From Cognitive to Neural Theories of Consciousness Efraïm Salari ABSTRACT There seems to be a tendency within the education of neuroscience and cognitive psychology to give little attention to what is not well understood. One subject that tends to get left out of education for this reason is the concept of consciousness. There have been, however, a number of studies that tried to address the question of how brain processes are related to consciousness. The main question of this thesis therefore is: what are, according to contemporary views, the cognitive and neuronal processes that are needed for, or correlated with conscious experience? Through high level cognitive theories a scientific understanding of consciousness was arrived. Based on this, lower level neural theories where developed, focusing on how neurons interact with each other, and which neurons/areas interact with each other during conscious and unconscious processing. Three important theories will be discussed to illustrate this. First, the global workspace hypothesis will be discussed which is a cognitive theory. It was developed to, for the first time after behaviourism, address questions on consciousness scientifically. It states that different forms of information are usually autonomously and unconsciously processed in different brain areas. Consciousness is however derived when brain areas share their output to a global ‘workspace’ onto which different areas are connected such as for instance memory, language and association areas. Second, the 40 Hz theory is discussed which was based on the same notions as the global workspace theory but which takes a neural point of view, and focuses on how remote brain areas communicate to form a unified conscious percept. It states that neurons in different brain areas that are involved in conscious percept, fire in synchronized oscillations in order to bind their information processing. Finally, the recurrent processing theory is considered which is also a neural theory of consciousness that is to some extend based on the notions of the global workspace hypotheses but focuses on which areas communicate during conscious percept and more specifically on the course of this communication. This theory deviates from previous theories of consciousness in its’ notion of how to measure consciousness. Most theories use an intuitive concept of consciousness, i.e. that people can report on what they are conscious of, and so reportability can be used as a measure of consciousness. However, the recurrent processing theory states that reportability might not always be sufficient for measuring consciousness (for instance in split brain and locked in patients) and states that a fully neural description of consciousness is needed. According to this theory, consciousness is arrived when recurrent processing between higher and lower order areas takes place, regardless of peoples reports. Keywords: Consciousness, global workspace, 40 Hz, recurrent processing “Conscious experience is at once the most familiar thing in the world and the most mysterious.” David J. Chalmers (1995)- The Conscious Mind: In Search of a Fundamental Theory INTRODUCTION The fields of neuroscience and cognitive psychology are concerned with one of the most fascinating questions in contemporary science: how does the brain work? The knowledge that is gained so far about the functioning of the brain is incredible. However, there seems to be a tendency within the education of these fields to give little attention to what is not well understood. One subject that tends to get left out of education is the concept of consciousness. In 1990 Francis Crick and Christof Koch already wrote: “IT IS REMARKABLE that most of the work in both cognitive science and the neurosciences makes no reference to consciousness (or 'awareness'), especially as many would regard consciousness as the major puzzle confronting the neural view of the mind and indeed at the present time it appears deeply mysterious to many people. This attitude is partly a legacy of behaviorism and partly because most workers in these areas cannot see any useful way of approaching the problem.” (Crick & Koch, 1990, p263) Although in the education of neuroscience the neuronal correlates of consciousness (NCC) are heavily overlooked, there have been a number of studies that tried to address the question of how brain processes are related to consciousness. Therefore the focus of this thesis will be on the following question: What are, according to contemporary views, the cognitive and neuronal processes that are needed for, or correlated with, conscious experience? One can debate however whether there is a relationship between the processes that are going on in the brain and consciousness in the first place. René Descartes, for example, thought that there is a duality between mind and body; that the mind originates from something non-physical. However, this not a very popular view anymore (but see Chalmers, 1995 for a contemporary dualistic approach) and in this thesis I will assume that a correlation between the physical state of the brain and conscious experiences exists. Indeed, a lot of evidence indicates that specific neuronal processes are associated with whether or not people are consciously aware of for instance a presented stimulus (Baars, 2005; Dehaene & Naccache, 2001; Crick & Koch, 1990, Lamme, 2006) What is the problem? For a lot of brain modalities, such as for instance motor processing or vision, there is a good understanding of both the cognitive (high level) and the underlying neural (low level) mechanisms of these functions. It is known for example which brain regions are responsible for these functions and what neural activity is involved. For instance, when a particular motor action is executed, neural activity originates in the primary motor areas, which then terminates in the muscles. In vision, light hits the retina, which activates ganglion cells from which neural activity travels towards V1, followed by higher order areas, integrating the information into a complete image of the visual field. However, which neuronal responses are accompanied by consciousness still remains to be a subject of debate and a lot of questions remain to be answered. For example, where along the visual stream does consciousness of a scene arise? There is no clear end point that has to be reached for consciousness to arise (such as a muscle in motion). Do specific cells, areas or networks need to be active for conscious awareness? Are there specific forms of cell communication involved? Why are we aware of some things and not of others? The reason why a lot of these questions have not been answered is due mostly to a lack of consensus on how to define consciousness and how to measure it. Because of that, early psychological research focused only on behavioural observations, without concerning about the processes in the mind that led to them. This movement, called behaviourism, did not address any questions on consciousness. However, with new imaging techniques, such as fMRI, the brain’s processes could be studied while functioning. This leaded to new insights, and cognitive models of consciousness where developed to address the problem of consciousness scientifically. These models where concerned with: what brain functions (i.e. attention, memory, vision) are involved in consciousness, and what brain areas are responsible for them. Subsequently, based on these higher level cognitive theories, lower level neural explanations where developed, focusing on what neural mechanisms are involved during conscious percept. To illustrate this, three important theories will be discussed. The first theory that is discussed is the global workspace hypothesis, which is a cognitive theory, describing the functions that are involved in consciousness and the brain areas related to that. It states that different forms of information are usually autonomously and unconsciously processed in different brain areas. Consciousness is however derived when brain areas share their output to a global ‘workspace’ onto which multiple areas are connected such as for instance memory, language and association areas. This theory is discussed here because it was one of the first theories that tried to break with behaviourism and start a scientific search for understanding consciousness. Also, it has been very influential for later theories, among which theories that focused on the lower level, neural mechanisms involved in consciousness. Two of these neural theories are described next: the 40 Hz theory and the recurrent processing theory. The second theory that is discussed is the 40 Hz theory, which focuses on how neurons involved in consciousness fire. It states that neurons in different brain areas that are involved in conscious percept, fire in synchronized oscillations in order to bind their information processing. This theory is discussed for two reasons. The first reason is that it was a reaction to the cognitive theories of consciousness, to come for the first time with a neural theory of consciousness, describing what neuronal mechanisms are involved during conscious precept. The second reason is that this theory is influenced by the global workspace theory and therefore comparisons can be made between these theories. Finally, the recurrent processing theory is considered which is also a neural theory of consciousness and focuses on which areas communicate during consciousness and more specifically on the course of this communication. This theory deviates from previous theories of consciousness. Most theories use an intuitive concept of consciousness, i.e. that people can report on what they are conscious of, and so reportability can be used as a measure of consciousness. However, the recurrent processing theory states that reportability might not always be sufficient for measuring consciousness (for instance in split brain and locked in patients) and states that a fully neural description of consciousness is needed. According to this theory, consciousness is arrived when recurrent processing between higher and lower order areas takes place, regardless of peoples reports. This theory is discussed for two reasons. The first reason is that it’s fully neural perspective on consciousness deviates from other theories in an interesting and unexpected way, which might be influential to our understanding of consciousness. The second reason is that it is a rather new theory, which is also influenced by the global workspace theory. All theories are built on the notion that for people to be able to report on their conscious content, different brain areas need to communicating with each other on a global scale, as suggested by the global workspace theory. The 40 Hz theory however, focuses on how different areas communicate during conscious content processing whereas the recurrent processing theory focuses on which areas communicate and in what way. What is consciousness? So far the term consciousness has been used without an explanation of what this actually means. Consciousness can be described in multiple ways. Baars (2005) for instance describes it in the following way: “It is broadly true that what we are conscious of, we can report with accuracy. Conscious brain events are therefore assessed by way of reportability.” Baars (2005, p.45). Lamme (2006) on the contrary argues that to use the term consciousness, it should not only be described in terms of other function. Reportability for instance relies on memory and language. Paralyzed patients however that cannot speak and move, still show forms of consciousness (as suggested by fMRI experiments; see for instance Owen et al., 2006) but have no way of expressing themselves. Lamme argues that consciousness should therefore also be described purely in terms of neural activity. Meaning that if specific neural activity is present, this should be considered a sign of consciousness, whether or not people can report awareness. Others, such as Crick & Koch (1990) have restrained themselves from a precise description of consciousness but use both reportability measures as well as neural evidence (for instance when using animal models). Because consciousness can be described in multiple ways, some have argued that there are multiple neural correlates of consciousness. Ned Block (2005) for instance argues that consciousness can be separated into access- and phenomenalconsciousness. Access-consciousness is the kind of consciousness that is involved when one can actively think about something (and so usually report on it). This arises when information is not only locally processed (for instance in visual areas only) but is also available to multiple brain areas such as those important for memory, evaluation and language. Access-consciousness, according to Block (2005), can be explained by the global-workspace hypothesis (Baars, 2005, Dehaene & Naccache, 2001). Phenomenal-consciousness is that what separates experiences from each other but without further evaluation of these experiences. In a visual scene for instance, there are different colours that one experiences, however one does not necessarily have to actively think about all the colours that are seen. According to Block (2005) this form of consciousness can be explained by the recurrent-processing theory (Lamme, 2006). Because different authors have used different definitions of consciousness, it is hard to give one description of the term ‘consciousness’. There is simply no consensus as to what consciousness conveys. However, one useful way of differentiating consciousness from unconsciousness is that the former comes with an ‘experience’ of some kind whereas the later does not (Chalmers, 1995). This ‘experience’ can be assessed in multiple ways, i.e. by reportability (Baars, 2005) or neural activity (see Lamme, 2006) and in this paper consciousness is assessed as whether or not the presentation of external stimuli are accompanied by an experience. Possible other forms of consciousness, such as thoughts that are not directly related to something in the environments are not discussed in this thesis to restrict the scope of this paper. An important and useful implication of describing consciousness as the presence of experience, is that some animals by definition need to have consciousness (although possible in a different form than humans). Monkeys for instance can be trained to report whether or not they saw a stimulus. Therefore animals can be used to investigate some forms of consciousness. Also, when using neural measures of consciousness, other animals such as cats can be used to study the neural correlates of consciousness (see for instance Crick & Koch, 1990). Because consciousness can conveys many things, this thesis focuses on theories that try to explain consciousness only in the form of experiencing and being aware of external input (stimuli perception). The terms consciousness, awareness and conscious percept are used interchangeably (but see Chalmers, 1995, for a different notion). The purpose of consciousness: Related to the question of what consciousness is, is the question of what the purpose of consciousness might be. Some researchers have addressed this question. Lamme for instance (2006) has argued that consciousness is necessary for learning and memory. This is because the neural activity that according to Lamme correlates with consciousness (recurrent processing) involves an optimal situation for brain plasticity. Dehaene et al. (2001) have also suggested that consciousness is necessary for particular mental functions. Like Lamme, they argue that consciousness is necessary for information maintenance. That is, studies have shown that only information that is consciously attended to can last longer than iconic memory. Studies for instance show that the judgement of a target word is influenced by a previously presented masked prime word, which was unconsciously processed but not consciously perceived. This was the case however only if they are separated less than a hundred milliseconds in time (Greenwald, 1996). This suggests that unconscious information decays very quickly. This result is supported by single cell recordings in monkeys (Rolls & Tovee, 1994). The neural activity of masked stimuli last only for as long as the stimulus is presented (ca. 50 ms) whereas the activity of equally long presented non-masked stimuli last for up to 300 ms. Other functions of consciousness, according to Dehaene and Naccache (2001), are its requirement for novel operations and also for intentional behaviour. However, all these purposes of consciousness do not give an answer to the philosophical question as to why some brain processing has to be accompanied by experience at all. One could argue for instance that consciousness has no purpose, and is mere an epiphenomenon (Chalmers, 1995). Why are people (and probably animals) not mere like machines, processing some input and producing some output (behaviour), without experience? That is, if you could have (in theory) a brain operating exactly the same with or without consciousness, than what is the purpose of consciousness? The hard problem: David Chalmers (1995) makes a distinction between the ‘easy’ problems of consciousness and what he has termed ‘the hard problem’. The easy problems are those that are going to be discussed in this thesis, involving which brain processes are related to consciousness. Chalmers however claims that there is no reductionist explanation for experience. This means that there is no explanation with physics as we know it, to explain everything that needs to be explained about consciousness. We can explain which neural processes are correlated with consciousness with physics, and even why the processing of some input to the brain leads some behavioural output. However, why is this processing accompanied by an experience? That is, broadly speaking one could argue that there is a logical (not empirical) possibility of a brain executing the exact same neuronal responses as a conscious brain but without being conscious all together. Think of creatures that are in every particle identical to ourselves but that live in a different world without consciousness. Chalmers has termed these copies ‘zombies’. A zombie will behave exactly the same as the original person since all the neuronal responses are exactly the same, and it is therefore really different from zombies usually seen in movies. It would even claim to be conscious, except that it is not. The fact that some neuronal processes involve consciousness does not mean that these processes could not have been there without consciousness. However, in our world these processes are, according to Chalmers, accompanied by consciousness like a law of nature, and consciousness might be a fundamental building block of physics, equal to for instance mass, time and space. Chalmers claims that with physics as we know it, we will never be able to explain everything that needs to be explained about consciousness. Therefore we need radically different theories about consciousness than those that seem intuitive to us. Although different from the way dualism was specified by Descartes, Chalmers adopts a particular form of dualism for explaining consciousness. Others have looked at other forms of physics such as quantum physics to explain consciousness (Squires, 1990). Still others claim that all that there is to explain are the neural mechanisms of consciousness, and that there is no hard problem. Dehaene and Naccache (2001) for instance fall into this last category. “Many scientists and philosophers still adhere to an essentialist view of consciousness, according to which conscious states are ineffable experiences of a distinct nature that may never be amenable to a physical explanation. Such a view, which amounts to a Cartesian dualism of substance, has led some to search for the bases of consciousness in a different form of physics (Penrose, 1990). Others make the radical claim that two human brains can be identical, atom for atom, and yet one can be conscious while the other is a mere `zombie' without consciousness (Chalmers, 1995).” Dehaene & Naccahce (2001, p.4) The problem with the last sentence of this statement is that it is actually incorrect. Chalmers (1995) does not claim that in this world two brain could exist atom for atom identical, one being conscious and the other not. However, he claims that one could think of a brain without consciousness, performing all the processes that a conscious brain would perform. Therefore the question of why these processed are accompanied with consciousness needs to be explained. Lamme (2006) does not claim to solve the hard problem but thinks that the recurrent processing theory brings it closer than previous theories. Crick & Koch (1990) do not make any specific claims about the hard problem in their 40 Hz theory. THEORIES OF CONSCIOUSNESS In this paper, the hard problem is left aside and focus is lead upon the ‘easy’ problems because these questions are interesting on their own, and they can scientifically be addressed. These involve questions such as; what brain activity is correlated with the conscious perception of a stimulus and how is this different from unconscious perception? Multiple theories on this matter have been developed and three of them are discussed here. First the Global Workspace hypothesis will be discussed followed by the 40 Hz theory and finally the recurrent processing theory. For each theory, first a short description will be given, followed by scientific findings supporting the theory. Global Workspace The global workspace hypothesis was developed in the nineteen eighty’s, mostly by Bernard Baars, as a rejection of behaviourism (Blackmore, 2005). It is one of the first contemporary theories to address the problem of consciousness scientifically. Description Baars (2005) describes the global workspace hypothesis using a theatre metaphor in which all of the theatre is dark except for the part that is within a spotlight. Everything that happens in the dark is not known to the rest of the system and therefore unconscious. Only the part that is within the spotlight is apparent to the whole system and therefore in consciousness. The spotlight is controlled by attention processes, which determine what the content of consciousness will be. The global workspace hypothesis is build upon six hypothesis and assumptions. These will first be described briefly and later support for them will follow. - (1) The brain is modular According to the global workspace hypothesis, the brain can be seen as a set of parallel processors each tuned to its own specific function or modality (e.g. perception, memory, motor planning) as indicated by imaging studies. This means that different functions/modalities are processed in different parts of the brain. - (2) Information can be processed unconsciously Each of these modalities process information more or less autonomously and in an unconscious manner, separate from the rest of the brain. - (3) Information needs to be globally available for consciousness Consciousness is derived however, at the moment information is shared and exchanged between modalities (Baars, 2005; Dehaene & Naccache, 2001). There is a centred and shared memory capacity, the global workspace, onto which otherwise separated modalities are connected and through which information can be shared to the rest of the brain. This makes information available not only at the local level but also at a global level. - (4) Minimum neuronal activation and attention attenuation is needed The modality that can send information to this global workspace is determined by topdown attention (Dehaene & Naccache, 2001). Because of this top-down regulation of which information enters the global workspace, information is shared with the system as a whole in a controlled and coordinated fashion (Baars, 2005). For a modular process to enter the global workspace -and therefore consciousness- neural activity needs to be amplified by top-down attention for a sufficient duration (see support section; Dehaene & Naccache, 2001). Subsequently, the information is thought to be sent to the rest of the modalities via corticocortical and corticothalamic fibers (Baars, 2005). - (5) Where in the brain is the Global Workspace located? Because the global workspace is thought to be a form of memory capacity, a logical question to ask would be where this workspace might be in the brain. However, there is no fixed set of regions that is connected to the global workspace, as different brain regions can be ‘mobilized’ and ‘demobilized’ (Dehaene and Naccache, 2001). According to these authors the brain contains ‘long-distance’ neurons, which connect remote brain areas and contribute to the global workspace. However, only some of these neurons are active at any particular time. Therefore they conclude that there is no particular need to point to one specific location in the brain that is active for all conscious content. However, these kinds of long distance neurons seem to mainly originate from prefrontal cortices (PFCs) and anterior cingulate cortex (ACC; Dehaene & Naccache, 2001; see support section). Furthermore, frontal and parietal association areas have consistently been indicated in processing of sensory information during conscious, but not during unconscious states (Baars, 2005). Baars (2005) suggests that there seems to be a special role for frontoparietal association areas in the formation of consciousness, even though these areas themselves are not responsible for processing for instance visual or sensory content. He proposes that these areas are important in the formation of a ‘self’ as these areas have been shown to be involved in interpreting sensory input and comparing it to expectations (Gazzaniga 1998; Baars, 2005) which is important for maintaining a sense of a ‘self’ that is consistent over time. - (6) Which modalities contribute to the Global Workspace? Because different brain areas can contribute to the global workspace but we do not become aware of all processes in the brain (e.g. you don’t have to be conscious to breath) the global workspace theory predicts that some modalities can contribute to the global workspace whereas others cannot. This would explain why we are conscious of only some content that is processed in the brain. According to Dehaene and Naccache (2001) at least five main modalities have the possibility to engage with the workspace. “[P]erceptual circuits that inform about the present state of the environment; motor circuits that allow the preparation and controlled execution of actions; long- term memory circuits that can reinstate past workspace states; evaluation circuits that attribute them a valence in relation to previous experience; and attentional or top-down circuits that selectively gate the focus of interest.” (Dehaene & Naccache, 2001, p 14). They argue that the global interconnectedness between these modalities makes conscious experience a unitary feeling. Furthermore, connections to language and motor areas make the content of workspace available for reportability. Support The previous assumptions will now be discussed again and support for them are provided. - (1) The brain is modular The global workspace hypothesis poses that the brain works in a modular fashion, i.e. that different functions are processed in different parts of the brain. This has been suggested by many imaging and lesion studies. For instance, different aspects of visual information are processed by different visual areas (V4 for colour, MT/V5 for motion etcetera; McKeefry & Zeki, 1997; Antal et al., 2005) and lesions to specific areas only disrupt specific features of vision (Livingstone & Hubel, 1988). In language for example, Broca’s area has been related to speech production (Alexander et al., 1990) whereas Wernicke’s area has been related to speech comprehension (Naeser et al., 1987). Again, disruption of these areas show specific deficits in speech production and speech comprehension respectively (Kertesz, Harlock & Coates, 1979). This suggests that certain computational operations in the brain are processed in different parts of the brain and are to some extend independent. - (2) Information can be processed unconsciously Besides that the global workspace hypothesis poses that different modalities are processed in different areas, it also claims that a lot of these modular processes take place unconsciously. Support for this statement comes from three different patient studies. These patients have in common that they are unaware of a particular form of visual input, however still show the ability to process this information unconsciously. First, people suffering from hemianopsic scotoma -blindness in only a part of the visual field- sometimes show (depending on the lesion) to be able to react above chance level to stimuli in the part of the visual field that is not consciously perceived (Weiskrantz, 1996). This has been called the ‘blindsight-phenomena’. Second, people suffering from prosopagnosia -a condition in which patients are not able to recognize faces- show different electrophysiological reactions to familiar faces than to unfamiliar faces although they claim not to recognize any of these faces (Renault et al., 1989). Third, people with hemispatial neglect tend to disregard their left visual field. They, however still show an enhanced performance on a lexical decision task if a related visual prime is shown in the left and visually unconscious field (McGlinchey-Berroth et al., 1993). Hemianopsic scotoma-, prosopagnosia - and neglect-patients all have in common their lack of conscious awareness of a particular form of information, while still being able to show behavioural or neural responses to this information. Therefore, processing of this information needs to have been taken place in an unconscious manner. Other forms of evidence that information can be processed unconsciously come from studies in healthy subjects. Some studies have shown for instance that if isolated stimuli are presented for very brief periods of time (29-50 ms), people can report seeing them and name them correctly (Deheane et al., 2001). However, when these stimuli are preceded and followed by a mask, they will not be consciously perceived. Still, these stimuli are unconsciously processed. If unmasked words are preceded by a masked prime word, reaction times where significantly faster if the prime was the same word as the unmasked target word than when it was a different word (Dehaene et al., 2001). This indicates that semantic information of unconsciously perceived words has been processed. A lot of processing is thus taking place unconsciously. - (3) Information needs to be globally available for consciousness As shown before, information can be processed in an unconscious manner, however, the global workspace hypothesis poses that this information enters consciousness at the moment it is globally available. Indeed, studies have shown that unconsciously perceived stimuli affect the primary cortices of the corresponding modalities but consciously perceived stimuli on the contrary also affect other areas such as the frontal and parietal cortex (Baars, 2005). Sleep experiments for instance show that during unconscious states only the primary auditory cortex is activated after auditory stimulation (Portas et al., 2000) but not frontal or parietal areas. Furthermore, studies with people in vegetative state also show that in these patients only the primary sensory cortices are active after presentation of auditory and tactile stimuli (Laureys et al., 2000, 2002) similar to healthy controls. However, the healthy controls also show activation in association cortices whereas the patients did not. Unconscious processed stimuli therefore solely activate the primary cortices. Indeed, word masking studies by Dehaene et al. (2001) have indicated that words followed by a mask where only processed by word recognition areas. However, when the same words were consciously perceived, other areas such as frontal and parietal regions where also active. This indicates that people do not become conscious of a stimulus when it is only locally processed but that they do when this information is also globally available. - (4) Minimum neuronal activation and attention attenuation is needed The global workspace hypothesis claims that information enters consciousness when it is globally available but only some of the neuronal processing that takes place enters consciousness. It has been suggested that for content to enter consciousness, first, there needs to be at least a minimum duration of neuronal activity (predicted 100 ms; Dehaene & Naccache, 2001; Lamme, 2006) and, second, this activity needs to be amplified by attention. First, the need for a minimum duration of neural activity will be considered. Indeed, single unit studies (Rolls and Tovee, 1994) show that masked, and therefore unconsciously perceived stimuli that were presented for only 20-30 milliseconds, initiate neuronal firing for as long as the presentation time. However, neurons remain active far longer than the stimulus presentation time for non-masked and consciously perceived stimuli, up to 200-300 milliseconds. This extended neural activity has been attributed to the influence of memory (Rolls and Tovee, 1994). This suggests that a minimum duration of neural activity duration is needed for awareness. Second, the need of attentional attenuation of this activity is discussed. For a stimulus to be consciously experienced it needs to be presented for a minimum duration and with sufficient clarity (Dehaene & Naccache, 2001). However, this is not enough for it to be perceived consciously. Directed attention is needed to amplify the neuronal activity that is encoding the stimulus (Dehaene & Naccache, 2001; Baars, 2005). Evidence for this can be found in hemispatial neglect patients as well as healthy participants. Neglect is thought to result mainly from attentional deficits (due to parietal cortex lesions) that draw the focus of attention completely to the right side of the visual field. If two stimuli are presented, one on the left and one on the right, patients will only report seeing the right stimulus. When, however, only the left stimulus is presented people do report seeing it. This means that if attention is not available for the left stimulus (because of the presence of the stimulus on the right), people will not consciously perceive this stimulus. However, when attention is available for the stimulus on the left (when there is no stimulus on the right), people will perceive it. This shows that attention is needed to become consciously aware of a stimulus. Furthermore, studies in healthy participants show that attention is needed for conscious processing. One interesting phenomenon that supports this is called ‘inattentional blindness’ (Mack, 2003). In studies investigating this phenomenon, people are instructed to direct their attention to a particular set of stimuli and to perform a demanding cognitive task with these stimuli. Most of the participants however failed to perceive an extra (un-attended) stimulus at the point where they were looking. This un-attended stimulus had sufficient contrast and duration to be perceived if the other stimuli where not displayed or if no task directing attention away from them was involved. Again, this shows that attention is needed to become consciously aware of a stimulus. - (5) Where in the brain is the Global Workspace located? The brain modalities that are attenuated by attention differ according to the situation in which someone finds itself. There is therefore not one set of areas that is always involved in the global workspace but different areas may contribute to the workspace at different points in time. Support for this comes from for instance imaging studies indicating that different brain areas are active during different experiences. Therefore, depending on the experience, relevant areas contribute to the in global workspace (Dehaene & Naccache, 2001). For instance, during the experience of a face, the fusiform face area (Kanwisher, McDermott & Chun, 1997) will be contributing to the global workspace whereas during the perception of motion, area V5 (Antal et al., 2005) will be mobilized. Furthermore, the PFC and ACC have also been suggested to play a crucial role for consciousness as indicated by brain imaging studies, even though they are not directly involved in sensory processing (Dehaene & Naccache, 2001). One interesting study for example comes from McIntosh, Rajah, and Lobaugh (1999) where they showed people multiple stimuli that were related to each other, which was only detected by some of the participants. Interestingly they found that people that became aware of this relationship showed increased activation in the left PFC. Moreover, in accordance with the workspace hypothesis, they found long-distance functional connections appearing at the moment people became conscious of the relationship. - (6) Which modalities contribute to the Global Workspace? Although the areas that contribute to the global workspace can differ, only some modalities have the ability to contribute to the global workspace whereas others have not. Anatomical studies with monkeys show that the PFC contains long-distance neurons connecting this area to temporal and parietal areas (important for perceptual categorization and attention), motor and language related areas (for action upon the conscious content) and the hippocampus (for memory; Goldman-Rakic, 1988 in; Dehaene & Naccache, 2001). This indicates that these modalities can contribute to the global workspace. Summery The global workspace theory is one of the first contemporary theories that tried to address the problem of consciousness scientifically. It poses that information is processed in a different parts of the brain in an autonomous way. Information that is processed in a particular brain area can however be shared with other brain areas through a shared memory space: the global workspace. People stay unconscious of the information that is not shared, however information that is shared with other brain areas (for instance areas important for memory and language etc.) via this workspace will enter consciousness. Long-distance neurons connect different brain areas to this workspace. Which information is to be globally available is influenced by attention processes. Although the workspace is not specifically located in one place and dependents on the set of neurons that contributes to it at a given moment in time, the PFC and ACC might have an important role in the formation of conscious experience. Possibly these areas are involved in the formation of a sense of a ‘self’ that is continuous and consistent over time. Personal opinion There is a lot to say in favour of the Global Workspace model. It is a simple concept and it conceptually has been influential for other theories of consciousness. Furthermore, there is a lot of evidence that confirms the proposed ideas. I think the global workspace hypotheses lays down a good cognitive understanding of consciousness. However, in my opinion the role of attention has not been fully explored in this theory. Furthermore, these authors have stated that the global workspace theory explains consciousness, denying the hard problem of consciousness. A notion on which I disagree with the authors. Therefore, here follow two personal critiques. First, a big role is reserved for attention in determining what information enters consciousness. However, there is no clear specification on how attention works and how it makes ‘decisions’ as to what enters consciousness. Although this might need a new theory altogether I think it is an important issue to address since, without an explanation of how attention works there is still some part of the mechanisms behind consciousness that is not explained. Furthermore, because no real description of the attention is given, it remains unclear how attention and consciousness are different concepts. Second, Dehaene & Naccache (2001) claim that the question of consciousness can be solved with cognitive psychology and neuroscience. Furthermore, Baars (2005) states that: “the primary functional role of consciousness [is]: to allow a theater architecture to operate in the brain, in order to integrate, provide access, and coordinate the functioning of very large numbers of specialized networks that otherwise operate autonomously.” (Baars, 2005, p47-48). However, in my opinion ‘the hard problem’ is still a valid problem. That the brain might work in a global workspace fashion, with information globally available, might only be because it is efficient. Indeed, computer simulations suggest that effortful processing of information is accompanied by global information availability (Dehaene, Kerszberg & Changeux, 1998). This does not mean however that information processing needs to be accompanied by ‘awareness’ or ‘experience’ and the global workspace theory therefore does not provide an answer as to why this is the case. 40 Hz Oscillations Whereas the global workspace hypothesis is a high level cognitive model, Franscis Crick and Christof Koch (1999) have argued that a low level model on the neural basis of consciousness is needed. They build upon similar ideas as the global workspace hypothesis but they focus on how neurons involved in consciousness communicate. Their focus is mainly on the visual domain as that is well-studied domain. Description Many ideas that were postulated in the global workspace hypothesis are also seen in the 40 Hz oscillations hypothesis. For instance, Franscis Crick and Christof Koch (1999), believe that that the brain can be seen as a set of parallel processors and that during an experience, in multiple areas, neurons relevant to that experience are active. In other words, that the brain is modular with different brain areas responsible for the processing of different forms of information. Furthermore, they pose that consciousness can arise when neural activity is global and unified and that “some form of rather short-term memory and also […] some form of serial attentional mechanism” is needed (Crick & Koch, 1990, p263). This ‘form of short-term memory’ is comparable to the ‘global workspace’ on to which different brain areas are connected to share information. They further assume that attention mechanisms are needed to select what content is to get into consciousness. These are all higher order cognitive notions comparable to the global workspace hypothesis. The 40 Hz theory, however, emphasizes on what lower level neural mechanism exactly is used to create this global unity. There are a couple of questions that need to be answered according to these authors, with respect to neurons that give rise to conscious experiences and they give possible explanations for some of them. The first and most prominent question for which Crick and Koch give an explanation is: how do neurons responsible for consciousness fire differently (if at all) than other neurons? Second, what is the location of these neurons in the brain? Third, are they of any specific neuronal type? According to these authors, the brain does not have the capacity to store/represent every experience in a single cell. Although there are cells that have a very specific tuning to particular stimuli (like the grandmother cells; Gross, 2002), most experiences arise from the interaction of multiple neurons. However neurons that are responsible for different parts of one experience are located in distant areas. For instance in vision, neurons that are responsible for colour processing and neurons that are responsible for motion processing need to act in unity to result in a unified experience, even though they are located in different areas. How the brain creates this unity is what has been called the ‘binding problem’ (Treisman, 1996). To be able to experience a new ‘scene’, some sort of binding is necessary between neurons in different areas. This needs to happen fast and dynamically (however, if learning is involved however, bindings needs to be permanent). Crick and Koch argue that, with respect to their first question, neurons involved in visual consciousness fire in synchronization to create unified processing (see support section). Attention is thought to select relevant neurons throughout the brain that need to fire in synchronization. This synchronization is established when neurons in different areas fire in oscillations at the same frequency. The frequencies with which this may occur may vary between 35-75 Hz but they named this whole range the ‘40 Hz oscillations’ (in this thesis it is sometimes referred to as the gamma band oscillations). Furthermore, they argue that, with respect to their second question, neurons involved in consciousness are mainly located in the neocortex and that, with respect to their third question, these neurons are ‘complex cells’ (as opposed to simple cells; see support section). Support - (1) Neurons that are responsible for consciousness fire in synchrony To become conscious of for example a visual scene, different features (colour, motion, etc.) of that scene need to be bound together. The proposed mechanism responsible for binding different features of an experience is to synchronize the firing of neurons responsible for the processing of those features. It is believed that this synchronization mainly happens at oscillations in the gamma range (according to these authors and others; Crick & Koch, 1990; Koch & Rees, 2002). Animal research has shown for instance that recordings in different neurons of the visual cortex responded in an oscillatory fashion when stimulated by moving light bars in a particular direction (Gray et al., 1989). These oscillations were mainly seen in the gamma range. Recordings of two different neurons with the same preferred movement orientation show that when two bars were presented, one in each neurons’ receptive field but moving in opposite direction, these oscillations where not synchronized. When both bars moved in the same direction, only a weak synchronization was found. However, when one bar (spanning both receptive fields) was presented, the synchronization was enhanced remarkably. This suggests that the processing of an object in different parts of the brain is being unified through synchronized oscillations. Moreover, in humans, gamma oscillations have been shown to be related to conscious percept. Rodriguez et al. (1999) presented people stimuli that either could be perceived as meaningless shapes or as faces. Long distance gamma oscillation synchronization became apparent at the moment people start to see a face out of the shapes. Another line of research indicates that synchronized oscillatory neuronal firing is important for consciousness comes from attention studies. Attention is thought to be responsible for selecting what is to emerge into consciousness (based on the global workspace hypothesis, see that section for support) and it was found that selective attention facilitates synchronized neuronal activity in visual and somatosensory areas (Rees, Kreiman & Koch, 2002). A study with monkeys showed for example that neurons responding to an attended stimulus exhibited increased gamma oscillations compared to nearby neurons responding to distracter stimuli (Fries et al. 2001). The amplification of gamma oscillations with attention suggests that gamma oscillations are correlated with conscious percept. Even though the way attention influences consciousness is not fully known, Crick and Koch (1990) provide some ideas about how exogenous visual attention might work. There could be a topographical salience map (possibly located in the thalamus) indicating where important information is located in the visual field. This salience map would then be a combination of individual feature maps (e.g. where does color stand out, where is there motion). The salient locations are those locations in a visual scene where these features are different from the background. A particular location to attend to is selected in a winner-take-all fashion. Once a location is selected, feedback pathways (form the salience map to the involved visual neurons) will synchronize the activity in all visual neurons that correspond to that location. There might be a couple of reasons why synchronized oscillations would be useful for unifying the processing of stimuli in different parts of the brain. One of the reasons might be that synchronized activity means that incoming signals arrive at a neuron at the same time which will lead to larger effects than random spike activity of the same amount (Crick and Koch, 1990). Furthermore, according to Fries (2005) two different brain areas communicate to each other most efficiently if the neurons in those areas are both in their excitable phase. The excitability of the receiving neuron is known when the two areas oscillate coherently. Therefore, synchronized oscillations between different areas that process different features of a stimulus are thought to be used for communication between these areas. - (2) Where in the brain are the neurons responsible for consciousness? Crick and Koch (1990) argue that neurons that are responsible for consciousness are mainly located in the neocortex and paleocortex because damage to cerebral cortex is often accompanied by disturbances in consciousness. - (3) Are these neurons of any particular neuronal type? Research of the visual cortex showed that only a subset of neurons show to fire in oscillation. Gray et al. (1990) found that these neurons where mainly complex neurons as opposed to simple neurons. Simple neurons process information about a particular small part of the visual scene (think of it as the information of a pixel on a computer screen, which in itself doesn’t provide much information about the shape or form that pixel is part of). However, complex neurons integrate information from different simple cells to derive at concepts of visual forms (by combining the information in multiple pixels, forms and shapes can be made). Complex cells are therefore logical candidates to fire in an oscillatory fashion and to be involved in consciousness. Summary The 40 Hz theory was a reaction to cognitive theories of consciousness and was one of the first theories to explain consciousness on a neural level. It agrees with most of the ideas of the global workspace but focuses on what neuronal mechanisms are involved during conscious experience. First, it poses that there are simply not enough neurons to represent every experience by a different neuron and so neuronal interactions are needed to build up an experience. Second, different features of an experience are processed in distant brain areas and this information needs to be bound together to create a unified experience. Third, the mechanism that is responsible for this binding is the synchronization of neuronal oscillations (in the gamma range) in these distant areas. Attention is thought to be responsible for synchronizing these oscillations. Cells responsible for consciousness are mainly complex neurons located in the neocortex. Personal opinion The 40 Hz theory has tried to give a neuronal explanation for consciousness. Although the theory indeed has more of a low level focus than the global workspace theory, more proof for it is needed in my opinion. Although some studies (as outlined above) suggests that synchronized oscillations do involve consciousness (at least for visual awareness), there is still debate on this matter. For instance, disruption of synchrony in oscillations does not seem to impair perception in primates and therefore a causal role for synchronous oscillations in consciousness is not yet found (Rees, Kreiman & Koch, 2002). Moreover, Crick and Koch themselves argue that “Our tentative theory, most of the elements of which have already been proposed by others, is a program for research rather than a detailed model”. (Crick & Koch, 1990, p273). Therefore, more research and evidence is needed. Furthermore, the question of why gamma oscillations in particular are necessary for consciousness (and not just oscillations in general) is not addressed for instance. Besides, although the authors indicate that their ideas on how attention works is still incomplete, one important questions is left out all together, namely; how does attention select a winner out of multiple locations to attend to if those locations have the same level of salience. Furthermore, it only explains stimulus driven (exogenous) attention but not internally driven (endogenous) attention. Moreover, this idea of how attention works remains rather high level (cognitive) and a better low level (neuronal) understanding of it is not provided. Recurrent processing Just like Crick and Koch, others have also argued for a neuronal exploration of consciousness. One of them is Victor Lamme, who goes as far as stating that our intuitive notions of what consciousness is (i.e., people can report what they are conscious of), is constraining our understanding of it. In his 2006 paper ‘Towards a true neural stance on consciousness’, he argues that both behavioural observations (what people report as to what their conscious content is) as well as neural arguments should be considered as equally contributing to the scientific understanding of consciousness. Description There are conditions in which consciousness might exist, yet people fail to report it. To illustrate this, Lamme (2006) refers to the behaviour of split-brain patients (see also Gazzaniga, 1998). These are patients in which the corpus callosum, the massive collection of neuronal tracts that connect both hemispheres, is surgically been disconnected. These patients cannot verbally report on anything that is presented solely to the left visual field. However, they still show forms of recognition as indicated by non-verbal reports with the left hand. The reason for this is that left visual content is processed mostly by the right hemisphere but language is usually located mostly in the left hemisphere. Since communication between hemispheres is disrupted, no verbal reports can be made and therefore these patients claim not to be conscious of the stimulus. However, they can draw with their left hand what they ‘saw’ because the left hand is controlled mostly by the right hemisphere. Does this mean that people are not consciousness only because there is no language that can report on it? In considering this question, Lamme argues that neuronal arguments should account for as much as proof of consciousness as do behavioural measures, and uses research on ‘recurrent processing’ to illustrate how this can be done. What is recurrent processing? Just like the global workspace hypothesis and the 40 Hz oscillations theory, the recurrent processing (RP) theory notices that the brain acts in a modular fashion with different brain areas devoted to different functions. Furthermore, it also states that information exchange between different brain areas is needed for consciousness, which is comparable to the previous two discussed theories. However, according to this theory, this communication should be mutual in order for consciousness to arise. For instance, when an image hits the retina, this information travels very rapidly (10 ms per processing level) along the brain hierarchy from lower brain areas to the higher ones (Lamme, 2006). This information flow is called the ‘feed-forward sweep’. Within 100-150 ms this information has reached all involved brain areas. This feed forward of information alone, however seems not to be enough for conscious awareness. Communication back from higher order areas (e.g. v4, v5/MT) to lower order areas (e.g. v1) seems to be an important factor in the establishment of consciousness. This form of communication has been called recurrent processing. Recurrent processing can happen locally and globally. Local recurrent processing refers to communication between areas of a particular domain, such as the different visual areas. Global recurrent processing means that sensory information is also shared with non-sensory areas such as frontoparietal association areas. People are thought to be able to report on, for example a visual scene, especially when not only local information exchange but also global information exchange is established (in agreement with the global workspace theory). When only local recurrent processing is present, people do not generally seem to claim to be aware of a stimulus. However, this processing can still influence some forms of behaviour as indicated by masked priming studies (see support section). Lamme argues that we should take this as much as prove for consciousness as reports of awareness, even though this might seem counterintuitive. The reason for this is that it might help us understand what the function of consciousness is and separate it from other cognitive functions. To show that recurrent processing is involved conscious percept, four questions need to be addressed. First, is recurrent processing indeed involved during sensory processing? Second, is it involved during conscious percept? Third, is it also sufficient for consciousness? Fourth, what are the advantages of a pure neuronal understanding of consciousness? Most of the support studies have focused on awareness of visual stimuli. Support - (1) Is recurrent processing involved in visual processing? Recurrent processing means that not only information is traveling from lower to higher order areas (by means of the feed-forward sweep) but higher order areas are also communicating back to lower order areas. This form of processing has been indicated during visual perception and is indicated by neuronal activity after the feedforward sweep is finished. The feed-forward sweep reaches all higher order areas within 100 ms (Lamme & Roelfsema, 2000) but neurons in both lower and higher visual areas can stay active after the feed-forward sweep is finished. Studies have suggested that this might be due to recurrent processing. Neurons show an initial peak response to stimuli presented in their receptive field (RF), which reflects the feed-forward sweep (Lamme & Roelfsema, 2000; Supèr et al. 2001). However, the activity that is seen after the feed forward sweep is finished has been shown to be influenced by visual information outside a neurons receptive field (Allman et al, 1985). This is information outside a neurons receptive field that determines the salience of the stimulus (responsible for figure-background separation). For instance, if a line is surrounded by similar oriented lines, it is less salient than when it is surrounded by differently oriented lines. And this surrounding information (indicating how salience a stimulus is) influences the later neural activity. For instance, when the same stimulus is placed on two different backgrounds (different surroundings) in which it is more salient in one compared to the other, the fast neuronal response will cause equal activity in both conditions, since the stimulus is the same. However, the slower neuronal response is enhanced if the stimulus is more salient due to the background. The influence of a neurons activity by its surrounding is called context modulation. Because this late neural activity is modulated by information that is processed by other neurons, it must be due to recurrent processing. - (2) Is recurrent processing necessary for conscious experiences? Studies have suggested that feed-forward sweep alone is not sufficient for reportable consciousness experiences and that recurrent processing is necessary (Lamme, 2006). Backward masking studies show for instance that the presentation of a mask after a short delay (<100) to the stimulus can disrupt awareness of the stimulus. The feed forward sweep still seems to reach all areas (Lamme & Roelfsema, 2000) and it has therefore been suggested that masking suppresses the recurrent processing. Furthermore, transcranial magnetic stimulation (TMS) experiments in which visual areas where disrupted after the feed-forward sweep is finished show impaired perception (Lamme & Roelfsema, 2000). Therefore the decreased stimulus awareness has to be due to disrupting the recurrent processing. Furthermore, studies with monkeys suggest that recurrent processing is necessary for the monkeys to report the presence of a target stimulus (Supèr et al. 2001). Both detected targets and un-detected targets produced fast neuronal responses in V1 caused by the feed-forward sweep, showing that the detection failure is not caused because signals do not the reach visual cortex. However, a slower recurrent processing response (due to context modulation outside a neurons receptive field) seems to be enhanced when targets are detected. This suggests that when someone reports seeing a stimulus, recurrent processing is present. - (3) Is recurrent processing sufficient for consciousness? Although recurrent processing is present when people and monkeys report seeing a stimulus, RP can also be seen when people or monkeys fail to report any awareness of an object. Studies in which monkeys had to make a saccade to a target stimulus and ignore other stimuli (catch trials), have shown for instance that raising the number of catch trialsand so the decision threshold for detecting a target- induces instances of RP although the monkeys did not make a saccade (Supèr et al. 2001). In other words, the monkeys did not report to be aware of the stimulus but still RP was seen. Furthermore, human studies show that when an object is shown but no attention is paid to it (e.g. during inattentional blindness) and so the object is not reported to be seen, recurrent processing is present (Lamme, 2006). However, it is present only locally between visual areas. In this case there is no global recurrent processing (between visual areas and frontoparietal areas). On the contrary, when global recurrent processing is present, people usually report to be aware of the object (Lamme, 2006). This seems to falsify the statement that recurrent processing is sufficient for conscious experience but Lamme argues that this is because we only take consciousness as that what people/monkeys report to be aware of. This however relies on other modalities, such as language, attention and working memory. For instance, in the monkey study it could be that the monkey did in fact see the target but because of the many catch trials decided not to report it (Lamme, 2006). In the case of intentional blindness, people could be forgetting the stimuli that where not attended and therefore not report them although they might have been ‘conscious’ of them at the moment of perception (Lamme, 2006). Lamme (2006), therefore suggests that recurrent processing should be taken as an indication of consciousness. - (4) What are the advantages of a neural definition of consciousness? It might seem counter intuitive to state that people can be conscious a stimulus even if they don’t report on it. However there might be some advantages for doing so. Because of the reliance of behavioral report on other modalities, Lamme argues that our understanding of consciousness cannot be complete. He argues that if we attribute neural evidence for consciousness (such as the existence of RP) equal weight as behavioral reports, we might gain some insights. The first advantage is that we can describe consciousness without the use of other cognitive functions such as attention, language and memory etcetera, which are needed for reportability. This enables one to study consciousness as a concept on its own (Lamme, 2006), which is very useful in cases where all forms of communication are lost (such as locked-in syndrome) or not available in the first place (animals). Then a neural concept of consciousness is the only way to reach consensus on whether or not consciousness is present. Although it seems problematic that the neural concept of consciousness does not always accompany reportability, this can still be explained very well. In the cases where there is no reportablity (inattentional blindness, split brain), cognitive functions other than consciousness might be responsible for this (e.g., memory, language). For instance, because someone forgot about a perception or because, in split brain patients, language areas cannot be reached. This means that consciousness can be seen as separate from other functions without losing an explanation for why sometimes people cannot report being aware of something. The second advantage is that a neural definition for consciousness might help us understanding what the direct function of consciousness is. Because during recurrent processing, pre-and postsynaptic neurons are active at the same time, the Hebb rule is met. This rule states that the more two neurons fire simultaneously, the more they form a solid connection, which is important for learning. Therefore recurrent processing might be a basis for learning (Lamme, 2006). Stimuli that only produce a feed-forward sweep do not produce long lasting changes to the brain. Consciousness can thus be seen as fundamental for learning. Summary The recurrent processing theory focuses, like the 40 Hz theory, on the neural mechanisms of consciousness. However it deviates from an intuitive understanding of consciousness that is based on whether or not people can report to be conscious, and tries to explain consciousness fully by the presence of particular neural activity. It poses that consciousness can be measured by whether or not recurrent processing is present. Furthermore, if this is the case only locally, people are likely not to report on an experience whereas if recurrent processing is also seen globally, people are likely to report the experience. The advantage of seeing consciousness fully in neural terms is that it can be studied in cases where no reportabilty exists. Furthermore, it might give some insights in the purpose of conscious. Personal opinion Taking a neural concept of consciousness without linking it to reportabilty feels counter intuitive, however there are some clear advantages in doing so. It will give some insights in how consciousness is neuraly different from other functions and what the function of consciousness is, which makes the theory valuable in my opinion. There are however some personal critiques on how recurrent processing is measured and how consciousness as a modality is defined. First, there are studies that support the idea that recurrent processes are involved in consciousness. However, these studies involve assumptions or indirect measures of RP (e.g. context modulation). Studies measuring recurrent processing directly would therefore be a good addition, for instance by simultaneous single cell recordings of two cells. Second, whether consciousness can be completely separated from other function such as language, attention and memory can be debated. The two types of consciousness, with and without reportability, are neuraly different and so using the same terminology (as Lamme does) might not be the most efficient in my opinion. Ned Block (2005) may give a somewhat clearer distinction. He claims, as mentioned before, that there are different types of consciousness: phenomenal- and accessconsciousness (see introduction; “What is consciousness?”). Phenomenal consciousness, he links to Lammes’ recurrent processing and access consciousness to the global workspace. This distinction might do more justice to the different forms of consciousness. Third, Lamme seems to equate consciousness to some extent to learning but is not clear whether he thinks these processes are the same or still different. He argues that to use a term such as consciousness it should be separated from other functions. However, when saying that learning relies on consciousness (in both it’s reportable and unreportable form) it is not clear what makes the distinction between learning as a function and consciousness as a function. Conclusion/Summary In this thesis I gave an overview of three influential theories on the cognitive and neural correlates of consciousness. First, the Global Workspace theory was explained which states that consciousness arises if information is not only locally but also globally available. This was an early cognitive theory that tried to find a scientific explanation for the problem of consciousness and was influential for neural theories of consciousness, two of which have been discussed in this thesis. Second, the 40 Hz hypothesis was considered which was developed to move from the cognitive theories of consciousness to a more neural understanding. It builds upon the same ideas as the global workspace hypothesis but focuses on the neural mechanisms involved during conscious awareness. It states that different parts in the brain process different features of, for instance a visual scene, and these features are bound together if one becomes conscious of the scene. The neural mechanism for this binding is through to be the synchronization of gamma oscillations. Finally, the Recurrent Processing hypothesis was considered, which also gives a neural explanation of consciousness. However, it goes as far to state that consciousness should not be considered as only that what people report on but also as what kind of neural activity is present. According to this theory, recurrent processing can be viewed as a sign of consciousness even though people might not report to be conscious. Although these theories focus on different aspects they all share some common notions. All theories originate from the cognitive perspective that the brain works to some extent in a modular fashion, meaning that different functions are processed in different parts of the brain. Also, all theories state that when people report to become conscious, brain activity is seen on a global scale with different areas communicating with each other. The 40 Hz theory however, focuses on how different areas communicate during conscious content processing (through oscillations) whereas the recurrent processing theory focuses on which areas communicate (local recurrent interactions or global recurrent interactions) and in what direction this communication goes. As has been shown by the theories discussed in this thesis, a scientific understanding of consciousness was arrived through cognitive theories. Based on this, neural theories where developed, which focused on how neurons interact with each other and which neurons/areas interact with each other during conscious and unconscious processing. Furthermore, our perception of consciousness, its role and purpose, might even change because of these theories. Because of the many debates that are still going on about consciousness, this thesis was not meant to give an answer to the questions on consciousness. However, it discussed some of the ideas and theories concerning consciousness because these have had (in my opinion) to little attention in the education of neuroscience and cognitive psychology. There remains a lot of research and debate as a lot of questions are still not clearly answered. Consensus on the definition and purpose of consciousness is not yet settled and the differences between consciousness, attention and awareness etcetera are not yet clear. This makes consciousness still a fascinating subject of research. Acknowledgement Henk Aarts for the first directional advice, Albert Postma for finding the right people to contact, Lenny Ramsey and Ravi Chacko for there comments and advice, my fellow students for the discussions on the topic, with in particular Joost Haarsma and Victor Manuel Saenger, Jan Brascamp for reviewing and Serge Dumoulin for his advice and examination. References Alexander, M.P., Naeser, M.A. & Palumbo, P. (1990). Broca’s area aphasias: Aphasia after lesions including the frontal operculim. Neuology, 40, 353-362. Allman, J., Miezind, F., & McGuinness, E. (1985). RESPONSES FROM BEYOND FIELD : Neurophysiological Mechanisms for Local-Global Comparisons in Visual Neurons. Annual Reviews of Neuroscience, 8, 407-430. Antal, A., Nitsche, M. a, Kruse, W., Kincses, T. Z., Hoffmann, K.-P., & Paulus, W. (2004). Direct current stimulation over V5 enhances visuomotor coordination by improving motion perception in humans. Journal of Cognitive Neuroscience, 16(4), 521–7. doi:10.1162/089892904323057263 Baars, B. J. (2005). Global workspace theory of consciousness: toward a cognitive neuroscience of human experience. Progress in Brain Research, 150, 45–53. doi:10.1016/S0079-6123(05)50004-9 Blackmore. S. (2005). Conversations on Consciousness. Oxford, England: Oxford University Press. Block, N. (2005). Two neural correlates of consciousness. Trends in Cognitive Sciences, 9(2), 46–52. doi:10.1016/j.tics.2004.12.006 Chalmers, D.J. (1995)- The Conscious Mind: In Search of a Fundamental Theory. Oxford, England: Oxford University Press. Crick, F., & Koch, C. (1990). Towards a neurobiological theory of consciousness, 2, 263–275. Dehaene, S., Kerszberg, M., & Changeux, J. P. (1998). A neuronal model of a global workspace in effortful cognitive tasks. Proceedings of the National Academy of Sciences of the United States of America, 95(24), 14529–34. Dehaene, S., & Naccache, L. (2001). Towards a cognitive neuroscience of consciousness: basic evidence and a workspace framework. Cognition, 79(1-2), 1–37. Retrieved from http://www.ncbi.nlm.nih.gov/pubmed/11164022 Dehaene, S., Naccache, L., Cohen, L., Bihan, D. L., Mangin, J. F., Poline, J. B., & Rivière, D. (2001). Cerebral mechanisms of word masking and unconscious repetition priming. Nature Neuroscience, 4(7), 752–8. doi:10.1038/89551 Dennett, D. (2001). Are we explaining consciousness yet? Cognition, 79(1-2), 221–37. Retrieved from http://www.ncbi.nlm.nih.gov/pubmed/11164029 Fries, P., Reynolds, J. H., Rorie, a E., & Desimone, R. (2001). Modulation of oscillatory neuronal synchronization by selective visual attention. Science (New York, N.Y.), 291(5508), 1560–3. doi:10.1126/science.291.5508.1560 Fries, P. (2005). A mechanism for cognitive dynamics: neuronal communication through neuronal coherence. Trends in Cognitive Sciences, 9(10), 474–80. doi:10.1016/j.tics.2005.08.011 Gazzaniga, M. S. (1998). The split brain revisited. Scientific American, 279(1), 50–5. Retrieved from http://www.ncbi.nlm.nih.gov/pubmed/9648298 Gray, C. M., Engel, A. K., König, P., & Singer, W. (1990). Stimulus-Dependent Neuronal Oscillations in Cat Visual Cortex: Receptive Field Properties and Feature Dependence. The European Journal of Neuroscience, 2(7), 607–619. Retrieved from http://www.ncbi.nlm.nih.gov/pubmed/12106295 Gray, M.C., Köning, P., Engel, A.K. & Singer, W. (1989) Oscillatory responses in cat visual cortex exhibit inter-columnar synchronization which reflects global stimulus proporties. Nature, 338, 334-337. Greenwald, a G., Draine, S. C., & Abrams, R. L. (1996). Three cognitive markers of unconscious semantic activation. Science (New York, N.Y.), 273(5282), 1699–702. Retrieved from http://www.ncbi.nlm.nih.gov/pubmed/8781230 Gross, C. G. (2002). Genealogy of the “Grandmother Cell.” The Neuroscientist, 8(5), 512–518. doi:10.1177/107385802237175 Kanwisher, N., McDermott, J., & Chun, M. M. (1997). The fusiform face area: a module in human extrastriate cortex specialized for face perception. The Journal of Neuroscience : The Official Journal of the Society for Neuroscience, 17(11), 4302–11. Retrieved from http://www.ncbi.nlm.nih.gov/pubmed/9151747 Kouider, S., & Dehaene, S. (2007). Levels of processing during non-conscious perception: a critical review of visual masking. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences, 362(1481), 857–75. doi:10.1098/rstb.2007.2093 Kouider, S., Eger, E., Dolan, R., & Henson, R. N. (2009). Activity in face-responsive brain regions is modulated by invisible, attended faces: evidence from masked priming. Cerebral Cortex (New York, N.Y. : 1991), 19(1), 13–23. doi:10.1093/cercor/bhn048 Lamme, V. a F. (2006). Towards a true neural stance on consciousness. Trends in Cognitive Sciences, 10(11), 494–501. doi:10.1016/j.tics.2006.09.001 Lamme, V. a, & Roelfsema, P. R. (2000). The distinct modes of vision offered by feedforward and recurrent processing. Trends in Neurosciences, 23(11), 571–9. Retrieved from http://www.ncbi.nlm.nih.gov/pubmed/11074267 Laureys, S., Faymonville, M. E., Degueldre, C., Fiore, G. D., Damas, P., Lambermont, B., … Maquet, P. (2000). Auditory processing in the vegetative state. Brain : A Journal of Neurology, 123 ( Pt 8), 1589–601. Retrieved from http://www.ncbi.nlm.nih.gov/pubmed/10908189 Laureys, S., Faymonville, M. E., Peigneux, P., Damas, P., Lambermont, B., Del Fiore, G., … Maquet, P. (2002). Cortical Processing of Noxious Somatosensory Stimuli in the Persistent Vegetative State. NeuroImage, 17(2), 732–741. doi:10.1006/nimg.2002.1236 Livingstone, M., & Hubel, D. (1988). Segregation of Form, Color, Movement, and Depth: Anatomy, Physiology, and Perception. Science, 240, 740–749. Retrieved from http://www.sciencemag.org/content/240/4853/740.short Mack, A. (2003). Inattentional Blindness : Looking Without Seeing. Current Directions in Psychological Science, 12(1982), 180–185. McGlinchey-Berroth, R., Milberg, W.P., Verfaellie, M., Alaxander, M., Kilduff, P.T. (1993) Semantic processing in the neglected visual field: evidence from a lexical decision task. Cognitive Neuropsychology, 10, 79-108. McIntosh, A.R., Rajah, M.N. and Lobaugh, N.J. (1999) Interactions of prefrontal cortex in relation to awareness in sensory learning. Science, 284, 1531-1533 DOI: 10.1126/science.284.5419.1531 McKeefry, D. J., & Zeki, S. (1997). The position and topography of the human colour centre as revealed by functional magnetic resonance imaging. Brain : A Journal of Neurology, 120 ( Pt 12), 2229–42. Retrieved from http://www.ncbi.nlm.nih.gov/pubmed/9448578 Naeser, M.A., Helm-Estabrooks, N., Haas, G., Auerbach, S., Srinivasan, M. (1987). Relationship between lesoin extent in ‘Wernicke’s area’ on computed tomographic scan and predicting recovery of comprehension in wernicke’s aphasia. Archives of Neurology, 44, 73-82. Owen, A. M., Coleman, M. R., Boly, M., Davis, M. H., Laureys, S., & Pickard, J. D. (2006). Detecting awareness in the vegetative state. Science (New York, N.Y.), 313(5792), 1402. doi:10.1126/science.1130197 Physik, F., Physik, A., & Marburg, D.-. (1988). Biological Cybernetics Multiple Electrode and Correlation Analyses in the Cat, 130, 121–130. Portas, C. M., Krakow, K., Allen, P., Josephs, O., Armony, J. L., & Frith, C. D. (2000). Auditory processing across the sleep-wake cycle: simultaneous EEG and fMRI monitoring in humans. Neuron, 28(3), 991–9. Retrieved from http://www.ncbi.nlm.nih.gov/pubmed/11163282 Rees, G., Kreiman, G., & Koch, C. (2002). Neural correlates of consciousness in humans. Nature Reviews. Neuroscience, 3(4), 261–70. doi:10.1038/nrn783 Renault, B., Signoret, J. L., Debruille, B., Breton, F., & Bolgert, F. (1989). Brain potentials reveal covert facial recognition in prosopagnosia. Neuropsychologia, 27(7), 905–12. Retrieved from http://www.ncbi.nlm.nih.gov/pubmed/2771029 Rodriguez, E., & George, N. (1999). Perception ’ s shadow : long- distance synchronization of human brain activity. Nature, 397, 430–433. Roefe, M., & Beech, S. (1979). Computer Tomographic Prognosis in Aphasia Lesion Size , and and Nonverbal impairment, 50, 34–50. Scholte, H. S., Witteveen, S. C., Spekreijse, H., & Lamme, V. a F. (2006). The influence of inattention on the neural correlates of scene segmentation. Brain Research, 1076(1), 106–15. doi:10.1016/j.brainres.2005.10.051 Squires. E. (1990). Conscious mind in the physical world. Bristol, England: IOP Publising Ltd. Supèr, H., Spekreijse, H., & Lamme, V. a. (2001). Two distinct modes of sensory processing observed in monkey primary visual cortex (V1). Nature Neuroscience, 4(3), 304–10. doi:10.1038/85170 Toveet, J. (2014). speed in the cerebral cortex and the Processing of visual masking neurophysiology, 257(1348), 9–15. Treisman, a. (1996). The binding problem. Current Opinion in Neurobiology, 6(2), 171–8. Retrieved from http://www.ncbi.nlm.nih.gov/pubmed/8725958 Wegner, D. M. (2003). The mind’s best trick: how we experience conscious will. Trends in Cognitive Sciences, 7(2), 65–69. doi:10.1016/S1364-6613(03)00002-0 Weiskrantz, L. (1996). Blindsight revisited. Current Opinion in Neurobiology, 6(2), 215–20. Retrieved from http://www.ncbi.nlm.nih.gov/pubmed/8725963