Distinct ventral stream and prefrontal cortex representational dynamics during sustained conscious visual perception

Abstract:

Instances of sustained stationary sensory input are ubiquitous. However, previous work focused almost exclusively on transient onset responses. This presents a critical challenge for neural theories of consciousness, which should account for the full temporal extent of experience. To address this question, we use intracranial recordings from ten human patients with epilepsy to view diverse images of multiple durations. We reveal that, in sensory regions, despite dramatic changes in activation magnitude, the distributed representation of categories and exemplars remains sustained and stable. In contrast, in frontoparietal regions, we find transient content representation at stimulus onset. Our results highlight the connection between the anatomical and temporal correlates of experience. To the extent perception is sustained, it may rely on sensory representations and to the extent perception is discrete, centered on perceptual updating, it may rely on frontoparietal representations.

Authors:

• Gal Vishne

• Edden M. Gerber

• Robert T. Knight

• Leon Y. Deouell

Date: 2023

DOI: https://doi.org/10.1016/j.celrep.2023.112752

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Systematic comparison between a wireless EEG system with dry electrodes and a wired EEG system with wet electrodes

Abstract:

Recent advances in dry electrodes technology have facilitated the recording of EEG in situations not previously possible, thanks to the relatively swift electrode preparation and avoidance of applying gel to subject's hair. However, to become a true alternative, these systems should be compared to state-of-the-art wet EEG systems commonly used in clinical or research applications. In our study, we conducted a systematic comparison of electrodes application speed, subject comfort, and most critically electrophysiological signal quality between the conventional and wired Biosemi EEG system using wet active electrodes and the compact and wireless F1 EEG system consisting of dry passive electrodes. All subjects (n = 27) participated in two recording sessions on separate days, one with the wet EEG system and one with the dry EEG system, in which the session order was counterbalanced across subjects. In each session, we recorded their EEG during separate rest periods with eyes open and closed followed by two versions of a serial visual presentation target detection task. Each task component allows for a neural measure reflecting different characteristics of the data, including spectral power in canonical low frequency bands, event-related potential components (specifically, the P3b), and single trial classification based on machine learning. The performance across the two systems was similar in most measures, including the P3b amplitude and topography, as well as low frequency (theta, alpha, and beta) spectral power at rest. Both EEG systems performed well above chance in the classification analysis, with a marginal advantage of the wet system over the dry. Critically, all aforementioned electrophysiological metrics showed significant positive correlations (r = 0.54–0.89) between the two EEG systems. This multitude of measures provides a comprehensive comparison that captures different aspects of EEG data, including temporal precision, frequency domain as well as multivariate patterns of activity. Taken together, our results indicate that the dry EEG system used in this experiment can effectively record electrophysiological measures commonly used across the research and clinical contexts with comparable quality to the conventional wet EEG system.

Authors:

• Julia W.Y. Kam

• Sandon Griffin

• Alan Shen

• Shawn Patel

• Hermann Hinrichs

• Hans-Jochen Heinze

• Leon Y. Deouell

• Robert T. Knight

Date: 2019

DOI: 10.1016/j.neuroimage.2018.09.012

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Human posterior parietal cortex responds to visual stimuli as early as peristriate occipital cortex

ABSTRACT

Much of what is known about the timing of visual processing in the brain is inferred from intracranial studies in monkeys, with human data limited to mainly noninvasive methods with lower spatial resolution. Here, we estimated visual onset latencies from electrocorticographic (ECoG) recordings in a patient who was implanted with 112 subdural electrodes, distributed across the posterior cortex of the right hemisphere, for presurgical evaluation of intractable epilepsy. Functional MRI prior to surgery was used to determine boundaries of visual areas. The patient was presented with images of objects from several categories. Event‐related potentials (ERPs) were calculated across all categories excluding targets, and statistically reliable onset latencies were determined, using a bootstrapping procedure over the single trial baseline activity in individual electrodes. The distribution of onset latencies broadly reflected the known hierarchy of visual areas, with the earliest cortical responses in primary visual cortex, and higher areas showing later responses. A clear exception to this pattern was a robust, statistically reliable and spatially localized, very early response, on the bank of the posterior intraparietal sulcus (IPS). The response in the IPS started nearly simultaneously with responses detected in peristriate visual areas, around 60 ms poststimulus onset. Our results support the notion of early visual processing in the posterior parietal lobe, not respecting traditional hierarchies, and give direct evidence for onset times of visual responses across the human cortex.

AUTHORS

• Tamar I. Regev 

• Jonathan Winawer 

• Edden M. Gerber 

• Robert T. Knight 

• Leon Y. Deouell

Date: 2018

DOI: 10.1111/ejn.14164

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Cortical Representation of Persistent Visual Stimuli

Abstract:

Research into visual neural activity has focused almost exclusively on onset- or change-driven responses and little is known about how information is encoded in the brain during sustained periods of visual perception. We used intracranial recordings in humans to determine the degree to which the presence of a visual stimulus is persistently encoded by neural activity. The correspondence between stimulus duration and neural response duration was strongest in early visual cortex and gradually diminished along the visual hierarchy, such that is was weakest in inferior-temporal category-selective regions. A similar posterior-anterior gradient was found within inferior temporal face-selective regions, with posterior but not anterior sites showing persistent face-selective activity. The results suggest that regions that appear uniform in terms of their category selectivity are dissociated by how they temporally represent a stimulus in support of ongoing visual perception, and delineate a large-scale organizing principle of the ventral visual stream.

Authors:

• Edden M. Gerber

• Tal Golan

• Robert T. Knight

• Leon Y. Deouell

Date: 2017

DOI: http://dx.doi.org/10.1016/j.neuroimage.2017.08.028

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Non-sinusoidal activity can produce cross-frequency coupling in cortical signals in the absence of functional interaction between neural sources

ABSTRACT

The analysis of cross-frequency coupling (CFC) has become popular in studies involving intracranial and scalp EEG recordings in humans. It has been argued that some cases where CFC is mathematically present may not reflect an interaction of two distinct yet functionally coupled neural sources with different frequencies. Here we provide two empirical examples from intracranial recordings where CFC can be shown to be driven by the shape of a periodic waveform rather than by a functional interaction between distinct sources. Using simulations, we also present a generalized and realistic scenario where such coupling may arise. This scenario, which we term waveform-dependent CFC, arises when sharp waveforms (e.g., cortical potentials) occur throughout parts of the data, in particular if they occur rhythmically. Since the waveforms contain both low- and high-frequency components, these components can be inherently phase-aligned as long as the waveforms are spaced with appropriate intervals. We submit that such behavior of the data, which seems to be present in various cortical signals, cannot be interpreted as reflecting functional modulation between distinct neural sources without additional evidence. In addition, we show that even low amplitude periodic potentials that cannot be readily observed or controlled for, are sufficient for significant CFC to occur.

AUTHORS

• Edden Gerber

• Boaz Sadeh

• Andrew Ward

• Robert T. Knight

• Leon Y. Deouell

Date: 2016

DOI: 10.1371/journal.pone.0167351

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Extracting duration information in a picture category decoding task using Hidden Markov Models

ABSTRACT

Adapting classifiers for the purpose of brain signal decoding is a major challenge in brain–computer-interface (BCI) research. In a previous study we showed in principle that hidden Markov models (HMM) are a suitable alternative to the well-studied static classifiers. However, since we investigated a rather straightforward task, advantages from modeling of the signal could not be assessed. Approach. Here, we investigate a more complex data set in order to find out to what extent HMMs, as a dynamic classifier, can provide useful additional information. We show for a visual decoding problem that besides category information, HMMs can simultaneously decode picture duration without an additional training required. This decoding is based on a strong correlation that we found between picture duration and the behavior of the Viterbi paths. Main results. Decoding accuracies of up to 80% could be obtained for category and duration decoding with a single classifier trained on category information only. Significance. The extraction of multiple types of information using a single classifier enables the processing of more complex problems, while preserving good training results even on small databases. Therefore, it provides a convenient framework for online real-life BCI utilizations.

AUTHORS

• Tim Pfeiffer

• Nicolai Heinze

• Robert Frysch

• Leon Y. Deouell

• Mircea Schoenfeld

• Robert T. Knight

• Georg Rose

Date: 2016

DOI: doi:10.1088/1741-2560/13/2/026010

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Brain activity during landmark and line bisection tasks

Authors:

• Metehan Cicek

• Leon Y. Deouell

• Robert T. Knight

Date: 2009

DOI: 10.3389/neuro.09.007.2009

PubMed: 19521543

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Abstract:

Neglect patients bisect lines far rightward of center whereas normal subjects typically bisect lines with a slight leftward bias supporting a right hemisphere bias for attention allocation. We used fMRI to assess the brain regions related to this function in normals, using two complementary tasks. In the Landmark task subjects were required to judge whether or not a presented line was bisected correctly. During the line bisection task, subjects moved a cursor and indicated when it reached the center of the line. The conjunction of BOLD activity for both tasks showed right lateralized intra-parietal sulcus and lateral peristriate cortex activity. The results provide evidence that predominantly right hemisphere lateralized processes are engaged in normal subjects during tasks that are failed in patients with unilateral neglect and highlight the importance of a right fronto-parietal network in attention allocation.

Executive function and higher-order cognition: EEG studies

Authors:

• Leon Y. Deouell

• Robert T. Knight

Date: 2009

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Abstract:

Multiple goals, from survival to pleasure, coexist at every given moment, and internal needs as well as external events act in a push–pull manner to bias behavior. To maintain optimal goal-directed behavior, a control (‘executive’) system is needed that will dynamically prioritize the processing of information as well as the planning of actions and their execution. In addition to direct motor planning, major components of this executive system are working memory, attention, and conflict/error monitoring. Working memory would allow maintaining a goal across time, as well as information required to achieve the goal. Selective attention (‘voluntary’ or ‘endogenous’ attention) would facilitate the processing of one stream of input and suppress another while involuntary attention mechanisms allow for changes in the environment, as well as changes in internal drives, to interfere with ongoing behavior in a rapid and flexible manner. A monitoring module is needed to assess the efficiency of the executed behavior so that behavior can be adjusted in an optimal way. Consequently, the executive system needs to interact with multiple sensory regions of the brain, as well as with motor output regions involved in orientation, locomotion, and speech. Event-related brain potentials (ERPs), recorded on the scalp (and recently also intracranially) have established scalp-recorded signatures of executive functions. The effect of brain lesions on these measures of electrical brain activity provides a window into the networks supporting the executive system. This article describes ERP studies conducted specifically with patients suffering from well-circumscribed brain lesions involving mainly the lateral prefrontal cortex (LPFC), a major hub of the executive system.

Detecting violations of sensory expectancies following cerebellar degeneration: a mismatch negativity study

Authors:

• Torgeir Moberget

• Christina M. Karns

• Leon Y. Deouell

• Magnus Lindgren

• Robert T. Knight

• Richard B. Ivry

Date: 2008

DOI: 10.1016/j.neuropsychologia.2008.03.016

PubMed: 18486157

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Abstract:

Two hypotheses concerning cerebellar function and predictive behavior are the sensory prediction hypothesis and the timing hypothesis. The former postulates that the cerebellum is critical in generating expectancies regarding forthcoming sensory information. The latter postulates that this structure is critical in generating expectancies that are precisely timed; for example, the expected duration of an event or the time between events. As such, the timing hypothesis constitutes a more specific form of prediction. The present experiment contrasted these two hypotheses by examining the mismatch negativity (MMN) response in patients with cerebellar cortical atrophy and matched controls. While watching a silent movie, a stream of task-irrelevant sounds was presented. A standard sound was presented 60% of the time, whereas the remaining sounds deviated from the standard on one of four dimensions: duration, intensity, pitch, or location. The timing between stimuli was either periodic or aperiodic. Based on the sensory prediction hypothesis, the MMN for the patients should be abnormal across all four dimensions. In contrast, the timing hypothesis would predict a selective impairment of the duration MMN. Moreover, the timing hypothesis would also predict that the enhancement of the MMN observed in controls when the stimuli are presented periodically should be attenuated in the patients. Compared to controls, the patients exhibited a delayed latency in the MMN to duration deviants and a similar trend for the intensity deviants, while pitch and location MMNs did not differ between groups. Periodicity had limited and somewhat inconsistent effects. The present results are at odds with a general role for the cerebellum in sensory prediction and provide partial support for the timing hypothesis.

No disillusions in auditory extinction: perceiving a melody comprised of unperceived notes

Authors:

• Leon Y. Deouell

• D. Deutsch

• Nachum Soroker

• Robert T. Knight

Date: 2008

DOI: 10.3389/neuro.09.015.2007

PubMed: 18958228

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Abstract:

The formation of coherent percepts requires grouping together spatio-temporally disparate sensory inputs. Two major questions arise: (1) is awareness necessary for this process; and (2) can non-conscious elements of the sensory input be grouped into a conscious percept? To address this question, we tested two patients suffering from severe left auditory extinction following right hemisphere damage. In extinction, patients are unaware of the presence of left side stimuli when they are presented simultaneously with right side stimuli. We used the ‘scale illusion’ to test whether extinguished tones on the left can be incorporated into the content of conscious awareness. In the scale illusion, healthy listeners obtain the illusion of distinct melodies, which are the result of grouping of information from both ears into illusory auditory streams. We show that the two patients were susceptible to the scale illusion while being consciously unaware of the stimuli presented on their left. This suggests that awareness is not necessary for auditory grouping and non-conscious elements can be incorporated into a conscious percept.

Spatio-temporal dynamics of neural mechanisms underlying component operations in working memory

Authors:

• Brian T. Miller

• Leon Y. Deouell

• Cathrine Dam

• Robert T. Knight

• Mark D'Esposito

Date: 2008

DOI: 10.1016/j.brainres.2008.01.059

PubMed: 18358455

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Abstract:

Neuroimaging and neurophysiology evidence suggests that component operations in working memory (WM) emerge from the coordinated interaction of posterior perceptual cortices with heteromodal regions in the prefrontal and parietal cortices. Still, little is known about bottom– up and top–down signaling during the formation and retrieval of WM representations. In the current set of experiments, we combine complementary fMRI and EEG measures to obtain high-resolution spatial and temporal measures of neural activity during WM encoding and retrieval processes. Across both experiments, participants performed a face delayed recognition WM task in which the nature of sensory input across stages was held constant. In experiment 1, we utilized a latency-resolved fMRI approach to assess temporal parameters of the BOLD response during stage-specific encoding and retrieval waveforms. Relative to the latency at encoding, the PFC exhibited an earlier peak of fMRI activity at retrieval showing stage-specific differences in the temporal dynamics of PFC engagement across WM operations. In experiment 2, we analyzed the first 200 ms of the ERP response during this WM task providing a more sensitive temporal measure of these differences. Divergence of the ERP pattern during encoding and retrieval began as early as 60 ms post-stimulus. The parallel fMRI and ERP results during memory-guided decisions support a key role of the PFC in top–down biasing of perceptual processing and reveal rapid differences across WM component operations in the presence of identical bottom–up sensory input.

Cerebral responses to change in spatial location of unattended sounds

Authors:

• Leon Y. Deouell

• Aaron S. Heller

• Rafael Malach

• Mark D'Esposito

• Robert T. Knight

Date: 2007

DOI: 10.1016/j.neuron.2007.08.019

PubMed: 17880900

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Abstract:

The neural basis of spatial processing in the auditory cortex has been controversial. Human fMRI studies suggest that a part of the planum temporale (PT) is involved in auditory spatial processing, but it was recently argued that this region is active only when the task requires voluntary spatial localization. If this is the case, then this region cannot harbor an ongoing spa- tial representation of the acoustic environment. In contrast, we show in three fMRI experiments that a region in the human medial PT is sensitive to background auditory spatial changes, even when subjects are not engaged in a spatial lo- calization task, and in fact attend the visual mo- dality. During such times, this area responded to rare location shifts, and even more so when spatial variation increased, consistent with spatially selective adaptation. Thus, acoustic space is represented in the human PT even when sound processing is not required by the ongoing task.

Spatial location is accurately tracked by human auditory sensory memory: evidence from the mismatch negativity

Authors:

• Leon Y. Deouell

• Ariel Parnes

• Natasha Pickard

• Robert T. Knight

Date: 2006

PubMed: 16987229

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Abstract:

The nature of spatial representation in human auditory cortex remains elusive. In particular, although humans can discriminate the locations of sounds as close as 1-10 degrees apart, such resolution has not been shown in auditory cortex of humans or animals. We used the mismatch negativity (MMN) event related brain potential to measure the neural response to spatial change in humans in narrow 10 degree spatial steps. Twelve participants were tested using a dense array EEG setup while watching a silent movie and ignoring the sounds. The MMN was reliably elicited by infrequent changes of spatial location of sounds in free field. The MMN amplitude was linearly related to the degree of spatial change with a resolution of at least 10 degrees. These electrophysiological responses occurred within a window of 100-200 milliseconds from stimulus onset, and were localized to the posterior superior temporal gyrus. We conclude that azimuthal spatial displacement is rapidly, accurately and automatically represented in auditory sensory memory in humans, at the level of the auditory cortex.

High gamma activity in response to deviant auditory stimuli recorded directly from human cortex

Authors:

• Erik Edwards

• Maryam Soltani

• Leon Y. Deouell

• Mitchel S. Berger

• Robert T. Knight

Date: 2005

PubMed: 16093343

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Abstract:

We recorded electrophysiological responses from the left frontal and temporal cortex of awake neurosurgical patients to both repetitive background and rare deviant auditory stimuli. Prominent sensory event-related potentials (ERPs) were recorded from auditory association cortex of the temporal lobe and adjacent regions surrounding the posterior Sylvian fissure. Deviant stimuli generated an additional longer latency mismatch response, maximal at more anterior temporal lobe sites. We found low gamma (30-60 Hz) in auditory association cortex, and we also show the existence of high-frequency oscillations above the traditional gamma range (high gamma, 60-250 Hz). Sensory and mismatch potentials were not reliably observed at frontal recording sites. We suggest that the high gamma oscillations are sensory-induced neocortical ripples, similar in physiological origin to the well-studied ripples of the hippocampus.

ERP measures of multiple attention deficits following prefrontal damage

Authors:

• Leon Y. Deouell

• Robert T. Knight

Date: 2005

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Abstract:

Maintaining a goal-directed behavior requires selectively attending to a subset of the sensory input at the expense of the rest of the input. At the same time, a surveillance mechanism must be in operation, so that deviant or novel events may bring about reorientation of attention and avoidance of potential hazards. Event- related potential (ERP) studies in patients with lateral prefrontal damage, due mainly to stroke, reveal deficits in all of these components of the attentional system. These deficits may explain some pervasive neuropsychological impairment in these patients.

Spatial asymmetries of auditory event-synthesis in humans

Authors:

• Leon Y. Deouell

• Christina M. Karns

• Tamara B. Harrison

• Robert T. Knight

Date: 2003

PubMed: 12531460

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Abstract:

We used the mismatch negativity event-related potential to examine how spatial location and feature variation affect the capacity of the auditory system to automatically respond to pairs of rapid (180 ms apart) acoustic changes within a single tone. When a tone first deviated from a standard tone in source location and then in its duration, we found independent responses to both deviations for right but not left field stimuli. In contrast, when the first deviation was in pitch and the second in duration, only the first deviation elicited a response, regardless of presentation side. These results suggest that information from either side of space is asymmetrically processed even in a free-field, and that the extent of the temporal window of integration is not a fixed property of the auditory system.

Electrophysiological methods and trans-cranial magnetic stimulation in behavioral neurology and neuropsychology

Authors:

• Leon Y. Deouell

• Richard B. Ivry

• Robert T. Knight

Date: 2003

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