Vicarious touch: Overlapping neural patterns between seeing and feeling touch.

Simulation theories propose that vicarious touch arises when seeing someone else being touched triggers corresponding representations of being touched. Prior electroencephalography (EEG) findings show that seeing touch modulates both early and late somatosensory responses (measured with or without direct tactile stimulation). Functional Magnetic Resonance Imaging (fMRI) studies have shown that seeing touch increases somatosensory cortical activation. These findings have been taken to suggest that when we see someone being touched, we simulate that touch in our sensory systems. The somatosensory overlap when seeing and feeling touch differs between individuals, potentially underpinning variation in vicarious touch experiences. Increases in amplitude (EEG) or cerebral blood flow response (fMRI), however, are limited in that they cannot test for the information contained in the neural signal: seeing touch may not activate the same information as feeling touch. Here, we use time-resolved multivariate pattern analysis on whole-brain EEG data from people with and without vicarious touch experiences to test whether seen touch evokes overlapping neural representations with the first-hand experience of touch. Participants felt touch to the fingers (tactile trials) or watched carefully matched videos of touch to another person's fingers (visual trials). In both groups, EEG was sufficiently sensitive to allow decoding of touch location (little finger vs. thumb) on tactile trials. However, only in individuals who reported feeling touch when watching videos of touch could a classifier trained on tactile trials distinguish touch location on visual trials. This demonstrates that, for people who experience vicarious touch, there is overlap in the information about touch location held in the neural patterns when seeing and feeling touch. The timecourse of this overlap implies that seeing touch evokes similar representations to later stages of tactile processing. Therefore, while simulation may underlie vicarious tactile sensations, our findings suggest this involves an abstracted representation of directly felt touch.

The nature of neural object representations during dynamic occlusion.

Objects disappearing briefly from sight due to occlusion is an inevitable occurrence in everyday life. Yet we generally have a strong experience that occluded objects continue to exist, despite the fact that they objectively disappear. This indicates that neural object representations must be maintained during dynamic occlusion. However, it is unclear what the nature of such representation is and in particular whether it is perception-like or more abstract, for example, reflecting limited features such as position or movement direction only. In this study, we address this question by examining how different object features such as object shape, luminance, and position are represented in the brain when a moving object is dynamically occluded. We apply multivariate decoding methods to Magnetoencephalography (MEG) data to track how object representations unfold over time. Our methods allow us to contrast the representations of multiple object features during occlusion and enable us to compare the neural responses evoked by visible and occluded objects. The results show that object position information is represented during occlusion to a limited extent while object identity features are not maintained through the period of occlusion. Together, this suggests that the nature of object representations during dynamic occlusion is different from visual representations during perception.

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.

Reward Rapidly Enhances Visual Perception.

Rewards exert a deep influence on our cognition and behavior. Here, we used a paradigm in which reward information was provided at either encoding or retrieval of a brief, masked stimulus to show that reward can also rapidly modulate perceptual encoding of visual information. Experiment 1 (n = 30 adults) showed that participants' response accuracy was enhanced when a to-be-encoded grating signaled high reward relative to low reward, but only when the grating was presented very briefly and participants reported that they were not consciously aware of it. Experiment 2 (n = 29 adults) showed that there was no difference in participants' response accuracy when reward information was instead provided at the stage of retrieval, ruling out an explanation of the reward-modulation effect in terms of differences in motivated retrieval. Taken together, our findings provide behavioral evidence consistent with a rapid reward modulation of visual perception, which may not require consciousness.

Characteristics of expert search behavior in volumetric medical image interpretation.

Purpose: Experienced radiologists have enhanced global processing ability relative to novices, allowing experts to rapidly detect medical abnormalities without performing an exhaustive search. However, evidence for global processing models is primarily limited to two-dimensional image interpretation, and it is unclear whether these findings generalize to volumetric images, which are widely used in clinical practice. We examined whether radiologists searching volumetric images use methods consistent with global processing models of expertise. In addition, we investigated whether search strategy (scanning/drilling) differs with experience level. Approach: Fifty radiologists with a wide range of experience evaluated chest computed-tomography scans for lung nodules while their eye movements and scrolling behaviors were tracked. Multiple linear regressions were used to determine: (1) how search behaviors differed with years of experience and the number of chest CTs evaluated per week and (2) which search behaviors predicted better performance. Results: Contrary to global processing models based on 2D images, experience was unrelated to measures of global processing (saccadic amplitude, coverage, time to first fixation, search time, and depth passes) in this task. Drilling behavior was associated with better accuracy than scanning behavior when controlling for observer experience. Greater image coverage was a strong predictor of task accuracy. Conclusions: Global processing ability may play a relatively small role in volumetric image interpretation, where global scene statistics are not available to radiologists in a single glance. Rather, in volumetric images, it may be more important to engage in search strategies that support a more thorough search of the image.

Concurrent neuroimaging and neurostimulation reveals a causal role for dlPFC in coding of task-relevant information.

Dorsolateral prefrontal cortex (dlPFC) is proposed to drive brain-wide focus by biasing processing in favour of task-relevant information. A longstanding debate concerns whether this is achieved through enhancing processing of relevant information and/or by inhibiting irrelevant information. To address this, we applied transcranial magnetic stimulation (TMS) during fMRI, and tested for causal changes in information coding. Participants attended to one feature, whilst ignoring another feature, of a visual object. If dlPFC is necessary for facilitation, disruptive TMS should decrease coding of attended features. Conversely, if dlPFC is crucial for inhibition, TMS should increase coding of ignored features. Here, we show that TMS decreases coding of relevant information across frontoparietal cortex, and the impact is significantly stronger than any effect on irrelevant information, which is not statistically detectable. This provides causal evidence for a specific role of dlPFC in enhancing task-relevant representations and demonstrates the cognitive-neural insights possible with concurrent TMS-fMRI-MVPA.

Neural signatures of vigilance decrements predict behavioural errors before they occur.

There are many monitoring environments, such as railway control, in which lapses of attention can have tragic consequences. Problematically, sustained monitoring for rare targets is difficult, with more misses and longer reaction times over time. What changes in the brain underpin these 'vigilance decrements'? We designed a multiple-object monitoring (MOM) paradigm to examine how the neural representation of information varied with target frequency and time performing the task. Behavioural performance decreased over time for the rare target (monitoring) condition, but not for a frequent target (active) condition. This was mirrored in neural decoding using magnetoencephalography: coding of critical information declined more during monitoring versus active conditions along the experiment. We developed new analyses that can predict behavioural errors from the neural data more than a second before they occurred. This facilitates pre-empting behavioural errors due to lapses in attention and provides new insight into the neural correlates of vigilance decrements.

Temporal dissociation of neural activity underlying synesthetic and perceptual colors.

Grapheme-color synesthetes experience color when seeing achromatic symbols. We examined whether similar neural mechanisms underlie color perception and synesthetic colors using magnetoencephalography. Classification models trained on neural activity from viewing colored stimuli could distinguish synesthetic color evoked by achromatic symbols after a delay of ∼100 ms. Our results provide an objective neural signature for synesthetic experience and temporal evidence consistent with higher-level processing in synesthesia.

Perception of visual-tactile asynchrony, bodily perceptual aberrations, and bodily illusions in schizophrenia.

Body perception can be altered in individuals with schizophrenia resulting in experiences of undefined boundaries, loss of ownership, and size changes. These individuals may also be more susceptible to the rubber hand illusion (RHI: an illusion of body perception that can also be induced in neurotypical populations), but the findings are mixed. Furthermore, the perception of multisensory timing, which is thought to be fundamental for body perception, is altered in schizophrenia. We tested whether altered perception of the temporal relationship between visual and tactile signals in schizophrenia predicts self-reported perceptual aberrations and RHI susceptibility. We found that the sensitivity to detect temporal asynchronies is reduced in schizophrenia and this is a significant predictor for bodily perceptual symptoms. In contrast, we found no evidence for a direct relationship between asynchrony detection sensitivity and RHI susceptibility. Instead, our findings suggest that experiencing more bodily perceptual symptoms increases the likelihood of endorsing unusual bodily experiences, resulting in higher RHI self-ratings but not higher proprioceptive drift scores. Our findings provide new insight into factors that may underlie the report of unusual body perceptions in schizophrenia.

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.

Toward an Individualized Neural Assessment of Receptive Language in Children.

Purpose We aimed to develop a noninvasive neural test of language comprehension to use with nonspeaking children for whom standard behavioral testing is unreliable (e.g., minimally verbal autism). Our aims were threefold. First, we sought to establish the sensitivity of two auditory paradigms to elicit neural responses in individual neurotypical children. Second, we aimed to validate the use of a portable and accessible electroencephalography (EEG) system, by comparing its recordings to those of a research-grade system. Third, in light of substantial interindividual variability in individuals' neural responses, we assessed whether multivariate decoding methods could improve sensitivity. Method We tested the sensitivity of two child-friendly covert N400 paradigms. Thirty-one typically developing children listened to identical spoken words that were either strongly predicted by the preceding context or violated lexical-semantic expectations. Context was given by a cue word (Experiment 1) or sentence frame (Experiment 2), and participants either made an overall judgment on word relatedness or counted lexical-semantic violations. We measured EEG concurrently from a research-grade system, Neuroscan's SynAmps2, and an adapted gaming system, Emotiv's EPOC+. Results We found substantial interindividual variability in the timing and topology of N400-like effects. For both paradigms and EEG systems, traditional N400 effects at the expected sensors and time points were statistically significant in around 50% of individuals. Using multivariate analyses, detection rate increased to 88% of individuals for the research-grade system in the sentences paradigm, illustrating the robustness of this method in the face of interindividual variations in topography. Conclusions There was large interindividual variability in neural responses, suggesting interindividual variation in either the cognitive response to lexical-semantic violations and/or the neural substrate of that response. Around half of our neurotypical participants showed the expected N400 effect at the expected location and time points. A low-cost, accessible EEG system provided comparable data for univariate analysis but was not well suited to multivariate decoding. However, multivariate analyses with a research-grade EEG system increased our detection rate to 88% of individuals. This approach provides a strong foundation to establish a neural index of language comprehension in children with limited communication. Supplemental Material https://doi.org/10.23641/asha.12606311.

Motor neuroplasticity: A MEG-fMRI study of motor imagery and execution in healthy ageing.

Age-related decline in motor function is associated with over-activation of the sensorimotor circuitry. Using a multimodal MEG-fMRI paradigm, we investigated whether this neural over-recruitment in old age would be related to changes in movement-related beta desynchronization (MRBD), a correlate of the inhibitory neurotransmitter γ-aminobutyric acid (GABA), and whether it would characterize compensatory recruitment or a reduction in neural specialization (dedifferentiation). We used MEG to assess age-related changes in beta band oscillations in primary motor cortices, fMRI to localize age-related changes in brain activity, and the Finger Configuration Task to measure task performance during overt and covert motor processing: motor execution (ME) and motor imagery (MI). The results are threefold: first, showing age-related neuroplasticity during ME of older adults, compared to young adults, as evidenced by increased MRBD in motor cortices and over-recruitment of sensorimotor areas; second, showing similar age-related neuroplastic changes during MI; and finally, showing signs of dedifferentiation during ME in older adults whose performance negatively correlated with connectivity to bilateral primary motor cortex. Together, these findings demonstrate that elevated MRBD levels, reflecting greater GABAergic inhibitory activity, and over-activation of the sensorimotor network during ME may not be compensatory, but rather might reflect an age-related decline of the quality of the underlying neural signal.

Late disruption of central visual field disrupts peripheral perception of form and color.

Evidence from neuroimaging and brain stimulation studies suggest that visual information about objects in the periphery is fed back to foveal retinotopic cortex in a separate representation that is essential for peripheral perception. The characteristics of this phenomenon have important theoretical implications for the role fovea-specific feedback might play in perception. In this work, we employed a recently developed behavioral paradigm to explore whether late disruption to central visual space impaired perception of color. In the first experiment, participants performed a shape discrimination task on colored novel objects in the periphery while fixating centrally. Consistent with the results from previous work, a visual distractor presented at fixation ~100ms after presentation of the peripheral stimuli impaired sensitivity to differences in peripheral shapes more than a visual distractor presented at other stimulus onset asynchronies. In a second experiment, participants performed a color discrimination task on the same colored objects. In a third experiment, we further tested for this foveal distractor effect with stimuli restricted to a low-level feature by using homogenous color patches. These two latter experiments resulted in a similar pattern of behavior: a central distractor presented at the critical stimulus onset asynchrony impaired sensitivity to peripheral color differences, but, importantly, the magnitude of the effect was stronger when peripheral objects contained complex shape information. These results show a behavioral effect consistent with disrupting feedback to the fovea, in line with the foveal feedback suggested by previous neuroimaging studies.

The structure of illusory conjunctions reveals hierarchical binding of multipart objects.

The world around us is filled with complex objects, full of color, motion, shape, and texture, and these features seem to be represented separately in the early visual system. Anne Treisman pointed out that binding these separate features together into coherent conscious percepts is a serious challenge, and she argued that selective attention plays a critical role in this process. Treisman also showed that, consistent with this view, outside the focus of attention we suffer from illusory conjunctions: misperceived pairings of features into objects. Here we used Treisman's logic to study the structure of pre-attentive representations of multipart, multicolor objects, by exploring the patterns of illusory conjunctions that arise outside the focus of attention. We found consistent evidence of some pre-attentive binding of colors to their parts, and weaker evidence of binding multiple colors of the same object. The extent to which such hierarchical binding occurs seems to depend on the geometric structure of multipart objects: Objects whose parts are easier to separate seem to exhibit greater pre-attentive binding. Together, these results suggest that representations outside the focus of attention are not entirely a "shapeless bundles of features," but preserve some meaningful object structure.

Visual body form and orientation cues do not modulate visuo-tactile temporal integration.

Body ownership relies on spatiotemporal correlations between multisensory signals and visual cues specifying oneself such as body form and orientation. The mechanism for the integration of bodily signals remains unclear. One approach to model multisensory integration that has been influential in the multisensory literature is Bayesian causal inference. This specifies that the brain integrates spatial and temporal signals coming from different modalities when it infers a common cause for inputs. As an example, the rubber hand illusion shows that visual form and orientation cues can promote the inference of a common cause (one's body) leading to spatial integration shown by a proprioceptive drift of the perceived location of the real hand towards the rubber hand. Recent studies investigating the effect of visual cues on temporal integration, however, have led to conflicting findings. These could be due to task differences, variation in ecological validity of stimuli and/or small samples. In this pre-registered study, we investigated the influence of visual information on temporal integration using a visuo-tactile temporal order judgement task with realistic stimuli and a sufficiently large sample determined by Bayesian analysis. Participants viewed videos of a touch being applied to plausible or implausible visual stimuli for one's hand (hand oriented plausibly, hand rotated 180 degrees, or a sponge) while also being touched at varying stimulus onset asynchronies. Participants judged which stimulus came first: viewed or felt touch. Results show that visual cues do not modulate visuo-tactile temporal order judgements. This is not in line with the idea that bodily signals indicating oneself influence the integration of multisensory signals in the temporal domain. The current study emphasises the importance of rigour in our methodologies and analyses to advance the understanding of how properties of multisensory events affect the encoding of temporal information in the brain.

Seeing versus knowing: The temporal dynamics of real and implied colour processing in the human brain.

Colour is a defining feature of many objects, playing a crucial role in our ability to rapidly recognise things in the world around us and make categorical distinctions. For example, colour is a useful cue when distinguishing lemons from limes or blackberries from raspberries. That means our representation of many objects includes key colour-related information. The question addressed here is whether the neural representation activated by knowing that something is red is the same as that activated when we actually see something red, particularly in regard to timing. We addressed this question using neural timeseries (magnetoencephalography, MEG) data to contrast real colour perception and implied object colour activation. We applied multivariate pattern analysis (MVPA) to analyse the brain activation patterns evoked by colour accessed via real colour perception and implied colour activation. Applying MVPA to MEG data allows us here to focus on the temporal dynamics of these processes. Male and female human participants (N = 18) viewed isoluminant red and green shapes and grey-scale, luminance-matched pictures of fruits and vegetables that are red (e.g., tomato) or green (e.g., kiwifruit) in nature. We show that the brain activation pattern evoked by real colour perception is similar to implied colour activation, but that this pattern is instantiated at a later time. These results suggest that a common colour representation can be triggered by activating object representations from memory and perceiving colours.

Do target detection and target localization always go together? Extracting information from briefly presented displays.

The human visual system is capable of processing an enormous amount of information in a short time. Although rapid target detection has been explored extensively, less is known about target localization. Here we used natural scenes and explored the relationship between being able to detect a target (present vs. absent) and being able to localize it. Across four presentation durations (~ 33-199 ms), participants viewed scenes taken from two superordinate categories (natural and manmade), each containing exemplars from four basic scene categories. In a two-interval forced choice task, observers were asked to detect a Gabor target inserted in one of the two scenes. This was followed by one of two different localization tasks. Participants were asked either to discriminate whether the target was on the left or the right side of the display or to click on the exact location where they had seen the target. Targets could be detected and localized at our shortest exposure duration (~ 33 ms), with a predictable improvement in performance with increasing exposure duration. We saw some evidence at this shortest duration of detection without localization, but further analyses demonstrated that these trials typically reflected coarse or imprecise localization information, rather than its complete absence. Experiment 2 replicated our main findings while exploring the effect of the level of "openness" in the scene. Our results are consistent with the notion that when we are able to extract what objects are present in a scene, we also have information about where each object is, which provides crucial guidance for our goal-directed actions.

Limits on visual awareness of object targets in the context of other object category masks: Investigating bottlenecks in the continuous flash suppression paradigm with hand and tool stimuli.

The continuous flash suppression (CFS) task can be used to investigate what limits our capacity to become aware of visual stimuli. In this task, a stream of rapidly changing mask images to one eye initially suppresses awareness for a static target image presented to the other eye. Several factors may determine the breakthrough time from mask suppression, one of which is the overlap in representation of the target/mask categories in higher visual cortex. This hypothesis is based on certain object categories (e.g., faces) being more effective in blocking awareness of other categories (e.g., buildings) than other combinations (e.g., cars/chairs). Previous work found mask effectiveness to be correlated with category-pair high-level representational similarity. As the cortical representations of hands and tools overlap, these categories are ideal to test this further as well as to examine alternative explanations. For our CFS experiments, we predicted longer breakthrough times for hands/tools compared to other pairs due to the reported cortical overlap. In contrast, across three experiments, participants were generally faster at detecting targets masked by hands or tools compared to other mask categories. Exploring low-level explanations, we found that the category average for edges (e.g., hands have less detail compared to cars) was the best predictor for the data. This low-level bottleneck could not completely account for the specific category patterns and the hand/tool effects, suggesting there are several levels at which object category-specific limits occur. Given these findings, it is important that low-level bottlenecks for visual awareness are considered when testing higher-level hypotheses.