Lexical Processing Strongly Affects Reading Times But Not Skipping During Natural Reading.

In a typical text, readers look much longer at some words than at others, even skipping many altogether. Historically, researchers explained this variation via low-level visual or oculomotor factors, but today it is primarily explained via factors determining a word's lexical processing ease, such as how well word identity can be predicted from context or discerned from parafoveal preview. While the existence of these effects is well established in controlled experiments, the relative importance of prediction, preview and low-level factors in natural reading remains unclear. Here, we address this question in three large naturalistic reading corpora (n = 104, 1.5 million words), using deep neural networks and Bayesian ideal observers to model linguistic prediction and parafoveal preview from moment to moment in natural reading. Strikingly, neither prediction nor preview was important for explaining word skipping-the vast majority of explained variation was explained by a simple oculomotor model, using just fixation position and word length. For reading times, by contrast, we found strong but independent contributions of prediction and preview, with effect sizes matching those from controlled experiments. Together, these results challenge dominant models of eye movements in reading, and instead support alternative models that describe skipping (but not reading times) as largely autonomous from word identification, and mostly determined by low-level oculomotor information.

Cortical activity during naturalistic music listening reflects short-range predictions based on long-term experience.

Expectations shape our experience of music. However, the internal model upon which listeners form melodic expectations is still debated. Do expectations stem from Gestalt-like principles or statistical learning? If the latter, does long-term experience play an important role, or are short-term regularities sufficient? And finally, what length of context informs contextual expectations? To answer these questions, we presented human listeners with diverse naturalistic compositions from Western classical music, while recording neural activity using MEG. We quantified note-level melodic surprise and uncertainty using various computational models of music, including a state-of-the-art transformer neural network. A time-resolved regression analysis revealed that neural activity over fronto-temporal sensors tracked melodic surprise particularly around 200ms and 300-500ms after note onset. This neural surprise response was dissociated from sensory-acoustic and adaptation effects. Neural surprise was best predicted by computational models that incorporated long-term statistical learning-rather than by simple, Gestalt-like principles. Yet, intriguingly, the surprise reflected primarily short-range musical contexts of less than ten notes. We present a full replication of our novel MEG results in an openly available EEG dataset. Together, these results elucidate the internal model that shapes melodic predictions during naturalistic music listening.

Reconstructing the predictive architecture of the mind and brain.

Predictive processing has become an influential framework in cognitive neuroscience. However, it often lacks specificity and direct empirical support. How can we probe the nature and limits of the predictive brain? We highlight the potential of recent advances in artificial intelligence (AI) for providing a richer and more computationally explicit test of this theory of cortical function.

A hierarchy of linguistic predictions during natural language comprehension.

Understanding spoken language requires transforming ambiguous acoustic streams into a hierarchy of representations, from phonemes to meaning. It has been suggested that the brain uses prediction to guide the interpretation of incoming input. However, the role of prediction in language processing remains disputed, with disagreement about both the ubiquity and representational nature of predictions. Here, we address both issues by analyzing brain recordings of participants listening to audiobooks, and using a deep neural network (GPT-2) to precisely quantify contextual predictions. First, we establish that brain responses to words are modulated by ubiquitous predictions. Next, we disentangle model-based predictions into distinct dimensions, revealing dissociable neural signatures of predictions about syntactic category (parts of speech), phonemes, and semantics. Finally, we show that high-level (word) predictions inform low-level (phoneme) predictions, supporting hierarchical predictive processing. Together, these results underscore the ubiquity of prediction in language processing, showing that the brain spontaneously predicts upcoming language at multiple levels of abstraction.

Dampened sensory representations for expected input across the ventral visual stream.

Expectations, derived from previous experience, can help in making perception faster, more reliable and informative. A key neural signature of perceptual expectations is expectation suppression, an attenuated neural response to expected compared with unexpected stimuli. While expectation suppression has been reported using a variety of paradigms and recording methods, it remains unclear what neural modulation underlies this response attenuation. Sharpening models propose that neural populations tuned away from an expected stimulus are particularly suppressed by expectations, thereby resulting in an attenuated, but sharper population response. In contrast, dampening models suggest that neural populations tuned toward the expected stimulus are most suppressed, thus resulting in a dampened, less redundant population response. Empirical support is divided, with some studies favoring sharpening, while others support dampening. A key limitation of previous neuroimaging studies is the ability to draw inferences about neural-level modulations based on population (e.g. voxel) level signals. Indeed, recent simulations of repetition suppression showed that opposite neural modulations can lead to comparable population-level modulations. Forward models provide one solution to this inference limitation. Here, we used forward models to implement sharpening and dampening models, mapping neural modulations to voxel-level data. We show that a feature-specific gain modulation, suppressing neurons tuned toward the expected stimulus, best explains the empirical fMRI data. Thus, our results support the dampening account of expectation suppression, suggesting that expectations reduce redundancy in sensory cortex, and thereby promote updating of internal models on the basis of surprising information.

Word contexts enhance the neural representation of individual letters in early visual cortex.

Visual context facilitates perception, but how this is neurally implemented remains unclear. One example of contextual facilitation is found in reading, where letters are more easily identified when embedded in a word. Bottom-up models explain this word advantage as a post-perceptual decision bias, while top-down models propose that word contexts enhance perception itself. Here, we arbitrate between these accounts by presenting words and nonwords and probing the representational fidelity of individual letters using functional magnetic resonance imaging. In line with top-down models, we find that word contexts enhance letter representations in early visual cortex. Moreover, we observe increased coupling between letter information in visual cortex and brain activity in key areas of the reading network, suggesting these areas may be the source of the enhancement. Our results provide evidence for top-down representational enhancement in word recognition, demonstrating that word contexts can modulate perceptual processing already at the earliest visual regions.

Confidence resets reveal hierarchical adaptive learning in humans.

Hierarchical processing is pervasive in the brain, but its computational significance for learning under uncertainty is disputed. On the one hand, hierarchical models provide an optimal framework and are becoming increasingly popular to study cognition. On the other hand, non-hierarchical (flat) models remain influential and can learn efficiently, even in uncertain and changing environments. Here, we show that previously proposed hallmarks of hierarchical learning, which relied on reports of learned quantities or choices in simple experiments, are insufficient to categorically distinguish hierarchical from flat models. Instead, we present a novel test which leverages a more complex task, whose hierarchical structure allows generalization between different statistics tracked in parallel. We use reports of confidence to quantitatively and qualitatively arbitrate between the two accounts of learning. Our results support the hierarchical learning framework, and demonstrate how confidence can be a useful metric in learning theory.

How Do Expectations Shape Perception?

Perception and perceptual decision-making are strongly facilitated by prior knowledge about the probabilistic structure of the world. While the computational benefits of using prior expectation in perception are clear, there are myriad ways in which this computation can be realized. We review here recent advances in our understanding of the neural sources and targets of expectations in perception. Furthermore, we discuss Bayesian theories of perception that prescribe how an agent should integrate prior knowledge and sensory information, and investigate how current and future empirical data can inform and constrain computational frameworks that implement such probabilistic integration in perception.

Attention-based perceptual learning does not affect access to awareness.

Visual information that is relevant for an observer gains prioritized access to awareness (Gayet, Van der Stigchel, & Paffen, 2014). Here we investigate whether information that was relevant for an extended duration is prioritized for access to awareness when it is no longer relevant. We applied a perceptual-learning paradigm, in which observers were trained for 3 days on a speed-discrimination task. This task used a stimulus consisting of two motion directions, of which one was relevant to the task and one irrelevant. Before and after training, we applied a motion-coherence task to validate whether perceptual learning took place, and a breaking continuous flash-suppression (b-CFS) paradigm to assess how training affected access to awareness. The results reveal that motion-coherence thresholds for the task-relevant motion direction selectively decreased after compared to before training, revealing that task-relevant perceptual learning took place. The results of the b-CFS task, however, reveal that access to awareness was not affected by task-relevant learning: Instead, detection times for motion undergoing CFS decreased, irrespective of its direction, after compared to before training. A follow-up experiment showed that the time to detect visual motion also decreased after 3 days without training, revealing that perceptual learning did not cause the general decrease in detection times. The latter is in line with results by Mastropasqua, Tse, and Turatto (2015) and has important consequences for studies applying b-CFS to assess access to awareness: Studies that intend to apply measurements involving b-CFS on different testing days should consider that breakthrough times will dramatically decrease from pre- to postmeasurement.

Great Expectations: Is there Evidence for Predictive Coding in Auditory Cortex?

Predictive coding is possibly one of the most influential, comprehensive, and controversial theories of neural function. While proponents praise its explanatory potential, critics object that key tenets of the theory are untested or even untestable. The present article critically examines existing evidence for predictive coding in the auditory modality. Specifically, we identify five key assumptions of the theory and evaluate each in the light of animal, human and modeling studies of auditory pattern processing. For the first two assumptions - that neural responses are shaped by expectations and that these expectations are hierarchically organized - animal and human studies provide compelling evidence. The anticipatory, predictive nature of these expectations also enjoys empirical support, especially from studies on unexpected stimulus omission. However, for the existence of separate error and prediction neurons, a key assumption of the theory, evidence is lacking. More work exists on the proposed oscillatory signatures of predictive coding, and on the relation between attention and precision. However, results on these latter two assumptions are mixed or contradictory. Looking to the future, more collaboration between human and animal studies, aided by model-based analyses will be needed to test specific assumptions and implementations of predictive coding - and, as such, help determine whether this popular grand theory can fulfill its expectations.

Visual input that matches the content of visual working memory requires less (not faster) evidence sampling to reach conscious access.

The content of visual working memory (VWM) affects the processing of concurrent visual input. Recently, it has been demonstrated that stimuli are released from interocular suppression faster when they match rather than mismatch a color that is memorized for subsequent recall. In order to investigate the nature of the interaction between visual representations elicited by VWM and visual representations elicited by retinal input, we modeled the perceptual processes leading up to this difference in suppression durations. We replicated the VWM modulation of suppression durations, and fitted sequential sampling models (linear ballistic accumulators) to the response time data. Model comparisons revealed that the data was best explained by a decrease in threshold for visual input that matches the content of VWM. Converging evidence was obtained by fitting similar sequential sampling models (shifted Wald model) to published datasets. Finally, to confirm that the previously observed threshold difference reflected processes occurring before rather than after the stimuli were released from suppression, we applied the same procedure to the data of an experiment in which stimuli were not interocularly suppressed. Here, we found no decrease in threshold for stimuli that match the content of VWM. We discuss our findings in light of a preactivation hypothesis, proposing that matching visual input taps into the same neural substrate that is already activated by a representation concurrently maintained in VWM, thereby reducing its threshold for reaching visual awareness.