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.

Split brain: divided perception but undivided consciousness.

In extensive studies with two split-brain patients we replicate the standard finding that stimuli cannot be compared across visual half-fields, indicating that each hemisphere processes information independently of the other. Yet, crucially, we show that the canonical textbook findings that a split-brain patient can only respond to stimuli in the left visual half-field with the left hand, and to stimuli in the right visual half-field with the right hand and verbally, are not universally true. Across a wide variety of tasks, split-brain patients with a complete and radiologically confirmed transection of the corpus callosum showed full awareness of presence, and well above chance-level recognition of location, orientation and identity of stimuli throughout the entire visual field, irrespective of response type (left hand, right hand, or verbally). Crucially, we used confidence ratings to assess conscious awareness. This revealed that also on high confidence trials, indicative of conscious perception, response type did not affect performance. These findings suggest that severing the cortical connections between hemispheres splits visual perception, but does not create two independent conscious perceivers within one brain.

The Uniformity Illusion.

Vision in the fovea, the center of the visual field, is much more accurate and detailed than vision in the periphery. This is not in line with the rich phenomenology of peripheral vision. Here, we investigated a visual illusion that shows that detailed peripheral visual experience is partially based on a reconstruction of reality. Participants fixated on the center of a visual display in which central stimuli differed from peripheral stimuli. Over time, participants perceived that the peripheral stimuli changed to match the central stimuli, so that the display seemed uniform. We showed that a wide range of visual features, including shape, orientation, motion, luminance, pattern, and identity, are susceptible to this uniformity illusion. We argue that the uniformity illusion is the result of a reconstruction of sparse visual information (from the periphery) based on more readily available detailed visual information (from the fovea), which gives rise to a rich, but illusory, experience of peripheral vision.

A social Bayesian brain: How social knowledge can shape visual perception.

A growing body of research suggests that social contextual factors such as desires and goals, affective states and stereotypes can shape early perceptual processes. We suggest that a generative Bayesian approach towards perception provides a powerful theoretical framework to accommodate how such high-level social factors can influence low-level perceptual processes in their earliest stages. We review experimental findings that show how social factors shape the perception and evaluation of people, behaviour, and socially relevant objects or information. Subsequently, we summarize the generative view of perception within the 'Bayesian brain', and show how such a framework can account for the pervasive effects of top-down social knowledge on social cognition. Finally, we sketch the theoretical and experimental implications of social predictive perception, indicating new directions for research on the effects and neurocognitive underpinnings of social cognition.

Expectations accelerate entry of visual stimuli into awareness.

How do expectations influence transitions between unconscious and conscious perceptual processing? According to the influential predictive processing framework, perceptual content is determined by predictive models of the causes of sensory signals. On one interpretation, conscious contents arise when predictive models are verified by matching sensory input (minimizing prediction error). On another, conscious contents arise when surprising events falsify current perceptual predictions. Finally, the cognitive impenetrability account posits that conscious perception is not affected by such higher level factors. To discriminate these positions, we combined predictive cueing with continuous flash suppression (CFS) in which the relative contrast of a target image gradually increases over time. In four experiments we established that expected stimuli enter consciousness faster than neutral or unexpected stimuli. These effects are difficult to account for in terms of response priming, pre-existing stimulus associations, or the attentional mechanisms that cause asynchronous temporal order judgments (of simultaneously presented stimuli). Our results further suggest that top-down expectations play a larger role when bottom-up input is ambiguous, in line with predictive processing accounts of perception. Taken together, our findings support the hypothesis that conscious access depends on verification of perceptual predictions.

Bottom-up and top-down attention are independent.

What is the relationship between top-down and bottom-up attention? Are both types of attention tightly interconnected, or are they independent? We investigated this by testing a large representative sample of the Dutch population on two attentional tasks: a visual search task gauging the efficiency of top-down attention and a singleton capture task gauging bottom-up attention. On both tasks we found typical performance--i.e., participants displayed a significant search slope on the search task and significant slowing caused by the unique, but irrelevant, object on the capture task. Moreover, the high levels of significance we observed indicate that the current set-up provided very high signal to noise ratios, and thus enough power to accurately unveil existing effects. Importantly, in this robust investigation we did not observe any correlation in performance between tasks. The use of Bayesian statistics strongly confirmed that performance on both tasks was uncorrelated. We argue that the current results suggest that there are two attentional systems that operate independently. We hypothesize that this may have implications beyond our understanding of attention. For instance, it may be that attention and consciousness are intertwined differently for top-down attention than for bottom-up attention.

Seeing Ɔ, remembering C: Illusions in short-term memory.

Perception can be shaped by our expectations, which can lead to perceptual illusions. Similarly, long-term memories can be shaped to fit our expectations, which can generate false memories. However, it is generally assumed that short-term memory for percepts formed just 1 or 2 seconds ago accurately represents the percepts as they were at the time of perception. Here 4 experiments consistently show that within this timeframe, participants go from reliably reporting what was there (perceptual inference accurately reflecting the bottom-up input), to erroneously but with high confidence reporting what they expected to be there (memory report strongly influenced by top-down expectations). Together, these experiments show that expectations can reshape perceptual representations over short time scales, leading to what we refer to as short-term memory (STM) illusions. These illusions appeared when participants saw a memory display which contained real and pseudo-letters (i.e. mirrored letters). Within seconds after the memory display disappeared, high confidence memory errors increased substantially. This increase in errors over time indicates that the high confidence errors do not (purely) result from incorrect perceptual encoding of the memory display. Moreover, high confidence errors occurred mainly for pseudo-to-real letter memories, and much less often for real-to-pseudo-letter memories, indicating that visual similarity is not the primary cause of this memory-bias. Instead 'world knowledge' (e.g., which orientation letters usually have) appear to drive these STM illusions. Our findings support a predictive processing view of the formation and maintenance of memory in which all memory stages, including STM, involve integration of bottom-up memory input with top-down predictions, such that prior expectations can shape memory traces.

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 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.

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.

The hard problem makes the easy problems hard - a reply to Doerig et al.

In their paper Doerig et al. argue that we should put the hard problem aside and focus on empirical data to solve the 'easy' problems of consciousness - finding the neural and functional correlates of consciousness. In other words 'shut up and measure'. This has worked well with other thorny issues, such as explaining life, so why not adopt this approach here? We argue that despite the popularity of this view it is not feasible. In order to collect any consciousness data one needs to take an implicit or explicit stance on the hard problem.

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.

The more often you see an object, the easier it becomes to track it.

Is it easier to track objects that you have seen repeatedly? We compared repeated blocks, where identities were the same from trial to trial, to unrepeated blocks, where identities varied. People were better in tracking objects that they saw repeatedly. We tested four hypotheses to explain this repetition benefit. First, perhaps the repeated condition benefits from consistent mapping of identities to target and distractor roles. However, the repetition benefit persisted even when both the repeated and the unrepeated conditions used consistent mapping. Second, repetition might improve the ability to recover targets that have been lost, or swapped with distractors. However, we observed a larger repetition benefit for color-color conjunctions, which do not benefit from such error recovery processes, than for unique features, which do. Furthermore, a repetition benefit was observed even in the absence of distractors. Third, perhaps repetition frees up resources by reducing memory load. However, increasing memory load by masking identities during the motion phase reduced the repetition benefit. The fourth hypothesis is that repetition facilitates identity tracking, which in turn improves location tracking. This hypothesis is consistent with all our results. Thus, our data suggest that identity and location tracking share a common resource.

Distinguishing between parallel and serial accounts of multiple object tracking.

Humans can track multiple moving objects. Is this accomplished by attending to all the objects at the same time or do we attend to each object in turn? We addressed this question using a novel application of the classic simultaneous-sequential paradigm. We considered a display in which objects moved for only part of the time. In one condition, the objects moved sequentially, whereas in the other condition they all moved and paused simultaneously. A parallel model would predict that the targets are tracked independently, so the tracking of one target should not be influenced by the movement of another target. Thus, one would expect equal performance in the two conditions. Conversely, a simple serial account of object tracking would predict that an observer's accuracy should be greater in the sequential condition because in that condition, at any one time, fewer targets are moving and thus need to be attended. In fact, in our experiments we observed performance in the simultaneous condition to be equal to or greater than the performance in the sequential condition. This occurred regardless of the number of targets or how the targets were positioned in the visual field. These results are more directly in line with a parallel account of multiple object tracking.

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.

No Evidence of Narrowly Defined Cognitive Penetrability in Unambiguous Vision.

The classical notion of cognitive impenetrability suggests that perceptual processing is an automatic modular system and not under conscious control. Near consensus is now emerging that this classical notion is untenable. However, as recently pointed out by Firestone and Scholl, this consensus is built on quicksand. In most studies claiming perception is cognitively penetrable, it remains unclear which actual process has been affected (perception, memory, imagery, input selection or judgment). In fact, the only available "proofs" for cognitive penetrability are proxies for perception, such as behavioral responses and neural correlates. We suggest that one can interpret cognitive penetrability in two different ways, a broad sense and a narrow sense. In the broad sense, attention and memory are not considered as "just" pre- and post-perceptual systems but as part of the mechanisms by which top-down processes influence the actual percept. Although many studies have proven top-down influences in this broader sense, it is still debatable whether cognitive penetrability remains tenable in a narrow sense. The narrow sense states that cognitive penetrability only occurs when top-down factors are flexible and cause a clear illusion from a first person perspective. So far, there is no strong evidence from a first person perspective that visual illusions can indeed be driven by high-level flexible factors. One cannot be cognitively trained to see and unsee visual illusions. We argue that this lack of convincing proof for cognitive penetrability in the narrow sense can be explained by the fact that most research focuses on foveal vision only. This type of perception may be too unambiguous for transient high-level factors to control perception. Therefore, illusions in more ambiguous perception, such as peripheral vision, can offer a unique insight into the matter. They produce a clear subjective percept based on unclear, degraded visual input: the optimal basis to study narrowly defined cognitive penetrability.

The Split-Brain Phenomenon Revisited: A Single Conscious Agent with Split Perception.

The split-brain phenomenon is caused by the surgical severing of the corpus callosum, the main route of communication between the cerebral hemispheres. The classical view of this syndrome asserts that conscious unity is abolished. The left hemisphere consciously experiences and functions independently of the right hemisphere. This view is a cornerstone of current consciousness research. In this review, we first discuss the evidence for the classical view. We then propose an alternative, the 'conscious unity, split perception' model. This model asserts that a split brain produces one conscious agent who experiences two parallel, unintegrated streams of information. In addition to changing our view of the split-brain phenomenon, this new model also poses a serious challenge for current dominant theories of consciousness.

Remapping high-capacity, pre-attentive, fragile sensory memory.

Humans typically make several saccades per second. This provides a challenge for the visual system as locations are largely coded in retinotopic (eye-centered) coordinates. Spatial remapping, the updating of retinotopic location coordinates of items in visuospatial memory, is typically assumed to be limited to robust, capacity-limited and attention-demanding working memory (WM). Are pre-attentive, maskable, sensory memory representations (e.g. fragile memory, FM) also remapped? We directly compared trans-saccadic WM (tWM) and trans-saccadic FM (tFM) in a retro-cue change-detection paradigm. Participants memorized oriented rectangles, made a saccade and reported whether they saw a change in a subsequent display. On some trials a retro-cue indicated the to-be-tested item prior to probe onset. This allowed sensory memory items to be included in the memory capacity estimate. The observed retro-cue benefit demonstrates a tFM capacity considerably above tWM. This provides evidence that some, if not all sensory memory was remapped to spatiotopic (world-centered, task-relevant) coordinates. In a second experiment, we show backward masks to be effective in retinotopic as well as spatiotopic coordinates, demonstrating that FM was indeed remapped to world-centered coordinates. Together this provides conclusive evidence that trans-saccadic spatial remapping is not limited to higher-level WM processes but also occurs for sensory memory representations.

Conscious visual memory with minimal attention.

Is conscious visual perception limited to the locations that a person attends? The remarkable phenomenon of change blindness, which shows that people miss nearly all unattended changes in a visual scene, suggests the answer is yes. However, change blindness is found after visual interference (a mask or a new scene), so that subjects have to rely on working memory (WM), which has limited capacity, to detect the change. Before such interference, however, a much larger capacity store, called fragile memory (FM), which is easily overwritten by newly presented visual information, is present. Whether these different stores depend equally on spatial attention is central to the debate on the role of attention in conscious vision. In 2 experiments, we found that minimizing spatial attention almost entirely erases visual WM, as expected. Critically, FM remains largely intact. Moreover, minimally attended FM responses yield accurate metacognition, suggesting that conscious memory persists with limited spatial attention. Together, our findings help resolve the fundamental issue of how attention affects perception: Both visual consciousness and memory can be supported by only minimal attention. (PsycINFO Database Record