Coordinating our actions with others changes how we behave and feel. Here, we provide evidence that interacting with others rests on a balance between self-other integration and segregation. Using a group walking paradigm, participants were instructed to synchronize with a metronome while listening to the sounds of 8 virtual partners. By manipulating the similarity and synchronicity of the partners’ steps to the participant’s own, our novel auditory task disentangles the effects of synchrony and self-other similarity and examines their contribution to both collective and individual awareness. We measured temporal coordination (step timing regularity and synchrony with the metronome), gait patterns, and subjective reports about sense of self and group cohesion. The main findings show that coordination is best when participants hear distinct but synchronous virtual others, leading to greater subjective feelings of agency, strength, dominance, and happiness.

Monitoring actions is essential for goal-directed behavior. However, as opposed to short-lasting, and regularly reinstating monitoring functions, the neural processes underlying continuous actionmonitoring are poorly understood. We investigate this using a pursuit-tracking paradigm. We show that beta band activity likelymaintains the sensorimotor program, while theta and alpha bands probably support attentional sampling and information gating, respectively. Alpha and beta band activity are most relevant during the initial tracking period, when sensorimotor calibrations are most intense. Theta band shifts from parietal to frontal cortices throughout tracking, likely reflecting a shift in the functional relevance from attentional sampling to action monitoring. This study shows that resource allocation mechanisms in prefrontal areas and stimulus-response mapping processes in the parietal cortex are crucial for adapting sensorimotor processes. It fills a knowledge gap in understanding the neural processes underlying action monitoring and suggests new directions for examining sensorimotor integration in more naturalistic experiments.

It is now widely accepted that the experience of pain is subject to cognitive influences that may determine the severity of subjectively perceived pain. Many of these top-down factors rely on memory-based processes, which in turn are related to prior experience, learned beliefs and behaviours about pain. As such, memory for pain heavily contributes to the physical pain experience. We posit that pain memory is bidirectional in that following each painful event a trace is stored and that these traces in turn may modify future pain perception prospectively. The following body of work explores aspects of what we have termed a memory template for pain. The results of these chapters taken together, examine these bidirectional aspects of short-term memory for pain employing a recall pain task. Specifically, we explore how, after an acute pain event, a short-term mental representation of the initial event persists. We show that during this time, sensory re-experiencing of the painful event is possible. Furthermore, we investigate aspects of recalled pain, namely intensity and vividness. Data suggests that the intensity and the vividness of this mental representation are determined by the intensity of the initial stimulus, as well as the time-to-test delay. We identify regions that characterise short-term memory for pain. Following on from studies in motor and visual imagery, we explore how pain imagery in the form of recall may affect subsequent pain perception. Our results demonstrate that the inclusion of pain-related imagery preceding physical pain events reduces affective qualities of pain. Testing healthy, naïve subjects, we replicate the effect observed in studies using attention management and imagery strategies, which normally require extensive training. Finally, in a cohort of neuropathic pain patients we show significant reductions in white matter connectivity between areas responsible for working and prospective memory. Collectively, these studies emphasise and elucidate the role of short-term memory of pain in physical pain perception. Acting both retrospectively and prospectively, cognitive reinforcement can increase or decrease the subjective feeling of pain, and therefore manipulating how pain is recalled may have therapeutic potential.

Behaviour is guided by accumulated experience, valuation and comparison. While many aspects associated with these functions are mediated by the frontal lobes, the precise contribution from particular regions remains debated. This thesis will deal with how an organism comes to select an option and will specifically focus on the role of the orbitofrontal cortex (OFC) in two mechanisms in this process: learning of outcome specificities and selecting between multiple options based on their expected values. Despite evidence emphasizing anatomical and connective heterogeneity within this structure, the OFC is often regarded as a uniform region. This thesis aims to resolve some of this uncertainty by assuming that the medial and lateral regions of the OFC contribute differentially to learning and decision-making. Two distinct methodologies were used in these investigations. First, the contribution of the medial OFC to social and emotional processing was examined. The findings from this study disprove previously held beliefs that the medial regions of the OFC guide social and emotional behaviours, but indicted a role for this region in value-guided decision-making. The second study examined functional differences between the lateral and medial OFC by making circumscribed lesions to either region in macaque monkeys. The animals performed a number of 3-armed bandit tasks which were designed to investigate different aspects of value assignment and comparison. The results show that while lateral OFC was required for "credit assignment" – the correct assignment of values to visual cues – medial OFC was critical for comparison of the cues' values during decision-making. In unchanging probabilistic environments, mOFC lesions induced decision-making impairments when value comparison was difficult without affecting credit assignment and associative learning. By contrast, lateral OFC lesions caused the opposite pattern of impairment. The final study used human-neuroimaging techniques to investigate the differential representation of outcome-specific contingency learning and found not only that the expectation of a unique outcome facilitated learning and memory recall but that this was supported by a neural network which included the lateral regions of the OFC and the anterior cingulate cortex. Activity in the mOFC did not correlate with outcome-specific contingency learning but instead reflected both the value associated with the receipt and expectation of a reward. Taken together, the results from this thesis suggest that specific parts of the OFC make markedly different contributions to these very different cognitive functions.

Overexpression of the NR2B subunit of the NMDA receptor in the forebrain has been shown to improve learning and memory in mice (Tang et al 1999), which provides exciting implications for the enhancement of human cognition. However, it was first essential to establish replicability, and since the Tang et al (1999) study used only male mice we wished to investigate possible sex differences. On the hidden platform watermaze, we found a trend for male NR2BOE mice to learn the task more quickly than male wildtype mice (as observed by Tang et al. 1999), but the opposite trend in female mice; female NR2BOE mice were slower to reach the hidden platform than female wildtype mice. This pattern of results was also observed on the spatial reference Y memory task and open field task (for anxiety), although not on the spatial working memory T maze task (despite a sex difference). However, wildtype and NR2BOE mice performed at similar levels on the novel object recognition task, the spatial novelty preference task, visible platform watermaze and visual discrimination task. A battery of tests considering some species typical behaviours of mice demonstrated that wildtype and NR2BOE mice were comparable on tests of motor ability, strength, co-ordination, anxiety, burrowing and nesting. This suggests that our behavioural results are not due to a general impairment or enhancement of species typical behaviours. We considered the possibility that the difference between the results of Tang et al (1999) and those we observed may be caused by age differences; hence we attempted to replicate our results on the hidden platform watermaze, spatial reference Y maze and open field test in age matched mice. However, the second cohort of NR2BOE mice performed at similar levels to wildtype mice, and at significantly improved levels compared to the mice of the first cohort. We also considered the effects of knocking out the NR2B subunit on learning and memory, and NR1 subunit deletion within the hippocampus. On the spatial working memory T maze, these mouse strains performed similarly to their respective wildtype strains. Similarly, on a two beacon watermaze (with one indicating the platform position), mice lacking the NR2B subunit were able to locate the platform in a similar length of time. To ensure that the null results we had observed in the second cohort were not due to loss of the NR2B protein overexpression in the forebrain, we performed polymerase chain reactions (PCR), quantitative real-time PCR, and Western blots. We ascertained that the transgene was indeed present and that NR2B mRNA and protein levels were elevated in the hippocampi of the NR2BOE mice. In conclusion, it is unclear why the behaviours we observed in the NR2BOE mice are different to those published in the literature. It is possible that they may be due to differences in environmental enrichment, but the cause of the genotype by sex differences observed in the mice of cohort 1 is unclear. Nonetheless, we have advanced our knowledge of the effects of modifications in the levels of the NR2B subunit of the NMDA receptor on learning and behaviour.

Selective attention is a mechanism used to allocate resources to information processing. Both mood states and emotionally salient stimuli can influence which information is selectively attended. This information is subsequently processed in a more elaborative manner and affects task performance. The experiments presented in this thesis explore the influence of mood and emotional stimuli on selective attention and consequently task performance. Mood induction procedures were used to induce transient neutral, sad, and happy mood states in healthy volunteers. A systematic review and meta-analysis of 41 studies using sad mood induction procedures showed cognitive impairments in performance in the context of task neutral stimuli. In contrast biases in attention towards mood-congruent negative stimuli led to improved task performance. A series of three behavioural experiments with 197 participants demonstrated that participants made decisions on the basis of less information when that information was preceded by emotional but not neutral stimuli. Induced mood state did not affect performance. The behavioural and neural correlates of visual attentional processing to emotional stimuli were explored using magnetoencephalography in 24 healthy participants following sad, happy, and neutral mood induction procedures. The M300, a component associated with selective attention, had greater amplitude following presentation of negative compared with positive stimuli, which was associated with improved task performance. Reduced M300 amplitude and impairments in performance occurred following sad mood induction procedures. The experiments presented in this thesis demonstrate prioritized processing of emotional information and provide some evidence for impaired performance following sad mood induction procedures.

The notion that we are voluntary agents who exercise power to choose and, in doing so, determine some of what happens in the world has been an important notion in certain theological accounts concerning our relationship with God (e.g. 'the freewill defence' for God's goodness and omnipotence in light of moral evil and accounts of human moral responsibility in relation to God). However, it has been claimed that the physicalism supported by contemporary neuroscience research calls into question human voluntary agency and, with it, human power to choose. Emergentist (or non-reductive physicalist) accounts of psychological phenomena have been presented as a way of reconciling the physicalism supported by contemporary neuroscience and the theologically important notion of human power to choose. But there are several issues that remain for the plausibility of the required kind of emergentist account; namely - Does recent neuroscience research show that voluntary agency is an illusion? and Is there evidence for neurophysiological causes which, along with neurophysiological conditions, determine all we do? In this dissertation I set out to address these issues and, in doing so, present an account of voluntary agency as power to choose in the state of being aware of alternatives. I argue that this account allows for the notion that human beings determine some of what happens in a way that is consistent with what contemporary neuroscience shows. Thus, contemporary neuroscience does not undermine this notion of human voluntary agency; or, then, the predominant theological view that we are morally responsible in our relationship with God.

In everyday life we regularly update our expectations about the locations at which sensory events may occur, and about the motor responses that are appropriate in a given situation. The experiments in this thesis investigated the neural correlates of perceptual processes and motor preparation during human decision making, and the regions that causally contribute to decision making in the human brain. In Chapter 3, I used electroencephalography (EEG) to investigate whether alpha-band (~8-14 Hz) oscillations provide a graded index of participants’ preparatory attentional states. Time-frequency analysis revealed that manipulating spatial certainty regarding the location of an upcoming visual target led to parametric changes in the lateralization of preparatory occipito-parietal alpha oscillations, and to parametric modulation of parieto-central beta-band (~15-25 Hz) power typically associated with response preparation. In Chapter 4, I used EEG to investigate whether evolution of lateralization of sensorimotor alpha- and beta-band activity reflected participants’ evolving expectations about an upcoming motor response. Lateralization of activity in both frequency bands varied parametrically with the available evidence, suggesting such lateralized activity correlates with participants’ internal decision variables. Further analysis identified unique contributions to lateralized and non-lateralized oscillatory activity due to the prior evidence, evidence update, and surprise related to the observed information at each stage of the task. In Chapter 5, I extended the paradigm developed in Chapter 4 for use with online repetitive transcranial magnetic stimulation (TMS) and concurrent EEG recording. Delivery of TMS during decision making allowed investigation of the causal role played by a left hemisphere medial intraparietal region that is the putative human homologue of the macaque medial intraparietal cortex (MIP). MIP stimulation disrupted decision-making behaviour by biasing participants’ decisions against contralateral-to-stimulation (i.e., right-handed) responses. Comparison of the magnitude of TMS-induced changes in behaviour and beta-band activity demonstrated that the intraparietal cortex plays a causal role both in decision making and in the appearance of beta-band activity over the motor cortex. In Chapter 6, the broader consequences of the experimental work presented in this thesis are discussed, in addition to promising directions for future research.

Accelerated long-term forgetting (ALF) is a recently described memory impairment associated with epilepsy. Patients with ALF appear to learn and initially retain new information normally, but forget it at an accelerated rate over subsequent days. ALF can have a profound impact on the lives of the people who suffer from it, but it is also of theoretical interest. In particular, the study of this disorder may provide insight into the mechanisms of memory consolidation. ALF is especially prevalent in transient epileptic amnesia (TEA), an epileptic syndrome in which the seizure focus is thought to be the medial temporal lobes (MTL). The MTL house the hippocampus and a number of other structures critical for declarative memory function. The aims of this doctoral thesis were to investigate which aspects of memory function are disrupted in patients with TEA-associated ALF, and to shed light on the neural basis of the memory impairment. Slow wave sleep (i.e. deep sleep) is known to exacerbate epileptic activity. It is also thought to play a key role in the consolidation of declarative memory. The most commonly posited explanation of ALF is the disruption of sleep- dependent memory consolidation. However, it remains possible that ALF is caused by a subtle problem with encoding that usually goes undetected until delayed memory tests. The results of this thesis demonstrate that sleep can actually benefit memory retention in TEA ALF patients just as much as it does in healthy people, and that it is not necessary for the retention interval to contain sleep in order for ALF to be seen. However, the relationship between slow wave sleep and memory was found to be abnormal in these patients. The amount of slow wave sleep, and the power in the slow oscillation frequency range, during the post-learning night correlated negatively with the benefit of that night of sleep for memory retention. Furthermore, resting-state brain activity patterns thought to reflect post-encoding memory reprocessing were found to correlate negatively with subsequent memory performance in these patients. Another chapter of this thesis provides evidence that TEA ALF patients encode memories abnormally; these patients showed reduced activity in the left hippocampus while viewing stimuli that they went on to forget. Furthermore, this encoding-related brain activity correlated with their long-term forgetting. The final experimental chapter reports a correlation in these patients between grey matter in the left hippocampus and long-term forgetting, which cannot entirely account for the encoding-related brain activity results. The hippocampus and its surrounding structures are thought to be critical to our ability to discriminate between similar stimuli and events. An intriguing hypothesis consistent with the pattern of results in this thesis is that ALF is caused by a functional impairment of the MTL that results in a diminished capacity to distinguish between similar experiences, ultimately causing memory problems; abnormally formed memories may interact with new material and memory consolidation processes in an aberrant manner, leading to retrieval deficits.

The natural world provides sensory systems with noisy and ambiguous information, which is often transformed into a more stable categorical percept. This thesis aims to investigate the nature of the neural representations in the visual system that support this transformation. To do so, we will employ a behavioural task that requires participants to average several independent sources of perceptual information. This task allows for the dissociation of two theoretically orthogonal sources of decision uncertainty: the mean distance of the perceptual information from a category boundary and the variability of the evidence under consideration. Behaviourally, both decreasing the mean distance to bound of information and increasing information variability are associated with increased errors and prolonged response times. We will present a computational model that can account for the independent behavioural effects of these two sources of uncertainty by assuming that categorical decisions are made on the basis of a probabilistic transformation of perceptual evidence. BOLD measurements demonstrate that these effects of mean and variability are supported by a partially dissociable network of brain regions. Electroencephalography demonstrates the differential influence of mean and variance in the pre- and post-decision period. Furthermore, we show that there is adaptation at the level of the perceptual representation to the information variance. Not only does this show that the visual system must represent information at the summary level, in addition to individual feature-based representation, but it also suggests that the costs associated with this form of perceptual uncertainty can be largely mitigated by the adoption of a more suitable representational range.