Physical Properties Of Stimuli Research Articles

Occipital TMS at phosphene detection threshold captures attention automatically

Strong stimuli may capture attention automatically, suggesting that attentional selection is determined primarily by physical stimulus properties. The mechanisms underlying capture remain controversial, in particular, whether feedforward subcortical processes are its main source. Also, it remains unclear whether only physical stimulus properties determine capture strength. Here, we demonstrate strong capture in the absence of feedforward input to subcortical structures such as the superior colliculus, by using transcranial magnetic stimulation (TMS) over occipital visual cortex as an attention cue. This implies that the feedforward sweep through subcortex is not necessary for capture to occur but rather provides an additional source of capture. Furthermore, seen cues captured attention more strongly than (physically identical) unseen cues, suggesting that the momentary state of the nervous system modulates attentional selection. In summary, we demonstrate the existence of several sources of attentional capture, and that both physical stimulus properties and the state of the nervous system influence capture.

Attentional cueing in numerical cognition.

Attentional cueing in numerical cognition.

Top-down modulation in human visual cortex predicts the stability of a perceptual illusion.

Conscious perception sometimes fluctuates strongly, even when the sensory input is constant. For example, in motion-induced blindness (MIB), a salient visual target surrounded by a moving pattern suddenly disappears from perception, only to reappear after some variable time. Whereas such changes of perception result from fluctuations of neural activity, mounting evidence suggests that the perceptual changes, in turn, may also cause modulations of activity in several brain areas, including visual cortex. In this study, we asked whether these latter modulations might affect the subsequent dynamics of perception. We used magnetoencephalography (MEG) to measure modulations in cortical population activity during MIB. We observed a transient, retinotopically widespread modulation of beta (12-30 Hz)-frequency power over visual cortex that was closely linked to the time of subjects' behavioral report of the target disappearance. This beta modulation was a top-down signal, decoupled from both the physical stimulus properties and the motor response but contingent on the behavioral relevance of the perceptual change. Critically, the modulation amplitude predicted the duration of the subsequent target disappearance. We propose that the transformation of the perceptual change into a report triggers a top-down mechanism that stabilizes the newly selected perceptual interpretation.

Online tracking of the contents of conscious perception using real-time fMRI.

Perception is an active process that interprets and structures the stimulus input based on assumptions about its possible causes. We use real-time functional magnetic resonance imaging (rtfMRI) to investigate a particularly powerful demonstration of dynamic object integration in which the same physical stimulus intermittently elicits categorically different conscious object percepts. In this study, we simulated an outline object that is moving behind a narrow slit. With such displays, the physically identical stimulus can elicit categorically different percepts that either correspond closely to the physical stimulus (vertically moving line segments) or represent a hypothesis about the underlying cause of the physical stimulus (a horizontally moving object that is partly occluded). In the latter case, the brain must construct an object from the input sequence. Combining rtfMRI with machine learning techniques we show that it is possible to determine online the momentary state of a subject's conscious percept from time resolved BOLD-activity. In addition, we found that feedback about the currently decoded percept increased the decoding rates compared to prior fMRI recordings of the same stimulus without feedback presentation. The analysis of the trained classifier revealed a brain network that discriminates contents of conscious perception with antagonistic interactions between early sensory areas that represent physical stimulus properties and higher-tier brain areas. During integrated object percepts, brain activity decreases in early sensory areas and increases in higher-tier areas. We conclude that it is possible to use BOLD responses to reliably track the contents of conscious visual perception with a relatively high temporal resolution. We suggest that our approach can also be used to investigate the neural basis of auditory object formation and discuss the results in the context of predictive coding theory.

Neural substrates of perceptual integration during bistable object perception

The way we perceive an object depends both on feedforward, bottom-up processing of its physical stimulus properties and on top-down factors such as attention, context, expectation, and task relevance. Here we compared neural activity elicited by varying perceptions of the same physical image--a bistable moving image in which perception spontaneously alternates between dissociated fragments and a single, unified object. A time-frequency analysis of EEG changes associated with the perceptual switch from object to fragment and vice versa revealed a greater decrease in alpha (8-12 Hz) accompanying the switch to object percept than to fragment percept. Recordings of event-related potentials elicited by irrelevant probes superimposed on the moving image revealed an enhanced positivity between 184 and 212 ms when the probes were contained within the boundaries of the perceived unitary object. The topography of the positivity (P2) in this latency range elicited by probes during object perception was distinct from the topography elicited by probes during fragment perception, suggesting that the neural processing of probes differed as a function of perceptual state. Two source localization algorithms estimated the neural generator of this object-related difference to lie in the lateral occipital cortex, a region long associated with object perception. These data suggest that perceived objects attract attention, incorporate visual elements occurring within their boundaries into unified object representations, and enhance the visual processing of elements occurring within their boundaries. Importantly, the perceived object in this case emerged as a function of the fluctuating perceptual state of the viewer.

Neuroanatomic correlates of the feature-salience hierarchy in face processing: An fMRI -adaptation study

Previous fMRI studies suggest that faces are represented holistically in human face processing regions. On the other hand, behavioral studies have also shown that some facial features are more salient than others for face recognition: the neural basis of this feature-salience hierarchy is not known. We used fMRI-adaptation together with a behavioral discrimination task and an ideal observer analysis to ask (1) whether different face parts contribute different amounts to the neural signal in face responsive regions, and (2) whether this response correlates more with the behavioral performance of human subjects or with the physical properties of the face stimuli. Twenty-three subjects performed a same/different discrimination experiment to characterize their ability to detect changes to different face parts. The same subjects underwent an fMRI-adaptation study, in which limited portions of the faces were repeated or changed between alternating stimuli. The behavioral study showed high efficiency in identity discrimination when the whole face, top half, or eyes changed, and low efficiency when the bottom half, nose, or mouth changed. During fMRI, there was a release of adaptation in the right and left fusiform face area (FFA) with changes to the whole face, top face-half, or the eyes. Changes to the bottom half, nose or mouth did not result in a significant release of adaptation in the right FFA, although bottom-half changes resulted in a release of adaptation in the left FFA. Adaptation in the right and left FFA and the right pSTS was correlated with human perceptual efficiency but not with ideal observer measures of the physical image differences between face parts. The feature-salience hierarchy of human face perception is therefore reflected in the activity in the right and left FFA and right pSTS, further supporting the key role of these structures in our perceptual experience of faces.

The inferior colliculus encodes the Franssen auditory spatial illusion

Illusions are effective tools for the study of the neural mechanisms underlying perception because neural responses can be correlated to the physical properties of stimuli and the subject's perceptions. The Franssen illusion (FI) is an auditory spatial illusion evoked by presenting a transient, abrupt tone and a slowly rising, sustained tone of the same frequency simultaneously on opposite sides of the subject. Perception of the FI consists of hearing a single sound, the sustained tone, on the side that the transient was presented. Both subcortical and cortical mechanisms for the FI have been proposed, but, to date, there is no direct evidence for either. The data show that humans and rhesus monkeys perceive the FI similarly. Recordings were taken from single units of the inferior colliculus in the monkey while they indicated the perceived location of sound sources with their gaze. The results show that the transient component of the Franssen stimulus, with a shorter first spike latency and higher discharge rate than the sustained tone, encodes the perception of sound location. Furthermore, the persistent erroneous perception of the sustained stimulus location is due to continued excitation of the same neurons, first activated by the transient, by the sustained stimulus without location information. These results demonstrate for the first time, on a trial-by-trial basis, a correlation between perception of an auditory spatial illusion and a subcortical physiological substrate.

Neural substrates of perceptual integration during bistable object perception

The way we perceive an object depends both on feedforward, bottom-up processing of its physical stimulus properties and on top-down factors such as attention, context, expectation, and task relevance. Here we compared neural activity elicited by varying perceptions of the same physical image—a bistable moving image in which perception spontaneously alternates between dissociated fragments and a single, unified object. A time-frequency analysis of EEG changes associated with the perceptual switch from object to fragment and vice versa revealed a greater decrease in alpha (8–12 Hz) accompanying the switch to object percept than to fragment percept. Recordings of event-related potentials elicited by irrelevant probes superimposed on the moving image revealed an enhanced positivity between 184 and 212 ms when the probes were contained within the boundaries of the perceived unitary object. The topography of the positivity (P2) in this latency range elicited by probes during object perception was distinct from the topography elicited by probes during fragment perception, suggesting that the neural processing of probes differed as a function of perceptual state. Two source localization algorithms estimated the neural generator of this object-related difference to lie in the lateral occipital cortex, a region long associated with object perception. These data suggest that perceived objects attract attention, incorporate visual elements occurring within their boundaries into unified object representations, and enhance the visual processing of elements occurring within their boundaries. Importantly, the perceived object in this case emerged as a function of the fluctuating perceptual state of the viewer.

Are auditory percepts determined by experience?

Audition–what listeners hear–is generally studied in terms of the physical properties of sound stimuli and physiological properties of the auditory system. Based on recent work in vision, we here consider an alternative perspective that sensory percepts are based on past experience. In this framework, basic auditory qualities (e.g., loudness and pitch) are based on the frequency of occurrence of stimulus patterns in natural acoustic stimuli. To explore this concept of audition, we examined five well-documented psychophysical functions. The frequency of occurrence of acoustic patterns in a database of natural sound stimuli (speech) predicts some qualitative aspects of these functions, but with substantial quantitative discrepancies. This approach may offer a rationale for auditory phenomena that are difficult to explain in terms of the physical attributes of the stimuli as such.

Integrative processing of invariant aspects of faces: Effect of gender and race processing on identity analysis

While separation of face identity and expression processing is favored by many face perception models, how the visual system analyzes identity and other face properties remains elusive. Here we investigated whether identity analysis is independent of or influenced by automatic processing of face gender and race. Participants searched for a target face among distractor faces whose gender or race was either the same as or different from the target face. Visual search was faster and more accurate when target and distractor faces differed in gender or race property than when not. The effect persisted for identification of both familiar and novel faces, and cannot be attributed to the low-level physical properties of stimuli or the earlier extraction of face gender/race information before identification. Together with complementary findings showing effects of identity analysis on gender and race categorization, these results indicate that invariant face properties are processed in an integrative way: visual analysis of one property involves, and is therefore affected by, automatic processing of the others. Implications for current theoretical models of face perception are discussed.

Internal signal correlates neural populations and biases perceptual decision reports

In perceptual decision-making tasks the activity of neurons in frontal and posterior parietal cortices covaries more with perceptual reports than with the physical properties of stimuli. This relationship is revealed when subjects have to make behavioral choices about weak or uncertain stimuli. If knowledge about stimulus onset time is available, decision making can be based on accumulation of sensory evidence. However, the time of stimulus onset or even its very presence is often ambiguous. By analyzing firing rates and correlated variability of frontal lobe neurons while monkeys perform a vibrotactile detection task, we show that behavioral outcomes are crucially affected by the state of cortical networks before stimulus onset times. The results suggest that sensory detection is partly due to a purely internal signal whereas the stimulus, if finally applied, adds a contribution to this initial processing later on. The probability to detect or miss the stimulus can thus be explained as the combined effect of this variable internal signal and the sensory evidence.

Augmented gamma band auditory steady-state responses: Support for NMDA hypofunction in schizophrenia

Individuals with schizophrenia (SZ) have deviations in auditory perception perhaps attributable to altered neural oscillatory response properties in thalamo-cortical and/or local cortico-cortical circuits. Previous EEG studies of auditory steady-state responses (aSSRs; a measure of sustained neuronal entrainment to repetitive stimulation) in SZ have indicated attenuated gamma range (≈40Hz) neural entrainment. Stimuli in most such studies have been relatively brief (500–1000ms) trains of 1ms clicks or amplitude modulated pure tones (1000Hz) with short, fixed interstimulus intervals (200–1000ms). The current study used extended (1500ms), more aurally dense broadband stimuli (500–4000Hz noise; previously demonstrated to elicit larger aSSRs) with longer, variable interstimulus intervals (2700–3300ms). Dense array EEG (256 sensor) was collected while 17 SZ and 16 healthy subjects passively listed to stimuli modulated at 15 different frequencies spanning beta and gamma ranges (16–44Hz in 2Hz steps). Results indicate that SZ have augmented aSSRs that were most extreme in the gamma range. Results also constructively replicate previous findings of attenuated low frequency auditory evoked responses (2–8Hz) in SZ. These findings (i) highlight differential characteristics of low versus high frequency and induced versus entrained oscillatory auditory responses in both SZ and healthy stimulus processing, (ii) provide support for an NMDA-receptor hypofunction-based pharmacological model of SZ, and (iii) report a novel pattern of aSSR abnormalities suggesting that gamma band neural entrainment deviations among SZ may be more complex than previously supposed, including possibly being substantially influenced by physical stimulus properties.

Identifying fragments of natural speech from the listener's MEG signals

It is a challenge for current signal analysis approaches to identify the electrophysiological brain signatures of continuous natural speech that the subject is listening to. To relate magnetoencephalographic (MEG) brain responses to the physical properties of such speech stimuli, we applied canonical correlation analysis (CCA) and a Bayesian mixture of CCA analyzers to extract MEG features related to the speech envelope. Seven healthy adults listened to news for an hour while their brain signals were recorded with whole-scalp MEG. We found shared signal time series (canonical variates) between the MEG signals and speech envelopes at 0.5-12 Hz. By splitting the test signals into equal-length fragments from 2 to 65 s (corresponding to 703 down to 21 pieces per the total speech stimulus) we obtained better than chance-level identification for speech fragments longer than 2-3 s, not used in the model training. The applied analysis approach thus allowed identification of segments of natural speech by means of partial reconstruction of the continuous speech envelope (i.e., the intensity variations of the speech sounds) from MEG responses, provided means to empirically assess the time scales obtainable in speech decoding with the canonical variates, and it demonstrated accurate identification of the heard speech fragments from the MEG data.

Auditory grouping mechanisms reflect a sound's relative position in a sequence

The human brain uses acoustic cues to decompose complex auditory scenes into its components. For instance to improve communication, a listener can select an individual “stream,” such as a talker in a crowded room, based on cues such as pitch or location. Despite numerous investigations into auditory streaming, few have demonstrated clear correlates of perception; instead, in many studies perception covaries with changes in physical stimulus properties (e.g., frequency separation). In the current report, we employ a classic ABA streaming paradigm and human electroencephalography (EEG) to disentangle the individual contributions of stimulus properties from changes in auditory perception. We find that changes in perceptual state—that is the perception of one versus two auditory streams with physically identical stimuli—and changes in physical stimulus properties are reflected independently in the event-related potential (ERP) during overlapping time windows. These findings emphasize the necessity of controlling for stimulus properties when studying perceptual effects of streaming. Furthermore, the independence of the perceptual effect from stimulus properties suggests the neural correlates of streaming reflect a tone's relative position within a larger sequence (1st, 2nd, 3rd) rather than its acoustics. By clarifying the role of stimulus attributes along with perceptual changes, this study helps explain precisely how the brain is able to distinguish a sound source of interest in an auditory scene.

Sequential Effects in Face-Attractiveness Judgment

A number of studies have shown that current-trial responses are biased toward the response of the preceding trial in perceptual decisionmaking tasks (the sequential effect-Holland and Lockhead, 1968 Perception & Psychophysics 3 409-414). The sequential effect has been widely observed in evaluation of the physical properties of stimuli as well as more complex properties. However, it is unclear whether subjective decisions (e.g., attractiveness judgments) are also susceptible to the sequential effect. Here, we examined whether the sequential effect would occur in face-attractiveness judgments. Forty-eight pictures of male and female faces were presented successively. Participants rated the attractiveness of each face on a 7-point scale. The results showed that the attractiveness rating of a given face assimilated toward the rating of the preceding trial. In a separate experiment, we provided the average attractiveness rating by others for each trial as feedback. The feedback weakened the sequential effect. These findings suggest that attractiveness judgment is also biased toward the preceding judgment, and hence the sequential effect can be extended into the domain of subjective decisionmaking.

Transient bold activity locked to perceptual reversals of auditory streaming in human auditory cortex and inferior colliculus

Our auditory system separates and tracks temporally interleaved sound sources by organizing them into distinct auditory streams. This streaming phenomenon is partly determined by physical stimulus properties but additionally depends on the internal state of the listener. As a consequence, streaming perception is often bistable and reversals between one- and two-stream percepts may occur spontaneously or be induced by a change of the stimulus. Here, we used functional MRI to investigate perceptual reversals in streaming based on interaural time differences (ITD) that produce a lateralized stimulus perception. Listeners were continuously presented with two interleaved streams, which slowly moved apart and together again. This paradigm produced longer intervals between reversals than stationary bistable stimuli but preserved temporal independence between perceptual reversals and physical stimulus transitions. Results showed prominent transient activity synchronized with the perceptual reversals in and around the auditory cortex. Sustained activity in the auditory cortex was observed during intervals where the ΔITD could potentially produce streaming, similar to previous studies. A localizer-based analysis additionally revealed transient activity time locked to perceptual reversals in the inferior colliculus. These data suggest that neural activity associated with streaming reversals is not limited to the thalamo-cortical system but involves early binaural processing in the auditory midbrain, already.

Expectations Change the Signatures and Timing of Electrophysiological Correlates of Perceptual Awareness

Previous experience allows the brain to predict what comes next. How these expectations affect conscious experience is poorly understood. In particular, it is unknown whether and when expectations interact with sensory evidence in granting access to conscious perception, and how this is reflected electrophysiologically. Here, we parametrically manipulate sensory evidence and expectations while measuring event-related potentials in human subjects to assess the time course of evoked responses that correlate with subjective visibility, the properties of the stimuli, and/or perceptual expectations. We found that expectations lower the threshold of conscious perception and reduce the latency of neuronal signatures differentiating seen and unseen stimuli. Without expectations, this differentiation occurs ∼300 ms and with expectations ∼200 ms after stimulus in occipitoparietal sensors. The amplitude of this differentiating response component (P2) decreases as visibility increases, regardless of whether this increase is attributable to enhanced sensory evidence and/or the gradual buildup of perceptual expectations. Importantly, at matched performance levels, responses to seen and unseen stimuli differed regardless of the physical stimulus properties. These findings indicate that the latency of the neuronal correlates of access to consciousness depend on whether access is driven by stimulus saliency or by a combination of expectations and sensory evidence.

Physical measurement of brain perception abilities. Foundations of a working methodology for the design of “intelligent” beings

Most of the important properties of the brain (thinking, consciousness, music, etc.) are severely ill-defined. They are not the direct output of biological sensors or their combinations but emerge from complex computations at the network level and are not necessarily represented in the sensory input or the activity of individual cells. They are emergent properties arising from dynamic interactions between neurons in the different relay stations of the sensory pathways where recognition of basic physical properties of incoming stimuli take place. Emergent properties and interactions between them range from physical properties of stimuli to cognitive operations as emotions or consciousness and gradually involve interactions between sensory pathways, associative cortexes, hippocampus, or the amygdala. Here we propose to build neural tissues from embryonic stem cells in “in vitro” controlled environments to determine the way physical inputs are transformed into what humans perceive and measure. We will start with “low complexity” tissues able to perform low level recognition of physical properties, to gradually increase the complexity of the tissue to investigate how the physical characteristics of the incoming stimuli correspond at higher levels to the emergent properties of the system. Mathematical methods based on networks theory, nonlinear dynamics, fractal theory and chaos among other will be used to determine and measure the emergent properties of the nervous tissue at different complexity levels. We expected to provide criteria and methodologies to measure human-like perception variables and use them for the design of future living artifacts (autonomous robots, intelligent sensors, hybrid systems, etc.).

Neural Systems Underlying Decisions about Affective Odors

Decision-making about affective value may occur after the reward value of a stimulus is represented and may involve different brain areas to those involved in decision-making about the physical properties of stimuli, such as intensity. In an fMRI study, we delivered two odors separated by a delay, with instructions on different trials to decide which odor was more pleasant or more intense or to rate the pleasantness and intensity of the second odor without making a decision. The fMRI signals in the medial prefrontal cortex area 10 (medial PFC) and in regions to which it projects, including the anterior cingulate cortex (ACC) and insula, were higher when decisions were being made compared with ratings, implicating these regions in decision-making. Decision-making about affective value was related to larger signals in the dorsal part of medial area 10 and the agranular insula, whereas decisions about intensity were related to larger activations in the dorsolateral prefrontal cortex (dorsolateral PFC), ventral premotor cortex, and anterior insula. For comparison, the mid orbitofrontal cortex (OFC) had activations related not to decision-making but to subjective pleasantness ratings, providing a continuous representation of affective value. In contrast, areas such as medial area 10 and the ACC are implicated in reaching a decision in which a binary outcome is produced.

Multiple serial picture presentation with millisecond resolution using a three-way LC-shutter-tachistoscope

Throughout recent years there has been an increasing interest in studying unconscious visual processes. Such conditions of unawareness are typically achieved by either a sufficient reduction of the stimulus presentation time or visual masking. However, there are growing concerns about the reliability of the presentation devices used. As all these devices show great variability in presentation parameters, the processing of visual stimuli becomes dependent on the display-device, e.g. minimal changes in the physical stimulus properties may have an enormous impact on stimulus processing by the sensory system and on the actual experience of the stimulus. Here we present a custom-built three-way LC-shutter-tachistoscope which allows experimental setups with both, precise and reliable stimulus delivery, and millisecond resolution. This tachistoscope consists of three LCD-projectors equipped with zoom lenses to enable stimulus presentation via a built-in mirror-system onto a back projection screen from an adjacent room. Two high-speed liquid crystal shutters are mounted serially in front of each projector to control the stimulus duration. To verify the intended properties empirically, different sequences of presentation times were performed while changes in optical power were measured using a photoreceiver. The obtained results demonstrate that interfering variabilities in stimulus parameters and stimulus rendering are markedly reduced. Together with the possibility to collect external signals and to send trigger-signals to other devices, this tachistoscope represents a highly flexible and easy to set up research tool not only for the study of unconscious processing in the brain but for vision research in general.