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UNLABELLED: Sensory signals are highly structured in both space and time. These regularities allow expectations about future stimulation to be formed, thereby facilitating decisions about upcoming visual features and objects. One such regularity is that the world is generally stable over short time scales. This feature of the world is exploited by the brain, leading to a bias in perception called serial dependence: previously seen stimuli bias the perception of subsequent stimuli, making them appear more similar to previous input than they really are. What are the neural processes that may underlie this bias in perceptual choice? Does serial dependence arise only in higher-level areas involved in perceptual decision-making, or does such a bias occur at the earliest levels of sensory processing? In this study, human subjects made decisions about the orientation of grating stimuli presented in the left or right visual field while activity patterns in their visual cortex were recorded using fMRI. In line with previous behavioral reports, reported orientation on the current trial was consistently biased toward the previously reported orientation. We found that the orientation signal in V1 was similarly biased toward the orientation presented on the previous trial. Both the perceptual decision and neural effects were spatially specific, such that the perceptual decision and neural representations on the current trial were only influenced by previous stimuli at the same location. These results suggest that biases in perceptual decisions induced by previous stimuli may result from neural biases in sensory cortex induced by recent perceptual history. SIGNIFICANCE STATEMENT: We perceive a stable visual scene, although our visual input is constantly changing. This experience may in part be driven by a bias in visual perception that causes images to be perceived as similar to those previously seen. Here, we provide evidence for a sensory bias that may underlie this perceptual effect. We find that neural representations in early visual cortex are biased toward previous perceptual decisions. Our results suggest a direct neural correlate of serial dependencies in visual perception. These findings elucidate how our perceptual decisions are shaped by our perceptual history.

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Stimulus expectation can modulate neural responses in early sensory cortical regions, with expected stimuli often leading to a reduced neural response. However, it is unclear whether this expectation suppression is an automatic phenomenon or is instead dependent on the type of task a subject is engaged in. To investigate this, human subjects were presented with visual grating stimuli in the periphery that were either predictable or non-predictable while they performed three tasks that differently engaged cognitive resources. In two of the tasks, the predictable stimulus was task-irrelevant and spatial attention was engaged at fixation, with a high load on either perceptual or working memory resources. In the third task, the predictable stimulus was task-relevant, and therefore spatially attended. We observed that expectation suppression is dependent on the cognitive resources engaged by a subjects' current task. When the grating was task-irrelevant, expectation suppression for predictable items was visible in retinotopically specific areas of early visual cortex (V1-V3) during the perceptual task, but it was abolished when working memory was loaded. When the grating was task-relevant and spatially attended, there was no significant effect of expectation in early visual cortex. These results suggest that expectation suppression is not an automatic phenomenon, but dependent on attentional state and type of available cognitive resources.

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The ability to recognize one's own successful cognitive processing, in e.g., perceptual or memory tasks, is often referred to as metacognition. How should we quantitatively measure such ability? Here we focus on a class of measures that assess the correspondence between trial-by-trial accuracy and one's own confidence. In general, for healthy subjects endowed with metacognitive sensitivity, when one is confident, one is more likely to be correct. Thus, the degree of association between accuracy and confidence can be taken as a quantitative measure of metacognition. However, many studies use a statistical correlation coefficient (e.g., Pearson's r) or its variant to assess this degree of association, and such measures are susceptible to undesirable influences from factors such as response biases. Here we review other measures based on signal detection theory and receiver operating characteristics (ROC) analysis that are "bias free," and relate these quantities to the calibration and discrimination measures developed in the probability estimation literature. We go on to distinguish between the related concepts of metacognitive bias (a difference in subjective confidence despite basic task performance remaining constant), metacognitive sensitivity (how good one is at distinguishing between one's own correct and incorrect judgments) and metacognitive efficiency (a subject's level of metacognitive sensitivity given a certain level of task performance). Finally, we discuss how these three concepts pose interesting questions for the study of metacognition and conscious awareness.

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Predictive coding models suggest that predicted sensory signals are attenuated (silencing of prediction error). These models, though influential, are challenged by the fact that prediction sometimes seems to enhance rather than reduce sensory signals, as in the case of attentional cueing experiments. One possible explanation is that in these experiments, prediction (i.e., stimulus probability) is confounded with attention (i.e., task relevance), which is known to boost rather than reduce sensory signal. However, recent theoretical work on predictive coding inspires an alternative hypothesis and suggests that attention and prediction operate synergistically to improve the precision of perceptual inference. This model posits that attention leads to heightened weighting of sensory evidence, thereby reversing the sensory silencing by prediction. Here, we factorially manipulated attention and prediction in a functional magnetic resonance imaging study and distinguished between these 2 hypotheses. Our results support a predictive coding model wherein attention reverses the sensory attenuation of predicted signals.

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Liberal acceptance, overconfidence, and increased activity of the neurotransmitter dopamine have been proposed to account for abnormal sensory experiences, for instance, hallucinations in schizophrenia. In normal subjects, increased sensory experience in Yoga Nidra meditation is linked to striatal dopamine release. We therefore hypothesize that the neurotransmitter dopamine may function as a regulator of subjective confidence of visual perception in the normal brain. Although much is known about the effect of stimulation by neurotransmitters on cognitive functions, their effect on subjective confidence of perception has never been recorded experimentally before. In a controlled study of 24 normal, healthy female university students with the dopamine agonist pergolide given orally, we show that dopaminergic activation increases confidence in seeing rapidly presented words. It also improves performance in a forced-choice word recognition task. These results demonstrate neurotransmitter regulation of subjective conscious experience of perception and provide evidence for a crucial role of dopamine.

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It was suggested over 20 years ago that the supplementary motor cortex is involved in self-generated behaviour. Since then, there have been many studies using electrophysiology and brain imaging of the role of the supplementary motor cortex and anterior cingulate cortex. In light of the findings, the proposal that these regions are crucial for self-generated action has recently been challenged. Here, we review the recent literature and argue that the proposal survives the findings. We further argue that it can be generalised to cover reflection on mental states. Finally, we suggest that the pattern of anatomical connections is consistent with the proposal that the medial frontal cortex is crucially involved in self-generated action and self-reflection.

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A very basic computational model is proposed to explain two puzzling findings in the time perception literature. First, spontaneous motor actions are preceded by up to 1-2 s of preparatory activity (Kornhuber and Deecke, 1965). Yet, subjects are only consciously aware of about a quarter of a second of motor preparation (Libet et al., 1983). Why are they not aware of the early part of preparation? Second, psychophysical findings (Spence et al., 2001) support the principle of attention prior entry (Titchener, 1908), which states that attended stimuli are perceived faster than unattended stimuli. However, electrophysiological studies reported no or little corresponding temporal difference between the neural signals for attended and unattended stimuli (McDonald et al., 2005; Vibell et al., 2007). We suggest that the key to understanding these puzzling findings is to think of onset detection in probabilistic terms. The two apparently paradoxical phenomena are naturally predicted by our signal detection theoretic model.

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Humans are unique in being able to reflect on their own performance. For example, we are more motivated to do well on a task when we are told that our abilities are being evaluated. We set out to study the effect of self-motivation on a working memory task. By telling one group of participants that we were assessing their cognitive abilities, and another group that we were simply optimizing task parameters, we managed to enhance the motivation to do well in the first group. We matched the performance between the groups. During functional magnetic resonance imaging, the motivated group showed enhanced activity when making errors. This activity was extensive, including the anterior paracingulate cortex, lateral prefrontal and orbitofrontal cortex. These areas showed enhanced interaction with each other. The anterior paracingulate activity correlated with self-image ratings, and overlapped with activity when participants explicitly reflected upon their performance. We suggest that the motivation to do well leads to treating errors as being in conflict with one's ideals for oneself.

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It is usually taken as given that consciousness involves superior or more elaborate forms of information processing. Contemporary models equate consciousness with global processing, system complexity, or depth or stability of computation. This is in stark contrast with the powerful philosophical intuition that being conscious is more than just having the ability to compute. I argue that it is also incompatible with current empirical findings. I present a model that is free from the strong assumption that consciousness predicts superior performance. The model is based on Bayesian decision theory, of which signal detection theory is a special case. It reflects the fact that the capacity for perceptual decisions is fundamentally limited by the presence and amount of noise in the system. To optimize performance, one therefore needs to set decision criteria that are based on the behaviour, i.e. the probability distributions, of the internal signals. One important realization is that the knowledge of how our internal signals behave statistically has to be learned over time. Essentially, we are doing statistics on our own brain. This 'higher-order' learning, however, may err, and this impairs our ability to set and maintain optimal criteria for perceptual decisions, which I argue is central to perception consciousness. I outline three possibilities of how conscious perception might be affected by failures of 'higher-order' representation. These all imply that one can have a dissociation between consciousness and performance. This model readily explains blindsight and hallucinations in formal terms, and is beginning to receive direct empirical support. I end by discussing some philosophical implications of the model.

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The author has previously claimed that neural activity in a medial frontal brain area represents conscious motor intentions. A. Machado and F. J. Silva attempted to challenge this claim by arguing that such intentions are not necessarily causally effective and must be caused by something else, so that they therefore cannot be the unmoved first movers in action. The author's reply is that he made no such claims about the causal status and origin of conscious intentions. In fact, he has elsewhere presented empirical evidence in support of the idea that intentions are not necessarily causal. But this does not stop researchers from studying the neural basis for the conscious impression that one intends and wills one's actions. One can feel and attend to such intentions, be they causal or not. The author's claim is that there is neural activity in the medial frontal wall that reflects such conscious feelings. Other recent empirical evidence that supports this view is described.

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Using functional magnetic resonance imaging, we tested whether unconscious information can influence the cognitive control system in the human prefrontal cortex. Volunteers had to prepare to perform either a phonological judgment or a semantic judgment on an upcoming word, based on the instruction given at the beginning of each trial. However, in some trials they were visually primed to prepare for the alternative (i.e., "wrong") task, and this impaired their performance. This priming effect is taken to depend on unconscious processes because the effect was present even when the volunteers could only discriminate the identity of the primes at chance level. Furthermore, the effect was stronger when the visibility of the prime was near zero than when the visibility of the prime was significantly higher. When volunteers were unconsciously primed to perform the alternative task, there was also decreased neural activity in the brain areas relevant to the instructed task and increased neural activity in the brain areas relevant to the alternative task, which shows that the volunteers were actually engaged in the wrong task, instead of simply being distracted. Activity in the mid-dorsolateral prefrontal cortex was also found to be associated with this unconscious priming effect. These results suggest that the cognitive control system in the prefrontal cortex is not exclusively driven by conscious information, as has been believed previously.

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Using transcranial magnetic stimulation (TMS), we have tested the time needed for the perceived onset of spontaneous motor intention to be fully determined. We found that TMS applied over the presupplementary motor area after the execution of a simple spontaneous action shifted the perceived onset of the motor intention backward in time, and shifted the perceived time of action execution forward in time. The size of the effect was similar regardless of whether TMS was applied immediately after the action or 200 msec after. The results of three control studies suggest that this effect is time-limited, specific to modality, and also specific to the anatomical site of stimulation. We conclude that the perceived onset of intention depends, at least in part, on neural activity that takes place after the execution of action. A model, which is based on the mechanism of cue integration under the presence of noise, is offered to explain the results. The implications for the conscious control of spontaneous actions are discussed.

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By using a paradigm based on metacontrast masking, we created experimental conditions in which the subjective report of consciousness differs but the objectively measured ability to discriminate visual targets does not. This approach allowed us to study the neural correlate of consciousness while having performance levels carefully matched in healthy human subjects. A comparison of the neural activity associated with these conditions as measured by functional MRI showed that conscious perception is associated with spatially specific activity in the mid-dorsolateral prefrontal cortex (area 46). Further analysis confirms that this activation is not only free from any performance confound, but is also not driven by differences in the timing of the physical stimuli. Our results suggest that the prefrontal cortex is important for the essentially subjective aspects of conscious perception.

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We investigated the neural mechanisms underlying the timing procedure that was devised by Libet et al. (1983) to measure the onset of conscious motor intentions in spontaneous actions. We previously showed that, when participants were required to estimate the onset of their intentions using this procedure, the activity in the presupplementary motor area (pre-SMA) was enhanced. Here, we show that when participants were required to estimate the onset of their motor executions (instead of their intentions), the activity in the cingulate motor area was enhanced. Across participants, the degree of this neural enhancement was correlated with the degree of perceptual bias: the higher the degree of enhancement, the earlier the perception. Analysis of data from a previous experiment suggests that the same principle holds true for the relationship between the perceived onset of intentions and the activity in the pre-SMA. We therefore argue that the timing method of Libet et al. (1983) is problematic, because the measuring process affects the neural representations of action and thus also the perceived onsets that the method is designed to measure.

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Filmmakers have long recognized the importance of editing techniques to guide the audiences' perceptions and enhance the impact of a scene. We demonstrate behaviorally that pairing identical faces with either neutral or emotionally salient contextual movies, an editing technique referred to as the 'Kuleshov Effect', results in both altered attributions of facial expression and mental-state. Using functional neuroimaging (fMRI), we show that faces paired with emotional movies enhance BOLD responses in the bilateral temporal pole, anterior cingulate cortices, amygdala and bilateral superior temporal sulcus relative to identical faces juxtaposed with neutral movies. An interaction was observed in the right amygdala when subtle happy and fear faces were juxtaposed with positive and negative movies, respectively. An interaction between happy faces and negative context was also observed in bilateral amygdala suggesting that the amygdala may act to prime or tag affective value to faces. A parametric modulation of BOLD signal by attribution ratings indicated a dissociation between ventrolateral and the ventromedial prefrontal cortex for negative and positive contextually evoked attributions, respectively. These prefrontal regions may act to guide appropriate choices across altering contexts. Together, these findings offer a neurobiological basis for contextual framing effects on social attributions.

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Intention is central to the concept of voluntary action. Using functional magnetic resonance imaging, we compared conditions in which participants made self-paced actions and attended either to their intention to move or to the actual movement. When they attended to their intention rather than their movement, there was an enhancement of activity in the pre-supplementary motor area (pre-SMA). We also found activations in the right dorsal prefrontal cortex and left intraparietal cortex. Prefrontal activity, but not parietal activity, was more strongly coupled with activity in the pre-SMA. We conclude that activity in the pre-SMA reflects the representation of intention.

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