whether performing a tailored, demanding, and engaging working memory task unrelated to pain (difficult) vs a task that requires less mental effort to be performed (easy), could lead to lower development of secondary hypersensitivity-a hallmark of central sensitization. Eighty-five healthy volunteers, randomly assigned to one of the 2 conditions, performed a visual task with a different cognitive load (difficult vs easy), while secondary hypersensitivity was induced on their nondominant forearm using high-frequency stimulation. To assess the development of secondary hypersensitivity, sensitivity to mechanical stimuli was measured 3 times: T0, for baseline and 20 (T1) and 40 (T2) minutes after the procedure. We did not observe any significant difference in the development of secondary hypersensitivity between the 2 groups, neither in terms of the intensity of mechanical sensitivity nor its spatial extent. Our results suggest that a top-down modulation through attention might not be sufficient to affect pain sensitization and the development of secondary hypersensitivity.

Attention to audiovisual speech shapes neural processing through feedback-feedforward loops between different nodes of the speech network.

Selective attention-related top-down modulation plays a significant role in separating relevant speech from irrelevant background speech when vocal attributes separating concurrent speakers are small and continuously evolving. Electrophysiological studies have shown that such top-down modulation enhances neural tracking of attended speech. Yet, the specific cortical regions involved remain unclear due to the limited spatial resolution of most electrophysiological techniques. To overcome such limitations, we collected both electroencephalography (EEG) (high temporal resolution) and functional magnetic resonance imaging (fMRI) (high spatial resolution), while human participants selectively attended to speakers in audiovisual scenes containing overlapping cocktail party speech. To utilise the advantages of the respective techniques, we analysed neural tracking of speech using the EEG data and performed representational dissimilarity-based EEG-fMRI fusion. We observed that attention enhanced neural tracking and modulated EEG correlates throughout the latencies studied. Further, attention-related enhancement of neural tracking fluctuated in predictable temporal profiles. We discuss how such temporal dynamics could arise from a combination of interactions between attention and prediction as well as plastic properties of the auditory cortex. EEG-fMRI fusion revealed attention-related iterative feedforward-feedback loops between hierarchically organised nodes of the ventral auditory object related processing stream. Our findings support models where attention facilitates dynamic neural changes in the auditory cortex, ultimately aiding discrimination of relevant sounds from irrelevant ones while conserving neural resources.

Temporal Properties of Self-Prioritization.

Using electroencephalogram (EEG), we tested the hypothesis that the association of a neutral stimulus with the self would elicit ultra-fast neural responses from early top-down feedback modulation to late feedforward periods for cognitive processing, resulting in self-prioritization in information processing. In two experiments, participants first learned three associations between personal labels (self, friend, stranger) and geometric shapes (Experiment 1) and three colors (Experiment 2), and then they judged whether the shape/color-label pairings matched. Stimuli in Experiment 2 were shown in a social communicative setting with two avatars facing each other, one aligned with the participant's view (first-person perspective) and the other with a third-person perspective. The color was present on the t-shirt of one avatar. This setup allowed for an examination of how social contexts (i.e., perspective taking) affect neural connectivity mediating self-related processing. Functional connectivity analyses in the alpha band (8-12 Hz) revealed that self-other discrimination was mediated by two distinct phases of neural couplings between frontal and occipital regions, involving an early phase of top-down feedback modulation from frontal to occipital areas followed by a later phase of feedforward signaling from occipital to frontal regions. Moreover, while social communicative settings influenced the later feedforward connectivity phase, they did not alter the early feedback coupling. The results indicate that regardless of stimulus type and social context, the early phase of neural connectivity represents an enhanced state of awareness towards self-related stimuli, whereas the later phase of neural connectivity may be associated with cognitive processing of socially meaningful stimuli.

Transduction, Calibration, and the Penetrability of Pain

Pains are subject to obvious, well-documented, and striking top-down influences. This is in stark contrast to visual perception, where the debate over cognitive penetrability tends to revolve around fairly subtle experimental effects. Several authors have recently taken up the question of whether top-down effects on pain count as cognitive penetrability, and what that might show us about traditional debates. I review some of the known mechanisms for top-down modulation of pain, and suggest that it reveals an issue with a relatively neglected part of the cognitive penetrability literature. Much of the debate inherits Pylyshyn’s stark contrast between transducers and cognition proper. His distinction grew out of his running fight with Gibson, and is far too strong to be defensible. I suggest that we might therefore view top-down influences on pain as a species of transducer calibration. This provides a novel but principled way to distinguish between several varieties of top-down effect according to their architectural features.

Rapid color categorization revealed by frequency-tagging-based EEG

There has been much debate on whether color categories affect how we perceive color. Recent theories have put emphasis on the role of top-down influence on color perception that the original continuous color space in the visual cortex may be transformed into categorical encoding due to top-down modulation. To test the influence of color categories on color perception, we adopted an RSVP paradigm, where color stimuli were presented at a fast speed of 100 ms per stimulus and were forward and backward masked by the preceding and following stimuli. Moreover, no explicit color naming or categorization was required. In theory, backward masking with such a short interval in a passive viewing task should constrain top-down influence from higher-level brain areas. To measure any potentially subtle differences in brain response elicited by different color categories, we embedded a sensitive frequency-tagging-based EEG paradigm within the RSVP stimuli stream where the oddball color stimuli were encoded with a different frequency from the base color stimuli. We showed that EEG responses to cross-category oddball colors at the frequency where the oddball stimuli were presented was significantly larger than the responses to within-category oddball colors. Our study suggested that the visual cortex can automatically and implicitly encode color categories when color stimuli are presented rapidly.

Electrophysiological (EEG) Correlates of Reward Effects on Early Sensory Perception in Humans

Existing research suggests that reward system and sensory perception networks function in concert and that activation in one may influence the other; however, the dynamics of these influences remains poorly understood. There is general agreement regarding the existence of an interaction between bottom-up and top-down signals in perception and attentional processing. While it’s not absolutely sure what stages of perception are influenced by cognition, it has been assumed that cognitive input influences later categorization stages of visual processing, and that earlier stages are solely involved in the pure bottom up extraction of basic features of sensory signals. Several recent experiments have challenged this idea by showing that top-down modulation by cognition may extend to early visual stages of perception. Recent Electrophysiological studies have begun to probe the neural correlates of the interaction between attention/reward, perception and cognitive control in humans. . We posit that selection of the value of our choices and actions from multiple alternatives may cause suppression of the sensory representations of unselected, low value stimuli while the selected, high value stimuli are enhanced. The present project has been proposed to study the dynamics of this selective process where effects of various reward categories on attention and early sensory perception are tracked using behavioral and electrophysiological (EEG) techniques. Methods from neuroscience, signal processing, psychophysics and EEG tools will be used in the project.

FMRI BOLD responses to film stimuli and their association with exhaled nitric oxide in asthma and health.

Little is known about central nervous system (CNS) responses to emotional stimuli in asthma. Nitric oxide in exhaled breath (FENO) is elevated in asthma due to allergic immune processes, but endogenous nitric oxide is also known to modulate CNS activity. We measured fMRI blood oxygen-dependent (BOLD) brain activation to negative (blood-injection-injury themes) and neutral films in 31 participants (15 with asthma). Regions-of-interest analysis was performed on key areas relevant to central adaptive control, threat processing, or salience networks, with dorsolateral prefrontal cortex (PFC), anterior insula, dorsal anterior cingulate cortex (dACC), amygdala, ventral striatum, ventral tegmentum, and periaqueductal gray, as well as top-down modulation of emotion, with ventrolateral and ventromedial PFC. Both groups showed less BOLD deactivation from fixation cross-baseline in the left anterior insula and bilateral ventromedial PFC for negative than neutral films, and for an additional number of areas, including the fusiform gyrus, for film versus recovery phases. Less deactivation during films followed by less recovery from deactivation was found in asthma compared to healthy controls. Changes in PCO2 did not explain these findings. FENO was positively related to BOLD activation in general, but more pronounced in healthy controls and more likely in neutral film processing. Thus, asthma is associated with altered processing of film stimuli across brain regions not limited to central adaptive control, threat processing, or salience networks. Higher levels of NO appear to facilitate CNS activity, but only in healthy controls, possibly due to allergy's masking effects on FENO.

Dynamic predictive coding: A model of hierarchical sequence learning and prediction in the neocortex.

We introduce dynamic predictive coding, a hierarchical model of spatiotemporal prediction and sequence learning in the neocortex. The model assumes that higher cortical levels modulate the temporal dynamics of lower levels, correcting their predictions of dynamics using prediction errors. As a result, lower levels form representations that encode sequences at shorter timescales (e.g., a single step) while higher levels form representations that encode sequences at longer timescales (e.g., an entire sequence). We tested this model using a two-level neural network, where the top-down modulation creates low-dimensional combinations of a set of learned temporal dynamics to explain input sequences. When trained on natural videos, the lower-level model neurons developed space-time receptive fields similar to those of simple cells in the primary visual cortex while the higher-level responses spanned longer timescales, mimicking temporal response hierarchies in the cortex. Additionally, the network's hierarchical sequence representation exhibited both predictive and postdictive effects resembling those observed in visual motion processing in humans (e.g., in the flash-lag illusion). When coupled with an associative memory emulating the role of the hippocampus, the model allowed episodic memories to be stored and retrieved, supporting cue-triggered recall of an input sequence similar to activity recall in the visual cortex. When extended to three hierarchical levels, the model learned progressively more abstract temporal representations along the hierarchy. Taken together, our results suggest that cortical processing and learning of sequences can be interpreted as dynamic predictive coding based on a hierarchical spatiotemporal generative model of the visual world.

Category-based attention facilitates memory search.

We often need to decide whether the object we look at is also the object we look for. When we look for one specific object, this process can be facilitated by feature-based attention. However, when we look for many objects at the same time (e.g., the products on our shopping list) such a strategy may no longer be possible, as research has shown that we can actively prepare to detect only one or two objects at a time. Therefore, looking for multiple objects additionally requires long-term memory search, slowing down decision making. Interestingly, however, previous research has shown that distractor objects can be efficiently rejected during memory search when they are from a different category than the items in the memory set. Here, using EEG, we show that this efficiency is supported by top-down attention at the category level. In Experiment 1, human participants (both sexes) performed a memory search task on individually presented objects from different categories, most of which were distractors. We observed category-level attentional modulation of distractor processing from ∼150 ms after stimulus onset, expressed both as an evoked response modulation and as an increase in decoding accuracy of same-category distractors. In Experiment 2, memory search was performed on two concurrently presented objects. When both objects were distractors, spatial attention (indexed by the N2pc component) was directed to the object that was of the same category as the objects in the memory set. Together, these results demonstrate how top-down attention can facilitate memory search.Significance statement When we are in the supermarket, we repeatedly decide whether a product we look at (e.g., a banana) is on our memorized shopping list (e.g., apples, oranges, kiwis). This requires searching our memory, which takes time. However, when the product is of an entirely different category (e.g., dairy instead of fruit), the decision can be made quickly. Here, we used EEG to show that this between-category advantage in memory search tasks is supported by top-down attentional modulation of visual processing: The visual response evoked by distractor objects was modulated by category membership, and spatial attention was quickly directed to the location of within-category (vs. between-category) distractors. These results demonstrate a close link between attention and memory.

Neural Decoding of the Speech Envelope: Effects of Intelligibility and Spectral Degradation.

During continuous speech perception, endogenous neural activity becomes time-locked to acoustic stimulus features, such as the speech amplitude envelope. This speech-brain coupling can be decoded using non-invasive brain imaging techniques, including electroencephalography (EEG). Neural decoding may provide clinical use as an objective measure of stimulus encoding by the brain-for example during cochlear implant listening, wherein the speech signal is severely spectrally degraded. Yet, interplay between acoustic and linguistic factors may lead to top-down modulation of perception, thereby complicating audiological applications. To address this ambiguity, we assess neural decoding of the speech envelope under spectral degradation with EEG in acoustically hearing listeners (n = 38; 18-35 years old) using vocoded speech. We dissociate sensory encoding from higher-order processing by employing intelligible (English) and non-intelligible (Dutch) stimuli, with auditory attention sustained using a repeated-phrase detection task. Subject-specific and group decoders were trained to reconstruct the speech envelope from held-out EEG data, with decoder significance determined via random permutation testing. Whereas speech envelope reconstruction did not vary by spectral resolution, intelligible speech was associated with better decoding accuracy in general. Results were similar across subject-specific and group analyses, with less consistent effects of spectral degradation in group decoding. Permutation tests revealed possible differences in decoder statistical significance by experimental condition. In general, while robust neural decoding was observed at the individual and group level, variability within participants would most likely prevent the clinical use of such a measure to differentiate levels of spectral degradation and intelligibility on an individual basis.

Top-down modulation of DLPFC in visual search: a study based on fMRI and TMS.

Effective visual search is essential for daily life, and attention orientation as well as inhibition of return play a significant role in visual search. Researches have established the involvement of dorsolateral prefrontal cortex in cognitive control during selective attention. However, neural evidence regarding dorsolateral prefrontal cortex modulates inhibition of return in visual search is still insufficient. In this study, we employed event-related functional magnetic resonance imaging and dynamic causal modeling to develop modulation models for two types of visual search tasks. In the region of interest analyses, we found that the right dorsolateral prefrontal cortex and temporoparietal junction were selectively activated in the main effect of search type. Dynamic causal modeling results indicated that temporoparietal junction received sensory inputs and only dorsolateral prefrontal cortex →temporoparietal junction connection was modulated in serial search. Such neural modulation presents a significant positive correlation with behavioral reaction time. Furthermore, theta burst stimulation via transcranial magnetic stimulation was utilized to modulate the dorsolateral prefrontal cortex region, resulting in the disappearance of the inhibition of return effect during serial search after receiving continuous theta burst stimulation. Our findings provide a new line of causal evidence that the top-down modulation by dorsolateral prefrontal cortex influences the inhibition of return effect during serial search possibly through the retention of inhibitory tagging via working memory storage.

Pathway-selective optogenetics reveals the functional anatomy of top-down attentional modulation in the macaque visual cortex.

Spatial attention represents a powerful top-down influence on sensory responses in primate visual cortical areas. The frontal eye field (FEF) has emerged as a key candidate area for the source of this modulation. However, it is unclear whether the FEF exerts its effects via its direct axonal projections to visual areas or indirectly through other brain areas and whether the FEF affects both the enhancement of attended and the suppression of unattended sensory responses. We used pathway-selective optogenetics in rhesus macaques performing a spatial attention task to inhibit the direct input from the FEF to area MT, an area along the dorsal visual pathway specialized for the processing of visual motion information. Our results show that the optogenetic inhibition of the FEF input specifically reduces attentional modulation in MT by about a third without affecting the neurons' sensory response component. We find that the direct FEF-to-MT pathway contributes to both the enhanced processing of target stimuli and the suppression of distractors. The FEF, thus, selectively modulates firing rates in visual area MT, and it does so via its direct axonal projections.

Top-down modulation of dichotic listening affects interhemispheric connectivity: an electroencephalography study.

Dichotic listening (DL) has been extensively used as a task to investigate auditory processing and hemispheric lateralisation in humans. According to the "callosal relay model," the typical finding of a right ear advantage (REA) occurs because the information coming from the right ear has direct access to the left dominant hemisphere while the information coming from the left ear has to cross via the corpus callosum. The underlying neuroanatomical correlates and neurophysiological mechanisms have been described using diffusion tensor imaging (DTI) and lagged phase synchronization (LPS) of the interhemispheric auditory pathway. During the non-forced condition of DL, functional connectivity (LPS) of interhemispheric gamma-band coupling has been described as a relevant mechanism related to auditory perception in DL. In this study, we aimed to extend the previous results by exploring the effects of top-down modulation of DL (forced-attention condition) on interhemispheric gamma-band LPS. Right-handed healthy participants (n = 31; 17 females) performed three blocks of DL with different attention instructions (no-attention, left-ear attention, right-ear attention) during simultaneous EEG recording with 64 channels. Source analysis was done with exact low-resolution brain electromagnetic tomography (eLORETA) and functional connectivity between bilateral auditory areas was assessed as LPS in the gamma-band frequency range. Twenty-four participants (77%) exhibited a right-ear advantage in the no-attention block. The left- and right-attention conditions significantly decreased and increased right-ear reports, respectively. Similar to the previous studies, functional connectivity analysis (gamma-band LPS) showed significantly increased connectivity between left and right Brodmann areas (BAs) 41 and 42 during left ear reports in contrast with right ear reports. Our new findings notably indicated that the right-attention condition exhibited significantly higher connectivity between BAs 42 compared with the no-attention condition. This enhancement of connectivity was more pronounced during the perception of right ear reports. Our results are in line with previous reports describing gamma-band synchronization as a relevant neurophysiological mechanism involved in the interhemispheric connectivity according to the callosal relay model. Moreover, we newly added some evidence of attentional effects on this interhemispheric connectivity, consistent with the attention-executive model. Our results suggest that reciprocal inhibition could be involved in hemispheric lateralization processes.

Mismatch Responses to Speech Contrasts in Preschoolers with and without Developmental Language Disorder.

This study compared cortical responses to speech in preschoolers with typical language development (TLD) and with Developmental Language Disorder (DLD). We investigated whether top-down language effects modulate speech perception in young children in an adult-like manner. We compared cortical mismatch responses (MMRs) during the passive perception of speech contrasts in three groups of participants: preschoolers with TLD (n = 11), preschoolers with DLD (n = 16), and adults (n = 20). We also measured children's phonological skills and investigated whether they are associated with the cortical discrimination of phonemic changes involving different linguistic complexities. The results indicated top-down language effects in adults, with enhanced cortical discrimination of lexical stimuli but not of non-words. In preschoolers, the TLD and DLD groups did not differ in the MMR measures, and no top-down effects were detected. Moreover, we found no association between MMRs and phonological skills, even though the DLD group's phonological skills were significantly lower. Our findings suggest that top-down language modulations in speech discrimination may not be present during early childhood, and that children with DLD may not exhibit cortical speech perception deficits. The lack of association between phonological and MMR measures indicates that further research is needed to understand the link between language skills and cortical activity in preschoolers.

Spontaneous Modulation Doping in Semi-Crystalline Conjugated Polymers Leads to High Conductivity at Low Doping Concentration.

The possibility to control the charge carrier density through doping is one of the defining properties of semiconductors. For organic semiconductors, the doping process is known to come with several problems associated with the dopant compromising the charge carrier mobility by deteriorating the host morphology and/or introducing Coulomb traps. While for inorganic semiconductors these factors can be mitigated through (top-down) modulation doping, this concept has not been employed in organics. Here, this work shows that properly chosen host/dopant combinations can give rise to spontaneous, bottom-up modulation doping, in which the dopants preferentially sit in an amorphous phase, while the actual charge transport occurs predominantly in a crystalline phase with an unaltered microstructure, spatially separating dopants and mobile charges. Combining experiments and numerical simulations, this work shows that this leads to exceptionally high conductivities at relatively low dopant concentrations.

Action-specific feature processing in the human cortex: An fMRI study

Sensorimotor integration involves feedforward and reentrant processing of sensory input. Grasp-related motor activity precedes and is thought to influence visual object processing. Yet, while the importance of reentrant feedback is well established in perception, the top-down modulations for action and the neural circuits involved in this process have received less attention. Do action-specific intentions influence the processing of visual information in the human cortex? Using a cue-separation fMRI paradigm, we found that action-specific instruction processing (manual alignment vs. grasp) became apparent only after the visual presentation of oriented stimuli, and occurred as early as in the primary visual cortex and extended to the dorsal visual stream, motor and premotor areas. Further, dorsal stream area aIPS, known to be involved in object manipulation, and the primary visual cortex showed task-related functional connectivity with frontal, parietal and temporal areas, consistent with the idea that reentrant feedback from dorsal and ventral visual stream areas modifies visual inputs to prepare for action. Importantly, both the task-dependent modulations and connections were linked specifically to the object presentation phase of the task, suggesting a role in processing the action goal. Our results show that intended manual actions have an early, pervasive, and differential influence on the cortical processing of vision.

Neurophysiological oscillatory markers of hypoalgesia in conditioned pain modulation.

Conditioned pain modulation (CPM) is an experimental procedure that consists of an ongoing noxious stimulus attenuating the pain perception caused by another noxious stimulus. A combination of the CPM paradigm with concurrent electrophysiological recordings can establish whether an association exists between experimentally modified pain perception and modulations of neural oscillations. We aimed to characterize how CPM modifies pain perception and underlying neural oscillations. We also interrogated whether these perceptual and/or neurophysiological effects are distinct in patients affected by chronic pain. We presented noxious electrical stimuli to the right ankle before, during, and after CPM induced by an ice pack placed on the left forearm. Seventeen patients with chronic pain and 17 control participants rated the electrical pain in each experimental condition. We used magnetoencephalography to examine the anatomy-specific effects of CPM on the neural oscillatory responses to the electrical pain. Regardless of the participant groups, CPM induced a reduction in subjective pain ratings and neural responses (beta-band [15-35 Hz] oscillations in the sensorimotor cortex) to electrical pain. Our findings of pain-induced beta-band activity may be associated with top-down modulations of pain, as reported in other perceptual modalities. Therefore, the reduced beta-band responses during CPM may indicate changes in top-down pain modulations.