Extrastriate Research Articles

New Insights Into the Anatomy, Connectivity and Clinical Implications of the Middle Longitudinal Fasciculus.

The middle longitudinal fascicle (MdLF) is a long, associative white matter tract connecting the superior temporal gyrus (STG) with the parietal and occipital lobe. Previous studies show different cortical terminations, and a possible segmentation pattern of the tract. In this study, we performed a post-mortem white matter dissection of 12 human hemispheres and an in vivo deterministic fiber tracking of 24 subjects acquired from the Human Connectome Project to establish whether a constant organization of fibers exists among the MdLF subcomponents and to acquire anatomical information on each subcomponent. Moreover, two clinical cases of brain tumors impinged on MdLF territories are reported to further discuss the anatomical results in light of previously published data on the functional involvement of this bundle. The main finding is that the MdLF is consistently organized into two layers: an antero-ventral segment (aMdLF) connecting the anterior STG (including temporal pole and planum polare) and the extrastriate lateral occipital cortex, and a posterior-dorsal segment (pMdLF) connecting the posterior STG, anterior transverse temporal gyrus and planum temporale with the superior parietal lobule and lateral occipital cortex. The anatomical connectivity pattern and quantitative differences between the MdLF subcomponents along with the clinical cases reported in this paper support the role of MdLF in high-order functions related to acoustic information. We suggest that pMdLF may contribute to the learning process associated with verbal-auditory stimuli, especially on left side, while aMdLF may play a role in processing/retrieving auditory information already consolidated within the temporal lobe.

Early hearing loss induces plasticity within extra‐striate visual cortex

Supranormal perceptual performance has been observed within the intact senses of early-deaf or blind humans and animals. For cortical areas deprived of their normal sensory input, numerous studies have shown that the lesioned modality is replaced by that of the intact sensory modalities through a process termed crossmodal plasticity. In contrast, little is known about the effects of loss of a particular sensory modality on the cortical representations of the remaining, intact sensory modalities. In the present study, an area of extrastriate visual cortex from early-deaf adult cats was examined for features of dendritic plasticity known to occur after early-deafness. Using light-microscopy of Golgi-stained pyramidal neurons from the posterolateral lateral suprasylvian (PLLS) cortex, dendritic spine density significantly increased (~19%), while spine head size was slightly but significantly decreased (~9%) following early hearing loss. Curiously, these changes were not localized to regions of the visual PLLS known to receive auditory inputs, but instead showed a broad pattern more reflective of the distribution of the area's visual features. Whereas hearing loss results in crossmodal plasticity in auditory cortices, the same peripheral lesion can also induce intramodal plasticity within representations of the intact sensory systems that may also contribute to supranormal performance.

Source dipole analysis reveals a new brain response to visual symmetry

Visual regularity activates a network of brain regions in the extrastriate cortex. Previous EEG studies have found that this response scales parametrically with proportion of symmetry in symmetry + noise displays. The parametric symmetry response happens in many tasks, but it is enhanced during active regularity discrimination. However, the origins and time course of this selective enhancement are unclear. Here we answered remaining questions with new source dipole analysis. As assumed, the parametric symmetry response found at the sensor level was generated by a pair of dipoles in the left and right extrastriate cortex. This bilateral activity was itself enhanced during regularity discrimination. However, we identified a third, and later, symmetry response in the posterior cingulate during regularity discrimination. Unlike the extrastriate response, this previously unknown activation only indexes strong, task relevant regularity signals. This clarifies the neural circuits which mediate the perceptual and cognitive aspects of symmetry discrimination.

Rapid processing of fearful faces relies on the right amygdala: evidence from individuals undergoing unilateral temporal lobectomy

Facial expressions of emotions have been shown to modulate early ERP components, in particular the N170. The underlying anatomical structure producing these early effects are unclear. In this study, we examined the N170 enhancement for fearful expressions in healthy controls as well as epileptic patients after unilateral left or right amygdala resection. We observed a greater N170 for fearful faces in healthy participants as well as in individuals with left amygdala resections. By contrast, the effect was not observed in patients who had undergone surgery in which the right amygdala had been removed. This result demonstrates that the amygdala produces an early brain response to fearful faces. This early response relies specifically on the right amygdala and occurs at around 170 ms. It is likely that such increases are due to a heightened response of the extrastriate cortex that occurs through rapid amygdalofugal projections to the visual areas.

Behavioural performance improvement in visuomotor learning correlates with functional and microstructural brain changes

A better understanding of practice-induced functional and structural changes in our brains can help us design more effective learning environments that provide better outcomes. Although there is growing evidence from human neuroimaging that experience-dependent brain plasticity is expressed in measurable brain changes that are correlated with behavioural performance, the relationship between behavioural performance and structural or functional brain changes, and particularly the time course of these changes, is not well characterised.To understand the link between neuroplastic changes and behavioural performance, 15 healthy participants in this study followed a systematic eye movement training programme for 30 min daily at home, 5 days a week and for 6 consecutive weeks. Behavioural performance statistics and eye tracking data were captured throughout the training period to evaluate learning outcomes. Imaging data (DTI and fMRI) were collected at baseline, after two and six weeks of continuous training, and four weeks after training ended.Participants showed significant improvements in behavioural performance (faster task completion time, lower fixation number and fixation duration).Spatially overlapping reductions in microstructural diffusivity measures (MD, AD and RD) and functional activation increases (BOLD signal) were observed in two main areas: extrastriate visual cortex (V3d) and the frontal part of the cerebellum/Fastigial Oculomotor Region (FOR), which are both involved in visual processing. An increase of functional activity was also recorded in the right frontal eye field.Behavioural, structural and functional changes were correlated. Microstructural change is a better predictor for long-term behavioural change than functional activation is, whereas the latter is superior in predicting instantaneous performance. Structural and functional measures at week 2 of the training programme also predict performance at week 6 and 10, which suggests that imaging data at an early stage of training may be useful in optimising practice environments or rehabilitative training programmes.

Visual experience dependent plasticity in humans.

While sensitive periods in brain development have often been studied by investigating the recovery of visual functions after a congenital phase of visual deprivation in non-human animals, research in humans who had recovered sight after a transient phase of congenital blindness is still scarce. Here, we discuss the hypothesis put forward based on non-human primate work which states that the effects of experience increase downstream the visual processing hierarchy. Recent results from behavioral and neuroscience studies in sight recovery individuals are discussed in the context of research findings from permanently congenitally blind humans as well as from prospective studies in infants and children.

Depth-dependent functional MRI responses to chromatic and achromatic stimuli throughout V1 and V2

In the primate visual system, form (shape, location) and color information are processed in separate but interacting pathways. Recent access to high-resolution neuroimaging has facilitated the exploration of the structure of these pathways at the mesoscopic level in the human visual cortex. We used 7T fMRI to observe selective activation of the primary visual cortex to chromatic versus achromatic stimuli in five participants across two scanning sessions. Achromatic checkerboards with low spatial frequency and high temporal frequency targeted the color-insensitive magnocellular pathway. Chromatic checkerboards with higher spatial frequency and low temporal frequency targeted the color-selective parvocellular pathway. This work resulted in three main findings. First, responses driven by chromatic stimuli had a laminar profile biased towards superficial layers of V1, as compared to responses driven by achromatic stimuli. Second, we found stronger preference for chromatic stimuli in parafoveal V1 compared with peripheral V1. Finally, we found alternating, stimulus-selective bands stemming from the V1 border into V2 and V3. Similar alternating patterns have been previously found in both NHP and human extrastriate cortex. Together, our findings confirm the utility of fMRI for revealing details of mesoscopic neural architecture in human cortex.

A twisted visual field map in the primate dorsomedial cortex predicted by topographic continuity.

Adjacent neurons in visual cortex have overlapping receptive fields within and across area boundaries, an arrangement theorized to minimize wiring cost. This constraint is traditionally thought to create retinotopic maps of opposing field signs (mirror and nonmirror visual field representations) in adjacent areas, a concept that has become central in current attempts to subdivide the extrastriate cortex. We simulated the formation of retinotopic maps using a model that balances constraints imposed by smoothness in the representation within an area and by congruence between areas. As in the primate cortex, this model usually leads to alternating mirror and nonmirror maps. However, we found that it can also produce a more complex type of map, consisting of sectors with opposing field sign within a single area. Using fully quantitative electrode array recordings, we then demonstrate that this type of inhomogeneous map exists in the controversial dorsomedial region of the primate extrastriate cortex.

Arousal-Biased Competition Explains Reduced Distraction by Reward Cues Under Threat

Anxiety has consistently been found to potentiate the attentional processing of physically salient stimuli. However, a recent study demonstrated that experimentally-inducing anxiety reduces attentional capture by previously reward-associated stimuli, suggesting that anxiety does not globally increase distractibility but rather its influence depends on the nature of the eliciting stimulus. To decipher how the processing of threat interacts with the processing of visual reward cues, we probed the neural mechanisms of oculomotor capture by previously reward-associated stimuli with and without the threat of unpredictable electric shock (blockwise manipulation, within-subjects), combining the value-driven attentional capture paradigm with the translational threat of shock paradigm in an fMRI study. Our eye-movement data replicated prior behavioral findings demonstrating an interaction between threat and distractor condition, with oculomotor capture by previously reward-associated distractors being reduced under threat. In our neuroimaging data, significant main effects of threat and distractor condition on stimulus-evoked blood-oxygen-level-dependent (BOLD) activation were evident in the extrastriate visual cortex, frontal eye field, intraparietal sulcus, and caudate tail, collectively referred to as the value-driven attention network (VDAN). In addition, we found an interaction between distractor condition and threat within each region of the VDAN. Surprisingly, the direction of this interaction was opposite that of the behavioral interaction, with the previously reward-associated distractor evoking especially strong neural activation under threat. Our neuroimaging data are consistent with the Arousal-Biased Competition (ABC) model of information processing in the visual system, although our corresponding behavioral results suggest that, at least under certain circumstances involving reward cues, such biased competition can be leveraged to facilitate more efficient ignoring at the level of eye movements. Our findings inform our understanding of value-driven attention, emotion-attention interactions, and mechanisms of signal suppression.

Scotopic Vision Is Selectively Processed in Thick-Type Columns in Human Extrastriate Cortex.

In humans, visual stimuli can be perceived across an enormous range of light levels. Evidence suggests that different neural mechanisms process different subdivisions of this range. For instance, in the retina, stimuli presented at very low (scotopic) light levels activate rod photoreceptors, whereas cone photoreceptors are activated relatively more at higher (photopic) light levels. Similarly, different retinal ganglion cells are activated by scotopic versus photopic stimuli. However, in the brain, it remains unknown whether scotopic versus photopic information is: 1) processed in distinct channels, or 2) neurally merged. Using high-resolution functional magnetic resonance imaging at 7 T, we confirmed the first hypothesis. We first localized thick versus thin-type columns within areas V2, V3, and V4, based on photopic selectivity to motion versus color, respectively. Next, we found that scotopic stimuli selectively activated thick- (compared to thin-) type columns in V2 and V3 (in measurements of both overlap and amplitude) and V4 (based on overlap). Finally, we found stronger resting-state functional connections between scotopically dominated area MT with thick- (compared to thin-) type columns in areas V2, V3, and V4. We conclude that scotopic stimuli are processed in partially segregated parallel streams, emphasizing magnocellular influence, from retina through middle stages of visual cortex.

Spontaneous travelling cortical waves gate perception in behaving primates.

Perceptual sensitivity varies from moment to moment. One potential source of variability is spontaneous fluctuations in cortical activity that can travel as a wave1. Spontaneous traveling waves have been reported during anesthesia2–7, but it is not known whether spontaneous traveling waves play a role during waking perception. Using newly developed analytic techniques to characterize the moment-to-moment dynamics of noisy multielectrode data, we find spontaneous waves of activity in extrastriate visual cortex of awake, behaving marmosets (Callithrix jacchus). In monkeys trained to detect faint visual targets, the timing and position of spontaneous traveling waves, prior to target onset, predict the magnitude of target-evoked activity and the likelihood of target detection. In contrast, spatially disorganized fluctuations of neural activity are much less predictive. These results reveal an important role for spontaneous traveling waves in sensory processing through modulating neural and perceptual sensitivity.

Visual Corticocortical Inputs to Ferret Area 18.

Visual cortical areas in the adult mammalian brain are linked by a network of interareal feedforward and feedback circuits. We investigated the topography of feedback projections to ferret (Mustela putorius furo) area 18 from extrastriate areas 19, 21, and Ssy. Our objective was to characterize the anatomical organization of the extrastriate feedback pool to area 18. We also wished to determine if feedback projections to area 18 share similar features as feedback projections to area 17. We injected the tracer cholera toxin B subunit (CTb) into area 18 of adult ferrets to visualize the distribution and pattern of retrogradely labeled cells in extrastriate cortex. We find several similarities to the feedback projection to area 17: (i) Multiple visual cortical areas provide feedback to area 18: areas 19, 21, Ssy, and weaker inputs from posterior parietal and lateral temporal visual areas. Within each area a greater proportion of feedback projections arises from the infragranular than from the supragranular layers. (ii) The cortical area immediately rostral to area 18 provides the greatest proportion of total cortical feedback, and has the greatest peak density of cells providing feedback to area 18. (iii) The spacing (peak cell density and nearest neighbor distances) of cells in extrastriate cortex providing feedback to areas 17 and 18 are similar. However, peak density of feedback cells to area 18 is comparable in the supra- and infragranular layers, whereas peak density of feedback cells to area 17 is higher in the infragranular layers. Another prominent difference is that dorsal area 18 receives a cortical input that area 17 does not: from ventral cortex representing the upper visual field; this appears to be roughly 25% of the feedback input to area 18. Lastly, area 17 receives a greater proportion of cortical feedback from area 21 than from Ssy, whereas area 18 receives more feedback from Ssy than from area 21. While the organization of feedback projections from extrastriate cortex to areas 17 and 18 is broadly similar, the main difference in input topography might arise due to differences in visual field representations of the two areas.

Pigeon nidopallium caudolaterale, entopallium, and mesopallium ventrolaterale neural responses during categorisation of Monet and Picasso paintings

Pigeons can successfully discriminate between sets of Picasso and Monet paintings. We recorded from three pallial brain areas: the nidopallium caudolaterale (NCL), an analogue of mammalian prefrontal cortex; the entopallium (ENTO), an intermediary visual area similar to primate extrastriate cortex; and the mesopallium ventrolaterale (MVL), a higher-order visual area similar to primate higher-order extrastriate cortex, while pigeons performed an S+/S− Picasso versus Monet discrimination task. In NCL, we found that activity reflected reward-driven categorisation, with a strong left-hemisphere dominance. In ENTO, we found that activity reflected stimulus-driven categorisation, also with a strong left-hemisphere dominance. Finally, in MVL, we found that activity reflected stimulus-driven categorisation, but no hemispheric differences were apparent. We argue that while NCL and ENTO primarily use reward and stimulus information, respectively, to discriminate Picasso and Monet paintings, both areas are also capable of integrating the other type of information during categorisation. We also argue that MVL functions similarly to ENTO in that it uses stimulus information to discriminate paintings, although not in an identical way. The current study adds some preliminary evidence to previous literature which emphasises visual lateralisation during discrimination learning in pigeons.

Electrophysiological priming effects confirm that the extrastriate symmetry network is not gated by luminance polarity.

It is known that the extrastriate cortex is activated by visual symmetry. This activation generates an ERP component called the Sustained Posterior Negativity (SPN). SPN amplitude increases (i.e., becomes more negative) with repeated presentations. We exploited this SPN priming effect to test whether the extrastriate symmetry response is gated by element luminance polarity. On each trial, participants observed three stimuli (patterns of dots) in rapid succession (500ms. with 200ms. gaps). The patterns were either symmetrical or random. The dot elements were either black or white on a grey background. The triplet sequences either showed repeated luminance (black>black>black, or white>white>white) or changing luminance (black>white>black, or white>black>white). As predicted, SPN priming was comparable in repeated and changing luminance conditions. Therefore, symmetry with black elements is not processed independently from symmetry with white elements. Source waveform analysis confirmed that this priming happened within the extrastriate symmetry network. We conclude that the network pools information across luminance polarity channels.

Subcortical neural tracks play an important role in executive function in schizophrenia: An experimental study among patients with schizophrenia and healthy comparisons

The literature has long emphasized the involvement of cortical and subcortical networks in executive function impairments among patients with schizophrenia. However, previous studies have not examined the relative involvement of monocular (mostly subcortical) versus binocular (mostly cortical) neural tracks in patients' EF deficits.Patients with schizophrenia and healthy comparisons were administered a dichotic version of the Stroop task, in which eye-of-origin manipulation was employed to isolate the involvement of monocular (mostly subcortical; thalamic regions) versus binocular (mostly cortical; extrastriate cortex) visual pathways. The eye-of-origin manipulation, in which a color patch (e.g., a green patch) was presented to one eye, and a word (e.g., “RED”) to the other eye, enabled a split of the conflicting information between the two monocular channels. This results in the presentation of conflicting information to the higher cortical regions, but not to the lower subcortical structures. In the Stroop color task, when the monocular neural channels were not exposed to the conflicting information, the differences in task performance between the patients and the HCs significantly increased, and only the patients exhibited larger task conflict. When monocular neural channels were not exposed to the conflicting information, a robust dysfunction of the patients' group was observed. This abnormality might result from impairments in cortical regions or reduced computational power available for solving the conflict. However, additional studies that take into account the resolution of monocular and binocular neural channels are needed to enrich our understanding of the interplay between cortical and subcortical mechanisms in patients' EF deficits.

Electrocorticography Evidence of Tactile Responses in Visual Cortices

There is ongoing debate regarding the extent to which human cortices are specialized for processing a given sensory input versus a given type of information, independently of the sensory source. Many neuroimaging and electrophysiological studies have reported that primary and extrastriate visual cortices respond to tactile and auditory stimulation, in addition to visual inputs, suggesting these cortices are intrinsically multisensory. In particular for tactile responses, few studies have proven neuronal processes in visual cortex in humans. Here, we assessed tactile responses in both low-level and extrastriate visual cortices using electrocorticography recordings in a human participant. Specifically, we observed significant spectral power increases in the high frequency band (30–100 Hz) in response to tactile stimuli, reportedly associated with spiking neuronal activity, in both low-level visual cortex (i.e. V2) and in the anterior part of the lateral occipital–temporal cortex. These sites were both involved in processing tactile information and responsive to visual stimulation. More generally, the present results add to a mounting literature in support of task-sensitive and sensory-independent mechanisms underlying functions like spatial, motion, and self-processing in the brain and extending from higher-level as well as to low-level cortices.

Proposal for human visual pathway in the extrastriate cortex by fiber tracking method using diffusion-weighted MRI

The extrastriate cortex in the human visual cortex is divided into two distinct clusters: the “what-information” processing area and the “where-information” processing area. It is widely accepted that the “what-information” cluster is processed through the ventral stream to the temporal cortex, and the “where-information” cluster through the dorsal stream to the parietal cortex. In human neuroanatomy, fiber bundles for the ventral stream (such as the inferior longitudinal fasciculus) are well defined, whereas fibers for the dorsal stream are poorly understood. In this study, we attempted to trace the dorsal stream fibers using a fiber tracking method using 7.0T diffusion-weighted MRI. We used data from a healthy male subject as well as from an unbiasedly selected nine-subject dataset in the Human Connectome Project. The surface of the visual area, including V1, V2, V3, V4, MT, was determined from the Brainnetome atlas (Fan et al., 2016), which is the connectivity-based parcellation framework of the human brain. The resulting visual pathway indicated that the putative pathway for the classical dorsal stream is unlikely to exist. Instead, we demonstrated that fiber connections exist between the angular gyrus with MT in the visual cortex, and between the angular gyrus and IT in the temporal cortex. Through that, we composed a two-pathway model for where-information processing that passes through the angular gyrus. Finally, we proposed a modified human visual pathway model based on our fiber tracking results in this report. The modified where-information pathway will provide a new aspect for the study of human visual processing.

Negative content enhances stimulus-specific cerebral activity during free viewing of pictures, faces, and words.

Negative visual stimuli have been found to elicit stronger brain activation than do neutral stimuli. Such emotion effects have been shown for pictures, faces, and words alike, but the literature suggests stimulus‐specific differences regarding locus and lateralization of the activity. In the current functional magnetic resonance imaging study, we directly compared brain responses to passively viewed negative and neutral pictures of complex scenes, faces, and words (nouns) in 43 healthy participants (21 males) varying in age and demographic background. Both negative pictures and faces activated the extrastriate visual cortices of both hemispheres more strongly than neutral ones, but effects were larger and extended more dorsally for pictures, whereas negative faces additionally activated the superior temporal sulci. Negative words differentially activated typical higher‐level language processing areas such as the left inferior frontal and angular gyrus. There were small emotion effects in the amygdala for faces and words, which were both lateralized to the left hemisphere. Although pictures elicited overall the strongest amygdala activity, amygdala response to negative pictures was not significantly stronger than to neutral ones. Across stimulus types, emotion effects converged in the left anterior insula. No gender effects were apparent, but age had a small, stimulus‐specific impact on emotion processing. Our study specifies similarities and differences in effects of negative emotional content on the processing of different types of stimuli, indicating that brain response to negative stimuli is specifically enhanced in areas involved in processing of the respective stimulus type in general and converges across stimuli in the left anterior insula.

Rapid eye movement sleep patterns of brain activation and deactivation occur within unique functional networks.

Rapid eye movement (REM) sleep is a paradoxical state where the individual appears asleep while the electroencephalogram pattern resembles that of wakefulness. Regional differences in brain metabolism have been observed during REM sleep compared to wakefulness, but it is not known whether the spatial distribution of metabolic differences corresponds to known functional networks in the brain. Here, we use a combination of techniques to evaluate the networks associated with sites of REM sleep activation and deactivation from previously published positron emission tomography studies. We use seed‐based functional connectivity from healthy adults acquired during quiet rest to show that REM‐activation regions are functionally connected in a network that includes retrosplenial cingulate cortex, parahippocampal gyrus, and extrastriate visual cortices, corresponding to components of the default mode network and visual networks. Regions deactivated during REM sleep localize to right‐lateralized fronto‐parietal and salience networks. A negatively correlated relationship was observed between REM‐activation and deactivation networks. Together, these findings show that regional activation and deactivation patterns of REM sleep tend to occur in distinct functional connectivity networks that are present during wakefulness, providing insights regarding the differential contributions of brain regions to the distinct subjective experiences that occur during REM sleep (dreaming) relative to wakefulness.

Structural and functional footprint of visual snow syndrome.

Patients with visual snow syndrome suffer from a continuous pan-field visual disturbance, additional visual symptoms, tinnitus, and non-perceptional symptoms. The pathophysiology of visual symptoms might involve dysfunctional visual cortex. So far, the extra-visual system has not been investigated. We aimed at identifying structural and functional correlates for visual and non-visual symptoms in visual snow syndrome. Patients were compared to age- and sex-matched controls using 18F-2-fluoro-2-deoxy-d-glucose PET (n = 20 per group) and voxel-based morphometry (n = 17 per group). Guided by the PET results, region of interest analysis was done in voxel-based morphometry to identify structural-functional correspondence. Grey matter volume was assessed globally. Patients had corresponding hypermetabolism and cortical volume increase in the extrastriate visual cortex at the junction of the right lingual and fusiform gyrus. There was hypometabolism in the right superior temporal gyrus and the left inferior parietal lobule. Patients had grey matter volume increases in the temporal and limbic lobes and decrease in the superior temporal gyrus. The corresponding structural and functional alterations emphasize the relevance of the visual association cortex for visual snow syndrome. The broad structural and functional footprint, however, confirms the clinical impression that the disorder extends beyond the visual system.