Event Probabilities Have a Different Impact on Early and Late Electroencephalographic Measures Regarded as Metrics of Prediction.

The oddball protocol has been used to study the neural and perceptual consequences of implicit predictions in the human brain. The protocol involves presenting a sequence of identical repeated events that are eventually broken by a novel "oddball" presentation. Oddball presentations have been linked to increased neural responding and to an exaggeration of perceived duration relative to repeated events. Because the number of repeated events in such protocols is circumscribed, as more repeats are encountered, the conditional probability of a further repeat decreases-whereas the conditional probability of an oddball increases. These facts have not been appreciated in many analyses of oddballs; repeats and oddballs have rather been treated as binary event categories. Here, we show that the human brain is sensitive to conditional event probabilities in an active, visual oddball paradigm. P300 responses (a relatively late component of visually evoked potentials measured with EEG) tended to be greater for less likely oddballs and repeats. By contrast, P1 responses (an earlier component) increased for repeats as a goal-relevant target presentation neared, but this effect occurred even when repeat probabilities were held constant, and oddball P1 responses were invariant. We also found that later, more likely oddballs seemed to last longer, and this effect was largely independent of the number of preceding repeats. These findings speak against a repetition suppression account of the temporal oddball effect. Overall, our data highlight an impact of event probability on later, rather than earlier, electroencephalographic measures previously related to predictive processes-and the importance of considering conditional probabilities in sequential presentation paradigms.

Enhanced electrophysiological responses to explicitly predicted and pre-imagined inputs, with confirmation from online decoding with neuro-feedback.

Neural responses to sensory inputs can scale with the likelihood of encountering the input. This is consistent with the predictive coding framework, in that the human brain is expected to be less responsive to predicted inputs. Typically, however, prediction is not explicitly measured. It is inferred from the probability of encountering an event. When an input is explicitly predicted, responses to predicted inputs can be enhanced. Here, we ask if this effect can be ascribed to a generic priming effect, from pre-cogitating about one of two possible inputs. Consistent with this, we find that P300s (a relatively late event-related potential measured with electroencephalography) are greater for explicitly predicted audio and visual inputs, and that this effect cannot be distinguished from a priming effect from pre-imagining audio or visual presentations. Evidence indicates that participants engaged in pre-imagining presentations, as we were able to decode online what type of presentation (audio or visual) they were imagining with a high success rate (approx. 73%), and we encouraged compliance with neuro-feedback regarding this success rate. Our data confirm that human cortex can be more responsive to inputs that have been subject to pre-cogitation-including explicit predictions. This highlights that while anticipatory processes can reduce responding to likely inputs, they can also enhance responding to explicitly predicted inputs.

Commonalities between the Berger Rhythm and spectra differences driven by cross-modal attention and imagination.

When people close their eyes, the power of alpha-band oscillatory brain activity increases. We explored the possibility that this could be related to a suppression of visual processing, rather than being a default dynamic of the visual brain. We recorded brain activity while people meditated with their eyes open or closed, and when people attended to or imagined having auditory or visual experiences. We could decode the attended or imagined modality of experiences based on the spectra of brain activity that prevailed while meditating with open or closed eyes. We also found anecdotal evidence suggesting the strength of imagined sensory experiences may be predicted by the dynamics of neural networks that are responsive to inputs. Overall, our data suggest spectra changes when people close their eyes might relate to a targeted suppression of visual processing, as opposed to being a default state of idle visual brains.

On why we lack confidence in some signal-detection-based analyses of confidence.

Signal-detection theory (SDT) is one of the most popular frameworks for analyzing data from studies of human behavior - including investigations of confidence. SDT-based analyses of confidence deliver both standard estimates of sensitivity (d'), and a second estimate informed by high-confidence decisions - meta d'. The extent to which meta d' estimates fall short of d' estimates is regarded as a measure of metacognitive inefficiency, quantifying the contamination of confidence by additional noise. These analyses rely on a key but questionable assumption - that repeated exposures to an input will evoke a normally-shaped distribution of perceptual experiences (the normality assumption). Here we show, via analyses inspired by an experiment and modelling, that when distributions of experience do not conform with the normality assumption, meta d' can be systematically underestimated relative to d'. Our data highlight that SDT-based analyses of confidence do not provide a ground truth measure of human metacognitive inefficiency. We explain why deviance from the normality assumption is especially a problem for some popular SDT-based analyses of confidence, in contrast to other analyses inspired by the SDT framework, which are more robust to violations of the normality assumption.

Predictive extrapolation effects can have a greater impact on visual decisions, while visual adaptation has a greater impact on conscious visual experience.

Human vision is shaped by historic and by predictive processes. The lingering impact of visual adaptation, for instance, can act to exaggerate differences between past and present inputs, whereas predictive processes can promote extrapolation effects that allow us to anticipate the near future. It is unclear to what extent either of these effects manifest in changes to conscious visual experience. It is also unclear how these influences combine, when acting in concert or opposition. We had people make decisions about the sizes of inputs, and report on levels of decisional confidence. Tests were either selectively subject to size adaptation, to an extrapolation effect, or to both of these effects. When these two effects were placed in opposition, extrapolation had a greater impact on decision making. However, our data suggest the influence of extrapolation is primarily decisional, whereas size adaptation more fully manifests in changes to conscious visual awareness.

An observer model of tilt perception, sensitivity and confidence.

Humans experience levels of confidence in perceptual decisions that tend to scale with the precision of their judgements; but not always. Sometimes precision can be held constant while confidence changes-leading researchers to assume precision and confidence are shaped by different types of information (e.g. perceptual and decisional). To assess this, we examined how visual adaptation to oriented inputs changes tilt perception, perceptual sensitivity and confidence. Some adaptors had a greater detrimental impact on measures of confidence than on precision. We could account for this using an observer model, where precision and confidence rely on different magnitudes of sensory information. These data show that differences in perceptual sensitivity and confidence can therefore emerge, not because these factors rely on different types of information, but because they rely on different magnitudes of sensory information.

Sharpening vision by adapting to flicker.

Human vision is surprisingly malleable. A static stimulus can seem to move after prolonged exposure to movement (the motion aftereffect), and exposure to tilted lines can make vertical lines seem oppositely tilted (the tilt aftereffect). The paradigm used to induce such distortions (adaptation) can provide powerful insights into the computations underlying human visual experience. Previously spatial form and stimulus dynamics were thought to be encoded independently, but here we show that adaptation to stimulus dynamics can sharpen form perception. We find that fast flicker adaptation (FFAd) shifts the tuning of face perception to higher spatial frequencies, enhances the acuity of spatial vision-allowing people to localize inputs with greater precision and to read finer scaled text, and it selectively reduces sensitivity to coarse-scale form signals. These findings are consistent with two interrelated influences: FFAd reduces the responsiveness of magnocellular neurons (which are important for encoding dynamics, but can have poor spatial resolution), and magnocellular responses contribute coarse spatial scale information when the visual system synthesizes form signals. Consequently, when magnocellular responses are mitigated via FFAd, human form perception is transiently sharpened because "blur" signals are mitigated.

Bidirectional Gender Face Aftereffects: Evidence Against Normative Facial Coding.

Facial appearance can be altered, not just by restyling but also by sensory processes. Exposure to a female face can, for instance, make subsequent faces look more masculine than they would otherwise. Two explanations exist. According to one, exposure to a female face renormalizes face perception, making that female and all other faces look more masculine as a consequence-a unidirectional effect. According to that explanation, exposure to a male face would have the opposite unidirectional effect. Another suggestion is that face gender is subject to contrastive aftereffects. These should make some faces look more masculine than the adaptor and other faces more feminine-a bidirectional effect. Here, we show that face gender aftereffects are bidirectional, as predicted by the latter hypothesis. Images of real faces rated as more and less masculine than adaptors at baseline tended to look even more and less masculine than adaptors post adaptation. This suggests that, rather than mental representations of all faces being recalibrated to better reflect the prevailing statistics of the environment, mental operations exaggerate differences between successive faces, and this can impact facial gender perception.

What is learned when learning to point at "invisible" targets?

Binocular masking is a particularly interesting means of suppressing human visual awareness, as images rendered subjectively "invisible" via binocular masking nonetheless excite robust activity in human visual cortex. Recently, binocular masking has been leveraged to show that people can be trained to better interact with inputs that, subjectively, remain invisible. Here we ask what is learned in such circumstances. Do people become more adept at using weak encoded signals to guide hand movements, or is signal encoding enhanced, resulting in heightened objective sensitivity? To assess these possibilities, we had people train on five consecutive days, to reach toward and point at a target presented in one of three masked locations. Target intensity was set to a fraction of a detection threshold determined pretraining for each participant. We found that people became better at selecting the target location with training, even when insisting they could not see the target. More important, posttraining we found objective thresholds had improved by an amount that was commensurate with an improvement in subjective visibility. Our data therefore show that training to coordinate with subjectively invisible targets can result in enhanced encodings of binocularly masked images.

A model-based comparison of three theories of audiovisual temporal recalibration.

Observers change their audio-visual timing judgements after exposure to asynchronous audiovisual signals. The mechanism underlying this temporal recalibration is currently debated. Three broad explanations have been suggested. According to the first, the time it takes for sensory signals to propagate through the brain has changed. The second explanation suggests that decisional criteria used to interpret signal timing have changed, but not time perception itself. A final possibility is that a population of neurones collectively encode relative times, and that exposure to a repeated timing relationship alters the balance of responses in this population. Here, we simplified each of these explanations to its core features in order to produce three corresponding six-parameter models, which generate contrasting patterns of predictions about how simultaneity judgements should vary across four adaptation conditions: No adaptation, synchronous adaptation, and auditory leading/lagging adaptation. We tested model predictions by fitting data from all four conditions simultaneously, in order to assess which model/explanation best described the complete pattern of results. The latency-shift and criterion-change models were better able to explain results for our sample as a whole. The population-code model did, however, account for improved performance following adaptation to a synchronous adapter, and best described the results of a subset of observers who reported least instances of synchrony.

Face aftereffects involve local repulsion, not renormalization.

After looking at a photograph of someone for a protracted period (adaptation), a previously neutral-looking face can take on an opposite appearance in terms of gender, identity, and other attributes-but what happens to the appearance of other faces? Face aftereffects have repeatedly been ascribed to perceptual renormalization. Renormalization predicts that the adapting face and more extreme versions of it should appear more neutral after adaptation (e.g., if the adaptor was male, it and hyper-masculine faces should look more feminine). Other aftereffects, such as tilt and spatial frequency, are locally repulsive, exaggerating differences between adapting and test stimuli. This predicts that the adapting face should be little changed in appearance after adaptation, while more extreme versions of it should look even more extreme (e.g., if the adaptor was male, it should look unchanged, while hyper-masculine faces should look even more masculine). Existing reports do not provide clear evidence for either pattern. We overcame this by using a spatial comparison task to measure the appearance of stimuli presented in differently adapted retinal locations. In behaviorally matched experiments we compared aftereffect patterns after adapting to tilt, facial identity, and facial gender. In all three experiments data matched the predictions of a locally repulsive, but not a renormalizing, aftereffect. These data are consistent with the existence of similar encoding strategies for tilt, facial identity, and facial gender.

Evidence for tilt normalization can be explained by anisotropic orientation sensitivity.

Some data have been taken as evidence that after prolonged viewing, near-vertical orientations "normalize" to appear more vertical than they did previously. After almost a century of research, the existence of tilt normalization remains controversial. The most recent evidence for tilt normalization comes from data suggesting a measurable "perceptual drift" of near-vertical adaptors toward vertical, which can be nulled by a slight physical rotation away from vertical (Müller, Schillinger, Do, & Leopold, 2009). We argue that biases in estimates of perceptual stasis could, however, result from the anisotropic organization of orientation-selective neurons in V1, with vertically-selective cells being more narrowly tuned than obliquely-selective cells. We describe a neurophysiologically plausible model that predicts greater sensitivity to orientation displacements toward than away from vertical. We demonstrate the predicted asymmetric pattern of sensitivity in human observers by determining threshold speeds for detecting rotation direction (Experiment 1), and by determining orientation discrimination thresholds for brief static stimuli (Experiment 2). Results imply that data suggesting a perceptual drift toward vertical instead result from greater discrimination sensitivity around cardinal than oblique orientations (the oblique effect), and thus do not constitute evidence for tilt normalization.

An object-centered aftereffect of a latent material property: A squishiness visual aftereffect, not causality adaptation.

Visual aftereffects are characterized by a changed perceptual experience after exposure to a visual input. For instance, exposure to rightward motion can make a static input seem to drift leftward-the motion aftereffect. Such aftereffects have been integral to building our understanding of the neural mechanisms and computational processes that underlie perception. Increasingly complex characteristics have been found to be susceptible to visual aftereffects, such as the appearance of human faces, the apparent number of visual elements, and the glossiness of a surface. Here we report that the apparent elasticity, or squishiness, of an object is also subject to a visual aftereffect. This relationship can explain data previously interpreted in terms of a causality aftereffect.

Why the long face? The importance of vertical image structure for biological "barcodes" underlying face recognition.

Humans are experts at face recognition. The mechanisms underlying this complex capacity are not fully understood. Recently, it has been proposed that face recognition is supported by a coarse-scale analysis of visual information contained in horizontal bands of contrast distributed along the vertical image axis-a biological facial "barcode" (Dakin & Watt, 2009). A critical prediction of the facial barcode hypothesis is that the distribution of image contrast along the vertical axis will be more important for face recognition than image distributions along the horizontal axis. Using a novel paradigm involving dynamic image distortions, a series of experiments are presented examining famous face recognition impairments from selectively disrupting image distributions along the vertical or horizontal image axes. Results show that disrupting the image distribution along the vertical image axis is more disruptive for recognition than matched distortions along the horizontal axis. Consistent with the facial barcode hypothesis, these results suggest that human face recognition relies disproportionately on appropriately scaled distributions of image contrast along the vertical image axis.

Deep Aphantasia: a visual brain with minimal influence from priors or inhibitory feedback?

The authors are both self-described congenital aphantasics, who feel they have never been able to have volitional imagined visual experiences during their waking lives. In addition, Loren has atypical experiences of a number of visual phenomena that involve an extrapolation or integration of visual information across space. In this perspective, we describe Loren's atypical experiences of a number of visual phenomena, and we suggest these ensue because her visual experiences are not strongly shaped by inhibitory feedback or by prior expectations. We describe Loren as having Deep Aphantasia, and Derek as shallow, as for both a paucity of feedback might prevent the generation of imagined visual experiences, but for Loren this additionally seems to disrupt activity at a sufficiently early locus to cause atypical experiences of actual visual inputs. Our purpose in describing these subjective experiences is to alert others to the possibility of there being sub-classes of congenital aphantasia, one of which-Deep Aphantasia, would be characterized by atypical experiences of actual visual inputs.

Predicting the subjective intensity of imagined experiences from electrophysiological measures of oscillatory brain activity.

Most people can conjure images and sounds that they experience in their minds. There are, however, marked individual differences. Some people report that they cannot generate imagined sensory experiences at all (aphantasics) and others report that they have unusually intense imagined experiences (hyper-phantasics). These individual differences have been linked to activity in sensory brain regions, driven by feedback. We would therefore expect imagined experiences to be associated with specific frequencies of oscillatory brain activity, as these can be a hallmark of neural interactions within and across regions of the brain. Replicating a number of other studies, relative to a Resting-State we find that the act of engaging in auditory or in visual imagery is linked to reductions in the power of oscillatory brain activity across a broad range of frequencies, with prominent peaks in the alpha band (8-12 Hz). This oscillatory activity, however, did not predict individual differences in the subjective intensity of imagined experiences. For audio imagery, these were rather predicted by reductions within the theta (6-9 Hz) and gamma (33-38 Hz) bands, and by increases in beta (15-17 Hz) band activity. For visual imagery these were predicted by reductions in lower (14-16 Hz) and upper (29-32 Hz) beta band activity, and by an increase in mid-beta band (24-26 Hz) activity. Our data suggest that there is sufficient ground truth in the subjective reports people use to describe the intensity of their imagined sensory experiences to allow these to be linked to the power of distinct rhythms of brain activity. In future, we hope to combine this approach with better measures of the subjective intensity of imagined sensory experiences to provide a clearer picture of individual differences in the subjective intensity of imagined experiences, and of why these eventuate.

Attentional-tracking acuity is modulated by illusory changes in perceived speed.

Many activities, such as driving or playing sports, require simultaneous monitoring of multiple, often moving, objects. Such situations tap people's ability to attend selected objects without tracking them with their eyes--this is known as attentional tracking. It has been established that attentional tracking can be affected by the physical speed of a moving target. In the experiments reported here, we showed that this effect is primarily due to apparent speeds, as opposed to physical speeds. We used sensory adaptation--in this case, prolonged exposure to adapting stimuli moving faster or slower than standard test stimuli--to modulate perceived speed. We found performance decrements and increments for apparently sped and slowed test stimuli when participants attempted attentional tracking. Our data suggest that both perceived speed and the acuity of attention for moving objects reflect a ratio of responses in low-pass and band-pass temporal-frequency channels in human vision.

The temporal visual oddball effect is not caused by repetition suppression.

The oddball paradigm is commonly used to investigate human time perception. Trains of identical repeated events ('standards') are presented, only to be interrupted by a different 'oddball' that seems to have a relatively protracted duration. One theoretical account has been that this effect is driven by repetition suppression for repeated standards. The idea is that repeated events seem shorter as they incur a progressively reduced neural response, which is supported by the finding that oddball perceived duration increases linearly with the number of preceding repeated standards. However, typical oddball paradigms confound the probability of oddball presentations with variable numbers of standard repetitions on each trial, allowing people to increasingly anticipate an oddball presentation as more standards are presented. We eliminated this by making participants aware of what fixed number of standards they would encounter before a final test input and tested different numbers of standards in separate experimental sessions. The final event of sequences, the test event, was equally likely to be an oddball or another repeat. We found a positive linear relationship between the number of preceding repeated standards and the perceived duration of oddball test events. However, we also found this for repeat tests events, which speaks against the repetition suppression account of the temporal oddball effect.

The best fitting of three contemporary observer models reveals how participants' strategy influences the window of subjective synchrony.

When experimenters vary the timing between two intersensory events, and participants judge their simultaneity, an inverse-U-shaped psychometric function is obtained. Typically, this simultaneity function is first fitted with a model for each participant separately, before best-fitting parameters are utilized (e.g., compared across conditions) in the second stage of a two-step inferential procedure. Often, simultaneity-function width is interpreted as representing sensitivity to asynchrony, and/or ascribed theoretical equivalence to a window of multisensory temporal binding. Here, we instead fit a single (principled) multilevel model to data from the entire group and across several conditions at once. By asking 20 participants to sometimes be more conservative in their judgments, we demonstrate how the width of the simultaneity function is prone to strategic change and thus questionable as a measure of either sensitivity to asynchrony or multisensory binding. By repeating our analysis with three different models (two implying a decision based directly on subjective asynchrony, and a third deriving this decision from the correlation between filtered responses to sensory inputs) we find that the first model, which hypothesizes, in particular, Gaussian latency noise and difficulty maintaining the stability of decision criteria across trials, is most plausible for these data. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

Neural correlates of visual acuity for fine text.

Human sensitivity to visual input often scales with the magnitude of evoked responses in the brain. Here, we demonstrate an exception. We record electroencephalography (EEG) while people attempt to resolve fine print - similar to people attempting to read eye charts (the world's most popular means of testing vision). We find that the ability to resolve fine print is associated with smaller evoked responses recorded by large clusters of occipital-parietal sensors ∼150 ms after people see words. Moreover, we find that a better ability to resolve fine print is associated with enhanced alpha-band oscillatory brain activity immediately prior to word presentations. These investigations were inspired by psychophysical data, which suggested the ability to resolve fine print can be enhanced by pre-adaptation to flicker, which should encourage a reduced neural response to inputs. We included this manipulation in this study, and our results are broadly consistent with this conjecture. As alpha-band activity has been linked to inhibitory interactions in visual cortex, we regard our data as evidence that smaller neural responses to fine print can be promoted by inhibitory processes that target unhelpful blur-related signals, which thereby sharpen subsequent visual experiences.