Alterations in resting-state functional connectivity after brain posterior lesions reflect the functionality of the visual system in hemianopic patients.

Emerging evidence suggests a role of the posterior cortices in regulating alpha oscillatory activity and organizing low-level processing in non-alpha frequency bands. Therefore, posterior brain lesions, which damage the neural circuits of the visual system, might affect functional connectivity patterns of brain rhythms. To test this hypothesis, eyes-closed resting state EEG signal was acquired from patients with hemianopia with left and right posterior lesions, patients without hemianopia with more anterior lesions and healthy controls. Left-lesioned hemianopics showed reduced intrahemispheric connectivity in the range of upper alpha only in the lesioned hemisphere, whereas right-lesioned hemianopics exhibited reduced intrahemispheric alpha connectivity in both hemispheres. In terms of network topology, these impairments were characterized by reduced local functional segregation, with no associated change in global functional integration. This suggests a crucial role of posterior cortices in promoting functional connectivity in the range of alpha. Right-lesioned hemianopics revealed also additional impairments in the theta range, with increased connectivity in this frequency band, characterized by both increased local segregated activity and decreased global integration. This indicates that lesions to right posterior cortices lead to stronger impairments in alpha connectivity and induce additional alterations in local and global low-level processing, suggesting a specialization of the right hemisphere in generating alpha oscillations and in coordinating complex interplays with lower frequency bands. Importantly, hemianopic patient's visual performance in the blind field was linked to alpha functional connectivity, corroborating the notion that alpha oscillatory patterns represent a biomarker of the integrity and the functioning of the underlying visual system.

Spectral Distribution Dynamics across Different Attentional Priority States.

Anticipatory covert spatial attention improves performance on tests of visual detection and discrimination, and shifts are accompanied by decreases and increases of α band power at electroencephalography (EEG) electrodes corresponding to the attended and unattended location, respectively. Although the increase at the unattended location is often interpreted as an active mechanism (e.g., inhibiting processing at the unattended location), most experiments cannot rule out the alternative possibility that it is a secondary consequence of selection elsewhere. To adjudicate between these accounts, we designed a Posner-style visual cueing task in which male and female human participants made orientation judgments of targets appearing at one of four locations: up, down, right, or left. Critically, trials were blocked such that within a block the locations along one meridian alternated in status between attended and unattended, and targets never appeared at the other two, making them irrelevant. Analyses of the concurrently measured EEG signal were conducted on "traditional" narrowband α (8-14 Hz), as well as on two components resulting from the decomposition of this signal: "periodic" α; and the slope of the aperiodic 1/f-like component. Although data from right-left blocks replicated the familiar pattern of lateralized asymmetry in narrowband α power, with neither α signal could we find evidence for any difference in the time course at unattended versus irrelevant locations, an outcome consistent with the secondary-consequence interpretation of attention-related dynamics in the α band. Additionally, 1/f slope was shallower at attended and unattended locations, relative to irrelevant, suggesting a tonic adjustment of physiological state.SIGNIFICANCE STATEMENT Visual spatial attention, the prioritization of one location in the visual field, is critical for guiding behavior in cluttered environments. Although influential theories posit an important role for α band oscillations in the inhibition of processing at unattended locations, we used a novel procedure to find evidence for an alternative interpretation: selection of one location may simply result in a return to physiological baseline at all others. In addition to determining one way that attention does not work (important for future progress in this field), we also discovered novel evidence for one way that it does work: by modifying the tonic physiological state (indexed by an aperiodic component of the electroencephalography (EEG)] at locations where spatial selection is likely to occur.

Hemispheric differences in altered reactivity of brain oscillations at rest after posterior lesions.

A variety of evidence supports the dominance of the right hemisphere in perceptual and visuo-spatial processing. Although growing evidence shows a strong link between alpha oscillations and the functionality of the visual system, asymmetries in alpha oscillatory patterns still need to be investigated. Converging findings indicate that the typical alpha desynchronization occurring in the transition from the eyes-closed to the eyes-open resting state might represent an index of reactivity of the visual system. Thus, investigating hemispheric asymmetries in EEG reactivity at the opening of the eyes in brain-lesioned patients may shed light on the contribution of specific cortical sites and each hemisphere in regulating the oscillatory patterns reflecting the functionality of the visual system. To this aim, EEG signal was recorded during eyes-closed and eyes-open resting state in hemianopic patients with posterior left or right lesions, patients without hemianopia with anterior lesions and healthy controls. Hemianopics with both left and right posterior lesions showed a reduced alpha reactivity at the opening of the eyes, suggesting that posterior cortices have a pivotal role in the functionality of alpha oscillations. However, right-lesioned hemianopics showed a greater dysfunction, demonstrated by a reactivity reduction more distributed over the scalp, compared to left-lesioned hemianopics. Moreover, they also revealed impaired reactivity in the theta range. This favors the hypothesis of a specialized role of the right hemisphere in orchestrating oscillatory patterns, both coordinating widespread alpha oscillatory activity and organizing focal processing in the theta range, to support visual processing at the opening of the eyes.

Alpha oscillations reveal implicit visual processing of motion in hemianopia.

After lesion or deafferentation of the primary visual cortex, hemianopic patients experience loss of conscious vision in their blind field. However, due to the spared colliculo-extrastriate pathway, they might retain the ability to implicitly process motion stimuli through the activation of spared dorsal-extrastriate areas, despite the absence of awareness. To test this hypothesis, Electroencephalogram (EEG) was recorded from a group of hemianopic patients without blindsight (i.e., who performed at chance in different forced-choice tasks), while motion stimuli, static stimuli or no stimuli (i.e., blank condition) were presented either in their intact or in their blind visual field. EEG analyses were performed in the time-frequency domain. The presentation of both motion and static stimuli in the intact field induced synchronization in the theta band and desynchronization both in the alpha and the beta band. In contrast, for stimuli presented in the blind field, significantly greater desynchronization in the alpha range was observed only after the presentation of motion stimuli, compared to the blank condition, over posterior parietal-occipital electrodes in the lesioned hemisphere, at a late time window (500-800 msec). No alpha desynchronization was elicited by static stimuli. These results show that hemianopic patients can process only visual signals relying on the activation of the dorsal pathway (i.e., motion stimuli) in the absence of awareness and suggest different patterns of electrophysiological activity for conscious and unconscious visual processing. Specifically, visual processing in the absence of awareness elicits an activity limited to the alpha range, most likely reflecting a "local" process, occurring within the extrastriate areas and not participating in inter-areal communication. This also suggests a response specificity in this frequency band for implicit visual processing. In contrast, visual awareness evokes changes in different frequency bands, suggesting a "global" process, accomplished by activity in a wide range of frequencies, probably within and across cortical areas.

Aberrant reward prediction error during Pavlovian appetitive learning in alexithymia.

Extensive literature shows that alexithymia, a subclinical trait defined by difficulties in identifying and describing feelings, is characterized by multifaceted impairments in processing emotional stimuli. Nevertheless, its underlying mechanisms remain elusive. Here, we hypothesize that alexithymia may be characterized by an alteration in learning the emotional value of encountered stimuli and test this by assessing differences between individuals with low (LA) and high (HA) levels of alexithymia in the computation of reward prediction errors (RPEs) during Pavlovian appetitive conditioning. As a marker of RPE, the amplitude of the feedback-related negativity (FRN) event-related potential was assessed while participants were presented with two conditioned stimuli (CS) associated with expected or unexpected feedback, indicating delivery of reward or no-reward. No-reward (vs reward) feedback elicited the FRN both in LA and HA. However, unexpected (vs expected) feedback enhanced the FRN in LA but not in HA, indicating impaired computation of RPE in HA. Thus, although HA show preserved sensitivity to rewards, they cannot use this response to update the value of CS that predict them. This impairment may hinder the construction of internal representations of emotional stimuli, leaving individuals with alexithymia unable to effectively recognize, respond and regulate their response to emotional stimuli.

Posterior brain lesions selectively alter alpha oscillatory activity and predict visual performance in hemianopic patients.

Alpha oscillatory frequency and amplitude have been linked to visual processing and to the excitability of the visual cortex at rest. Therefore, posterior brain lesions, which damage the neural circuits of the visual system might induce alterations in the alpha oscillatory activity. To investigate this hypothesis, EEG activity was recorded during eyes-closed resting state in patients with hemianopia with posterior brain lesions, patients without hemianopia with anterior brain lesions and age-matched healthy controls. Patients with posterior lesions revealed a selective slowdown of individual alpha frequency in both the intact and the lesioned hemisphere and a reduction of alpha amplitude in the lesioned hemisphere, resulting in an interhemispheric imbalanced oscillatory alpha activity, while no significant alterations in the alpha range were found in patients with anterior lesions. This suggests a crucial role of posterior cortices in coordinating alpha oscillations in the visual system. Moreover, right posterior lesions had a more severe reduction of individual alpha frequency and altering of the interhemispheric distribution of the alpha amplitude, in line with the notion of the prominence of the right posterior cortices in balancing the interhemispheric functioning. Crucially, the duration of the in individual alpha frequency and the interhemispheric imbalance in alpha amplitude were directly linked to visuo-spatial performance across all participants and to impaired visual detection abilities in hemianopics, therefore supporting a functional role of alpha oscillations in visual processing and suggesting that activity in this frequency range at rest represents a neurophysiological marker reliably reflecting the integrity and the functionality of the visual system in humans.

Pulvinar Lesions Disrupt Fear-Related Implicit Visual Processing in Hemianopic Patients.

The processing of emotional stimuli in the absence of awareness has been widely investigated in patients with lesions to the primary visual pathway since the classical studies on affective blindsight. In addition, recent evidence has shown that in hemianopic patients without blindsight only unseen fearful faces can be implicitly processed, inducing enhanced visual encoding (Cecere et al., 2014) and response facilitation (Bertini et al., 2013, 2017) to stimuli presented in their intact field. This fear-specific facilitation has been suggested to be mediated by activity in the spared visual subcortical pathway, comprising the superior colliculus (SC), the pulvinar and the amygdala. This suggests that the pulvinar might represent a critical relay structure, conveying threat-related visual information through the subcortical visual circuit. To test this hypothesis, hemianopic patients, with or without pulvinar lesions, performed a go/no-go task in which they had to discriminate simple visual stimuli, consisting in Gabor patches, displayed in their intact visual field, during the simultaneous presentation of faces with fearful, happy, and neutral expressions in their blind visual field. In line with previous evidence, hemianopic patients without pulvinar lesions showed response facilitation to stimuli displayed in the intact field, only while concurrent fearful faces were shown in their blind field. In contrast, no facilitatory effect was found in hemianopic patients with lesions of the pulvinar. These findings reveal that pulvinar lesions disrupt the implicit visual processing of fearful stimuli in hemianopic patients, therefore suggesting a pivotal role of this structure in relaying fear-related visual information from the SC to the amygdala.