On the perception of probable things: neural substrates of associative memory, imagery, and perception

Perception is influenced both by the immediate pattern of sensory inputs and by memories acquired through prior experiences with the world. Throughout much of its illustrious history, however, study of the cellular basis of perception has focused on neuronal structures and events that underlie the detection and discrimination of sensory stimuli. Relatively little attention has been paid to the means by which memories interact with incoming sensory signals. Building upon recent neurophysiological/behavioral studies of the cortical substrates of visual associative memory, I propose a specific functional process by which stored information about the world supplements sensory inputs to yield neuronal signals that can account for visual perceptual experience. This perspective represents a significant shift in the way we think about the cellular bases of perception.     

General validity of Levelt's propositions reveals common computational mechanisms for visual rivalry

The mechanisms underlying conscious visual perception are often studied with either binocular rivalry or perceptual rivalry stimuli. Despite existing research into both types of rivalry, it remains unclear to what extent their underlying mechanisms involve common computational rules. Computational models of binocular rivalry mechanisms are generally tested against Levelt's four propositions, describing the psychophysical relation between stimulus strength and alternation dynamics in binocular rivalry. Here we use a bistable rotating structure-from-motion sphere, a generally studied form of perceptual rivalry, to demonstrate that Levelt's propositions also apply to the alternation dynamics of perceptual rivalry. Importantly, these findings suggest that bistability in structure-from-motion results from active cross-inhibition between neural populations with computational principles similar to those present in binocular rivalry. Thus, although the neural input to the computational mechanism of rivalry may stem from different cortical neurons and different cognitive levels the computational principles just prior to the production of visual awareness appear to be common to the two types of rivalry.     

Blinded by the load: attention,awareness and the role of perceptual load

What is the relationship between attention and conscious awareness? Awareness sometimes appears to be restricted to the contents of focused attention, yet at other times irrelevant distractors will dominate awareness. This contradictory relationship has also been reflected in an abundance of discrepant research findings leading to an enduring controversy in cognitive psychology. Lavie''s load theory of attention suggests that the puzzle can be solved by considering the role of perceptual load. Although distractors will intrude upon awareness in conditions of low load, awareness will be restricted to the content of focused attention when the attended information involves high perceptual load. Here, we review recent evidence for this proposal with an emphasis on the various subjective blindness phenomena, and their neural correlates, induced by conditions of high perceptual load. We also present novel findings that clarify the role of attention in the response to stimulus contrast. Overall, this article demonstrates a critical role for perceptual load across the spectrum of perceptual processes leading to awareness, from the very early sensory responses related to contrast detection to explicit recognition of semantic content.     

Enhanced perception in savant syndrome: patterns,structure and creativity

According to the enhanced perceptual functioning (EPF) model, autistic perception is characterized by: enhanced low-level operations; locally oriented processing as a default setting; greater activation of perceptual areas during a range of visuospatial, language, working memory or reasoning tasks; autonomy towards higher processes; and superior involvement in intelligence. EPF has been useful in accounting for autistic relative peaks of ability in the visual and auditory modalities. However, the role played by atypical perceptual mechanisms in the emergence and character of savant abilities remains underdeveloped. We now propose that enhanced detection of patterns, including similarity within and among patterns, is one of the mechanisms responsible for operations on human codes, a type of material with which savants show particular facility. This mechanism would favour an orientation towards material possessing the highest level of internal structure, through the implicit detection of within- and between-code isomorphisms. A second mechanism, related to but exceeding the existing concept of redintegration, involves completion, or filling-in, of missing information in memorized or perceived units or structures. In the context of autistics'' enhanced perception, the nature and extent of these two mechanisms, and their possible contribution to the creativity evident in savant performance, are explored.     

View from the top: hierarchies and reverse hierarchies in the visual system

We propose that explicit vision advances in reverse hierarchical direction, as shown for perceptual learning. Processing along the feedforward hierarchy of areas, leading to increasingly complex representations, is automatic and implicit, while conscious perception begins at the hierarchy's top, gradually returning downward as needed. Thus, our initial conscious percept--vision at a glance--matches a high-level, generalized, categorical scene interpretation, identifying "forest before trees." For later vision with scrutiny, reverse hierarchy routines focus attention to specific, active, low-level units, incorporating into conscious perception detailed information available there. Reverse Hierarchy Theory dissociates between early explicit perception and implicit low-level vision, explaining a variety of phenomena. Feature search "pop-out" is attributed to high areas, where large receptive fields underlie spread attention detecting categorical differences. Search for conjunctions or fine discriminations depends on reentry to low-level specific receptive fields using serial focused attention, consistent with recently reported primary visual cortex effects.     

Short-Term Memory Affects Color Perception in Context

Color-based object selection — for instance, looking for ripe tomatoes in the market — places demands on both perceptual and memory processes: it is necessary to form a stable perceptual estimate of surface color from a variable visual signal, as well as to retain multiple perceptual estimates in memory while comparing objects. Nevertheless, perceptual and memory processes in the color domain are generally studied in separate research programs with the assumption that they are independent. Here, we demonstrate a strong failure of independence between color perception and memory: the effect of context on color appearance is substantially weakened by a short retention interval between a reference and test stimulus. This somewhat counterintuitive result is consistent with Bayesian estimation: as the precision of the representation of the reference surface and its context decays in memory, prior information gains more weight, causing the retained percepts to be drawn toward prior information about surface and context color. This interaction implies that to fully understand information processing in real-world color tasks, perception and memory need to be considered jointly.     

Is Attention Based on Spatial Contextual Memory Preferentially Guided by Low Spatial Frequency Signals?

A popular model of visual perception states that coarse information (carried by low spatial frequencies) along the dorsal stream is rapidly transmitted to prefrontal and medial temporal areas, activating contextual information from memory, which can in turn constrain detailed input carried by high spatial frequencies arriving at a slower rate along the ventral visual stream, thus facilitating the processing of ambiguous visual stimuli. We were interested in testing whether this model contributes to memory-guided orienting of attention. In particular, we asked whether global, low-spatial frequency (LSF) inputs play a dominant role in triggering contextual memories in order to facilitate the processing of the upcoming target stimulus. We explored this question over four experiments. The first experiment replicated the LSF advantage reported in perceptual discrimination tasks by showing that participants were faster and more accurate at matching a low spatial frequency version of a scene, compared to a high spatial frequency version, to its original counterpart in a forced-choice task. The subsequent three experiments tested the relative contributions of low versus high spatial frequencies during memory-guided covert spatial attention orienting tasks. Replicating the effects of memory-guided attention, pre-exposure to scenes associated with specific spatial memories for target locations (memory cues) led to higher perceptual discrimination and faster response times to identify targets embedded in the scenes. However, either high or low spatial frequency cues were equally effective; LSF signals did not selectively or preferentially contribute to the memory-driven attention benefits to performance. Our results challenge a generalized model that LSFs activate contextual memories, which in turn bias attention and facilitate perception.     

Time-of-day effects on cognition in preadolescents: a trails study

Cognitive performance fluctuates during the day due to diurnal variations in alertness level. This study examined: (1) whether cognitive performance in school-aged children is affected by time-of-day; (2) which functional domains are particularly vulnerable to time-of-day effects; and (3) whether the effects are more pronounced for cognitively more demanding tasks or task conditions. Children, aged 10-12 yrs, were randomly assigned to a test session starting either at 08:30 (n?=?802), 10:00 (n?=?713), or 13:00?h (n?=?652). Speed and accuracy of information processing were evaluated by tasks that assess input-related cognitive processes (e.g., stimulus encoding), central cognitive processes (e.g., working memory, sustained attention), and output-related processes (e.g., response organization) using the Amsterdam Neuropsychological Tasks program. Time-of-day effects in children were identified in specific neurocognitive domains, such as visuospatial processing and working memory, but only under cognitively more demanding task conditions. Sustained attention showed a speed-accuracy tradeoff with increased slowness and lapses in the early morning, but with better feedback responsiveness and perceptual sensitivity than in the early afternoon. Furthermore, there was a significant interaction of time-on-task with time-of-day for tempo, with the afternoon group increasing in tempo with time-on-task, and the early-morning group first showing a slowing of tempo with time-on-task, followed at the end of the task by a speed increase towards the initial levels. To conclude, the authors found time-of-day effects in preadolescents, which were confined to cognitively more demanding tasks tapping input-related and central cognitive processes.     

Neuronal mechanisms for the perception of ambiguous stimuli

Ambiguous figures that may take on the appearance of two or more distinct forms have fascinated philosophers and psychologists for generations. Recently, several laboratories have studied the neuronal basis of perceptual appearance at the level of single neurons in the cerebral cortex. Experiments that integrate neuronal recording with analyses based on sensory detection theory reveal a remarkable degree of specificity in these neuronal responses. The new challenges are to understand how cognitive processes, such as attention and memory, interact with perception to generate these neuronal signals.     

Non-Attended Representations are Perceptual Rather than Unconscious in Nature

Introspectively we experience a phenomenally rich world. In stark contrast, many studies show that we can only report on the few items that we happen to attend to. So what happens to the unattended objects? Are these consciously processed as our first person perspective would have us believe, or are they – in fact – entirely unconscious? Here, we attempt to resolve this question by investigating the perceptual characteristics of visual sensory memory. Sensory memory is a fleeting, high-capacity form of memory that precedes attentional selection and working memory. We found that memory capacity benefits from figural information induced by the Kanizsa illusion. Importantly, this benefit was larger for sensory memory than for working memory and depended critically on the illusion, not on the stimulus configuration. This shows that pre-attentive sensory memory contains representations that have a genuinely perceptual nature, suggesting that non-attended representations are phenomenally experienced rather than unconscious.     

Perceptions as hypotheses

Perceptions may be compared with hypotheses in science. The methods of acquiring scientific knowledge provide a working paradigm for investigating processes of perception. Much as the information channels of instruments, such as radio telescopes, transmit signals which are processed according to various assumptions to give useful data, so neural signals are processed to give data for perception. To understand perception, the signal codes and the stored knowledge or assumptions used for deriving perceptual hypotheses must be discovered. Systematic perceptual errors are important clues for appreciating signal channel limitations, and for discovering hypothesis-generating procedures. Although this distinction between 'physiological' and 'cognitive' aspects of perception may be logically clear, it is in practice surprisingly difficult to establish which are responsible even for clearly established phenomena such as the classical distortion illusions. Experimental results are presented, aimed at distinguishing between and disconvering what happens when there is mismatch with the neural signal channel, and when neural signals are processed inappropriately for the current situation. This leads us to make some distinctions between perceptual and scientific hypotheses, which raise in a new form the problem: What are 'objects'?     

Cognitive outcome in adult women affected by congenital adrenal hyperplasia due to 21-hydroxylase deficiency

BACKGROUND: Some research suggests that girls with congenital adrenal hyperplasia (CAH), who are exposed to higher than normal levels of prenatal androgens, perform better on spatial tasks, worse on verbal tasks and have a greater incidence of left-handedness than unaffected controls, all of which suggests the development of a more male-typical cognitive pattern. However, research in all three areas has produced inconsistent findings. OBJECTIVES: To determine if prenatal androgen exposure has an organizing effect on female cognitive development and to what extent. METHODS: 24 women, 21-71 years, with either the salt-losing (SL) or simple virilizing (SV) forms of CAH due to 21-hydroxylase deficiency, and 18 controls, 21-73 years, who were unaffected female relatives or women with polycystic ovary syndrome, were assessed with IQ, handedness, executive function, verbal learning and memory, non-verbal learning and memory, perceptual speed, visuospatial processing and visuomotor ability measures. The battery included tests known to elicit sex differences and control measures. RESULTS: The findings did not support the hypothesis that women with CAH develop a more male-typical cognitive pattern. CONCLUSION: This study differs from others in the older age of its participants, grouping by SL/SV diagnosis and assessment of medical treatment and compliance as determined through hormone assays. Our findings provide additional support for the conclusion that, in adult women with CAH, previous prenatal androgen exposure does not enhance spatial abilities, impair verbal abilities nor alter hand preferences in a long-lasting way.     

A cortical-hippocampal system for declarative memory

Recent neurobiological studies have begun to reveal the cognitive and neural coding mechanisms that underlie declarative memory--our ability to recollect everyday events and factual knowledge. These studies indicate that the critical circuitry involves bidirectional connections between the neocortex, the parahippocampal region and the hippocampus. Each of these areas makes a unique contribution to memory processing. Widespread high-order neocortical areas provide dedicated processors for perceptual, motor or cognitive information that is influenced by other components of the system. The parahippocampal region mediates convergence of this information and extends the persistence of neocortical memory representations. The hippocampus encodes the sequences of places and events that compose episodic memories, and links them together through their common elements. Here I describe how these mechanisms work together to create and re-create fully networked representations of previous experiences and knowledge about the world.     

Neuronal mechanisms for visual stability: progress and problems

How our vision remains stable in spite of the interruptions produced by saccadic eye movements has been a repeatedly revisited perceptual puzzle. The major hypothesis is that a corollary discharge (CD) or efference copy signal provides information that the eye has moved, and this information is used to compensate for the motion. There has been progress in the search for neuronal correlates of such a CD in the monkey brain, the best animal model of the human visual system. In this article, we briefly summarize the evidence for a CD pathway to frontal cortex, and then consider four questions on the relation of neuronal mechanisms in the monkey brain to stable visual perception. First, how can we determine whether the neuronal activity is related to stable visual perception? Second, is the activity a possible neuronal correlate of the proposed transsaccadic memory hypothesis of visual stability? Third, are the neuronal mechanisms modified by visual attention and does our perceived visual stability actually result from neuronal mechanisms related primarily to the central visual field? Fourth, does the pathway from superior colliculus through the pulvinar nucleus to visual cortex contribute to visual stability through suppression of the visual blur produced by saccades?     

The Competitive Influences of Perceptual Load and Working Memory Guidance on Selective Attention

The perceptual load theory in selective attention literature proposes that the interference from task-irrelevant distractor is eliminated when perceptual capacity is fully consumed by task-relevant information. However, the biased competition model suggests that the contents of working memory (WM) can guide attentional selection automatically, even when this guidance is detrimental to visual search. An intriguing but unsolved question is what will happen when selective attention is influenced by both perceptual load and WM guidance. To study this issue, behavioral performances and event-related potentials (ERPs) were recorded when participants were presented with a cue to either identify or hold in memory and had to perform a visual search task subsequently, under conditions of low or high perceptual load. Behavioural data showed that high perceptual load eliminated the attentional capture by WM. The ERP results revealed an obvious WM guidance effect in P1 component with invalid trials eliciting larger P1 than neutral trials, regardless of the level of perceptual load. The interaction between perceptual load and WM guidance was significant for the posterior N1 component. The memory guidance effect on N1 was eliminated by high perceptual load. Standardized Low Resolution Electrical Tomography Analysis (sLORETA) showed that the WM guidance effect and the perceptual load effect on attention can be localized into the occipital area and parietal lobe, respectively. Merely identifying the cue produced no effect on the P1 or N1 component. These results suggest that in selective attention, the information held in WM could capture attention at the early stage of visual processing in the occipital cortex. Interestingly, this initial capture of attention by WM could be modulated by the level of perceptual load and the parietal lobe mediates target selection at the discrimination stage.     

Musical experience and the aging auditory system: implications for cognitive abilities and hearing speech in noise

Much of our daily communication occurs in the presence of background noise, compromising our ability to hear. While understanding speech in noise is a challenge for everyone, it becomes increasingly difficult as we age. Although aging is generally accompanied by hearing loss, this perceptual decline cannot fully account for the difficulties experienced by older adults for hearing in noise. Decreased cognitive skills concurrent with reduced perceptual acuity are thought to contribute to the difficulty older adults experience understanding speech in noise. Given that musical experience positively impacts speech perception in noise in young adults (ages 18-30), we asked whether musical experience benefits an older cohort of musicians (ages 45-65), potentially offsetting the age-related decline in speech-in-noise perceptual abilities and associated cognitive function (i.e., working memory). Consistent with performance in young adults, older musicians demonstrated enhanced speech-in-noise perception relative to nonmusicians along with greater auditory, but not visual, working memory capacity. By demonstrating that speech-in-noise perception and related cognitive function are enhanced in older musicians, our results imply that musical training may reduce the impact of age-related auditory decline.     

Human areas V3A and V6 compensate for self-induced planar visual motion

Little is known about mechanisms mediating a stable perception of the world during pursuit eye movements. Here, we used fMRI to determine to what extent human motion-responsive areas integrate planar retinal motion with nonretinal eye movement signals in order to discard self-induced planar retinal motion and to respond to objective ("real") motion. In?contrast to other areas, V3A lacked responses to?self-induced planar retinal motion but responded strongly to head-centered motion, even when retinally canceled by pursuit. This indicates a near-complete multimodal integration of visual with nonvisual planar motion signals in V3A. V3A could be mapped selectively and robustly in every single subject on this basis. V6 also reported head-centered planar motion, even when 3D flow was added to it, but was suppressed by retinal planar motion. These findings suggest a dominant contribution of human areas V3A and V6 to head-centered motion perception and to perceptual stability during eye movements.     

Dissociated mechanisms of extracting perceptual information into visual working memory

Background

The processing mechanisms of visual working memory (VWM) have been extensively explored in the recent decade. However, how the perceptual information is extracted into VWM remains largely unclear. The current study investigated this issue by testing whether the perceptual information was extracted into VWM via an integrated-object manner so that all the irrelevant information would be extracted (object hypothesis), or via a feature-based manner so that only the target-relevant information would be extracted (feature hypothesis), or via an analogous processing manner as that in visual perception (analogy hypothesis).

Methodology/Principal Findings

High-discriminable information which is processed at the parallel stage of visual perception and fine-grained information which is processed via focal attention were selected as the representatives of perceptual information. The analogy hypothesis predicted that whereas high-discriminable information is extracted into VWM automatically, fine-grained information will be extracted only if it is task-relevant. By manipulating the information type of the irrelevant dimension in a change-detection task, we found that the performance was affected and the ERP component N270 was enhanced if a change between the probe and the memorized stimulus consisted of irrelevant high-discriminable information, but not if it consisted of irrelevant fine-grained information.

Conclusions/Significance

We conclude that dissociated extraction mechanisms exist in VWM for information resolved via dissociated processes in visual perception (at least for the information tested in the current study), supporting the analogy hypothesis.     

The Influence of a Working Memory Task on Affective Perception of Facial Expressions

In a dual-task paradigm, participants performed a spatial location working memory task and a forced two-choice perceptual decision task (neutral vs. fearful) with gradually morphed emotional faces (neutral ∼ fearful). Task-irrelevant word distractors (negative, neutral, and control) were experimentally manipulated during spatial working memory encoding. We hypothesized that, if affective perception is influenced by concurrent cognitive load using a working memory task, task-irrelevant emotional distractors would bias subsequent perceptual decision-making on ambiguous facial expression. We found that when either neutral or negative emotional words were presented as task-irrelevant working-memory distractors, participants more frequently reported fearful face perception - but only at the higher emotional intensity levels of morphed faces. Also, the affective perception bias due to negative emotional distractors correlated with a decrease in working memory performance. Taken together, our findings suggest that concurrent working memory load by task-irrelevant distractors has an impact on affective perception of facial expressions.     

Self-awareness in dreaming.

The process by which self-awareness or subjective experience (consciousness) is maintained has been conceptually and phenomenologically associated primarily with the waking state. In the present study the authors investigate, through introspective reports, the existence and variety of self-awareness in dreaming and whether this phenomenon could, under certain conditions, be distinguished into primary and reflective consciousness. For consistency with theory and dreaming research, instead of reflective consciousness we used the term 'reflective awareness.' Findings indicate that self-awareness in dreaming can be found in its both primary and reflective modes. Phenomenologically, primary consciousness exists in four basic modalities: perceptual, experiential, cognitive, and memory-based recognition. Expressions of these primary consciousness modalities are accessible through introspective interviews during waking. Based on participants' statements, reflective consciousness (awareness) in dreaming was initiated by noticing positive and negative feelings and by personally defined oddities. These findings point to a possible oscillation between primary and reflective consciousness in dreaming. (PsycINFO Database Record (c) 2010 APA, all rights reserved)