The link between brain learning, attention, and consciousness

The processes whereby our brains continue to learn about a changing world in a stable fashion throughout life are proposed to lead to conscious experiences. These processes include the learning of top-down expectations, the matching of these expectations against bottom-up data, the focusing of attention upon the expected clusters of information, and the development of resonant states between bottom-up and top-down processes as they reach an attentive consensus between what is expected and what is there in the outside world. It is suggested that all conscious states in the brain are resonant states and that these resonant states trigger learning of sensory and cognitive representations. The models which summarize these concepts are therefore called Adaptive Resonance Theory, or ART, models. Psychophysical and neurobiological data in support of ART are presented from early vision, visual object recognition, auditory streaming, variable-rate speech perception, somatosensory perception, and cognitive-emotional interactions, among others. It is noted that ART mechanisms seem to be operative at all levels of the visual system, and it is proposed how these mechanisms are realized by known laminar circuits of visual cortex. It is predicted that the same circuit realization of ART mechanisms will be found in the laminar circuits of all sensory and cognitive neocortex. Concepts and data are summarized concerning how some visual percepts may be visibly, or modally, perceived, whereas amodal percepts may be consciously recognized even though they are perceptually invisible. It is also suggested that sensory and cognitive processing in the What processing stream of the brain obey top-down matching and learning laws that are often complementary to those used for spatial and motor processing in the brain's Where processing stream. This enables our sensory and cognitive representations to maintain their stability as we learn more about the world, while allowing spatial and motor representations to forget learned maps and gains that are no longer appropriate as our bodies develop and grow from infanthood to adulthood. Procedural memories are proposed to be unconscious because the inhibitory matching process that supports these spatial and motor processes cannot lead to resonance.     

ERP and MEG correlates of visual consciousness: The second decade

The first decade of event-related potential (ERP) research had established that the most consistent correlates of the onset of visual consciousness are the early visual awareness negativity (VAN), a posterior negative component in the N2 time range, and the late positivity (LP), an anterior positive component in the P3 time range. Two earlier extensive reviews ten years ago had concluded that VAN is the earliest and most reliable correlate of visual phenomenal consciousness, whereas LP probably reflects later processes associated with reflective/access consciousness. This article provides an update to those earlier reviews. ERP and MEG studies that have appeared since 2010 and directly compared ERPs between aware and unaware conditions are reviewed, and important new developments in the field are discussed. The result corroborates VAN as the earliest and most consistent signature of visual phenomenal consciousness, and casts further doubt on LP as an ERP correlate of phenomenal consciousness.     

情绪面孔的意识神经相关物及其无意识自动加工：来自事件相关电位的证据

意识的神经相关物尚有争议, 且个体能否无意识自动检测视觉环境变化尚不清楚。本研究采用非注意视盲范式操控视觉意识, 并引入具有社会信息的情绪面孔, 探讨意识的神经相关物以及视觉意识与自动检测变化机制的关系。在A阶段, 部分被试对任务无关的情绪面孔处于无意识水平; 在B阶段, 所有被试对任务无关的情绪面孔处于意识水平; 在C阶段, 所有被试对任务相关的情绪面孔处于意识水平。结果显示, 任务无关的情绪面孔的意识过程诱发视觉意识负波(visual awareness negativity, VAN)、晚期正成分(late positivity, LP)和晚期枕区正成分(late occipital positivity, LOP)。此外, 无意识的情绪面孔能诱发视觉失匹配负波(visual mismatch negativity, vMMN), 且其幅值不受意识影响, 但是受任务相关性调制。这些结果提示对情绪面孔的视觉意识在不同的时间进程上有不同的ERP指标——VAN反映早期知觉经验, 而LP和LOP反映晚期意识过程, 而且面孔情绪信息的自动加工独立于视觉意识, 但是受视觉注意调制。     

Testing the dorsal stream attention hypothesis: Electrophysiological correlates and the effects of ventral stream damage

The roles of dorsal and ventral processing streams in visual orienting and conscious perception were examined in two experiments. The first employed high density EEG with source localization. The second comprised a neuropsychological case study. Visual orienting was assessed with an attention procedure, where peripheral letters cued participants towards a target location. In the perception procedure participants responded to the same letters by performing an explicit conscious discrimination. In Experiment 1, the peripheral letters elicited rapid dorsal stream activation in the attention procedure, and this activation preceded top-down enhancement of target processing in occipital cortex. In the perception procedure early ventral stream activation was seen. In addition, peripheral letters elicited an “early directing attention negativity” (EDAN) over parietal recording sites in the attention procedure, but not in the perception procedure. In Experiment 2, a patient with a bilateral ventral stream lesion but preserved dorsal stream function showed clear disruption to performance in the perception procedure, whilst exhibiting a normal visual orienting effect in the attention procedure. Taken together these findings (1) highlight the distinct roles of the dorsal and ventral streams in attention and perception, and (2) suggest how these streams might interact, via reentrant effects of attention on perceptual processing.     

Attentional sensitization of unconscious visual processing: Top-down influences on masked priming

Classical theories of automaticity assume that automatic processes elicited by unconscious stimuli are autonomous and independent of higher-level cognitive influences. In contrast to these classical conceptions, we argue that automatic processing depends on attentional amplification of task-congruent processing pathways and propose an attentional sensitization model of unconscious visual processing: According to this model, unconscious visual processing is automatic in the sense that it is initiated without deliberate intention. However, unconscious visual processing is susceptible to attentional top-down control and is only elicited if the cognitive system is configured accordingly. In this article, we describe our attentional sensitization model and review recent evidence demonstrating attentional influences on subliminal priming, a prototypical example of an automatic process. We show that subliminal priming (a) depends on attentional resources, (b) is susceptible to stimulus expectations, (c) is influenced by action intentions, and (d) is modulated by task sets. These data suggest that attention enhances or attenuates unconscious visual processes in congruency with attentional task representations similar to conscious perception. We argue that seemingly paradoxical, hitherto unexplained findings regarding the automaticity of the underlying processes in many cognitive domains can be easily accommodated by our attentional sensitization model. We conclude this review with a discussion of future research questions regar-ding the nature of attentional control of unconscious visual processing.     

Conscious, preconscious, and subliminal processing: a testable taxonomy

Of the many brain events evoked by a visual stimulus, which are specifically associated with conscious perception, and which merely reflect non-conscious processing? Several recent neuroimaging studies have contrasted conscious and non-conscious visual processing, but their results appear inconsistent. Some support a correlation of conscious perception with early occipital events, others with late parieto-frontal activity. Here we attempt to make sense of these dissenting results. On the basis of the global neuronal workspace hypothesis, we propose a taxonomy that distinguishes between vigilance and access to conscious report, as well as between subliminal, preconscious and conscious processing. We suggest that these distinctions map onto different neural mechanisms, and that conscious perception is systematically associated with surges of parieto-frontal activity causing top-down amplification.     

A global workspace model for phenomenal and access consciousness

Both the global workspace theory and Block’s distinction between phenomenal and access consciousness, are central in the current debates about consciousness and the neural correlates of consciousness. In this article, a unifying global workspace model for phenomenal and access consciousness is proposed. In the model, recurrent neural interactions take place in distinct yet interacting access and phenomenal brain loops. The effectiveness of feedback signaling onto sensory cortical maps is emphasized for the neural correlates of phenomenal consciousness. Two forms of top-down attention, attention for perception and attention for access, play differential roles for phenomenal and access consciousness. The model is implemented in a neural network form, with the simulation of single and multiple visual object processing, and of the attentional blink.     

Top-down and bottom-up control of visual selection

The present paper argues for the notion that when attention is spread across the visual field in the first sweep of information through the brain visual selection is completely stimulus-driven. Only later in time, through recurrent feedback processing, volitional control based on expectancy and goal set will bias visual selection in a top-down manner. Here we review behavioral evidence as well as evidence from ERP, fMRI, TMS and single cell recording consistent with stimulus-driven selection. Alternative viewpoints that assume a large role for top-down processing are discussed. It is argued that in most cases evidence supporting top-down control on visual selection in fact demonstrates top-down control on processes occurring later in time, following initial selection. We conclude that top-down knowledge regarding non-spatial features of the objects cannot alter the initial selection priority. Only by adjusting the size of the attentional window, the initial sweep of information through the brain may be altered in a top-down way.     

Visual consciousness revisited: magnocellular and parvocellular contributions to conscious and nonconscious vision

Current theoretical approaches to consciousness and vision associate the dorsal cortical pathway, in which magnocellular (M) input is dominant, with nonconscious visual processing and the ventral cortical pathway, in which parvocellular (P) input is dominant, with conscious visual processing. We explored the known differences between M and P contrast-response functions to investigate the roles of these channels in vision. Simulations of contrast-dependent priming revealed that priming effects obtained with unmasked, visible primes were best modeled by equations characteristic of M channel responses, whereas priming effects obtained with masked, invisible primes were best modeled by equations characteristic of P channel responses. In the context of current theoretical approaches to conscious and nonconscious processing, our results indicate a surprisingly significant role of M channels in conscious vision. In a broader discussion of the role of M channels in vision, we propose a neurophysiologically plausible interpretation of the present results: M channels indirectly contribute to conscious object vision via top-down modulation of reentrant activity in the ventral object-recognition stream.     

Attention and consciousness: two distinct brain processes

The close relationship between attention and consciousness has led many scholars to conflate these processes. This article summarizes psychophysical evidence, arguing that top-down attention and consciousness are distinct phenomena that need not occur together and that can be manipulated using distinct paradigms. Subjects can become conscious of an isolated object or the gist of a scene despite the near absence of top-down attention; conversely, subjects can attend to perceptually invisible objects. Furthermore, top-down attention and consciousness can have opposing effects. Such dissociations are easier to understand when the different functions of these two processes are considered. Untangling their tight relationship is necessary for the scientific elucidation of consciousness and its material substrate.     

大脑视知觉调控的神经机制

大脑的知觉加工并非单纯由外部刺激驱动,而是存在自上而下的知觉调控。尽管这一现象被大量实验研究证实,但其神经机制仍然是认知神经科学研究的重要问题。本研究系统介绍了知觉调控的神经基础、实现形式、研究范式,及其理论模型,分析指出了当前研究面临的主要问题,并对未来的研究进行了展望,以期促进该问题研究的进一步开展。     

Conscious Vision for Action Versus Unconscious Vision for Action?

David Milner and Melvyn Goodale’s dissociation hypothesis is commonly taken to state that there are two functionally specialized cortical streams of visual processing originating in striate (V1) cortex: a dorsal, action‐related “unconscious” stream and a ventral, perception‐related “conscious” stream. As Milner and Goodale acknowledge, findings from blindsight studies suggest a more sophisticated picture that replaces the distinction between unconscious vision for action and conscious vision for perception with a tripartite division between unconscious vision for action, conscious vision for perception, and unconscious vision for perception. The combination excluded by the tripartite division is the possibility of conscious vision for action. But are there good grounds for concluding that there is no conscious vision for action? There is now overwhelming evidence that illusions and perceived size can have a significant effect on action ( Bruno & Franz, 2009 ; Dassonville & Bala, 2004 ; Franz & Gegenfurtner, 2008 ; McIntosh & Lashley, 2008 ). There is also suggestive evidence that any sophisticated visual behavior requires collaboration between the two visual streams at every stage of the process ( Schenk & McIntosh, 2010 ). I nonetheless want to make a case for the tripartite division between unconscious vision for action, conscious vision for perception, and unconscious vision for perception. My aim here is not to refute the evidence showing that conscious vision can affect action but rather to argue (a) that we cannot gain cognitive access to action‐guiding dorsal stream representations, and (b) that these representations do not correlate with phenomenal consciousness. This vindicates the semi‐conservative view that the dissociation hypothesis is best understood as a tripartite division.     

Visual event-related potentials in schizotypal personality disorder and schizophrenia

Schizophrenia (SZ) occurs among a spectrum of disorders with similar characteristics, including schizotypal personality disorder (SPD). Visual processing disturbances have been reported in both disorders, but it is not yet clear which processes are disturbed in both SZ and SPD, suggestive of a common endophenotype, and which appear only in SZ. In order to address this question, the authors evaluated visual event-related potentials (ERPs) elicited during a line-orientation discrimination task in control, SPD, and SZ participants. Visual ERPs allow specification of both the time course and physiological correlates of visual perception and cognition. SZ patients had smaller P100 and P300a amplitudes and prolonged P300b latency compared to the control group. SZ patients also had smaller N160, N200, P300a, and P300b amplitudes compared to the SPD group. SPD participants did not differ from control participants on any ERP measure. These data documented pervasive abnormalities in visual perception and attention in SZ but not in SPD, suggesting that these visual ERP disturbances may not represent a common endophenotype.     

The temporal evolution of pre-stimulus slow cortical potentials is associated with an upcoming stimulus’ access to visual consciousness

Slow cortical potentials (SCPs) have been proposed to be essential for the formation of conscious experience. To examine their temporal characteristics, we recorded electroencephalography during a visual backward-masking task, which required participants to localize the missing part of a target stimulus. A subsequent confidence rating was used as a proxy for the target’s access to consciousness. Event-related potentials (ERPs) of all correct trials were determined relative to a brief period immediately before the target and then compared among consciousness levels. In an interval ranging from 2 s prior to target presentation up to this period, a negative relationship between slowly fluctuating ERP values and the level of consciousness became evident. After target presentation, high conscious awareness was characterized by an enhanced visual awareness negativity, an increased P3 component, and associated positive SCPs. Together, these findings add new evidence to the proposed role of SCPs in the emergence of visual consciousness.     

Attention versus consciousness in the visual brain: differences in conception, phenomenology, behavior, neuroanatomy, and physiology.

A common confound between consciousness and attention makes it difficult to think clearly about recent advances in the understanding of the visual brain. Visual consciousness involves phenomenal experience of the visual world, but visual attention is more plausibly treated as a function that selects and maintains the selection of potential conscious contents, often unconsciously. In the same sense, eye movements select conscious visual events, which are not the same as conscious visual experience. According to common sense, visual experience is consciousness, and selective processes are labeled as attention. The distinction is reflected in very different behavioral measures and in very different brain anatomy and physiology. Visual consciousness tends to be associated with the "what" stream of visual feature neurons in the ventral temporal lobe. In contrast, attentional selection and maintenance are mediated by other brain regions, ranging from superior colliculi to thalamus, prefrontal cortex, and anterior cingulate. The author applied the common-sense distinction between attention and consciousness to the theoretical positions of M. I. Posner (1992, 1994) and D. LaBerge (1997, 1998) to show how it helps to clarify the evidence. He concluded that clarity of thought is served by calling a thing by its proper name.     

Visual consciousness of faces in the attentional blink: Knowledge-based effects of trustworthiness dominate over appearance-based impressions

How we perceive and evaluate other persons depends on appearance-based impressions as well as top-down information such as knowledge about someone’s character. To date, little is known about how these two sources of information affect the conscious perception of faces, about their relative contributions and possible interactions. Here, we directly compare how social-affective knowledge and visual cues of trustworthiness impact the access of faces to visual consciousness. Low- and average-trustworthy looking faces were associated with neutral or negative information about a person’s social behavior. Trustworthiness and facial expression ratings showed robust and independent effects of both sources of information during evaluations. Under conditions of reduced attention in an attentional blink task, prioritized detection of faces associated with negative as compared to neutral person knowledge was observed, whereas facial trustworthiness did not affect detection. Thus, while both factors influenced conscious perception and evaluation, only person knowledge impacted the access to consciousness.     

错误加工意识水平与错误相关脑电成分

错误加工的意识水平包括有意识水平、无意识水平以及介于两者之间的意识水平。意识水平的划分通常通过错误报告范式实现。根据其对意识水平的划分, 可以将相关研究分为二元意识水平研究和多元意识水平研究。已有研究, 主要关注于意识水平与错误诱发的Ne、Pe的关系。两类研究一致表明, Pe受意识水平的调节, 但是并没有得到fMRI研究的一致支持。对Ne的激活是否独立于意识, 研究者们仍存在争议。还有研究考察了注意对错误意识水平的影响, 结果支持Pe与意识水平共变的观点, 但是对注意如何影响错误意识水平还存在质疑。在对以上研究进行整理与分析的基础上, 对将来的研究提出了建议。     

The feeling of fluent perception: a single experience from multiple asynchronous sources

Zeki and co-workers recently proposed that perception can best be described as locally distributed, asynchronous processes that each create a kind of microconsciousness, which condense into an experienced percept. The present article is aimed at extending this theory to metacognitive feelings. We present evidence that perceptual fluency-the subjective feeling of ease during perceptual processing-is based on speed of processing at different stages of the perceptual process. Specifically, detection of briefly presented stimuli was influenced by figure-ground contrast, but not by symmetry (Experiment 1) or the font (Experiment 2) of the stimuli. Conversely, discrimination of these stimuli was influenced by whether they were symmetric (Experiment 1) and by the font they were presented in (Experiment 2), but not by figure-ground contrast. Both tasks however were related with the subjective experience of fluency (Experiments 1 and 2). We conclude that subjective fluency is the conscious phenomenal correlate of different processing stages in visual perception.     

Top-down control of visual perception: attention in natural vision

Top-down perceptual influences can bias (or pre-empt) perception. In natural scenes, the receptive fields of neurons in the inferior temporal visual cortex (IT) shrink to become close to the size of objects. This facilitates the read-out of information from the ventral visual system, because the information is primarily about the object at the fovea. Top-down attentional influences are much less evident in natural scenes than when objects are shown against blank backgrounds, though are still present. It is suggested that the reduced receptive-field size in natural scenes, and the effects of top-down attention contribute to change blindness. The receptive fields of IT neurons in complex scenes, though including the fovea, are frequently asymmetric around the fovea, and it is proposed that this is the solution the IT uses to represent multiple objects and their relative spatial positions in a scene. Networks that implement probabilistic decision-making are described, and it is suggested that, when in perceptual systems they take decisions (or 'test hypotheses'), they influence lower-level networks to bias visual perception. Finally, it is shown that similar processes extend to systems involved in the processing of emotion-provoking sensory stimuli, in that word-level cognitive states provide top-down biasing that reaches as far down as the orbitofrontal cortex, where, at the first stage of affective representations, olfactory, taste, flavour, and touch processing is biased (or pre-empted) in humans.     

Unconscious attention modulates the silencing effect of top-down predictions

The brain is considered to be proactive in that it continuously generates predictions about external environment stimuli. Recent Bayesian models of perception have demonstrated that prediction and attention operate synergistically to optimize stimulus processing. However, the relation between prediction and unconscious attention remains unclear given the relative neglect of unconscious attention in scholarly literatures. To investigate this issue, twenty participants (12 women) performed an orientation identification task in which a circular grating appeared either in the left or in the right visual field in a single 30–40 min session, during which 64-channel EEG data were acquired. Behavioral results showed an unconscious-attended effect and a facilitated effect. Importantly, prediction-related P1 and N1 silencing effects were observed in the unconscious-attended condition, probably reflecting that unconscious attention improves the precision of top-down predictions at an early stage of processing, thereby increasing the synaptic gain of predictor neurons. Moreover, unlike the early ERP components, P3 revealed a reversed pattern of results, which displayed a silencing effect of prediction only in the unattended condition, suggesting that the influence of unconscious attention on the silencing effect may change over time.