The future of metacognition research: Balancing construct breadth with measurement rigor.

Foundational work in the psychology of metacognition identified a distinction between metacognitive knowledge (stable beliefs about one's capacities) and metacognitive experiences (local evaluations of performance). More recently, the field has focused on developing tasks and metrics that seek to identify metacognitive capacities from momentary estimates of confidence in performance, and providing precise computational accounts of metacognitive failure. However, this notable progress in formalising models of metacognitive judgments may come at a cost of ignoring broader elements of the psychology of metacognition - such as how stable meta-knowledge is formed, how social cognition and metacognition interact, and how we evaluate affective states that do not have an obvious ground truth. We propose that construct breadth in metacognition research can be restored while maintaining rigour in measurement, and highlight promising avenues for expanding the scope of metacognition research. Such a research programme is well placed to recapture qualitative features of metacognitive knowledge and experience while maintaining the psychophysical rigor that characterises modern research on confidence and performance monitoring.

An Accumulating Neural Signal Underlying Binocular Rivalry Dynamics.

During binocular rivalry, conflicting images are presented one to each eye and perception alternates stochastically between them. Despite stable percepts between alternations, modeling suggests that neural signals representing the two images change gradually, and that the duration of stable percepts are determined by the time required for these signals to reach a threshold that triggers an alternation. However, direct physiological evidence for such signals has been lacking. Here, we identify a neural signal in the human visual cortex that shows these predicted properties. We measured steady-state visual evoked potentials (SSVEPs) in 84 human participants (62 females, 22 males) who were presented with orthogonal gratings, one to each eye, flickering at different frequencies. Participants indicated their percept while EEG data were collected. The time courses of the SSVEP amplitudes at the two frequencies were then compared across different percept durations, within participants. For all durations, the amplitude of signals corresponding to the suppressed stimulus increased and the amplitude corresponding to the dominant stimulus decreased throughout the percept. Critically, longer percepts were characterized by more gradual increases in the suppressed signal and more gradual decreases of the dominant signal. Changes in signals were similar and rapid at the end of all percepts, presumably reflecting perceptual transitions. These features of the SSVEP time courses are well predicted by a model in which perceptual transitions are produced by the accumulation of noisy signals. Identification of this signal underlying binocular rivalry should allow strong tests of neural models of rivalry, bistable perception, and neural suppression.SIGNIFICANCE STATEMENT During binocular rivalry, two conflicting images are presented to the two eyes and perception alternates between them, with switches occurring at seemingly random times. Rivalry is an important and longstanding model system in neuroscience, used for understanding neural suppression, intrinsic neural dynamics, and even the neural correlates of consciousness. All models of rivalry propose that it depends on gradually changing neural activity that on reaching some threshold triggers the perceptual switches. This manuscript reports the first physiological measurement of neural signals with that set of properties in human participants. The signals, measured with EEG in human observers, closely match the predictions of recent models of rivalry, and should pave the way for much future work.

Reducing and deducing the structures of consciousness through meditation.

According to many first-person accounts, consciousness comprises a subject-object structure involving a mental action or attitude starting from the "subjective pole" upon an object of experience. In recent years, many paradigms have been developed to manipulate and empirically investigate the object of consciousness. However, well-controlled investigation of subjective aspects of consciousness has been more challenging. One way, subjective aspects of consciousness are proposed to be studied is using meditation states that alter its subject-object structure. Most work to study consciousness in this way has been done using Buddhist meditation traditions and techniques. There is another meditation tradition that has been around for at least as long as early Buddhist traditions (if not longer) with the central goal of developing a fine-grained first-person understanding of consciousness and its constituents by its manipulation through meditation, namely the Tantric tradition of Yoga. However, due to the heavy reliance of Yogic traditions on the ancient Indian Samkhya philosophical system, their insights about consciousness have been more challenging to translate into contemporary research. Where such translation has been attempted, they have lacked accompanying phenomenological description of the procedures undertaken for making the precise subject-object manipulations as postulated. In this paper, I address these issues by first detailing how Tantric Yoga philosophy can be effectively translated as a systematic phenomenological account of consciousness spanning the entirety of the subject-object space divided into four "structures of consciousness" from subject to object. This follows from the work of the 20th century polymath and founder of the Tantric Yoga school of Ananda Marga, Prabhat Ranjan Sarkar, who expounded on the "cognitivization" of Samkhya philosophy. I then detail stepwise meditation procedures that make theoretical knowledge of these structures of consciousness a practical reality to a Tantric Yoga meditator in the first-person. This is achieved by entering meditative states through stepwise experiential reduction of the structures of consciousness from object to subject, as part of their meditative goal of "self-realization." I end by briefly discussing the overlap of these putative meditation states with proposed states from other meditation traditions, and how these states could help advance an empirical study of consciousness.

Characteristics of Kundalini-Related Sensory, Motor, and Affective Experiences During Tantric Yoga Meditation.

Traditional spiritual literature contains rich anecdotal reports of spontaneously arising experiences occurring during meditation practice, but formal investigation of such experiences is limited. Previous work has sometimes related spontaneous experiences to the Indian traditional contemplative concept of kundalini. Historically, descriptions of kundalini come out of Tantric schools of Yoga, where it has been described as a "rising energy" moving within the spinal column up to the brain. Spontaneous meditation experiences have previously been studied within Buddhist and Christian practices and within eclectic groups of contemplative practitioners. Prior explorations of kundalini have emphasized extreme experiences, sometimes having clinical consequences. We conducted a first such investigation of kundalini-related experiences within a sample of meditators from a single Tantric Yoga tradition (known as Ananda Marga) that emphasizes the role of kundalini. We developed a semi-structured questionnaire to conduct an exploratory pilot investigation of spontaneous sensory, motor and affective experiences during meditation practice. In addition to identifying the characteristics of subjective experiences, we measured quantity of meditation, supplemental practices, trait affect and trait mindfulness. We administered it to 80 volunteers at two Ananda Marga retreats. Among reported experiences, we found the highest prevalence for positive mood shifts, followed by motor and then sensory experiences. The frequency of spontaneous experiences was not related to the quantity of practiced meditation or trait measures of mindfulness and affect. Self-reports included multiple descriptions of rising sensations, sometimes being directly called kundalini. Experiences with rising sensations were complex and many included references to positive affect, including ecstatic qualities. There were also reports of spontaneous anomalous experiences. These experiences of rising sensations resemble prior clinical descriptions that were considered kundalini-related. The individuals who reported rising sensations could not be distinguished from other participants based on the incidence of experiences, quantity of meditation practice, or trait measures of mindfulness and affect. In contrast, greater amount of Tantric Yoga meditation practice was associated with greater positive affect, less negative affect and greater mindfulness. Further study of these exploratory findings and how they may be related to spiritual and well-being goals of meditation is warranted along with scientific investigation of purported kundalini phenomena.

Alpha and theta oscillations are inversely related to progressive levels of meditation depth.

Meditation training is proposed to enhance mental well-being by modulating neural activity, particularly alpha and theta brain oscillations, and autonomic activity. Although such enhancement also depends on the quality of meditation, little is known about how these neural and physiological changes relate to meditation quality. One model characterizes meditation quality as five increasing levels of 'depth': hindrances, relaxation, concentration, transpersonal qualities and nonduality. We investigated the neural oscillatory (theta, alpha, beta and gamma) and physiological (respiration rate, heart rate and heart rate variability) correlates of the self-reported meditation depth in long-term meditators (LTMs) and meditation-naïve controls (CTLs). To determine the neural and physiological correlates of meditation depth, we modelled the change in the slope of the relationship between self-reported experiential degree at each of the five depth levels and the multiple neural and physiological measures. CTLs reported experiencing more 'hindrances' than LTMs, while LTMs reported more 'transpersonal qualities' and 'nonduality' compared to CTLs, confirming the experiential manipulation of meditation depth. We found that in both groups, theta (4-6 Hz) and alpha (7-13 Hz) oscillations were related to meditation depth in a precisely opposite manner. The theta amplitude positively correlated with 'hindrances' and increasingly negatively correlated with increasing meditation depth levels. Alpha amplitude negatively correlated with 'hindrances' and increasingly positively with increasing depth levels. The increase in the inverse association between theta and meditation depth occurred over different scalp locations in the two groups-frontal midline in LTMs and frontal lateral in CTLs-possibly reflecting the downregulation of two different aspects of executive processing-monitoring and attention regulation, respectively-during deep meditation. These results suggest a functional dissociation of the two classical neural signatures of meditation training, namely, alpha and theta oscillations. Moreover, while essential for overcoming 'hindrances', executive neural processing appears to be downregulated during deeper meditation experiences.

Neural correlates of nonjudgmental perception induced through meditation.

Ambiguous sensory stimuli provide insight into the dynamics of the human mind. When viewing substantially different images in the two eyes (i.e., binocular rivalry (BR)), perception spontaneously fluctuates between the two images along with patch-like mixtures of the two, with limited ability to control such fluctuations. Previous studies have shown that long-term meditation training can enable a more stable perception by reducing such fluctuations. Using electroencephalography, we investigated the neural bases of perceptual stabilization in long-term meditators (LTMs) and age-matched meditation-naive control participants. We measured BR alternations before and after participants practiced meditation. We expected that perceptual stabilization through meditation could occur via one of two neurocognitive mechanisms: (1) a more engaged/effortful attention reflected by increased long-range phase synchronization between early visual sensory and higher-level brain regions, or (2) a disengaged/nonevaluative form of attention reflected by decreased phase synchronization. We found that compared with control participants, LTMs were in a significantly longer mixed perceptual state following concentrative meditation practice. The increase in mixed percepts across individuals was strongly correlated with reduced parietal-occipital gamma-band (30-50 Hz) phase synchrony. These findings suggest that concentrative meditation enables a nonevaluative perceptual stance supported by reduced communication between hierarchical visual brain regions.

Event-related potential and behavioural differences in affective self-referential processing in long-term meditators versus controls.

Contemplative practices are thought to modify one's experience of self and fundamentally change self-referential processing. However, few studies have examined the brain correlates of self-referential processing in long-term meditators. Here, we used the self-referential encoding task (SRET) to examine event-related potentials (ERP) during assessment of pleasant and unpleasant self-views in long-term meditators versus age-matched meditation-naïve control participants. Compared with controls, meditators endorsed significantly more pleasant and fewer unpleasant words as self-referential. We also found a between-group difference in the early component of the late-positive-potential (LPP) of the ERP characterized by a larger response to unpleasant versus pleasant words in controls and no difference in meditators. A cross-sectional design, such as the one used in the present study, has certain caveats like self-selectivity bias. If such caveats did not affect our results, these findings suggest that a long-term contemplative lifestyle, of which meditation training is an integral part, alters self-referential processing towards a more adaptive view of self and neural equivalence towards pleasant and unpleasant self-views. These findings suggest that long-term meditation training may affect brain and behavioural mechanism that support a more flexible and healthy relationship to one's self.

Frequency of alpha oscillation predicts individual differences in perceptual stability during binocular rivalry.

When ambiguous visual stimuli have multiple interpretations, human perception can alternate between them, producing perceptual multistability. There is a large variation between individuals in how long stable percepts endure, on average, between switches, but the underlying neural basis of this individual difference in perceptual dynamics remains obscure. Here, we show that in one widely studied multistable paradigm-binocular rivalry-perceptual stability in individuals is predicted by the frequency of their neural oscillations within the alpha range (7-13 Hz). Our results suggest revising models of rivalry to incorporate effects of neural oscillations on perceptual alternations, and raise the possibility that a common factor may influence dynamics in many neural processes.

Conflict-sensitive neurons gate interocular suppression in human visual cortex.

Neural suppression plays an important role in cortical function, including sensory, memory, and motor systems. It remains, however, relatively poorly understood. A paradigmatic case arises when conflicting images are presented to the two eyes. These images can compete for awareness, and one is usually strongly suppressed. The mechanisms that resolve such interocular conflict remain unclear. Suppression could arise solely from "winner-take-all" competition between neurons responsive to each eye. Alternatively, suppression could also depend upon neurons detecting interocular conflict. Here, we provide physiological evidence in human visual cortex for the latter: suppression depends upon conflict-sensitive neurons. We recorded steady-state visual evoked potentials (SSVEP), and used the logic of selective adaptation. The amplitude of SSVEP responses at intermodulation frequencies strengthened as interocular conflict in the stimulus increased, suggesting the presence of neurons responsive to conflict. Critically, adaptation to conflict both reduced this SSVEP effect, and increased the amount of conflict needed to produce perceptual suppression. The simplest account of these results is that interocular-conflict-sensitive neurons exist in human cortex: adaptation likely reduced the responsiveness of these neurons which in turn raised the amount of conflict required to produce perceptual suppression. Similar mechanisms may be used to resolve other varieties of perceptual conflict.

Polar-angle representation of saccadic eye movements in human superior colliculus.

The superior colliculus (SC) is a layered midbrain structure involved in directing both head and eye movements and coordinating visual attention. Although a retinotopic organization for the mediation of saccadic eye-movements has been shown in monkey SC, in human SC the topography of saccades has not been confirmed. Here, a novel experimental paradigm was performed by five participants (one female) while high-resolution (1.2-mm) functional magnetic resonance imaging was used to measure activity evoked by saccadic eye movements within human SC. Results provide three critical observations about the topography of the SC: (1) saccades along the superior-inferior visual axis are mapped across the medial-lateral anatomy of the SC; (2) the saccadic eye-movement representation is in register with the retinotopic organization of visual stimulation; and (3) activity evoked by saccades occurs deeper within SC than that evoked by visual stimulation. These approaches lay the foundation for studying the organization of human subcortical - and enhanced cortical mapping - of eye-movement mechanisms.

Sustained Cortical and Subcortical Measures of Auditory and Visual Plasticity following Short-Term Perceptual Learning.

Short-term training can lead to improvements in behavioral discrimination of auditory and visual stimuli, as well as enhanced EEG responses to those stimuli. In the auditory domain, fluency with tonal languages and musical training has been associated with long-term cortical and subcortical plasticity, but less is known about the effects of shorter-term training. This study combined electroencephalography (EEG) and behavioral measures to investigate short-term learning and neural plasticity in both auditory and visual domains. Forty adult participants were divided into four groups. Three groups trained on one of three tasks, involving discrimination of auditory fundamental frequency (F0), auditory amplitude modulation rate (AM), or visual orientation (VIS). The fourth (control) group received no training. Pre- and post-training tests, as well as retention tests 30 days after training, involved behavioral discrimination thresholds, steady-state visually evoked potentials (SSVEP) to the flicker frequencies of visual stimuli, and auditory envelope-following responses simultaneously evoked and measured in response to rapid stimulus F0 (EFR), thought to reflect subcortical generators, and slow amplitude modulation (ASSR), thought to reflect cortical generators. Enhancement of the ASSR was observed in both auditory-trained groups, not specific to the AM-trained group, whereas enhancement of the SSVEP was found only in the visually-trained group. No evidence was found for changes in the EFR. The results suggest that some aspects of neural plasticity can develop rapidly and may generalize across tasks but not across modalities. Behaviorally, the pattern of learning was complex, with significant cross-task and cross-modal learning effects.

Neurons that detect interocular conflict during binocular rivalry revealed with EEG.

When the two eyes view incompatible images, perception alternates between them. What neural computations underlie this binocular rivalry? Perceptual alternations may simply reflect competition between the sets of monocular neurons that respond to each image, with the stronger driving perception. Here, we test an alternative hypothesis, that the computations that resolve rivalry make use of an active signal that reflects interocular conflict. Images presented to each eye were flickered at different frequencies while we measured steady-state visually evoked potentials (SSVEP). Signals at frequencies that are combinations of the two input frequencies can arise only from binocular neurons. In a first experiment, we measured energy at these "intermodulation" frequencies during binocular rivalry and found it to be highest immediately before rivalry restarted following a period of incomplete resolution of rivalry (a "mixed" percept). This suggests that the intermodulation signals may arise from neurons important for resolving the conflict between the two eyes' inputs. In a second experiment, we tested whether the intermodulation signal arose from neurons that measure interocular conflict by parametrically increasing conflict while simultaneously reducing image contrast. The activity of neurons that receive input from both eyes but are not sensitive to conflict should reduce monotonically as contrast decreases. The intermodulation response, however, peaked at intermediate levels of conflict, suggesting that it arises in part from neurons that respond to interocular conflict. Binocular rivalry appears to depend on an active mechanism that detects interocular conflict, whose levels of activity can be measured by the intermodulation frequencies of the SSVEP.

Endogenous attention signals evoked by threshold contrast detection in human superior colliculus.

Human superior colliculus (SC) responds in a retinotopically selective manner when attention is deployed on a high-contrast visual stimulus using a discrimination task. To further elucidate the role of SC in endogenous visual attention, high-resolution fMRI was used to demonstrate that SC also exhibits a retinotopically selective response for covert attention in the absence of significant visual stimulation using a threshold-contrast detection task. SC neurons have a laminar organization according to their function, with visually responsive neurons present in the superficial layers and visuomotor neurons in the intermediate layers. The results show that the response evoked by the threshold-contrast detection task is significantly deeper than the response evoked by the high-contrast speed discrimination task, reflecting a functional dissociation of the attentional enhancement of visuomotor and visual neurons, respectively. Such a functional dissociation of attention within SC laminae provides a subcortical basis for the oculomotor theory of attention.

High-resolution functional magnetic resonance imaging methods for human midbrain.

Functional MRI (fMRI) is a widely used tool for non-invasively measuring correlates of human brain activity. However, its use has mostly been focused upon measuring activity on the surface of cerebral cortex rather than in subcortical regions such as midbrain and brainstem. Subcortical fMRI must overcome two challenges: spatial resolution and physiological noise. Here we describe an optimized set of techniques developed to perform high-resolution fMRI in human SC, a structure on the dorsal surface of the midbrain; the methods can also be used to image other brainstem and subcortical structures. High-resolution (1.2 mm voxels) fMRI of the SC requires a non-conventional approach. The desired spatial sampling is obtained using a multi-shot (interleaved) spiral acquisition (1). Since, T(2)* of SC tissue is longer than in cortex, a correspondingly longer echo time (T(E) ~ 40 msec) is used to maximize functional contrast. To cover the full extent of the SC, 8-10 slices are obtained. For each session a structural anatomy with the same slice prescription as the fMRI is also obtained, which is used to align the functional data to a high-resolution reference volume. In a separate session, for each subject, we create a high-resolution (0.7 mm sampling) reference volume using a T(1)-weighted sequence that gives good tissue contrast. In the reference volume, the midbrain region is segmented using the ITK-SNAP software application (2). This segmentation is used to create a 3D surface representation of the midbrain that is both smooth and accurate (3). The surface vertices and normals are used to create a map of depth from the midbrain surface within the tissue (4). Functional data is transformed into the coordinate system of the segmented reference volume. Depth associations of the voxels enable the averaging of fMRI time series data within specified depth ranges to improve signal quality. Data is rendered on the 3D surface for visualization. In our lab we use this technique for measuring topographic maps of visual stimulation and covert and overt visual attention within the SC (1). As an example, we demonstrate the topographic representation of polar angle to visual stimulation in SC.

Surface-based analysis methods for high-resolution functional magnetic resonance imaging.

Functional magnetic resonance imaging (fMRI) has become a popular technique for studies of human brain activity. Typically, fMRI is performed with >3-mm sampling, so that the imaging data can be regarded as two-dimensional samples that average through the 1.5-4-mm thickness of cerebral cortex. The increasing use of higher spatial resolutions, <1.5-mm sampling, complicates the analysis of fMRI, as one must now consider activity variations within the depth of the brain tissue. We present a set of surface-based methods to exploit the use of high-resolution fMRI for depth analysis. These methods utilize white-matter segmentations coupled with deformable-surface algorithms to create a smooth surface representation at the gray-white interface and pial membrane. These surfaces provide vertex positions and normals for depth calculations, enabling averaging schemes that can increase contrast-to-noise ratio, as well as permitting the direct analysis of depth profiles of functional activity in the human brain.

Topography of covert visual attention in human superior colliculus.

Experiments were performed to examine the topography of covert visual attention signals in human superior colliculus (SC), both across its surface and in its depth. We measured the retinotopic organization of SC to direct visual stimulation using a 90° wedge of moving dots that slowly rotated around fixation. Subjects (n = 5) were cued to perform a difficult speed-discrimination task in the rotating region. To measure the retinotopy of covert attention, we used a full-field array of similarly moving dots. Subjects were cued to perform the same speed-discrimination task within a 90° wedge-shaped region, and only the cue rotated around fixation. High-resolution functional magnetic resonance imaging (fMRI, 1.2 mm voxels) data were acquired throughout SC. These data were then aligned to a high-resolution T1-weighted reference volume. The SC was segmented in this volume so that the surface of the SC could be computationally modeled and to permit calculation of a depth map for laminar analysis. Retinotopic maps were obtained for both direct visual stimulation and covert attention. These maps showed a similar spatial distribution to visual stimulation maps observed in rhesus macaque and were in registration with each other. Within the depth of SC, both visual attention and stimulation produced activity primarily in the superficial and intermediate layers, but stimulation activity extended significantly more deeply than attention.