Perceptual Expectations Are Reflected by Early Alpha Power Reduction.

The predictability of a stimulus can be characterized by its transitional probability. Perceptual expectations derived from the transitional probability of the stimulus were found to modulate the early alpha oscillations in the sensory regions of the brain when neural responses to expected versus unexpected stimuli were compared. The objective of our study was to find out the extent to which this low-frequency oscillation reflects stimulus predictability. We aimed to detect the alpha-power difference with smaller differences in transitional probabilities by comparing expected stimuli with neutral ones. We studied the effect of expectation on perception by applying an unsupervised visual statistical learning paradigm with expected and neutral stimuli embedded in an image sequence while recording EEG. Time-frequency analysis showed that expected stimuli elicit lower alpha power in the window of 8-12 Hz and 0-400 msec after stimulus presentation, appearing in the centroparietal region. Comparing previous findings of expectancy-based alpha-band modulation with our results suggests that early alpha oscillation shows an inverse relationship with stimulus predictability. Although current data are insufficient to determine the origin of the alpha power reduction, this could be a potential sign of expectation suppression in cortical oscillatory activity.

Online measurement of learning temporal statistical structure in categorization tasks.

The ability to grasp relevant patterns from a continuous stream of environmental information is called statistical learning. Although the representations that emerge during visual statistical learning (VSL) are well characterized, little is known about how they are formed. We developed a sensitive behavioral design to characterize the VSL trajectory during ongoing task performance. In sequential categorization tasks, we assessed two previously identified VSL markers: priming of the second predictable image in a pair manifested by a reduced reaction time (RT) and greater accuracy, and the anticipatory effect on the first image revealed by a longer RT. First, in Experiment 1A, we used an adapted paradigm and replicated these VSL markers; however, they appeared to be confounded by motor learning. Next, in Experiment 1B, we confirmed the confounding influence of motor learning. To assess VSL without motor learning, in Experiment 2 we (1) simplified the categorization task, (2) raised the number of subjects and image repetitions, and (3) increased the number of single unpaired images. Using linear mixed-effect modeling and estimated marginal means of linear trends, we found that the RT curves differed significantly between predictable paired and control single images. Further, the VSL curve fitted a logarithmic model, suggesting a rapid learning process. These results suggest that our paradigm in Experiment 2 seems to be a viable online tool to monitor the behavioral correlates of unsupervised implicit VSL.

LCD Monitors as an Alternative for Precision Demanding Visual Psychophysical Experiments.

Precise timing and presentation of stimuli is critical in vision research, still, the limiting factor in successful recognition is often the monitor itself that is used to present the stimuli. The most widespread method is the use of monitors controlled by personal computers. Traditionally, most experiments used cathode-ray tubes but they are more and more difficult to access, and instead, liquid-crystal displays are getting more and more popular. The two types have fundamentally different working principles and limitations in displaying the stimulus.In our experiments, the temporal precision of the stimulus presentation was in focus. We investigated whether liquid-crystal displays, which are not considered to be fit to display fast successive stimuli, can represent an alternative choice for cathode-ray tubes. We used the double flash and the flicker illusion to compare the technical capabilities of the two monitor types. These illusions not only do require a precise timing but also a very short exposure to the stimuli. At the same time, the interstimulus interval is also of extreme importance. In addition, these illusions require peripheral stimulation of the retina, which is more sensitive to the temporal aspects of the visual stimulus. On the basis of previous studies and our own psychophysical results, we suggest that liquid-crystal displays might be a good alternative for precise, frame-to-frame stimulus presentation even if parts of the stimuli are projected on the peripheral retina.

Illusory flashes and perception.

Information from the environment can be based on a single or several modalities. The simultaneous processing of information separated in space and/or time depends on multiple factors. Visual illusions serve as a good tool with which to investigate the parallel processing of information and their interactions. This study was designed to gain information about a unimodal illusion: a target that flashes once seems to flash more as a result of a simultaneously presented inducer flashing several times nearby. The first aim of this work was to understand whether the number of perceived flashes is merely a result of a bias in the criterion level or whether it is based on a real percept. We then clarified how the illusion finds its way into the percept. The final step was designed to establish the logic of the processing in the background by determining whether the modality appropriateness hypothesis, the information reliability hypothesis, or the discontinuity theory best explains the predominant role of the inducer.

Gamma oscillations in visual statistical learning correlate with individual behavioral differences.

Statistical learning is assumed to be a fundamentally general sensory process across modalities, age, other cognitive functions, and even species. Despite this general role, behavioral testing on regularity acquisition shows great variance among individuals. The current study aimed to find neural correlates of visual statistical learning showing a correlation with behavioral results. Based on a pilot study, we conducted an EEG study where participants were exposed to associated stimulus pairs; the acquisition was tested through a familiarity test. We identified an oscillation in the gamma range (40-70 Hz, 0.5-0.75 s post-stimulus), which showed a positive correlation with the behavioral results. This change in activity was located in a left frontoparietal cluster. Based on its latency and location, this difference was identified as a late gamma activity, a correlate of model-based learning. Such learning is a summary of several top-down mechanisms that modulate the recollection of statistical relationships such as the capacity of working memory or attention. These results suggest that, during acquisition, individual behavioral variance is influenced by dominant learning processes which affect the recall of previously gained information.

Transcranial Stimulation of the Orbitofrontal Cortex Affects Decisions about Magnocellular Optimized Stimuli.

Visual categorization plays an important role in fast and efficient information processing; still the neuronal basis of fast categorization has not been established yet. There are two main hypotheses known; both agree that primary, global impressions are based on the information acquired through the magnocellular pathway (MC). It is unclear whether this information is available through the MC that provides information (also) for the ventral pathway or through top-down mechanisms by connections between the dorsal pathway and the ventral pathway via the frontal cortex. To clarify this, a categorization task was performed by 48 subjects; they had to make decisions about objects' sizes. We created stimuli specific to the magno- and parvocellular pathway (PC) on the basis of their spatial frequency content. Transcranial direct-current stimulation was used to assess the role of frontal areas, a target of the MC. Stimulation did not bias the accuracy of decisions when stimuli optimized for the PC were used. In the case of stimuli optimized for the MC, anodal stimulation improved the subjects' accuracy in the behavioral test, while cathodal stimulation impaired accuracy. Our results support the hypothesis that fast visual categorization processes rely on top-down mechanisms that promote fast predictions through coarse information carried by MC via the orbitofrontal cortex.

Task-related modulation in the monkey inferotemporal cortex.

The latencies of the neuronal responses from the inferotemporal cortical cells were analyzed in animals performing a visual fixation task and a recognition task with the same stimulus set. A consistent reduction in response latencies of about 10 ms was observed in favor of the recognition task. It was found that behavioral relevance reduces the latency in the inferotemporal cortex and it was concluded that behavioral significance accelerates information processing. This effect has not been described previously.

Functional organization of visual cortex in the owl monkey.

In this study, we compared the organization of orientation preference in visual areas V1, V2, and V3. Within these visual areas, we also quantified the relationship between orientation preference and cytochrome oxidase (CO) staining patterns. V1 maps of orientation preference contained both pinwheels and linear zones. The location of CO blobs did not relate in a systematic way to maps of orientation; although, as in other primates, there were approximately twice as many pinwheels as CO blobs. V2 contained bands of high and low orientation selectivity. The bands of high orientation selectivity were organized into pinwheels and linear zones, but iso-orientation domains were twice as large as those in V1. Quantitative comparisons between bands containing high or low orientation selectivity and CO dark and light bands suggested that at least four functional compartments exist in V2, CO dense bands with either high or low orientation selectivity, and CO light bands with either high or low selectivity. We also demonstrated that two functional compartments exist in V3, with zones of high orientation selectivity corresponding to CO dense areas and zones of low orientation selectivity corresponding to CO pale areas. Together with previous findings, these results suggest that the modular organization of V1 is similar across primates and indeed across most mammals. V2 organization in owl monkeys also appears similar to that of other simians but different from that of prosimians and other mammals. Finally, V3 of owl monkeys shows a compartmental organization for orientation selectivity that remains to be demonstrated in other primates.

Achromatic shape processing in the inferotemporal cortex of the macaque.

The responses of single neurones in the inferior temporal cortex of awake macaque monkeys to chromatic and achromatic stimuli were investigated, with the aim of determining whether colour-independent processing occurs in this last unimodal area of the ventral visual pathway. There were no differences in the firing rate of the responses (responsiveness) or the selectivity of the inferior temporal neurons towards greyscale and coloured images. The latency of the responses was the same in the two conditions. These results stress the importance of the inferior temporal cortex in colour-independent object recognition.

Audio-visual integration through the parallel visual pathways.

Audio-visual integration has been shown to be present in a wide range of different conditions, some of which are processed through the dorsal, and others through the ventral visual pathway. Whereas neuroimaging studies have revealed integration-related activity in the brain, there has been no imaging study of the possible role of segregated visual streams in audio-visual integration. We set out to determine how the different visual pathways participate in this communication. We investigated how audio-visual integration can be supported through the dorsal and ventral visual pathways during the double flash illusion. Low-contrast and chromatic isoluminant stimuli were used to drive preferably the dorsal and ventral pathways, respectively. In order to identify the anatomical substrates of the audio-visual interaction in the two conditions, the psychophysical results were correlated with the white matter integrity as measured by diffusion tensor imaging.The psychophysiological data revealed a robust double flash illusion in both conditions. A correlation between the psychophysical results and local fractional anisotropy was found in the occipito-parietal white matter in the low-contrast condition, while a similar correlation was found in the infero-temporal white matter in the chromatic isoluminant condition. Our results indicate that both of the parallel visual pathways may play a role in the audio-visual interaction.

Static and dynamic views of visual cortical organization.

Without the aid of modern techniques Cajal speculated that cells in the visual cortex were connected in circuits. From Cajal's time until fairly recently, the flow of information within the cells and circuits of visual cortex has been described as progressing from input to output, from sensation to action. In this chapter we argue that a paradigm shift in our concept of the visual cortical neuron is under way. The most important change in our view concerns the neuron's functional role. Visual cortical neurons do not have static functional signatures but instead function dynamically depending on the ongoing activity of the networks to which they belong. These networks are not merely top-down or bottom-up unidirectional transmission lines, but rather represent machinery that uses recurrent information and is dynamic and highly adaptable. With the advancement of technology for analyzing the conversations of multiple neurons at many levels in the visual system and higher resolution imaging, we predict that the paradigm shift will progress to the point where neurons are no longer viewed as independent processing units but as members of subsets of networks where their role is mapped in space-time coordinates in relationship to the other neuronal members. This view moves us far from Cajal's original views of the neuron. Nevertheless, we believe that understanding the basic morphology and wiring of networks will continue to contribute to our overall understanding of the visual cortex.

A review on the inferior temporal cortex of the macaque.

In our review, we summarize recent advances in the research of the inferior temporal cortex (ITC) of the macaque monkey. This area of the cortex is known to have a crucial role in visual shape recognition and is regarded as being at the end stage of the so-called ventral visual pathway. In the last decade, several new findings appeared in the field without being integrated in a coherent view about the function, position, and operating principles of the area. During this decade, experimental techniques developed a great deal, and the way we look at the brain and brain function changed too. In this review, we try to integrate knowledge about the ITC to the changing view about the brain while outlining the work that has been done in the last decade.

Low-threshold Ca2+-associated bursts are rare events in the LGN of the awake behaving monkey.

It has been proposed that low-threshold Ca2+ (LT)-associated bursts in the lateral geniculate nucleus (LGN) of awake animals communicate significant or unexpected visual events to cortex. The present study investigated this hypothesis by examining the incidence of LT bursts in 146 cells recorded from the LGN of three macaque monkeys. Bursts were defined as clusters of two or more action potentials separated by not more than 4 ms and preceded by a > or = 100-ms quiescent interval. The incidence of bursts was examined in several intensive-training Go-NoGo and target selection tasks as well as in training-free tasks where natural scenes with both familiar and novel contents were shown. Our chief findings were as follows. 1) Bursts occur in the majority of cells under every condition tested, 2) burst incidence is very low (<1 burst every 10 s), 3) bursts occur in association with a receptive field stimulus on average only once every 23 times in 65% of cells tested, 4) cells responding with bursts to the stimulus also tended to exhibit higher levels of spontaneous bursting, 5) the presence of bursts did not depend on the novelty of the stimulus or its behavioral relevance. When the monkeys explored static natural scenes, 6) bursts were not correlated with short-term changes in the image sampled by the cell's receptive field during saccades. Burst incidence 7) did not increase when images were novel or when they evoked an emotional reaction, and 8) bursts did not decrease when images were familiar. 9) Bursts were not correlated with saccades in the dark, but 10) more spikes participated in bursts in the dark. Although these results confirm the occurrence of LT bursts in LGN cells of awake monkeys, they do not support the hypothesis that these bursts are a privileged means of transferring sensory information, that they signal unexpected or significant visual events, or that they are involved uniquely in the coding of natural scenes.

On the impact of attention and motor planning on the lateral geniculate nucleus.

Although the lateral geniculate nucleus (LGN) is one of the most thoroughly characterized thalamic nuclei, its functional role remains controversial. Traditionally, the LGN in primates has been viewed as the lowest level of a set of feedforward parallel visual pathways to cortex. These feedforward pathways are pictured as connected hierarchies of areas designed to construct the visual image gradually - adding more complex features as one marches through successive levels of the hierarchy. In terms of synapse number and circuitry, the anatomy suggests that the LGN can be viewed also as the ultimate terminus in a series of feedback pathways that originate at the highest cortical levels. Since the visual system is dynamic, a more accurate picture of image construction might be one in which information flows bidirectionally, through both the feedforward and feedback pathways constantly and simultaneously. Based upon evidence from anatomy, physiology, and imaging, we argue that the LGN is more than a simple gate for retinal information. Here, we review evidence that suggests that one function of the LGN is to enhance relevant visual signals through circuits related to both motor planning and attention. Specifically, we argue that major extraretinal inputs to the LGN may provide: (1) eye movement information to enhance and bind visual signals related to new saccade targets and (2) top-down and bottom-up information about target relevance to selectively enhance visual signals through spatial attention.

Human brain regions involved in visual categorization.

Categorization of dot patterns is a frequently used paradigm in the behavioral study of natural categorization. To determine the human brain regions involved in categorization, we used Positron Emission Tomography to compare regional Cerebral Blood Flow patterns in two tasks employing patterns that consisted of nine dots. In the categorization task, subjects categorized novel exemplars of two categories, generated by distorting two prototypes, and other random dot patterns. In the control task, subjects judged the position of similarly distorted patterns. Each task was presented at two matched levels of difficulty. Fixation of the fixation target served as baseline condition. The categorization task differentially activated the orbitofrontal cortex and two dorsolateral prefrontal regions. These three prefrontal regions were equally weakly active in the position discrimination task and the baseline condition. The intraparietal sulcus was activated in both tasks, albeit significantly less in the position discrimination than in the categorization task. A similar activation pattern was present in the neostriatum. Task difficulty had no effect. These functional imaging results show that the dot-pattern categorization task strongly engages prefrontal and parietal cortical areas. The activation of prefrontal cortex during visual categorization in humans agrees with the recent finding of category-related responses in macaque prefrontal neurons.