Mapping the dynamics of visual feature coding: Insights into perception and integration.

The basic computations performed in the human early visual cortex are the foundation for visual perception. While we know a lot about these computations, a key missing piece is how the coding of visual features relates to our perception of the environment. To investigate visual feature coding, interactions, and their relationship to human perception, we investigated neural responses and perceptual similarity judgements to a large set of visual stimuli that varied parametrically along four feature dimensions. We measured neural responses using electroencephalography (N = 16) to 256 grating stimuli that varied in orientation, spatial frequency, contrast, and colour. We then mapped the response profiles of the neural coding of each visual feature and their interactions, and related these to independently obtained behavioural judgements of stimulus similarity. The results confirmed fundamental principles of feature coding in the visual system, such that all four features were processed simultaneously but differed in their dynamics, and there was distinctive conjunction coding for different combinations of features in the neural responses. Importantly, modelling of the behaviour revealed that every stimulus feature contributed to perceptual judgements, despite the untargeted nature of the behavioural task. Further, the relationship between neural coding and behaviour was evident from initial processing stages, signifying that the fundamental features, not just their interactions, contribute to perception. This study highlights the importance of understanding how feature coding progresses through the visual hierarchy and the relationship between different stages of processing and perception.

Properties of imagined experience across visual, auditory, and other sensory modalities.

Little is known about the perceptual characteristics of mental images nor how they vary across sensory modalities. We conducted an exhaustive survey into how mental images are experienced across modalities, mainly targeting visual and auditory imagery of a single stimulus, the letter "O", to facilitate direct comparisons. We investigated temporal properties of mental images (e.g. onset latency, duration), spatial properties (e.g. apparent location), effort (e.g. ease, spontaneity, control), movement requirements (e.g. eye movements), real-imagined interactions (e.g. inner speech while reading), beliefs about imagery norms and terminologies, as well as respondent confidence. Participants also reported on the five traditional senses and their prominence during thinking, imagining, and dreaming. Overall, visual and auditory experiences dominated mental events, although auditory mental images were superior to visual mental images on almost every metric tested except regarding spatial properties. Our findings suggest that modality-specific differences in mental imagery may parallel those of other sensory neural processes.

Modelling perception as a hierarchical competition differentiates imagined, veridical, and hallucinated percepts.

Mental imagery is a process by which thoughts become experienced with sensory characteristics. Yet, it is not clear why mental images appear diminished compared to veridical images, nor how mental images are phenomenologically distinct from hallucinations, another type of non-veridical sensory experience. Current evidence suggests that imagination and veridical perception share neural resources. If so, we argue that considering how neural representations of externally generated stimuli (i.e. sensory input) and internally generated stimuli (i.e. thoughts) might interfere with one another can sufficiently differentiate between veridical, imaginary, and hallucinatory perception. We here use a simple computational model of a serially connected, hierarchical network with bidirectional information flow to emulate the primate visual system. We show that modelling even first approximations of neural competition can more coherently explain imagery phenomenology than non-competitive models. Our simulations predict that, without competing sensory input, imagined stimuli should ubiquitously dominate hierarchical representations. However, with competition, imagination should dominate high-level representations but largely fail to outcompete sensory inputs at lower processing levels. To interpret our findings, we assume that low-level stimulus information (e.g. in early visual cortices) contributes most to the sensory aspects of perceptual experience, while high-level stimulus information (e.g. towards temporal regions) contributes most to its abstract aspects. Our findings therefore suggest that ongoing bottom-up inputs during waking life may prevent imagination from overriding veridical sensory experience. In contrast, internally generated stimuli may be hallucinated when sensory input is dampened or eradicated. Our approach can explain individual differences in imagery, along with aspects of daydreaming, hallucinations, and non-visual mental imagery.

Visual Representations: Insights from Neural Decoding.

Patterns of brain activity contain meaningful information about the perceived world. Recent decades have welcomed a new era in neural analyses, with computational techniques from machine learning applied to neural data to decode information represented in the brain. In this article, we review how decoding approaches have advanced our understanding of visual representations and discuss efforts to characterize both the complexity and the behavioral relevance of these representations. We outline the current consensus regarding the spatiotemporal structure of visual representations and review recent findings that suggest that visual representations are at once robust to perturbations, yet sensitive to different mental states. Beyond representations of the physical world, recent decoding work has shone a light on how the brain instantiates internally generated states, for example, during imagery and prediction. Going forward, decoding has remarkable potential to assess the functional relevance of visual representations for human behavior, reveal how representations change across development and during aging, and uncover their presentation in various mental disorders.

Capacity for movement is an organisational principle in object representations.

The ability to perceive moving objects is crucial for threat identification and survival. Recent neuroimaging evidence has shown that goal-directed movement is an important element of object processing in the brain. However, prior work has primarily used moving stimuli that are also animate, making it difficult to disentangle the effect of movement from aliveness or animacy in representational categorisation. In the current study, we investigated the relationship between how the brain processes movement and aliveness by including stimuli that are alive but still (e.g., plants), and stimuli that are not alive but move (e.g., waves). We examined electroencephalographic (EEG) data recorded while participants viewed static images of moving or non-moving objects that were either natural or artificial. Participants classified the images according to aliveness, or according to capacity for movement. Movement explained significant variance in the neural data over and above that of aliveness, showing that capacity for movement is an important dimension in the representation of visual objects in humans.

Are you for real? Decoding realistic AI-generated faces from neural activity.

Can we trust our eyes? Until recently, we rarely had to question whether what we see is indeed what exists, but this is changing. Artificial neural networks can now generate realistic images that challenge our perception of what is real. This new reality can have significant implications for cybersecurity, counterfeiting, fake news, and border security. We investigated how the human brain encodes and interprets realistic artificially generated images using behaviour and brain imaging. We found that we could reliably decode AI generated faces using people's neural activity. However, while at a group level people performed near chance classifying real and realistic fakes, participants tended to interchange the labels, classifying real faces as realistic fakes and vice versa. Understanding this difference between brain and behavioural responses may be key in determining the 'real' in our new reality. Stimuli, code, and data for this study can be found at https://osf.io/n2z73/.

The time-course of feature-based attention effects dissociated from temporal expectation and target-related processes.

Selective attention prioritises relevant information amongst competing sensory input. Time-resolved electrophysiological studies have shown stronger representation of attended compared to unattended stimuli, which has been interpreted as an effect of attention on information coding. However, because attention is often manipulated by making only the attended stimulus a target to be remembered and/or responded to, many reported attention effects have been confounded with target-related processes such as visual short-term memory or decision-making. In addition, attention effects could be influenced by temporal expectation about when something is likely to happen. The aim of this study was to investigate the dynamic effect of attention on visual processing using multivariate pattern analysis of electroencephalography (EEG) data, while (1) controlling for target-related confounds, and (2) directly investigating the influence of temporal expectation. Participants viewed rapid sequences of overlaid oriented grating pairs while detecting a "target" grating of a particular orientation. We manipulated attention, one grating was attended and the other ignored (cued by colour), and temporal expectation, with stimulus onset timing either predictable or not. We controlled for target-related processing confounds by only analysing non-target trials. Both attended and ignored gratings were initially coded equally in the pattern of responses across EEG sensors. An effect of attention, with preferential coding of the attended stimulus, emerged approximately 230 ms after stimulus onset. This attention effect occurred even when controlling for target-related processing confounds, and regardless of stimulus onset expectation. These results provide insight into the effect of feature-based attention on the dynamic processing of competing visual information.

Linking the Brain with Behavior: The Neural Dynamics of Success and Failure in Goal-directed Behavior.

The human brain is extremely flexible and capable of rapidly selecting relevant information in accordance with task goals. Regions of frontoparietal cortex flexibly represent relevant task information such as task rules and stimulus features when participants perform tasks successfully, but less is known about how information processing breaks down when participants make mistakes. This is important for understanding whether and when information coding recorded with neuroimaging is directly meaningful for behavior. Here, we used magnetoencephalography to assess the temporal dynamics of information processing and linked neural responses with goal-directed behavior by analyzing how they changed on behavioral error. Participants performed a difficult stimulus-response task using two stimulus-response mapping rules. We used time-resolved multivariate pattern analysis to characterize the progression of information coding from perceptual information about the stimulus, cue and rule coding, and finally, motor response. Response-aligned analyses revealed a ramping up of perceptual information before a correct response, suggestive of internal evidence accumulation. Strikingly, when participants made a stimulus-related error, and not when they made other types of errors, patterns of activity initially reflected the stimulus presented, but later reversed, and accumulated toward a representation of the "incorrect" stimulus. This suggests that the patterns recorded at later time points reflect an internally generated stimulus representation that was used to make the (incorrect) decision. These results illustrate the orderly and overlapping temporal dynamics of information coding in perceptual decision-making and show a clear link between neural patterns in the late stages of processing and behavior.

Human EEG recordings for 1,854 concepts presented in rapid serial visual presentation streams.

The neural basis of object recognition and semantic knowledge has been extensively studied but the high dimensionality of object space makes it challenging to develop overarching theories on how the brain organises object knowledge. To help understand how the brain allows us to recognise, categorise, and represent objects and object categories, there is a growing interest in using large-scale image databases for neuroimaging experiments. In the current paper, we present THINGS-EEG, a dataset containing human electroencephalography responses from 50 subjects to 1,854 object concepts and 22,248 images in the THINGS stimulus set, a manually curated and high-quality image database that was specifically designed for studying human vision. The THINGS-EEG dataset provides neuroimaging recordings to a systematic collection of objects and concepts and can therefore support a wide array of research to understand visual object processing in the human brain.

Overfitting the Literature to One Set of Stimuli and Data.

A large number of papers in Computational Cognitive Neuroscience are developing and testing novel analysis methods using one specific neuroimaging dataset and problematic experimental stimuli. Publication bias and confirmatory exploration will result in overfitting to the limited available data. We highlight the problems with this specific dataset and argue for the need to collect more good quality open neuroimaging data using a variety of experimental stimuli, in order to test the generalisability of current published results, and allow for more robust results in future work.

The Influence of Object-Color Knowledge on Emerging Object Representations in the Brain.

The ability to rapidly and accurately recognize complex objects is a crucial function of the human visual system. To recognize an object, we need to bind incoming visual features, such as color and form, together into cohesive neural representations and integrate these with our preexisting knowledge about the world. For some objects, typical color is a central feature for recognition; for example, a banana is typically yellow. Here, we applied multivariate pattern analysis on time-resolved neuroimaging (MEG) data to examine how object-color knowledge affects emerging object representations over time. Our results from 20 participants (11 female) show that the typicality of object-color combinations influences object representations, although not at the initial stages of object and color processing. We find evidence that color decoding peaks later for atypical object-color combinations compared with typical object-color combinations, illustrating the interplay between processing incoming object features and stored object knowledge. Together, these results provide new insights into the integration of incoming visual information with existing conceptual object knowledge.SIGNIFICANCE STATEMENT To recognize objects, we have to be able to bind object features, such as color and shape, into one coherent representation and compare it with stored object knowledge. The MEG data presented here provide novel insights about the integration of incoming visual information with our knowledge about the world. Using color as a model to understand the interaction between seeing and knowing, we show that there is a unique pattern of brain activity for congruently colored objects (e.g., a yellow banana) relative to incongruently colored objects (e.g., a red banana). This effect of object-color knowledge only occurs after single object features are processed, demonstrating that conceptual knowledge is accessed relatively late in the visual processing hierarchy.

Decoding Images in the Mind's Eye: The Temporal Dynamics of Visual Imagery.

Mental imagery is the ability to generate images in the mind in the absence of sensory input. Both perceptual visual processing and internally generated imagery engage large, overlapping networks of brain regions. However, it is unclear whether they are characterized by similar temporal dynamics. Recent magnetoencephalography work has shown that object category information was decodable from brain activity during mental imagery, but the timing was delayed relative to perception. The current study builds on these findings, using electroencephalography to investigate the dynamics of mental imagery. Sixteen participants viewed two images of the Sydney Harbour Bridge and two images of Santa Claus. On each trial, they viewed a sequence of the four images and were asked to imagine one of them, which was cued retroactively by its temporal location in the sequence. Time-resolved multivariate pattern analysis was used to decode the viewed and imagined stimuli. Although category and exemplar information was decodable for viewed stimuli, there were no informative patterns of activity during mental imagery. The current findings suggest stimulus complexity, task design and individual differences may influence the ability to successfully decode imagined images. We discuss the implications of these results in the context of prior findings of mental imagery.

Untangling featural and conceptual object representations.

How are visual inputs transformed into conceptual representations by the human visual system? The contents of human perception, such as objects presented on a visual display, can reliably be decoded from voxel activation patterns in fMRI, and in evoked sensor activations in MEG and EEG. A prevailing question is the extent to which brain activation associated with object categories is due to statistical regularities of visual features within object categories. Here, we assessed the contribution of mid-level features to conceptual category decoding using EEG and a novel fast periodic decoding paradigm. Our study used a stimulus set consisting of intact objects from the animate (e.g., fish) and inanimate categories (e.g., chair) and scrambled versions of the same objects that were unrecognizable and preserved their visual features (Long et al., 2018). By presenting the images at different periodic rates, we biased processing to different levels of the visual hierarchy. We found that scrambled objects and their intact counterparts elicited similar patterns of activation, which could be used to decode the conceptual category (animate or inanimate), even for the unrecognizable scrambled objects. Animacy decoding for the scrambled objects, however, was only possible at the slowest periodic presentation rate. Animacy decoding for intact objects was faster, more robust, and could be achieved at faster presentation rates. Our results confirm that the mid-level visual features preserved in the scrambled objects contribute to animacy decoding, but also demonstrate that the dynamics vary markedly for intact versus scrambled objects. Our findings suggest a complex interplay between visual feature coding and categorical representations that is mediated by the visual system's capacity to use image features to resolve a recognisable object.

The influence of image masking on object representations during rapid serial visual presentation.

Rapid image presentations combined with time-resolved multivariate analysis methods of EEG or MEG (rapid-MVPA) offer unique potential in assessing the temporal limitations of the human visual system. Recent work has shown that multiple visual objects presented sequentially can be simultaneously decoded from M/EEG recordings. Interestingly, object representations reached higher stages of processing for slower image presentation rates compared to fast rates. This fast rate attenuation is probably caused by forward and backward masking from the other images in the stream. Two factors that are likely to influence masking during rapid streams are stimulus duration and stimulus onset asynchrony (SOA). Here, we disentangle these effects by studying the emerging neural representation of visual objects using rapid-MVPA while independently manipulating stimulus duration and SOA. Our results show that longer SOAs enhance the decodability of neural representations, regardless of stimulus presentation duration, suggesting that subsequent images act as effective backward masks. In contrast, image duration does not appear to have a graded influence on object representations. Interestingly, however, decodability was improved when there was a gap between subsequent images, indicating that an abrupt onset or offset of an image enhances its representation. Our study yields insight into the dynamics of object processing in rapid streams, paving the way for future work using this promising approach.

The representational dynamics of visual objects in rapid serial visual processing streams.

In our daily lives, we are bombarded with a stream of rapidly changing visual input. Humans have the remarkable capacity to detect and identify objects in fast-changing scenes. Yet, when studying brain representations, stimuli are generally presented in isolation. Here, we studied the dynamics of human vision using a combination of fast stimulus presentation rates, electroencephalography and multivariate decoding analyses. Using a presentation rate of 5 images per second, we obtained the representational structure of a large number of stimuli, and showed the emerging abstract categorical organisation of this structure. Furthermore, we could separate the temporal dynamics of perceptual processing from higher-level target selection effects. In a second experiment, we used the same paradigm at 20Hz to show that shorter image presentation limits the categorical abstraction of object representations. Our results show that applying multivariate pattern analysis to every image in rapid serial visual processing streams has unprecedented potential for studying the temporal dynamics of the structure of representations in the human visual system.

Distinct neural processes for the perception of familiar versus unfamiliar faces along the visual hierarchy revealed by EEG.

Humans recognize faces with ease, despite the complexity of the task and of the visual system which underlies it. Different spatial regions, including both the core and extended face processing networks, and distinct temporal stages of processing have been implicated in face recognition, but there is ongoing controversy regarding the extent to which the mechanisms for recognizing a familiar face differ from those for an unfamiliar face. Here, we used electroencephalogram (EEG) and flicker SSVEP, a high signal-to-noise approach, and searchlight decoding methods to elucidate the mechanisms mediating the processing of familiar and unfamiliar faces in the time domain. Familiar and unfamiliar faces were presented periodically at 15 Hz, 6 Hz and 3.75 Hz either upright or inverted in separate blocks, with the rationale that faster frequencies require shorter processing times per image and tap into fundamentally different levels of visual processing. The 15 Hz trials, likely to reflect early visual processing, exhibited enhanced neural responses for familiar over unfamiliar face trials, but only when the faces were upright. In contrast, decoding methods revealed similar classification accuracies for upright and inverted faces for both familiar and unfamiliar faces. For the 6 Hz frequency, familiar faces had lower amplitude responses than unfamiliar faces, and decoding familiarity was more accurate for upright compared with inverted faces. Finally, the 3.75 Hz frequency revealed no main effects of familiarity, but decoding showed significant correlations with behavioral ratings of face familiarity, suggesting that activity evoked by this slow presentation frequency reflected higher-level, cognitive aspects of familiarity processing. This three-way dissociation between frequencies reveals that fundamentally different stages of the visual hierarchy are modulated by face familiarity. The combination of experimental and analytical approaches used here represent a novel method for elucidating spatio-temporal characteristics within the visual system.

More than Action: The Dorsal Pathway Contributes to the Perception of 3-D Structure.

An evolving view in cognitive neuroscience is that the dorsal visual pathway not only plays a key role in visuomotor behavior but that it also contributes functionally to the recognition of objects. To characterize the nature of the object representations derived by the dorsal pathway, we assessed perceptual performance in the context of the continuous flash suppression paradigm, which suppresses object processing in the ventral pathway while sparing computation in the dorsal pathway. In a series of experiments, prime stimuli, which were rendered imperceptible by the continuous flash suppression, still contributed to perceptual decisions related to the subsequent perceptible target stimuli. However, the contribution of the prime to perception was contingent on the prime's structural coherence, in that a perceptual advantage was observed only for targets primed by objects with legitimate 3-D structure. Finally, we obtained additional evidence to demonstrate that the processing of the suppressed objects was contingent on the magnocellular, rather than the parvocellular, system, further linking the processing of the suppressed stimuli to the dorsal pathway. Together, these results provide novel evidence that the dorsal pathway does not only support visuomotor control but, rather, that it also derives the structural description of 3-D objects and contributes to shape perception.

Differentiation of Types of Visual Agnosia Using EEG.

Visual recognition deficits are the hallmark symptom of visual agnosia, a neuropsychological disorder typically associated with damage to the visual system. Most research into visual agnosia focuses on characterizing the deficits through detailed behavioral testing, and structural and functional brain scans are used to determine the spatial extent of any cortical damage. Although the hierarchical nature of the visual system leads to clear predictions about the temporal dynamics of cortical deficits, there has been little research on the use of neuroimaging methods with high temporal resolution to characterize the temporal profile of agnosia deficits. Here, we employed high-density electroencephalography (EEG) to investigate alterations in the temporal dynamics of the visual system in two individuals with visual agnosia. In the context of a steady state visual evoked potential paradigm (SSVEP), individuals viewed pattern-reversing checkerboards of differing spatial frequency, and we assessed the responses of the visual system in the frequency and temporal domain. JW, a patient with early visual cortex damage, showed impaired SSVEP response relative to a control group and to the second patient (SM) who had right temporal lobe damage. JW also showed lower decoding accuracy for early visual responses (around 100 ms). SM, whose lesion is more anterior in the visual system, showed good decoding accuracy initially but low decoding after 500 ms. Overall, EEG and multivariate decoding methods can yield important insights into the temporal dynamics of visual responses in individuals with visual agnosia.

Very high density EEG elucidates spatiotemporal aspects of early visual processing.

Standard human EEG systems based on spatial Nyquist estimates suggest that 20-30 mm electrode spacing suffices to capture neural signals on the scalp, but recent studies posit that increasing sensor density can provide higher resolution neural information. Here, we compared "super-Nyquist" density EEG ("SND") with Nyquist density ("ND") arrays for assessing the spatiotemporal aspects of early visual processing. EEG was measured from 128 electrodes arranged over occipitotemporal brain regions (14 mm spacing) while participants viewed flickering checkerboard stimuli. Analyses compared SND with ND-equivalent subsets of the same electrodes. Frequency-tagged stimuli were classified more accurately with SND than ND arrays in both the time and the frequency domains. Representational similarity analysis revealed that a computational model of V1 correlated more highly with the SND than the ND array. Overall, SND EEG captured more neural information from visual cortex, arguing for increased development of this approach in basic and translational neuroscience.

Word and face processing engage overlapping distributed networks: Evidence from RSVP and EEG investigations.

Words and faces have vastly different visual properties, but increasing evidence suggests that word and face processing engage overlapping distributed networks. For instance, fMRI studies have shown overlapping activity for face and word processing in the fusiform gyrus despite well-characterized lateralization of these objects to the left and right hemispheres, respectively. To investigate whether face and word perception influences perception of the other stimulus class and elucidate the mechanisms underlying such interactions, we presented images using rapid serial visual presentations. Across 3 experiments, participants discriminated 2 face, word, and glasses targets (T1 and T2) embedded in a stream of images. As expected, T2 discrimination was impaired when it followed T1 by 200 to 300 ms relative to longer intertarget lags, the so-called attentional blink. Interestingly, T2 discrimination accuracy was significantly reduced at short intertarget lags when a face was followed by a word (face-word) compared with glasses-word and word-word combinations, indicating that face processing interfered with word perception. The reverse effect was not observed; that is, word-face performance was no different than the other object combinations. EEG results indicated the left N170 to T1 was correlated with the word decrement for face-word trials, but not for other object combinations. Taken together, the results suggest face processing interferes with word processing, providing evidence for overlapping neural mechanisms of these 2 object types. Furthermore, asymmetrical face-word interference points to greater overlap of face and word representations in the left than the right hemisphere. (PsycINFO Database Record