Perceptual Reports Research Articles

Target Selection Signals Influence Perceptual Decisions by Modulating the Onset and Rate of Evidence Accumulation

Computational and neurophysiological research has highlighted neural processes that accumulate sensory evidence for perceptual decisions. These processes have been studied in the context of highly simplified perceptual discrimination paradigms in which the physical evidence appears at times and locations that are either entirely predictable or exogenously cued (e.g., by the onset of the stimulus itself). Yet, we are rarely afforded such certainty in everyday life. For example, when driving along a busy motorway, we must continually monitor the movements of surrounding vehicles for events that call for a lane change. In such scenarios, it is unknown which of the continuously present information sources will become relevant or when. Although it is well established that evidence integration provides an effective mechanism for countering the impact of noise, the question of how this mechanism is implemented in the face of uncertain evidence onsets has yet to be answered. Here, we show that when monitoring two potential sources of information for evidence occurring unpredictably in both time and space, the human brain employs discrete, early target selection signals that significantly modulate the onset and rate of neural evidence accumulation, and thereby the timing and accuracy of perceptual reports. These selection signals share many of the key characteristics of the N2pc component highlighted in the literature on visual search yet are present even in the absence of distractors and under situations of low temporal and spatial uncertainty. These data provide novel insights into how target selection supports decision making in uncertain environments.

Why the “stimulus-error” did not go away

Psychologists in the early years of the discipline were much concerned with the stimulus-error. Roughly, this is the problem encountered in introspective experiments when subjects are liable to frame their perceptual reports in terms of what they know of the stimulus, instead of just drawing on their perceptual experiences as they are supposedly felt. “Introspectionist” psychologist E. B. Titchener and his student E. G. Boring both argued in the early 20th century that the stimulus-error is a serious methodological pit-fall. While many of the theoretical suppositions motivating Titchener and Boring have been unfashionable since the rise of behaviourism, the stimulus-error brings our attention to one matter of perennial importance to psychophysics and the psychology of perception. This is the fact that subjects are liable to give different kinds of perceptual reports in response to the same stimulus. I discuss attempts to control for variable reports in recent experimental work on colour and lightness constancy, and the disputes that have arisen over which kinds of reports are legitimate. Some contemporary psychologists do warn us against a stimulus-error, even though they do not use this terminology. I argue that concern over the stimulus-error is diagnostic of psychologists' deep theoretical commitments, such as their conception of sensation, or their demarcation of perception from cognition. I conclude by discussing the relevance of this debate to current philosophy of perception.

Trunk rotation affects temporal order judgments with direct saccades: Influence of handedness

Manipulation of the trunk midline has been shown to improve visuospatial performance in patients with unilateral visual neglect. The goal of the present study was to disentangle motor and perceptual components of egocentric midline manipulations and to investigate the contribution of individual hand preference. Two versions of visual temporal order judgment (TOJ) tasks were tested in healthy right- and left-handed subjects while trunk rotation was varied. In the congruent version, subjects were required to execute a saccade to the first of two horizontal stimuli presented with different stimulus onset asynchronies (SOA). In the incongruent version, subjects were required to perform a vertical saccade to a pre-learned color target, thereby dissociating motor response from the perceptual stimulus location. The main findings of this study are a trunk rotation and response direction specific impact on temporal judgments in form of a prior entry bias for right hemifield stimuli during rightward trunk rotation, but only in the congruent task. This trunk rotation-induced spatial bias was most pronounced in left-handed participants but had the same sign in the right-handed group. Results suggest that egocentric midline shifts in healthy subjects induce a spatially-specific motor, but not a perceptual, bias and underline the importance of taking individual differences in functional laterality such as handedness and mode of perceptual report into account when evaluating effects of trunk rotation in either healthy subjects or neurological patients.

Inattentional blindness reflects limitations on perception, not memory: Evidence from repeated failures of awareness

Perhaps the most striking phenomenon of visual awareness is inattentional blindness (IB), in which a surprisingly salient event right in front of you may go completely unseen when unattended. Does IB reflect a failure of perception, or only of subsequent memory? Previous work has been unable to answer this question, due to a seemingly intractable dilemma: ruling out memory requires immediate perceptual reports, but soliciting such reports fuels an expectation that eliminates IB. Here we introduce a way of evoking repeated IB in the same subjects and the same session: we show that observers fail to report seeing salient events’ not only when they have no expectation, but also when they have the wrong expectations about the events nature. This occurs when observers must immediately report seeing anything unexpected, even mid-event. Repeated IB thus demonstrates that IB is aptly named: it reflects a genuine deficit in moment-by-moment conscious perception, rather than a form of inattentional amnesia.

Evidence for a functional and anatomical dissociation in the use of size constancy for perceptual report and goal-directed grasping

Evidence for a functional and anatomical dissociation in the use of size constancy for perceptual report and goal-directed grasping

On the brain-imaging markers of neural correlates of consciousness.

For many years, since Baars (1988) explicitly formulated it, contrastive analysis has been the key methodological approach in experimental studies of consciousness. When certain properly chosen psychological experimental setups (allowing an invariant target stimulus either to be consciusly experienced or not) were combined with brain-imaging methods, contrastive analysis became a quite powerful tool of research (Crick, 1994; Koch, 2004). By subtracting markers of brain processes recorded in the conditions without conscious experience of the target from the markers recorded in the conditions where the same target is consciously experienced it was believed that the markers of neural correlates of consciousness (NCC) can be obtained. However, as it turned out in the subsequent theoretical and experimental analysis, the picture is not so clear and simple (Bachmann, 2000, 2009; Miller, 2007; Aru et al., 2012; de Graaf et al., 2012). For example, when in the invariant conditions of independent variables a masked visual stimulus was consciously perceived or not (consciousness of the target standed as a dependent variable), NCC which were measured as a spectral perturbation of EEG was present already before stimulus presentation (Aru and Bachmann, 2009). Thus, the neural correlate of consciousness of a stimulus was present earlier than the stimulus itself was presented. Now, a reader must not get excited here because instead of some paranormal explanations brain-science based explanations can be comfortably used. In order to overcome the conceptual crisis hitting the traditional contrastive analysis based NCC research it was suggested that unconscious prerequisite processes (NCCpr) emerging as a result of contrastive analysis of brain-process markers of consciousness and similarly unconscious consequent processes (NCCco) must be differentiated from the constitutive processes directly associated with conscious experience (Aru et al., 2012; de Graaf et al., 2012). Thus, new experimental approaches were in need to avoid the trap of distilling prerequisite, direct, and consequent processes as mutually confounded and empirically inseparable. Despite some first attempts in this direction (Aru and Bachmann, 2015), the specialist landscape in this domain has remained obscure and no breakthrough solutions have been in sight. Moreover, there seems to be a number of additional uncertainties when we try to disentangle the various sub-types of NCC. Even NCCpr and NCCco are not unitary in terms of their theoretical meaning and associated neural processes. First, as the contents on which the perceptual report is founded can be selective, the markers of unused conscious contents may be erroneously neglected as markers of unconscious processes. They actually belong to consciousness level processes, but related to contents of consciousness qualitatively different from the ones specified by NCC. Second, in measuring NCC we must be able to disentangle contributions of the general consciousness enabling mechanisms and the selective contents representing mechanisms because their markers can be different and thus confused. In what follows I will substantiate these two issues.

Shared sensory estimates for human motion perception and pursuit eye movements.

Are sensory estimates formed centrally in the brain and then shared between perceptual and motor pathways or is centrally represented sensory activity decoded independently to drive awareness and action? Questions about the brain's information flow pose a challenge because systems-level estimates of environmental signals are only accessible indirectly as behavior. Assessing whether sensory estimates are shared between perceptual and motor circuits requires comparing perceptual reports with motor behavior arising from the same sensory activity. Extrastriate visual cortex both mediates the perception of visual motion and provides the visual inputs for behaviors such as smooth pursuit eye movements. Pursuit has been a valuable testing ground for theories of sensory information processing because the neural circuits and physiological response properties of motion-responsive cortical areas are well studied, sensory estimates of visual motion signals are formed quickly, and the initiation of pursuit is closely coupled to sensory estimates of target motion. Here, we analyzed variability in visually driven smooth pursuit and perceptual reports of target direction and speed in human subjects while we manipulated the signal-to-noise level of motion estimates. Comparable levels of variability throughout viewing time and across conditions provide evidence for shared noise sources in the perception and action pathways arising from a common sensory estimate. We found that conditions that create poor, low-gain pursuit create a discrepancy between the precision of perception and that of pursuit. Differences in pursuit gain arising from differences in optic flow strength in the stimulus reconcile much of the controversy on this topic.

Action and perception are temporally coupled by a common mechanism that leads to a timing misperception.

We move our eyes to explore the world, but visual areas determining where to look next (action) are different from those determining what we are seeing (perception). Whether, or how, action and perception are temporally coordinated is not known. The preparation time course of an action (e.g., a saccade) has been widely studied with the gap/overlap paradigm with temporal asynchronies (TA) between peripheral target onset and fixation point offset (gap, synchronous, or overlap). However, whether the subjects perceive the gap or overlap, and when they perceive it, has not been studied. We adapted the gap/overlap paradigm to study the temporal coupling of action and perception. Human subjects made saccades to targets with different TAs with respect to fixation point offset and reported whether they perceived the stimuli as separated by a gap or overlapped in time. Both saccadic and perceptual report reaction times changed in the same way as a function of TA. The TA dependencies of the time change for action and perception were very similar, suggesting a common neural substrate. Unexpectedly, in the perceptual task, subjects misperceived lights overlapping by less than ∼100 ms as separated in time (overlap seen as gap). We present an attention-perception model with a map of prominence in the superior colliculus that modulates the stimulus signal's effectiveness in the action and perception pathways. This common source of modulation determines how competition between stimuli is resolved, causes the TA dependence of action and perception to be the same, and causes the misperception.

Inattentional blindness reflects limitations on perception, not memory: Evidence from repeated failures of awareness.

Perhaps the most striking phenomenon of visual awareness is inattentional blindness (IB), in which a surprisingly salient event right in front of you may go completely unseen when unattended. Does IB reflect a failure of perception, or only of subsequent memory? Previous work has been unable to answer this question, due to a seemingly intractable dilemma: ruling out memory requires immediate perceptual reports, but soliciting such reports fuels an expectation that eliminates IB. Here we introduce a way of evoking repeated IB in the same subjects and the same session: we show that observers fail to report seeing salient events' not only when they have no expectation, but also when they have the wrong expectations about the events nature. This occurs when observers must immediately report seeing anything unexpected, even mid-event. Repeated IB thus demonstrates that IB is aptly named: it reflects a genuine deficit in moment-by-moment conscious perception, rather than a form of inattentional amnesia.

Measuring perception without introspection.

Binocular rivalry, the perceptual alternation between incompatible monocular stimuli, is conventionally measured by asking the subject which percept is currently visible. This is problematic because the response is unverifiable, open to response bias, and falsely assumes that the perceptual experience is binary. We overcame these limitations in a new approach that does not require subjective reporting of perceptual state. A brief test stimulus was added to one eye's inducing stimulus at random times and contrasts. The test was presented at one of two spatial locations, the subject indicated which alternative had been shown, and the correctness of the response was recorded as a function of test contrast. Given the random timing of the test stimulus, it was sometimes delivered when the tested eye was dominant and, at other times, suppressed. Accordingly, the psychometric function recorded during rivalry should be a mixture of the dominance and suppression forms of the function. This was indeed the case: The probability of a correct response during rivalry was significantly less than that obtained with a binocularly congruent stimulus. The psychometric function during rivalry was well modeled as a weighted sum of the congruence curve with an assumed suppression curve. Optimal fitting provided estimates of both suppression depth and percept predominance that corresponded closely with estimates obtained with the conventional method. We have therefore characterized rivalry without the uncertainties introduced by the subject's perceptual report. This provides a model that may be applicable to the broader field of perceptual ambiguity.

Feature Predictability Flexibly Supports Auditory Stream Segregation or Integration

Many sound sources emit series of discrete sounds. Auditory perception must bind these sounds together (stream integration) while separating them from sounds emitted by other sources (stream segregation). One cue for identifying successive sounds that belong together is the predictability between their feature values. Previous studies have demonstrated that independent predictable patterns appearing separately in two interleaved sound sequences support perceptual segregation. The converse case, whether a joint predictable pattern in a mixture of interleaved sequences supports perceptual integration, has not yet been put to a rigorous empirical test. This was mainly due to difficulties in manipulating the predictability of the full sequence independently of the predictability of the interleaved subsequences. The present study implemented such an independent manipulation. Listeners continuously indicated whether they perceived a tone sequence as integrated or segregated, while predictable patterns set up to support one or the other percept were manipulated without the participants’ knowledge. Perceptual reports demonstrate that predictability supports stream segregation or integration depending on the type of predictable pattern that is present in the sequence. The effects of predictability were so pronounced as to qualitatively flip perception from predominantly (62%) integrated to predominantly (73%) segregated. These results suggest that auditory perception flexibly responds to encountered regular patterns, favoring predictable perceptual organizations over unpredictable ones. Besides underlining the role of predictability as a cue within auditory scene analysis, the present design also provides a general framework that accommodates previous investigations focusing on sub-comparisons within the present set of experimental manipulations. Results of intermediate conditions shed light on why some previous studies have obtained little to no effects of predictability on auditory scene analysis.

Perceptual hysteresis as a marker of perceptual inflexibility in schizophrenia

People with schizophrenia are known to exhibit difficulties in the updating of their current belief states even in the light of disconfirmatory evidence. In the present study we tested the hypothesis that people with schizophrenia could also manifest perceptual inflexibility, or difficulties in the updating of their current sensory states. The presence of perceptual inflexibility might contribute both to the patients' altered perception of reality and the formation of some delusions as well as to their social cognition deficits. Here, we addressed this issue with a protocol of auditory hysteresis, a direct measure of sensory persistence, on a population of stabilized antipsychotic-treated schizophrenia patients and a sample of control subjects. Trials consisted of emotional signals (i.e., screams) and neutral signals (i.e., spectrally-rotated versions of the emotional stimuli) progressively emerging from white noise - Ascending Sequences - or progressively fading away in white noise - Descending Sequences. Results showed that patients presented significantly stronger hysteresis effects than control subjects, as evidenced by a higher rate of perceptual reports in Descending Sequences. The present study thus provides direct evidence of perceptual inflexibility in schizophrenia.

Beta, but not gamma, band oscillations index visual form-motion integration.

Electrophysiological oscillations in different frequency bands co-occur with perceptual, motor and cognitive processes but their function and respective contributions to these processes need further investigations. Here, we recorded MEG signals and seek for percept related modulations of alpha, beta and gamma band activity during a perceptual form/motion integration task. Participants reported their bound or unbound perception of ambiguously moving displays that could either be seen as a whole square-like shape moving along a Lissajou's figure (bound percept) or as pairs of bars oscillating independently along cardinal axes (unbound percept). We found that beta (15–25 Hz), but not gamma (55–85 Hz) oscillations, index perceptual states at the individual and group level. The gamma band activity found in the occipital lobe, although significantly higher during visual stimulation than during base line, is similar in all perceptual states. Similarly, decreased alpha activity during visual stimulation is not different for the different percepts. Trial-by-trial classification of perceptual reports based on beta band oscillations was significant in most observers, further supporting the view that modulation of beta power reliably index perceptual integration of form/motion stimuli, even at the individual level.

Ongoing behavior predicts perceptual report of interval duration

The ability to estimate the passage of time is essential for adaptive behavior in complex environments. Yet, it is not known how the brain encodes time over the durations necessary to explain animal behavior. Under temporally structured reinforcement schedules, animals tend to develop temporally structured behavior, and interval timing has been suggested to be accomplished by learning sequences of behavioral states. If this is true, trial to trial fluctuations in behavioral sequences should be predictive of fluctuations in time estimation. We trained rodents in an duration categorization task while continuously monitoring their behavior with a high speed camera. Animals developed highly reproducible behavioral sequences during the interval being timed. Moreover, those sequences were often predictive of perceptual report from early in the trial, providing support to the idea that animals may use learned behavioral patterns to estimate the duration of time intervals. To better resolve the issue, we propose that continuous and simultaneous behavioral and neural monitoring will enable identification of neural activity related to time perception that is not explained by ongoing behavior.

Statistical and perceptual updating: correlated impairments in right brain injury

It has been hypothesized that many of the cognitive impairments commonly seen after right brain damage (RBD) can be characterized as a failure to build or update mental models. We (Danckert et al. in Neglect as a disorder of representational updating. NOVA Open Access, New York, 2012a; Cereb Cortex 22:2745-2760, 2012b) were the first to directly assess the association between RBD and updating and found that RBD patients were unable to exploit a strongly biased play strategy in their opponent in the children's game rock, paper, scissors. Given that this game required many other cognitive capacities (i.e., working memory, sustained attention, reward processing), RBD patients could have failed this task for various reasons other than a failure to update. To assess the generality of updating deficits after RBD, we had RBD, left brain-damaged (LBD) patients and healthy controls (HCs) describe line drawings that evolved gradually from one figure (e.g., rabbit) to another (e.g., duck) in addition to the RPS updating task. RBD patients took significantly longer to alter their perceptual report from the initial object to the final object than did LBD patients and HCs. Although both patient groups performed poorly on the RPS task, only the RBD patients showed a significant correlation between the two, very different, updating tasks. We suggest these data indicate a general deficiency in the ability to update mental representations following RBD.

Internal and External Influences on the Rate of Sensory Evidence Accumulation in the Human Brain

We frequently need to make timely decisions based on sensory evidence that is weak, ambiguous, or noisy resulting from conditions in the external environment (e.g., a cluttered visual scene) or within the brain itself (e.g., inattention, neural noise). Here we examine how externally and internally driven variations in the quality of sensory evidence affect the build-to-threshold dynamics of a supramodal "decision variable" signal and, hence, the timing and accuracy of decision reports in humans. Observers performed a continuous-monitoring version of the prototypical two-alternative dot-motion discrimination task, which is known to strongly benefit from sequential sampling and temporal accumulation of evidence. A centroparietal positive potential (CPP), which we previously established as a supramodal decision signal based on its invariance to motor or sensory parameters, exhibited two key identifying properties associated with the "decision variable" long described in sequential sampling models: (1) its buildup rate systematically scaled with sensory evidence strength across four levels of motion coherence, consistent with temporal integration; and (2) its amplitude reached a stereotyped level at the moment of perceptual report executions, consistent with a boundary-crossing stopping criterion. The buildup rate of the CPP also strongly predicted reaction time within coherence levels (i.e., independent of physical evidence strength), and this endogenous variation was linked with attentional fluctuations indexed by the level of parieto-occipital α-band activity preceding target onset. In tandem with the CPP, build-to-threshold dynamics were also observed in an effector-selective motor preparation signal; however, the buildup of this motor-specific process significantly lagged that of the supramodal process.

Essential kinematic information, athletic experience, and affordance perception for others

The present study investigated the role of different types of movement in affordance perception, as well as the influence of sports experience. Perception of another actor's maximum vertical jumping height and horizontal long-jumping distance was evaluated for basketball players, soccer players, and nonplayer controls after viewing point-light representations of the actors' movements. Perceptual reports were more accurate after jumping-related movements (walking and squatting) were viewed than after nonrelated movements (standing and twisting). Vertical jump reports were more accurate than horizontal jump reports. Basketball and soccer players demonstrated higher accuracy than did controls. This research establishes that point-light displays contain essential kinematic information sufficient to support accurate affordance perception, and athletes appear better attuned to kinematic information specifying affordances for others as a result of their sports experience.

The what, where and how of auditory-object perception

The fundamental perceptual unit in hearing is the 'auditory object'. Similar to visual objects, auditory objects are the computational result of the auditory system's capacity to detect, extract, segregate and group spectrotemporal regularities in the acoustic environment; the multitude of acoustic stimuli around us together form the auditory scene. However, unlike the visual scene, resolving the component objects within the auditory scene crucially depends on their temporal structure. Neural correlates of auditory objects are found throughout the auditory system. However, neural responses do not become correlated with a listener's perceptual reports until the level of the cortex. The roles of different neural structures and the contribution of different cognitive states to the perception of auditory objects are not yet fully understood.

Different roles of similarity and predictability in auditory stream segregation

Sound sources often emit trains of discrete sounds, such as a series of footsteps. Previously, two different principles have been suggested for how the human auditory system binds discrete sounds together into perceptual units. The feature similarity principle is based on linking sounds with similar characteristics over time. The predictability principle is based on linking sounds that follow each other in a predictable manner. The present study compared the effects of these two principles. Participants were presented with tone sequences and instructed to continuously indicate whether they perceived a single coherent sequence or two concurrent streams of sound. We investigated the influence of separate manipulations of similarity and predictability on these perceptual reports. Both grouping principles affected perception of the tone sequences, albeit with different characteristics. In particular, results suggest that whereas predictability is only analyzed for the currently perceived sound organization, f...

Population Responses to Contour Integration: Early Encoding of Discrete Elements and Late Perceptual Grouping

The neuronal mechanisms underlying perceptual grouping of discrete, similarly oriented elements are not well understood. To investigate this, we measured neural population responses using voltage-sensitive dye imaging in V1 of monkeys trained on a contour-detection task. By mapping the contour and background elements onto V1, we could study their neural processing. Population response early in time showed activation patches corresponding to the contour/background individual elements. However, late increased activity in the contour elements, along with suppressed activity in the background elements, enabled us to visualize in single trials a salient continuous contour "popping out" from a suppressed background. This modulated activity in the contour and in background extended beyond the cortical representation of individual contour or background elements. Finally, the late modulation was correlated with behavioral performance of contour saliency and the monkeys' perceptual report. Thus, opposing responses in the contour and background may underlie perceptual grouping in V1.