To increase fitness for survival, organisms not only passively react to environmental changes but also actively predict future events to prepare for potential hazards within their environment. Accumulating evidence indicates that the human brain is a remarkable predictive machine which constantly models causal relationships and predicts future events. This ‘predictive processing' framework, a prediction-based form of Bayesian inference, states that the brain continuously generates and updates predictions about incoming sensory signals. This framework has been showing notable explanatory power in understanding the mechanisms behind both human behaviour and neurophysiological data and elegantly specifies the underlying computational principles of the neural system. However, even though predictive processing has the potential to provide a unified theory of the brain (Karl Friston, 2010), we still have a limited understanding about fundamental aspects of this model, such as how it deals with different types of information, learns statistical regularities and perhaps most fundamentally of all what its relationship to conscious experience is. This thesis aims to investigate the major gaps in our current understanding of the predictive processing framework via a series of studies. Study 1 investigated the fundamental relationship between unconscious statistical inference reflected by predictive processing and conscious access. It demonstrated that predictions that are in line with sensory evidence accelerate conscious access. Study 2 investigated how low level information within the sensory hierarchy is dealt with by predictive processing and regularity learning mechanisms through “perceptual echo” in which the cross-correlation between a sequence of randomly fluctuating luminance values and occipital electrophysiological (EEG) signals exhibits a long-lasting periodic (~100ms cycle) reverberation of the input stimulus. This study identified a new form of regularity learning and the results demonstrate that the perceptual echo may reflect an iterative learning process, governed by predictive processing. Study 3 investigated how supra-modal predictive processing is capable of learning regularities of temporal duration and also temporal predictions about future events. This study revealed a supramodal temporal prediction mechanism which processes auditory and visual temporal information and integrates information from the duration and rhythmic structures of events. Together these studies provide a global picture of predictive processing and regularity learning across differing types of predictive information.

The sense of agency is the experience of being the initiator of our intentional actions and their outcomes. According to higher order thought theory, a representation becomes conscious when there is a higher order state about it. Thus conscious experience, including that of intentions, is metacognitive. The experience of involuntariness characteristic of hypnotic responding may be attributable to the formation and maintenance of inaccurate metacognitive higher order states of intending. Conversely, the practice of Buddhist mindfulness meditation may develop accurate metacognition, including higher order states of intending. Highly hypnotisable people and mindfulness meditators may therefore occupy two ends of a spectrum of metacognitive ability with regard to unconscious intentions. The presented research investigated predicted trait differences in cognitive tasks which directly or indirectly reflect metacognition of intentions: the timing of an experience of an intention to move and the compressed time interval between a voluntary action and its outcome, known as intentional binding. As an implicit measure of sense of agency, intentional binding was also employed to investigate the veridicality of reports of the experience of involuntariness in hypnotic responding. Additionally, while hypnosis presents a unique opportunity to investigate reliable changes in agentic experience, existing hypnosis screening instruments are time consuming and present a barrier to wider adoption of hypnosis as an instrument for studying consciousness. Here a revised, time-efficient hypnosis screening procedure (the SWASH) is presented. Consistent with predictions, highly hypnotisable groups reported later awareness of motor intentions than less hypnotisable groups and meditators earlier awareness than non-meditators. In an intentional binding task, high hypnotisables showed less binding of an action-outcome toward an action (outcome binding) than low hypnotisables and meditators more outcome binding than non-meditators. Outcome binding was reduced in post-hypnotic involuntary action compared to voluntary action. It is proposed that intentional binding is driven by a cue combination mechanism and that these differences reflect varying precision of motor intention related information in reported timing judgements. The SWASH was found to be a reliable hypnosis screening instrument.

Our thoughts and behaviours can sometimes be influenced by stimuli that we are not consciously aware of having seen. For example, the presentation of a word that is blocked from entering conscious visual perception through masking can subsequently influence the cognitive processing of a further target word. However, the idea that unconscious cognition is sophisticated enough to process the semantic meaning of subliminal stimuli is controversial. This thesis attempts to explore the extent of subliminal priming. Empirical research centering on subjective methods of measuring conscious knowledge is presented in a series of three articles. The first article investigates the subliminal priming of negation. A series of experiments demonstrates that unconscious processing can accurately discriminate between two nouns beyond chance performance when subliminally instructed to either pick or not pick a given noun. This article demonstrates not only semantic processing of the instructional word, but also unconscious cognitive control by following a two-word subliminal instruction to not choose the primed noun. The second article investigates subliminal priming of active versus passive verb voice by presenting a prime sentence denoting one of two characters as either active or passive and asking which of two pictorial representations best matches the prime. The series of experiments demonstrates that overall, participants were able to identify the correct image for both active and passive conditions beyond chance expectations. This article suggests that individuals are able to process the meaning of word combinations that they are not aware of seeing. The third article attempts to determine whether subliminal processing is sophisticated enough to allow for the activation of specific anxieties relating to relationships. Whilst the findings reveal a small subliminal priming effect on generalised anxiety, the evidence regarding the subliminal priming of very specific anxieties is insensitive. The unconscious is shown in these experiments to be more powerful than previously supposed in terms of the fine grained processing of the semantics of word combinations, though not yet in terms of the fine grained resolution of emotional priming.

The following work presents an exploration of human search behaviour both from biological and computational perspectives. Search behaviour is defined as the movements made by an organism while attempting to find a resource. This work describes some of the principal procedures used to record movement, methods for analysing the data and possible ways of interpreting the data. In order to obtain a database of searching behaviour, an experimental setup was built and tested to generate the search paths of human participants. The test arena occupied part of a football field and the targets consisted of an array of 20 golf balls. In the first set of experiments, a random and regular distribution of targets were tested. For each distribution, three distinct conspicuity levels were constructed: a cryptic level, in which targets were painted the same colour as the grass, a semi-conspicuous level in which targets were left white and a conspicuous condition in which the position of each target was marked by a red flag, protruding one metre from the ground. The subjects tested were 9-11 year old children and their search paths were collected using a GPS device. Subjects did not recognise the spatial cues regarding the way targets were spatially distributed. A minimal decision model, the bouncing search model, was built based on the characteristics of the childrens search paths. The model produced an outstanding fit of the children's behavioural data. In the second set of experiments, a new group of children were tested for two new distributions obtained by arranging the targets in patches. Again, children appeared unable to recognise spatial information during the collection processes. The children's behaviour once again produced a good match with that of the bouncing search model. This work introduces several new methodological aspects to be explored to further understand the decision processes involved when humans search. Also, it illustrates that integrating biology and computational science can result in innovative research.

Two key questions underpin the research presented here. Firstly, how does altered higher order awareness contribute to hypnotic experience? Secondly, how do meditation and hypnosis differ in terms of the role of higher order awareness? These questions are addressed here in the form of four papers. In the first paper I review the literatures of hypnosis and meditation in order to consider the similarities and differences between meditation and hypnosis in terms of the role of attentional skill and the neural underpinnings of each. I then draw conclusions regarding the contrasting role of higher order awareness and metacognition in meditation and hypnosis. Paper two explores higher order awareness in hypnosis by comparing the effects of alcohol, compared to placebo, on hypnotisability and associated frontal lobe executive functioning. Paper three compares meditation and hypnosis by investigating differences in higher order thoughts, mindfulness, absorption and perceptual encoding style as revealed by self-report measures. The final paper takes a broader look at higher order awareness and its relation to the experience of agency and involuntariness in hypnotic suggestion using a Libet type paradigm.

In this thesis I explore functions of the neuromodulator dopamine in the context of autonomous learning and behaviour. I first investigate dopaminergic influence within a simulated agent-based model, demonstrating how modulation of synaptic plasticity can enable reward-mediated learning that is both adaptive and self-limiting. I describe how this mechanism is driven by the dynamics of agentenvironment interaction and consequently suggest roles for both complex spontaneous neuronal activity and specific neuroanatomy in the expression of early, exploratory behaviour. I then show how the observed response of dopamine neurons in the mammalian basal ganglia may also be modelled by similar processes involving dopaminergic neuromodulation and cortical spike-pattern representation within an architecture of counteracting excitatory and inhibitory neural pathways, reflecting gross mammalian neuroanatomy. Significantly, I demonstrate how combined modulation of synaptic plasticity and neuronal excitability enables specific (timely) spike-patterns to be recognised and selectively responded to by efferent neural populations, therefore providing a novel spike-timing based implementation of the hypothetical ‘serial-compound' representation suggested by temporal difference learning. I subsequently discuss more recent work, focused upon modelling those complex spike-patterns observed in cortex. Here, I describe neural features likely to contribute to the expression of such activity and subsequently present novel simulation software allowing for interactive exploration of these factors, in a more comprehensive neural model that implements both dynamical synapses and dopaminergic neuromodulation. I conclude by describing how the work presented ultimately suggests an integrated theory of autonomous learning, in which direct coupling of agent and environment supports a predictive coding mechanism, bootstrapped in early development by a more fundamental process of trial-and-error learning.

One of the central tenets of the embodied theory of language comprehension is that the process of understanding prompts the same perceptuo-motor activity involved in actual perception and action. This activity is a component of comprehension that is not memory–dependent and is hypothesized to be intact in Alzheimer's Disease (AD). Each article in this thesis is aimed at answering the question whether individuals with probable AD, healthy older adults and younger adults show differences in their performance on tests where perceptual and motoric priming take place during language comprehension. The second question each article asks is whether language comprehension in AD can be facilitated by the specific use of this perceptual and motoric priming. Article I examines whether the way individuals with pAD represent verbs spatially matches the way healthy older and younger adults do, and how stable these representations are. It also explores in what way spatial representations may relate to verb comprehension, more specifically, whether representations matching the norms translate into a better quality of verb comprehension. Article II tests the interaction between the verbs' spatial representations taking place during comprehension and perceptual cues - compatible and incompatible to the representations - in order to investigate whether individuals with pAD show differences in susceptibility to perceptual cues, compared to healthy older and younger participants. The second aim of this article is to explore in what way performance on a word-picture verification task can be affected, with reference to the fact that in previous studies on young participants, both priming and interference have resulted from the interaction of linguistic and perceptual processing. Article III explores the Action Compatibility Effect (ACE) (Glenberg & Kaschak, 2002) with the aim of finding out whether the ACE exists for volunteers with pAD and whether it can facilitate language comprehension. The order of presentation of language and movement is manipulated to establish whether there is a reciprocal relationship between them. This information could be crucial in view of possible applications to individuals with pAD. These articles test, for the first time, the effects of the manipulation of the perceptuo-motor component during language comprehension in individuals with pAD; they are intended as a methodological exploration contributing to a better understanding of the potential of embodiment principles to support language comprehension changes associated with pAD. Embodiment effects need to be studied further with a view to putting them to use in either clinical or real-life applications.

It is often reported that after amputation people experience "a constant or inconstant ... sensory ghost ... faintly felt at time, but ready to be called up to [their] perception" (Mitchell, 1866). Perceptual abnormalities have been highlighted in amputees, such as sensations in the phantom when being stroked elsewhere (Ramachandran et al., 1992) or when observing someone in pain (Giummarra and Bradshaw, 2008). This thesis explored the perceptual changes that occur following amputation whist focusing on pain, vision and touch. A sample of over 100 amputees were recruited through the National Health Service. Despite finding no difference in phantom pain based on physical amputation details or nonpainful perceptual phenomena, results from Paper 1 indicated that phantom pain may be more intense, with sensations occurring more frequently, in amputees whose pain was triggerinduced. The survey in Paper 2 identified a group of amputees who in losing a limb acquired mirror-touch synaesthesia. Higher levels of empathy found in mirror-touch amputees might mean that some people are predisposed to develop synaesthesia, but that it takes sensory loss to bring dormant cross-sensory interactions into consciousness. Although the mirror-system may reach supra-threshold levels in some amputees, the experiments in Paper 3 suggested a relatively intact mirror-system in amputees overall. Specifically, in a task of apparent biological motion, amputees showed a similar, although weaker, pattern of results to normalbodied participants. The results of Paper 4 showed that tactile spatial acuity on the face was also largely not affected by amputation, as no difference was found between the sides ipsilateral and contralateral to the stump. In Paper 5 cross-modal cuing was used to investigate whether referred tactile sensations could prime a visually presented target in space occupied by the phantom limb. We conclude that perception is only moderately affected in most amputees, but that in some the sensory loss causes normally sub-threshold processing to enhance into conscious awareness.

Face adaptation techniques have been used extensively to investigate how faces are processed. It has even been suggested that face adaptation is functional in calibrating the visual system to the diet of faces to which an observer is exposed. Yet most adaptation studies to date have used unfamiliar faces: few have used faces with real world familiarity. Familiar faces have more abstractive representations than unfamiliar faces. The experiments in this thesis therefore examined face adaptation for familiar faces. Chapters 2 and 3 explored the role of explicit recognition of familiar faces in producing face identity after-effects (FIAEs). Chapter 2 used composite faces (the top half of a celebrity's face paired with the bottom half of an unfamiliar face) as adaptors and showed that only recognised composites produced significant adaptation. In Chapter 3 the adaptors were cryptic faces (unfamiliar faces subtly transformed towards a celebrity's face) and faces of celebrity's siblings. Unrecognised cryptic and sibling faces produced FIAEs for their related celebrity, but only when adapting and testing on the same viewpoint. Adaptation only transferred across viewpoint when a face was explicitly recognised. Chapter 4 demonstrated that face adaptation could occur for ecologically valid, personally familiar stimuli, a necessary pre-requisite if adaptation is functional in calibrating face processing mechanisms. A video of a lecturer's face produced FIAEs equivalent to that produced by static images. Chapters 5 and 6 used a different type of after-effect, the face distortion after-effect (FDAE), to explore the stability of our representations for personally familiar faces, and showed that even representations of highly familiar faces can be affected by exposure to distorted faces. The work presented here shows that it is important to take facial familiarity into account when investigating face adaptation effects, as well as increasing our understanding of how familiarity affects the representations of faces.

In human fear conditioning 'resistance to extinction' occurs when the removal of the aversive outcome fails to produce a reduction in conditioned responding. This phenomenon is important to understanding the persistence of anxiety disorders such as phobias. The research presented in this thesis examines factors that promote the acquisition and maintenance of learned fear response and attempts to differentiate between different explanations of the resistance to extinction phenomenon. To investigate the impact of different conditioning procedures (evaluative or classical conditioning) on the durability of the conditioned response (CR), event-related potential (ERP) methodology was employed. In addition, the role of the fear-relevance of the conditioned stimulus (CS), in supporting the acquisition and resistance to extinction of the CR, was explored. Evidence suggested that extinction effects are likely to reflect procedural differences in conditioning rather than different underlying learning processes. Extinction effects were dissociable across procedures, supporting the role of the type of unconditioned stimulus (US) in explaining past demonstrations of extinction when responses were indexed by physiological measures. Verbally transmitted, threat information heightened aversive US-expectancies and fear beliefs without the need for conditioning. Additionally, fear-beliefs were reduced without the need for extinction training when positive information was provided. Contrary to Davey's (1997) expectancy bias model, the results do not support the hypothesis that verbal information interacts with direct contingency experience to create fear responses; instead, information appears to be a direct pathway to fear. ERP measures for fear responses did not echo the effects of verbal information and contingency on fear-beliefs. However, the comparability of our ERP data, to other research using physiological measures of response, is discussed regarding the number of trials required to calculate the average ERP response. Due to averaging over a large number of trials the ERP measure may not be sensitive to fluctuations in response that may be dependent on information or contingency manipulations. In conclusion our data suggests the importance of verbal information as a pathway to fear and the role of cognitive factors in the prevention and treatment of fears.