Visual integration across fixation: automatic processes are split but conscious processes remain unified in the split-brain.

The classic view holds that when "split-brain" patients are presented with an object in the right visual field, they will correctly identify it verbally and with the right hand. However, when the object is presented in the left visual field, the patient verbally states that he saw nothing but nevertheless identifies it accurately with the left hand. This interaction suggests that perception, recognition and responding are separated in the two isolated hemispheres. However, there is now accumulating evidence that this interaction is not absolute; for instance, split-brain patients are able to detect and localise stimuli anywhere in the visual field verbally and with either hand. In this study we set out to explore this cross-hemifield interaction in more detail with the split-brain patient DDC and carried out two experiments. The aim of these experiments is to unveil the unity of deliberate and automatic processing in the context of visual integration across hemispheres. Experiment 1 suggests that automatic processing is split in this context. In contrast, when the patient is forced to adopt a conscious, deliberate, approach, processing seemed to be unified across visual fields (and thus across hemispheres). First, we looked at the confidence that DDC has in his responses. The experiment involved a simultaneous "same" versus "different" matching task with two shapes presented either within one hemifield or across fixation. The results showed that we replicated the observation that split brain patients cannot match across fixation, but more interesting, that DDC was very confident in the across-fixation condition while performing at chance-level. On the basis of this result, we hypothesised a two-route explanation. In healthy subjects, the visual information from the two hemifields is integrated in an automatic, unconscious fashion via the intact splenium, and this route has been severed in DDC. However, we know from previous experiments that some transfer of information remains possible. We proposed that this second route (perhaps less visual; more symbolic) may become apparent when he is forced to use a deliberate, consciously controlled approach. In an experiment where he is informed, by a second stimulus presented in one hemifield, what to do with the first stimulus that was presented in the same or the opposite hemifield, we showed that there was indeed interhemispheric transfer of information. We suggest that this two-route model may help in clarifying some of the controversial issues in split-brain research.

Mid-range visual functions in relation to higher-order visual functions after stroke.

INTRODUCTION: We aimed to investigate whether associations between deficits in "mid-range" visual functions and deficits in higher-order visual cognitive functions in stroke patients are more in line with a hierarchical, two-pathway model of the visual brain, or with a patchwork model, which assumes a parallel organization with many processing routes and cross-talk. METHODS: A group of 182 ischemic stroke patients was assessed with a new diagnostic set-up for the investigation of a comprehensive range of visuosensory mid-range functions: color, shape, location, orientation, correlated motion, contrast and texture. With logistic regression analyses we investigated the predictive value of these mid-range functions for deficits in visuoconstruction (Copy of the Rey-Complex Figure Test), visual emotion recognition (Ekman 60 Faces Test of the FEEST) and visual memory (computerized Doors-test). RESULTS: Results showed that performance on most mid-range visual tasks could not predict performance on higher-order visual cognitive tasks. Correlations were low to weak. Impaired visuoconstruction and visual memory were only modestly predicted by a worse location perception. Impaired emotion perception was modestly predicted by a worse orientation perception. In addition, double dissociations were found: there were patients with selective deficits in mid-range visual functions without higher-order visual deficits and vice versa. CONCLUSIONS: Our findings are not in line with the hierarchical, two-pathway model. Instead, the findings are more in line with alternative "patchwork" models, arguing for a parallel organization with many processing routes and cross-talk. However, future studies are needed to test these alternative models.

Mid-range visual deficits after stroke: Prevalence and co-occurrence.

Visual deficits are common after stroke and are powerful predictors for the chronic functional outcome. However, while basic visual field and recognition deficits are relatively easy to assess with standardized methods, selective deficits in visual primitives, such as shape or motion, are harder to identify, as they often require a symmetrical bilateral posterior lesion in order to provoke full field deficits. Therefore, we do not know how often they occur. Nevertheless, they can have severe repercussions for daily-life functioning. We aimed to investigate the prevalence and co-occurrence of hemifield "mid-range" visual deficits (i.e. color, shape, location, orientation, correlated motion, contrast, texture and glossiness), using a novel experimental set-up with a gaze-contingent presentation of the stimuli. To this end, a prospective cohort of 220 ischemic (sub)cortical stroke patients and a healthy control group was assessed with this set-up. When comparing performance of patients with controls, the results showed that deficits in motion-perception were most prevalent (26%), followed by color (22%), texture (22%), location (21%), orientation (18%), contrast (14%), shape (14%) and glossiness (13%). 63% of the stroke patients showed one or more mid-range visual deficits. Overlap of deficits was small; they mostly occurred in isolation or co-occurred with only one or two other deficits. To conclude, it was found that deficits in "mid-range" visual functions were very prevalent. These deficits are likely to affect the chronic post-stroke condition. Since we found no strong patterns of co-occurrences, we suggest that an assessment of deficits at this level of visual processing requires screening the full range of visual functions.

Accelerated Long-Term Forgetting: Prolonged Delayed Recognition as Sensitive Measurement for Different Profiles of Long-Term Memory and Metacognitive Confidence in Stroke Patients.

OBJECTIVE: Deficits in episodic memory are frequently reported after ischemic stroke. In standard clinical care, episodic memory is assessed after a 20-30 min delay, with abnormal memory decay over this period being characterized as rapid forgetting (RF). Previous studies have shown abnormal forgetting over a prolonged interval (days to weeks) despite normal acquisition, referred to as accelerated long-term forgetting (ALF). METHOD: We examined whether ALF is present in stroke patients (N = 91) using immediate testing (T1), testing after a short delay (20-30 min, T2), and testing after a prolonged delay (one week, T3). Based on performance compared to matched controls (N = 85), patients were divided into (1) patients without forgetting, (2) patients with RF between T1 and T2, and (3) patients with ALF at T3. Furthermore, confidence ratings were assessed. RESULTS: ALF was present in a moderate amount of stroke patients (17%), but ALF was even more prevalent in our stroke sample than RF after a 20-30 min delay (which was found in only 13% of our patients). Patients reported a lower confidence for their responses, independent of their actual performance. CONCLUSIONS: Adding a one-week delayed measurement may potentially assist in identifying patients with memory decrements that may otherwise go undetected.

Visual features drive the category-specific impairments on categorization tasks in a patient with object agnosia.

Object and scene recognition both require mapping of incoming sensory information to existing conceptual knowledge about the world. A notable finding in brain-damaged patients is that they may show differentially impaired performance for specific categories, such as for "living exemplars". While numerous patients with category-specific impairments have been reported, the explanations for these deficits remain controversial. In the current study, we investigate the ability of a brain injured patient with a well-established category-specific impairment of semantic memory to perform two categorization experiments: 'natural' vs. 'manmade' scenes (experiment 1) and objects (experiment 2). Our findings show that the pattern of categorical impairment does not respect the natural versus manmade distinction. This suggests that the impairments may be better explained by differences in visual features, rather than by category membership. Using Deep Convolutional Neural Networks (DCNNs) as 'artificial animal models' we further explored this idea. Results indicated that DCNNs with 'lesions' in higher order layers showed similar response patterns, with decreased relative performance for manmade scenes (experiment 1) and natural objects (experiment 2), even though they have no semantic category knowledge, apart from a mapping between pictures and labels. Collectively, these results suggest that the direction of category-effects to a large extent depends, at least in MS' case, on the degree of perceptual differentiation called for, and not semantic knowledge.

On the necessity of recurrent processing during object recognition: it depends on the need for scene segmentation.

While feed-forward activity may suffice for recognizing objects in isolation, additional visual operations that aid object recognition might be needed for real-world scenes. One such additional operation is figure-ground segmentation; extracting the relevant features and locations of the target object while ignoring irrelevant features. In this study of 60 human participants (female and male), we show objects on backgrounds of increasing complexity to investigate whether recurrent computations are increasingly important for segmenting objects from more complex backgrounds. Three lines of evidence show that recurrent processing is critical for recognition of objects embedded in complex scenes. First, behavioral results indicated a greater reduction in performance after masking objects presented on more complex backgrounds; with the degree of impairment increasing with increasing background complexity. Second, electroencephalography (EEG) measurements showed clear differences in the evoked response potentials (ERPs) between conditions around time points beyond feed-forward activity and exploratory object decoding analyses based on the EEG signal indicated later decoding onsets for objects embedded in more complex backgrounds. Third, Deep Convolutional Neural Network performance confirmed this interpretation; feed-forward and less deep networks showed a higher degree of impairment in recognition for objects in complex backgrounds compared to recurrent and deeper networks. Together, these results support the notion that recurrent computations drive figure-ground segmentation of objects in complex scenes.SIGNIFICANCE STATEMENTThe incredible speed of object recognition suggests that it relies purely on a fast feed-forward build-up of perceptual activity. However, this view is contradicted by studies showing that disruption of recurrent processing leads to decreased object recognition performance. Here we resolve this issue by showing that how object recognition is resolved, and whether recurrent processing is crucial, depends on the context in which it is presented. For objects presented in isolation or in 'simple' environments, feed-forward activity could be sufficient for successful object recognition. However, when the environment is more complex, additional processing seems necessary to select the elements that belong to the object, and by that segregate them from the background.

Are visual working memory and episodic memory distinct processes? Insight from stroke patients by lesion-symptom mapping.

Working memory and episodic memory are two different processes, although the nature of their interrelationship is debated. As these processes are predominantly studied in isolation, it is unclear whether they crucially rely on different neural substrates. To obtain more insight in this, 81 adults with sub-acute ischemic stroke and 29 elderly controls were assessed on a visual working memory task, followed by a surprise subsequent memory test for the same stimuli. Multivariate, atlas- and track-based lesion-symptom mapping (LSM) analyses were performed to identify anatomical correlates of visual memory. Behavioral results gave moderate evidence for independence between discriminability in working memory and subsequent memory, and strong evidence for a correlation in response bias on the two tasks in stroke patients. LSM analyses suggested there might be independent regions associated with working memory and episodic memory. Lesions in the right arcuate fasciculus were more strongly associated with discriminability in working memory than in subsequent memory, while lesions in the frontal operculum in the right hemisphere were more strongly associated with criterion setting in subsequent memory. These findings support the view that some processes involved in working memory and episodic memory rely on separate mechanisms, while acknowledging that there might also be shared processes.

The neural underpinnings of facial emotion recognition in ischemic stroke patients.

Deficits in facial emotion recognition occur frequently after stroke, with adverse social and behavioural consequences. The aim of this study was to investigate the neural underpinnings of the recognition of emotional expressions, in particular of the distinct basic emotions (anger, disgust, fear, happiness, sadness and surprise). A group of 110 ischaemic stroke patients with lesions in (sub)cortical areas of the cerebrum was included. Emotion recognition was assessed with the Ekman 60 Faces Test of the FEEST. Patient data were compared to data of 162 matched healthy controls (HC's). For the patients, whole brain voxel-based lesion-symptom mapping (VLSM) on 3-Tesla MRI images was performed. Results showed that patients performed significantly worse than HC's on both overall recognition of emotions, and specifically of disgust, fear, sadness and surprise. VLSM showed significant lesion-symptom associations for FEEST total in the right fronto-temporal region. Additionally, VLSM for the distinct emotions showed, apart from overlapping brain regions (insula, putamen and Rolandic operculum), also regions related to specific emotions. These were: middle and superior temporal gyrus (anger); caudate nucleus (disgust); superior corona radiate white matter tract, superior longitudinal fasciculus and middle frontal gyrus (happiness) and inferior frontal gyrus (sadness). Our findings help in understanding how lesions in specific brain regions can selectively affect the recognition of the basic emotions.

Singularity and consciousness: A neuropsychological contribution.

In common sense experience based on introspection, consciousness is singular. There is only one 'me' and that is the one that is conscious. This means that 'singularity' is a defining aspect of 'consciousness'. However, the three main theories of consciousness, Integrated Information, Global Workspace and Recurrent Processing theory, are generally not very clear on this issue. These theories have traditionally relied heavily on neuropsychological observations and have interpreted various disorders, such as anosognosia, neglect and split-brain as impairments in conscious awareness without any reference to 'the singularity'. In this review, we will re-examine the theoretical implications of these impairments in conscious awareness and propose a new way how to conceptualize consciousness of singularity. We will argue that the subjective feeling of singularity can coexist with several disunified conscious experiences. Singularity awareness may only come into existence due to environmental response constraints. That is, perceptual, language, memory, attentional and motor processes may largely proceed unintegrated in parallel, whereas a sense of unity only arises when organisms need to respond coherently constrained by the affordances of the environment. Next, we examine from this perspective psychiatric disorders and psycho-active drugs. Finally, we present a first attempt to test this hypothesis with a resting state imaging experiment in a split-brain patient. The results suggest that there is substantial coherence of activation across the two hemispheres. These data show that a complete lesioning of the corpus callosum does not, in general, alter the resting state networks of the brain. Thus, we propose that we have separate systems in the brain that generate distributed conscious. The sense of singularity, the experience of a 'Me-ness', emerges in the interaction between the world and response-planning systems, and this leads to coherent activation in the different functional networks across the cortex.

Consequence of stroke for feature recall and binding in visual working memory.

Visual memory for objects involves the integration, or binding, of individual features into a coherent representation. We used a novel approach to assess feature binding, using a delayed-reproduction task in combination with computational modeling and lesion analysis. We assessed stroke patients and neurotypical controls on a visual working memory task in which spatial arrays of colored disks were presented. After a brief delay, participants either had to report the color of one disk cued by its location or the location of one disk cued by its color. Our results demonstrate that, in the controls, report imprecision and swap errors (non-target reports) can be explained by a single source of variability. Stroke patients showed an overall decrease in memory precision for both color and location, with only limited evidence for deviations from the predicted relationship between report precision and swap errors. These deviations were primarily deficits in reporting items rather than selecting items based on the cue. Atlas-based lesion-symptom mapping showed that selection and reporting deficits, precision in reporting color, and precision in reporting location were associated with different lesion profiles. Deficits in binding are associated with lesions in the left somatosensory cortex, deficits in the precision of reporting color with bilateral fronto-parietal regions, and no anatomical substrates were identified for precision in reporting location. Our results converge with previous reports that working memory representations are widely distributed in the brain and can be found across sensory, parietal, temporal, and prefrontal cortices. Stroke patients demonstrate mostly subtle impairments in visual working memory, perhaps because representations from different areas in the brain can partly compensate for impaired encoding in lesioned areas. These findings contribute to understanding of the relation between memorizing features and their bound representations.

Emotion Recognition and Traffic-Related Risk-Taking Behavior in Patients with Neurodegenerative Diseases.

OBJECTIVES: Neurodegenerative diseases (NDDs), such as Alzheimer's disease, frontotemporal dementia, dementia with Lewy bodies, and Huntington's disease, inevitably lead to impairments in higher-order cognitive functions, including the perception of emotional cues and decision-making behavior. Such impairments are likely to cause risky daily life behavior, for instance, in traffic. Impaired recognition of emotional expressions, such as fear, is considered a marker of impaired experience of emotions. Lower fear experience can, in turn, be related to risk-taking behavior. The aim of our study was to investigate whether impaired emotion recognition in patients with NDD is indeed related to unsafe decision-making in risky everyday life situations, which has not been investigated yet. METHODS: Fifty-one patients with an NDD were included. Emotion recognition was measured with the Facial Expressions of Emotions: Stimuli and Test (FEEST). Risk-taking behavior was measured with driving simulator scenarios and the Action Selection Test (AST). Data from matched healthy controls were used: FEEST (n = 182), AST (n = 36), and driving simulator (n = 18). RESULTS: Compared to healthy controls, patients showed significantly worse emotion recognition, particularly of anger, disgust, fear, and sadness. Furthermore, patients took significantly more risks in the driving simulator rides and the AST. Only poor recognition of fear was related to a higher amount of risky decisions in situations involving a direct danger. CONCLUSIONS: To determine whether patients with an NDD are still fit to drive, it is crucial to assess their ability to make safe decisions. Measuring emotion recognition may be a valuable contribution to this judgment.

Post-Stroke Working Memory Dysfunction: A Meta-Analysis and Systematic Review.

This review investigates the severity and nature of post-stroke working memory deficits with reference to the multi-component model of working memory. We conducted a systematic search in PubMed up to March 2019 with search terms for stroke and memory. Studies on adult stroke patients, that included a control group, and assessed working memory function, were selected. Effect sizes (Hedges' g) were extracted from 50 studies (in total 3,084 stroke patients) based on the sample size, mean and standard deviation of patients and controls. Performance of stroke patients was compared to healthy controls on low-load (i.e. capacity) and high-load (executively demanding) working memory tasks, grouped by modality (verbal, non-verbal). A separate analysis compared patients in the sub-acute and the chronic stage. Longitudinal studies and effects of lesion location were systematically reviewed. Stroke patients demonstrated significant deficits in working memory with a moderate effect size for both low-load (Hedges' g = -.58 [-.82 to -.43]) and high-load (Hedges' g = -.59 [-.73 to -.45]) tasks. The effect sizes were comparable for verbal and non-verbal material. Systematically reviewing the literature showed that working memory deficits remain prominent in the chronic stage of stroke. Lesions in a widespread fronto-parietal network are associated with working memory deficits. Stroke patients show decrements of moderate magnitude in all subsystems of working memory. This review clearly demonstrates the global nature of the impairment in working memory post-stroke.

Depth in convolutional neural networks solves scene segmentation.

Feed-forward deep convolutional neural networks (DCNNs) are, under specific conditions, matching and even surpassing human performance in object recognition in natural scenes. This performance suggests that the analysis of a loose collection of image features could support the recognition of natural object categories, without dedicated systems to solve specific visual subtasks. Research in humans however suggests that while feedforward activity may suffice for sparse scenes with isolated objects, additional visual operations ('routines') that aid the recognition process (e.g. segmentation or grouping) are needed for more complex scenes. Linking human visual processing to performance of DCNNs with increasing depth, we here explored if, how, and when object information is differentiated from the backgrounds they appear on. To this end, we controlled the information in both objects and backgrounds, as well as the relationship between them by adding noise, manipulating background congruence and systematically occluding parts of the image. Results indicate that with an increase in network depth, there is an increase in the distinction between object- and background information. For more shallow networks, results indicated a benefit of training on segmented objects. Overall, these results indicate that, de facto, scene segmentation can be performed by a network of sufficient depth. We conclude that the human brain could perform scene segmentation in the context of object identification without an explicit mechanism, by selecting or "binding" features that belong to the object and ignoring other features, in a manner similar to a very deep convolutional neural network.

Somatosensation in the Brain: A Theoretical Re-evaluation and a New Model.

The somatosensory system is important for many functions, such as tactile recognition, the perception of our body, and motor actions. We present a comprehensive review of the human and animal literature on somatosensory processing over the past 10 years and evaluate how well existing models can accommodate the new observations. Based on these observations and a survey of the brain structures involved in somatosensation, we suggest that a new model is needed that describes multiple networks involved in separate subfunctions. These networks are highly interconnected and often multimodal in nature. The model includes basic somatosensory processing and five higher-order networks involved in haptic object recognition and memory, body perception, body ownership, affective processing, and action.

Split-Brain: What We Know Now and Why This is Important for Understanding Consciousness.

Recently, the discussion regarding the consequences of cutting the corpus callosum ("split-brain") has regained momentum (Corballis, Corballis, Berlucchi, & Marzi, Brain, 141(6), e46, 2018; Pinto et al., Brain, 140(5), 1231-1237, 2017a; Pinto, Lamme, & de Haan, Brain, 140(11), e68, 2017; Volz & Gazzaniga, Brain, 140(7), 2051-2060, 2017; Volz, Hillyard, Miller, & Gazzaniga, Brain, 141(3), e15, 2018). This collective review paper aims to summarize the empirical common ground, to delineate the different interpretations, and to identify the remaining questions. In short, callosotomy leads to a broad breakdown of functional integration ranging from perception to attention. However, the breakdown is not absolute as several processes, such as action control, seem to remain unified. Disagreement exists about the responsible mechanisms for this remaining unity. The main issue concerns the first-person perspective of a split-brain patient. Does a split-brain harbor a split consciousness or is consciousness unified? The current consensus is that the body of evidence is insufficient to answer this question, and different suggestions are made with respect to how future studies might address this paucity. In addition, it is suggested that the answers might not be a simple yes or no but that intermediate conceptualizations need to be considered.

Impairments in Emotion Recognition and Risk-Taking Behavior After Isolated, Cerebellar Stroke.

An increasing amount of research has shown a cerebellar involvement in higher order cognitive functions, including emotional processing and decision-making. However, it has not been investigated whether impairments in facial emotion recognition, which could be a marker of impaired emotional experiences, are related to risky decision-making in these patients. Therefore, we aimed to investigate facial emotion recognition and risky decision-making in these patients as well as to investigate a relationship between these constructs. Thirteen patients with a discrete, isolated, cerebellar lesion as a consequence of a stroke were included in the study. Emotion recognition was assessed with the Facial Expressions of Emotions-Stimuli and Test (FEEST). Risk-taking behavior was assessed with the Action Selection Test (AST). Furthermore, 106 matched healthy controls performed the FEEST and 20 matched healthy controls performed the AST. Compared with healthy controls, patients were significantly worse in the recognition of emotional expressions and they took significantly more risks. In addition, a worse ability to recognize fearful facial expressions was strongly related to an increase in risky decisions in the AST. Therefore, we suggest that tests of emotion recognition should be incorporated into the neuropsychological assessment after cerebellar stroke to boost detection and treatment of these impairments in these patients.

Unified tactile detection and localisation in split-brain patients.

In 'split-brain' patients, the corpus callosum has been surgically severed to alleviate medically intractable, severe epilepsy. The classic claim is that after removal of the corpus callosum an object presented in the right visual field will be identified correctly verbally and with the right hand but not with the left hand. When the object is presented in the left visual field the patient verbally states that he saw nothing but nevertheless identifies it accurately with the left hand. This interaction suggests that perception, recognition and responding are separated in the two isolated hemispheres. However, there is now accumulating evidence that this interaction is not absolute. Recently, we (Pinto et al., 2017) showed that accurate detection and location of stimuli anywhere in the visual field could be performed with both hands. In this study, we explored detection and localisation of tactile stimulation on the body. In line with our previous results, we observed that split-brain patients can signal detection and localisation with either hand anywhere on the body (be it the arm or the leg) but they remain unable to match positions touched on both arms or legs simultaneously. These results add to the evidence suggesting that the effects of removal of the corpus callosum may be less severe than sometimes claimed.

Plasticity versus chronicity: Stable performance on category fluency 40 years post-onset.

What is the long-term trajectory of semantic memory deficits in patients who have suffered structural brain damage? Memory is, per definition, a changing faculty. The traditional view is that after an initial recovery period, the mature human brain has little capacity to repair or reorganize. More recently, it has been suggested that the central nervous system may be more plastic with the ability to change in neural structure, connectivity, and function. The latter observations are, however, largely based on normal learning in healthy subjects. Here, we report a patient who suffered bilateral ventro-medial damage after presumed herpes encephalitis in 1971. He was seen regularly in the eighties, and we recently had the opportunity to re-assess his semantic memory deficits. On semantic category fluency, he showed a very clear category-specific deficit performing better that control data on non-living categories and significantly worse on living items. Recent testing showed that his impairments have remained unchanged for more than 40 years. We suggest cautiousness when extrapolating the concept of brain plasticity, as observed during normal learning, to plasticity in the context of structural brain damage.