Rapid withdrawal from a threatening animal is movement-specific and mediated by reflex-like neural processing.

Responses to potentially dangerous stimuli are among the most basic animal behaviors. While research has shown that threats automatically capture the attention of human participants, research has failed to demonstrate automatic behavioral responses to threats in humans. Using a novel naturalistic paradigm, we show that two species of animals humans often report fearing trigger rapid withdrawal responses: participants withdrew their arm from photos of snakes and spiders faster, and with higher acceleration when compared to bird and butterfly stimuli. The behavior was specific to withdrawal as approach movements or button-press/release tasks failed to detect a similar difference. Moreover, between-participant differences in how aversive they found the stimuli predicted the participant's withdrawal speed, indicating that the paradigm was also sensitive to trait-level differences between individuals. Using electroencephalography (EEG), we show that the fast withdrawal was mediated by two attentional processes. First, fast withdrawal responses were associated with early amplification of sensory signals (40-110 ms after stimulus). Second, a later correlate of feature-based attention (early posterior negativity, EPN, 200-240 ms after stimulus) revealed the opposite pattern: Stronger EPN was associated with slower behavioral responses, suggesting that the deployment of attention towards the threatening stimulus features, or failure to "disengage" attention from the stimulus, was detrimental for withdrawal speed. Altogether, the results suggest that rapid behavioral withdrawal from a threatening animal is mediated by reflex-like attentional processing, and later, conscious attention to stimulus features may hinder escaping the treat.

Promise and challenges for discovering transcranial magnetic stimulation induced "numbsense"-Commentary on Ro & Koenig (2021).

The notion that behavioral responses to stimuli can be mediated by separate unconscious and conscious sensory pathways remains popular, but also hotly debated. Recently, Ro and Koenig (2021) reported that when activity in somatosensory cortex was interfered with transcranial magnetic stimulation (TMS), participants could discriminate tactile stimuli they reported not consciously feeling. The study launches an interesting new area of research, helping to uncover mechanisms of unconscious perception that possibly generalize across different sensory modalities. However, we argue here that the study by Ro and Koenig also has several significant shortcomings, and it fails to provide evidence that pathways bypassing primary somatosensory cortex enable unconscious tactile discrimination. By referring to numerous studies investigating TMS-induced blindsight, we outline challenges in demonstrating unconscious sensory pathways using TMS. By facing to these challenges, research investigating TMS-induced numbsense has potential to stimulate progress in stubborn debates and reveal modality-general mechanisms of unconscious perception.

Subliminal perception is continuous with conscious vision and can be predicted from prestimulus electroencephalographic activity.

Individuals are able to discriminate visual stimuli they report not consciously seeing. This phenomenon is known as "subliminal perception." Such capacity is often assumed to be relatively automatic in nature and rely on stimulus-driven activity in low-level cortical areas. Instead, here we asked to what extent neural activity before stimulus presentation influences subliminal perception. We asked participants to discriminate the location of a briefly presented low-contrast visual stimulus and then rate how well they saw the stimulus. Consistent with previous studies, participants correctly discriminated with slightly above chance-level accuracy the location of a stimulus they reported not seeing. Signal detection analyses indicated that while subjects categorized their percepts as "unconscious," their capacity to discriminate these stimuli lay on the same continuum as conscious vision. We show that the accuracy of discriminating the location of a subliminal stimulus could be predicted with relatively high accuracy (AUC = 0.70) based on lateralized electroencephalographic (EEG) activity before the stimulus, the hemifield where the stimulus was presented, and the accuracy of previous trial's discrimination response. Altogether, our results suggest that rather than being a separate unconscious capacity, subliminal perception is based on similar processes as conscious vision.

Functional Magnetic Resonance Imaging during Visual Perception Tasks in Adolescents Born Prematurely.

OBJECTIVES: Impairments in visual perception are among the most common developmental difficulties related to being born prematurely, and they are often accompanied by problems in other developmental domains. Neural activation in participants born prematurely and full-term during tasks that assess several areas of visual perception has not been studied. To better understand the neural substrates of the visual perceptual impairments, we compared behavioral performance and brain activations during visual perception tasks in adolescents born very preterm (birth weight ≤1500 g or gestational age <32 weeks) and full-term. METHODS: Tasks assessing visual closure, discrimination of a deviating figure, and discrimination of figure and ground from the Motor-Free Visual Perception Test, Third Edition were performed by participants born very preterm (n = 37) and full-term (n = 34) at 12 years of age during functional magnetic resonance imaging. RESULTS: Behavioral performance in the visual perception tasks did not differ between the groups. However, during the visual closure task, brain activation was significantly stronger in the group born very preterm in a number of areas including the frontal, anterior cingulate, temporal, and posterior medial parietal/cingulate cortices, as well as in parts of the cerebellum, thalamus, and caudate nucleus. CONCLUSIONS: Differing activations during the visual closure task potentially reflect a compensatory neural process related to premature birth or lesser neural efficiency or may be a result of the use of compensatory behavioral strategies in the study group born very preterm.

V1 activity during feedforward and early feedback processing is necessary for both conscious and unconscious motion perception.

The study of blindsight has revealed a seminal dissociation between conscious vision and visually guided behavior: some patients who are blind due to V1 lesions seem to be able to employ unconscious visual information in their behavior. The standard assumption is that these findings generalize to the neurologically healthy. We tested whether unconscious processing of motion is possible without the contribution of V1 in neurologically healthy participants by disturbing activity in V1 using transcranial magnetic stimulation (TMS). Unconscious processing was measured with redundant target effect (RTE), a phenomenon where participants respond faster to two stimuli than to one stimulus, when the task is just to respond as fast as possible when one stimulus or two simultaneous stimuli are presented. We measured the RTE caused by a motion stimulus. V1 activity was interfered with different stimulus onset asynchronies (SOA) to test whether TMS delivered in a specific time window suppresses conscious perception (participant reports seeing only one of the two stimuli) but does not affect unconscious processing (RTE). We observed that at each SOA, when TMS suppressed conscious perception of the stimulus, the RTE was also eliminated. However, when visibility of the redundant target was suppressed with a visual mask, we found unconscious processing of motion. This suggests that unconscious processing of motion depends on V1 in neurologically healthy humans. We conclude that the neural mechanisms that enable motion processing in blindsight are modulated by neuroplastic changes in connectivity between subcortical areas and the visual cortex after the V1 lesion. Neurologically healthy observers cannot process motion unconsciously without functioning of V1.

Consciousness as a concrete physical phenomenon.

The typical empirical approach to studying consciousness holds that we can only observe the neural correlates of experiences, not the experiences themselves. In this paper we argue, in contrast, that experiences are concrete physical phenomena that can causally interact with other phenomena, including observers. Hence, experiences can be observed and scientifically modelled. We propose that the epistemic gap between an experience and a scientific model of its neural mechanisms stems from the fact that the model is merely a theoretical construct based on observations, and distinct from the concrete phenomenon it models, namely the experience itself. In this sense, there is a gap between any natural phenomenon and its scientific model. On this approach, a neuroscientific theory of the constitutive mechanisms of an experience is literally a model of the subjective experience itself. We argue that this metatheoretical framework provides a solid basis for the empirical study of consciousness.

Dynamic Changes in Cortical Effective Connectivity Underlie Transsaccadic Integration in Humans.

Due to saccadic eye movements the retinal image is abruptly displaced 2-4 times a second, yet we experience a stable and continuous stream of vision. It is known that saccades modulate neural processing in various local brain areas, but the question of how saccades influence neural communication between different areas in the thalamo-cortical system has remained unanswered. By combining transcranial magnetic stimulation with electroencephalography, we found that saccades were accompanied by dynamic changes in causal communication between different brain areas in humans. These changes were anticipatory; they began before the actual eye movement. Compared with fixation, communication between posterior cortical areas was first briefly enhanced during saccades, but subsequently peri-saccadic information did not ignite sustained activity in fronto-parietal cortices. This suggests that the brain constructs a spatially stable and temporally continuous stream of conscious vision from discrete fixations by restricting the access of peri-saccadic visual information to sustained processing in fronto-parietal cortices.

Early processing in primary visual cortex is necessary for conscious and unconscious vision while late processing is necessary only for conscious vision in neurologically healthy humans.

The neural mechanisms underlying conscious and unconscious visual processes remain controversial. Blindsight patients may process visual stimuli unconsciously despite their V1 lesion, promoting anatomical models, which suggest that pathways bypassing the V1 support unconscious vision. On the other hand, physiological models argue that the major geniculostriate pathway via V1 is involved in both unconscious and conscious vision, but in different time windows and in different types of neural activity. According to physiological models, feedforward activity via V1 to higher areas mediates unconscious processes whereas feedback loops of recurrent activity from higher areas back to V1 support conscious vision. With transcranial magnetic stimulation (TMS) it is possible to study the causal role of a brain region during specific time points in neurologically healthy participants. In the present study, we measured unconscious processing with redundant target effect, a phenomenon where participants respond faster to two stimuli than one even when one of the stimuli is not consciously perceived. We tested the physiological feedforward-feedback model of vision by suppressing conscious vision by interfering selectively either with early or later V1 activity with TMS. Our results show that early V1 activity (60ms) is necessary for both unconscious and conscious vision. During later processing stages (90ms), V1 contributes selectively to conscious vision. These findings support the feedforward-feedback-model of consciousness.

Transcranial magnetic stimulation of early visual cortex suppresses conscious representations in a dichotomous manner without gradually decreasing their precision.

Transcranial magnetic stimulation (TMS) of early visual cortex can suppresses visual perception at early stages of processing. The suppression can be measured both with objective forced-choice tasks and with subjective ratings of visual awareness, but there is lack of objective evidence on how and whether the TMS influences the quality of representations. Does TMS decrease the precision of representations in graded manner, or does it lead to dichotomous, "all-or-nothing" suppression. We resolved this question by using a continuous measure of the perceptual error: the observers had to perceive the orientation of a target (Landort-C) and to adjust the orientation of a probe to match that of the target. Mixture modeling was applied to estimate the probability of guess trials and the standard deviation of the non-guess trials. TMS delivered 60-150 ms after stimulus-onset influenced only the guessing rate, whereas the standard deviation (i.e., precision) was not affected. This suggests that TMS suppressed representations dichotomously without affecting their precision. The guessing probability correlated with subjective visibility ratings, suggesting that it measured visual awareness. In a control experiment, manipulation of the stimulus contrast affected the standard deviation of the errors, indicating that contrast has a gradual influence on the precision of representations. The findings suggest that TMS of early visual cortex suppresses perception in dichotomous manner by decreasing the signal-to-noise ratio by increasing the noise level, whereas reduction of the signal level (i.e., contrast) decreases the precision of representations.

Judging Total Volumes Of Silhouetted Spheres In Different Numerosities.

Volume and number are addressed separately rather than combined in perception research. Yet, our everyday problems often involve summed continuous volumes of countable solid objects with partial depth cues (e.g., food items). The participants were presented with a set of black-and-white silhouettes of spheres that independently varied in numerosity (from 1 to 6) and total volume (2, 4, 6, or 8), and an adjacent silhouette of a partially filled cylinder. They judged how much the silhouetted sphere(s) in the set would raise the level of the cylinder content if the spheres were immersed into that content. Higher total volumes and numerosities of the spheres were judged slower and underestimated. Lower total volumes and numerosities were judged faster and overestimated. These effects strongly reflected the total silhouette area of the spheres in a set. The discontinuous effect of numerosity on judgment accuracy and speed suggested separate judgment modes below and above Numerosity 3.

Rapid and accurate processing of multiple objects in briefly presented scenes.

Humans can detect multiple objects in briefly presented natural visual scenes, but the mechanisms through which the objects are segmented from the background and consciously accessed remain open. By asking participants to report how many humans natural photos presented for 50 ms contain, we show that up to three items can be rapidly enumerated from natural scenes without compromising speed or accuracy. In contrast to standard parallel and serial models of object selection, our results revealed that the participants were fastest in enumerating two objects; even enumerating one single item required additional processing time. Also enumeration accuracy slightly increased in the subitizing range as number increased. Our results suggest that the visual system is tuned to process multiple items, which may underlie spatial and numerical cognition, and be beneficial in real-world situations that often require dealing with more than one object at a time.

Overlapping activity periods in early visual cortex and posterior intraparietal area in conscious visual shape perception: a TMS study.

Parietal cortex is often activated in brain imaging studies on conscious visual processing, but its causal role and timing in conscious and nonconscious perception are poorly understood. We studied the role of posterior intraparietal sulcus (IPS) and early visual areas (V1/V2) in conscious and nonconscious vision by interfering with their functioning with MRI-guided transcranial magnetic stimulation (TMS). The observers made binary forced-choice decisions concerning the shape or color of the metacontrast masked targets and rated the quality of their conscious perception. TMS was applied 30, 60, 90, or 120ms after stimulus-onset. In the shape discrimination task, TMS of V1/V2 impaired conscious perception at 60, 90, and 120ms and nonconscious perception at 90ms. TMS of IPS impaired only conscious shape perception, also around 90ms. Conscious color perception was facilitated or suppressed depending on the strength of the TMS-induced electric field in V1/V2 at 90ms. The results suggest that simultaneous activity in V1/V2 and IPS around 90ms is necessary for visual awareness of shape but not for nonconscious perception. The overlapping activity periods of IPS and V1/V2 may reflect recurrent interaction between parietal cortex and V1 in conscious shape perception.

Low-Level Auditory Processing Correlates With Language Abilities: An ERP Study Investigating Sequence Learning and Auditory Processing in School-Aged Children.

Auditory processing and procedural learning deficits have been associated with language learning difficulties. We investigated the relationship of these skills and school-age language abilities in children with and without a history of late talking using auditory event related potentials (ERPs). Late talking (i.e., slow early language development) increases the risk of persistent language difficulties, but its causes remain unknown. Participants in this study were children with varying language abilities (n = 60). Half of the participants (n = 30) had a history of late talking. We measured procedural learning by manipulating the predictability of sine tone stimuli in a passive auditory ERP paradigm. Auditory processing was tested by examining how the presence of noise (increasing perceptual demands) affected the ERPs. Contrary to our hypotheses on auditory processing and language development, the effect of noise on ERPs did not correlate with school-age language abilities in children with or without a history of late talking. Our paradigm failed to reveal interpretable effects of predictability leaving us unable to assess the effects of procedural learning. However, better language abilities were related to weaker responses in a 75-175 ms time window, and stronger responses in a 150-250 ms time window. We suggest that the weak early responses in children with better language ability reflect efficient processing of low-level auditory information, allowing deeper processing of later, high-level auditory information. We assume that these differences reflect variation in brain maturation between individuals with varying language abilities.

Introducing the Intra-Individual Variability Hypothesis in Explaining Individual Differences in Language Development.

PURPOSE: Response times (RTs) are commonly used in studying language acquisition. However, previous research utilizing RT in the context of language has largely overlooked the intra-individual variability (IIV) of RTs, which could hold significant information about the processes underlying language acquisition. METHOD: We explored the association between language abilities and RT variability in visuomotor tasks using two data sets from previously published studies. The participants were 7- to 10-year-old children (n = 77). RESULTS: Our results suggest that increased variability in RTs is associated with weaker language abilities. Specifically, this within-participant variability in visuomotor RTs, especially the proportion of unusually slow responses, predicted language abilities better than mean RTs, a factor often linked to language skills in past research. CONCLUSIONS: Based on our findings, we introduce the IIV hypothesis in explaining individual differences in language development. According to our hypothesis, inconsistency in the timing of cognitive processes, reflected by increased IIV in RTs, degrades learning different aspects of language, and results in individual differences in language abilities. Future studies should further examine the relationship between IIV and language abilities, and test the extent to which the possible relationship is causal.

Neurologically Healthy Humans' Ability to Make Saccades Toward Unseen Targets.

Some patients with a visual field loss due to a lesion in the primary visual cortex (V1) can shift their gaze to stimuli presented in their blind visual field. The extent to which a similar "blindsight" capacity is present in neurologically healthy individuals remains unknown. Using retinotopically navigated transcranial magnetic stimulation (TMS) of V1 (Experiment 1) and metacontrast masking (Experiment 2) to suppress conscious vision, we examined neurologically healthy humans' ability to make saccadic eye movements toward visual targets that they reported not seeing. In the TMS experiment, the participants were more likely to initiate a saccade when a stimulus was presented, and they reported not seeing it, than in trials which no stimulus was presented. However, this happened only in a very small proportion (∼8%) of unseen trials, suggesting that saccadic reactions were largely based on conscious perception. In both experiments, saccade landing location was influenced by unconscious information: When the participants denied seeing the target but made a saccade, the saccade was made toward the correct location (TMS: 68%, metacontrast: 63%) more often than predicted by chance. Signal detection theoretic measures suggested that in the TMS experiment, saccades toward unseen targets may have been based on weak conscious experiences. In both experiments, reduced visibility of the target stimulus was associated with slower and less precise gaze shifts. These results suggest that saccades made by neurologically healthy humans may be influenced by unconscious information, although the initiation of saccades is largely based on conscious vision.

Budget-Based Classification of Parkinson's Disease From Resting State EEG.

Early detection is vital for future neuroprotective treatments of Parkinson's disease (PD). Resting state electroencephalographic (EEG) recording has shown potential as a cost-effective means to aid in detection of neurological disorders such as PD. In this study, we investigated how the number and placement of electrodes affects classifying PD patients and healthy controls using machine learning based on EEG sample entropy. We used a custom budget-based search algorithm for selecting optimized sets of channels for classification, and iterated over variable channel budgets to investigate changes in classification performance. Our data consisted of 60-channel EEG collected at three different recording sites, each of which included observations collected both eyes open (total N = 178) and eyes closed (total N = 131). Our results with the data recorded eyes open demonstrated reasonable classification performance (ACC = .76; AUC = .76) with only 5 channels placed far away from each other, the selected regions including right-frontal, left-temporal and midline-occipital sites. Comparison to randomly selected subsets of channels indicated improved classifier performance only with relatively small channel-budgets. The results with the data recorded eyes closed demonstrated consistently worse classification performance (when compared to eyes open data), and classifier performance improved more steadily as a function of number of channels. In summary, our results suggest that a small subset of electrodes of an EEG recording can suffice for detecting PD with a classification performance on par with a full set of electrodes. Furthermore our results demonstrate that separately collected EEG data sets can be used for pooled machine learning based PD detection with reasonable classification performance.

Recurrent processing in V1/V2 contributes to categorization of natural scenes.

Humans are able to categorize complex natural scenes very rapidly and effortlessly, which has led to an assumption that such ultra-rapid categorization is driven by feedforward activation of ventral brain areas. However, recent accounts of visual perception stress the role of recurrent interactions that start rapidly after the activation of V1. To study whether or not recurrent processes play a causal role in categorization, we applied fMRI-guided transcranial magnetic stimulation on early visual cortex (V1/V2) and lateral occipital cortex (LO) while the participants categorized natural images as containing animals or not. The results showed that V1/V2 contributed to categorization speed and to subjective perception during a long activity period before and after the contribution of LO had started. This pattern of results suggests that recurrent interactions in visual cortex between areas along the ventral stream and striate cortex play a causal role in categorization and perception of natural scenes.

Unconscious and conscious processing of color rely on activity in early visual cortex: a TMS study.

Chromatic information is processed by the visual system both at an unconscious level and at a level that results in conscious perception of color. It remains unclear whether both conscious and unconscious processing of chromatic information depend on activity in the early visual cortex or whether unconscious chromatic processing can also rely on other neural mechanisms. In this study, the contribution of early visual cortex activity to conscious and unconscious chromatic processing was studied using single-pulse TMS in three time windows 40-100 msec after stimulus onset in three conditions: conscious color recognition, forced-choice discrimination of consciously invisible color, and unconscious color priming. We found that conscious perception and both measures of unconscious processing of chromatic information depended on activity in early visual cortex 70-100 msec after stimulus presentation. Unconscious forced-choice discrimination was above chance only when participants reported perceiving some stimulus features (but not color).

Unconscious response priming by shape depends on geniculostriate visual projection.

It has been suggested that unconscious visual processing of some stimulus features might occur without the contribution of early visual cortex (V1/V2). In the present study, the causal role of V1/V2 in unconscious processing of simple shapes in intact human brain was studied by applying transcranial magnetic stimulation (TMS) on early visual cortex or lateral occipital cortex (LO) while observers performed a metacontrast-masked response priming task with arrow figures as visual stimuli. Magnetic stimulation of V1/V2 impaired masked priming 30-90 ms after the onset of the prime. Stimulation of LO reduced the magnitude of masked priming at 90-120 ms, but this effect occurred only in the early parts of the priming experiment. A control task measuring the visibility of masked primes indicated that the orientation of masked primes could not be consciously discriminated and that TMS did not influence the conscious visibility of the primes indirectly by reducing the effectiveness of the mask in the critical time windows. We conclude that feedforward sweep of processing from V1/V2 (30-90 ms) to LO (90 ms and above) is necessary for unconscious priming of shape, whereas conscious perception requires also the contribution of recurrent (feedback) processing.

Modality-specific and modality-general electrophysiological correlates of visual and auditory awareness: Evidence from a bimodal ERP experiment.

To date, most studies on the event-related potential (ERP) correlates of conscious perception have examined a single perceptual modality. We compared electrophysiological correlates of visual and auditory awareness in the same experiment to test whether there are modality-specific and modality-general correlates of conscious perception. We used near threshold stimulation and analyzed event-related potentials in response to aware and unaware trials in visual, auditory and bimodal conditions. The results showed modality-specific negative amplitude correlates of conscious perception between 200 and 300 ms after stimulus onset. A combination of these auditory and visual awareness negativities was observed in the bimodal condition. A later positive amplitude difference, whose early part was modality-specific, possibly reflecting access to global workspace, and later part shared modality-general features, possibly indicating higher level cognitive processing involving the decision making, was also observed.