Is anterior N2 enhancement a reliable electrophysiological index of concealed information?

Concealed information tests (CITs) are used to determine whether an individual possesses information about an item of interest. Event-related potential (ERP) measures in CITs have focused almost exclusively on the P3b component, showing that this component is larger when lying about the item of interest (probe) than telling the truth about control items (irrelevants). Recent studies have begun to examine other ERP components, such as the anterior N2, with mixed results. A seminal CIT study found that visual probes elicit a larger anterior N2 than irrelevants (Gamer and Berti, 2010) and suggested that this component indexes cognitive control processes engaged when lying about probes. However, this study did not control for potential intrinsic differences among the stimuli: the same probe and irrelevants were used for all participants, and there was no control condition composed of uninformed participants. Here, first we show that the N2 effect found in the study by Gamer and Berti (2010) was in large part due to stimulus differences, as the effect observed in a concealed information condition was comparable to that found in two matched control conditions without any concealed information (Experiments 1 and 2). Next, we addressed the issue of the generality of the N2 findings by counterbalancing a new set of stimuli across participants and by using a control condition with uninformed participants (Experiment 3). Results show that the probe did not elicit a larger anterior N2 than the irrelevants under these controlled conditions. These findings suggest that caution should be taken in using the N2 as an index of concealed information in CITs. Furthermore, they are a reminder that results of CIT studies (not only with ERPs) performed without stimulus counterbalancing and suitable control conditions may be confounded by differential intrinsic properties of the stimuli employed.

The N170, not the P1, indexes the earliest time for categorical perception of faces, regardless of interstimulus variance.

A negative event-related potential (ERP) at occipito-temporal sites peaking around 150-170 ms after stimulus onset (N170) is typically larger for faces than other object categories. Most theories interpret this finding as due to face-selective processing in occipito-temporal and temporal cortex. However, a controversial account recently attributed the N170 effect to differences in interstimulus variance (ISV) among the images typically used for face and object conditions and proposed that the earlier P1 instead indexes the categorical processes generally attributed to the N170. This ERP study aimed to test this account definitively by using conditions in which the same face and object were shown repeatedly, eliminating both physical and perceptual ISV. Fourier amplitude spectra of faces and objects were matched to equate basic low-level visual properties that may affect early ERPs such as the P1. Results demonstrate that i) face selectivity of the N170 is largely preserved across many object categories after abolishing ISV, and ii) stimulus category does not modulate the P1. This conclusively refutes the ISV account while strongly supporting category as a critical factor driving N170 face selectivity.

Lying in the scanner: covert countermeasures disrupt deception detection by functional magnetic resonance imaging.

Functional magnetic resonance imaging (fMRI) studies have documented differences between deceptive and honest responses. Capitalizing on this research, companies marketing fMRI-based lie detection services have been founded, generating methodological and ethical concerns in scientific and legal communities. Critically, no fMRI study has examined directly the effect of countermeasures, methods used by prevaricators to defeat deception detection procedures. An fMRI study was conducted to fill this research gap using a concealed information paradigm in which participants were trained to use countermeasures. Robust group fMRI differences between deceptive and honest responses were found without, but not with countermeasures. Furthermore, in single participants, deception detection accuracy was 100% without countermeasures, using activation in ventrolateral and medial prefrontal cortices, but fell to 33% with countermeasures. These findings show that fMRI-based deception detection measures can be vulnerable to countermeasures, calling for caution before applying these methods to real-world situations.

Finding meaning in novel geometric shapes influences electrophysiological correlates of repetition and dissociates perceptual and conceptual priming.

Repeatedly viewing an object can engender fluency-related implicit memory for perceptual and conceptual attributes, as indexed in tests of perceptual and conceptual priming, respectively. Stimuli with minimal pre-experimental meaning allow direct comparisons between these two types of priming and explorations of whether corresponding neural mechanisms differ. We therefore examined electrophysiological correlates of perceptual and conceptual priming for minimalist geometric shapes (squiggles). Response time measures of conceptual priming were evident for squiggles rated by individual subjects as most meaningful, but not for those rated least meaningful. Conceptual-priming magnitude was proportional across individuals to the amplitude of FN400 brain potentials, but only for meaningful squiggles. Perceptual priming was evident for squiggles irrespective of meaningfulness, and perceptual-priming magnitude was proportional to the amplitude of frontal P170 potentials. These findings therefore show that a single exposure to a novel stimulus can lead to neural processing accompanying conceptual priming that is distinct from that accompanying perceptual priming (FN400 potentials vs. P170 potentials, respectively). Overall, this evidence is also relevant to the current debate over the neural correlates of familiarity-based recognition, and runs counter to the prominent supposition that familiarity can be generically indexed by FN400 potentials.

Object knowledge during entry-level categorization is activated and modified by implicit memory after 200 ms.

The timing of activating memory about visual objects is important for theories of human cognition but largely unknown, especially for tasks like entry level categorization that activate semantic memory. We tested an implicit memory-categorization "equivalence" hypothesis of multiple memory systems theory that a cortical system that stores structural knowledge to support entry level categorization also stores long-term, perceptual implicit memory, resulting in priming of this knowledge. Event-related brain potentials (ERPs) were recorded to impoverished pictures of new and repeated objects that were similar in perceptual properties but differed in categorization success. The cortical dynamics of object knowledge were defined using categorization ratings and naming. As predicted, rating, naming, and repetition effects on a frontocentral N350 show that implicit memory modifies the object knowledge network supporting categorization. This ERP is a complex of components between 200 and 500 ms indexing temporally overlapping substates from more perceptual to more conceptual knowledge. A frontopolar N350 subcomponent defines the first substate of a process of object model selection from occipitotemporal cortex based on shape similarity, and indicates that implicit memory in this system is greater with better categorization success. Afterwards, parietal positivity and a slow wave index secondary, post-model selection processes, like evaluating the success of a decision or memory match, and working memory for overt report, respectively. Altogether, ERP findings support the equivalence hypothesis and a two-state interactive account of visual object knowledge, and delineate the timing of multiple memory systems.

Role of a lateralized parietal-basal ganglia circuit in hierarchical pattern perception: evidence from Parkinson's disease.

The role of corticostriatal circuits in hierarchical pattern perception was examined in Parkinson's disease. The hypothesis was tested that patients with right-side onset of motor symptoms (RPD, left hemisphere dysfunction) would be impaired at local level processing because the left posterior temporoparietal junction (TP) emphasizes processing of local information. By contrast, left-side onset patients (LPD; right hemisphere dysfunction) would show impaired global processing because right TP emphasizes global processing. Participants identified targets at local or global levels without and with attention biased toward those levels. Despite normal attentional control between levels, LPD patients showed a single dissociation, demonstrating abnormal global level processing under all conditions, whereas RPD patients showed abnormal local level processing mainly when attention was biased toward the local level. These findings link side of motor symptom onset to visuospatial cognitive abilities that depend upon the contralateral TP, highlighting that side of onset can predict visuospatial impairments, and provide evidence that an inferior parietal-basal ganglia pathway involving the caudate head and the hemispherically asymmetrical TP region is necessary for hierarchical pattern perception.

Where vision meets memory: prefrontal-posterior networks for visual object constancy during categorization and recognition.

Objects seen from unusual relative to more canonical views require more time to categorize and recognize, and, according to object model verification theories, additionally recruit prefrontal processes for cognitive control that interact with parietal processes for mental rotation. To test this using functional magnetic resonance imaging, people categorized and recognized known objects from unusual and canonical views. Canonical views activated some components of a default network more on categorization than recognition. Activation to unusual views showed that both ventral and dorsal visual pathways, and prefrontal cortex, have key roles in visual object constancy. Unusual views activated object-sensitive and mental rotation (and not saccade) regions in ventrocaudal intraparietal, transverse occipital, and inferotemporal sulci, and ventral premotor cortex for verification processes of model testing on any task. A collateral-lingual sulci "place" area activated for mental rotation, working memory, and unusual views on correct recognition and categorization trials to accomplish detailed spatial matching. Ventrolateral prefrontal cortex and object-sensitive lateral occipital sulcus activated for mental rotation and unusual views on categorization more than recognition, supporting verification processes of model prediction. This visual knowledge framework integrates vision and memory theories to explain how distinct prefrontal-posterior networks enable meaningful interactions with objects in diverse situations.

Sensory Processing Sensitivity in the context of Environmental Sensitivity: A critical review and development of research agenda.

Sensory Processing Sensitivity (SPS) is a common, heritable and evolutionarily conserved trait describing inter-individual differences in sensitivity to both negative and positive environments. Despite societal interest in SPS, scientific knowledge is lagging behind. Here, we critically discuss how SPS relates to other theories, how to measure SPS, whether SPS is a continuous vs categorical trait, its relation to other temperament and personality traits, the underlying aetiology and neurobiological mechanisms, and relations to both typical and atypical development, including mental and sensory disorders. Drawing on the diverse expertise of the authors, we set an agenda for future research to stimulate the field. We conclude that SPS increases risk for stress-related problems in response to negative environments, but also provides greater benefit from positive and supportive experiences. The field requires more reliable and objective assessment of SPS, and deeper understanding of its mechanisms to differentiate it from other traits. Future research needs to target prevention of adverse effects associated with SPS, and exploitation of its positive potential to improve well-being and mental health.

An FMRI study of the role of the medial temporal lobe in implicit and explicit sequence learning.

fMRI was used to investigate the neural substrates supporting implicit and explicit sequence learning, focusing especially upon the role of the medial temporal lobe. Participants performed a serial reaction time task (SRTT). For implicit learning, they were naive about a repeating pattern, whereas for explicit learning, participants memorized another repeating sequence. fMRI analyses comparing repeating versus random sequence blocks demonstrated activation of frontal, parietal, cingulate, and striatal regions implicated in previous SRTT studies. Importantly, mediotemporal lobe regions were active in both explicit and implicit SRTT learning. Moreover, the results provide evidence of a role for the hippocampus and related cortices in the formation of higher order associations under both implicit and explicit learning conditions, regardless of conscious awareness of sequence knowledge.

Visual mental imagery and perception produce opposite adaptation effects on early brain potentials.

Event-related potentials (ERPs) were recorded during a rapid adaptation paradigm to determine whether visual perception and visual mental imagery of faces recruit the same early perceptual processes. The early effect of face and object adaptors, either perceived or visualized, on test stimuli, was assessed by measuring the amplitude of the N170/VPP complex, typically much larger for faces than for other object categories. Faces elicited a robust N170/VPP complex, localized to posterior ventrolateral occipitotemporal cortex. Both visualized and perceived adaptors affected the N170/VPP complex to test faces from 120 ms post-stimulus, reflecting effects on neural populations supporting early perceptual face categorization. Critically, while perceived adaptors suppressed the amplitude of the N170/VPP, visualized adaptors enhanced it. We suggest that perceived adaptors affect neural populations in the neocortex supporting early perceptual processing of faces via bottom-up mechanisms, whereas visualized adaptors affect them via top-down mechanisms. Similar enhancement effects were found on the N170/VPP complex to non-face objects, suggesting such effects are a general consequence of visual imagery on processing of faces and other object categories. These findings support image-percept equivalence theories and may explain, in part, why visual percepts and visual mental images are not routinely confused, even though both engage similar neural populations in the visual system.

HIV infection affects parietal-dependent spatial cognition: evidence from mental rotation and hierarchical pattern perception.

Human immunodeficiency virus (HIV) in the asymptomatic phase of the infection impairs some aspects of cognition, but little is known about how visuospatial functions are affected. In the present study, performance on tasks of mental rotation and hierarchical pattern perception was investigated in 14 HIV-positive men and 12 age- and education-matched HIV-negative men. Processes related to mental rotation of objects and hands were impaired in HIV-positive participants as compared to the HIV-negative group. The HIV-positive group was also impaired on hierarchical pattern perception of local targets under global biasing conditions. Consistent with these results, the HIV-positive participants showed impaired performance on standard clinical neuropsychological tests of visuospatial function. These findings indicate that the detrimental effects of HIV on cognition appear even in asymptomatic individuals and affect diverse visuospatial functions that depend upon the integrity of parietal brain regions.

Frontostriatal circuits are necessary for visuomotor transformation: mental rotation in Parkinson's disease.

The mental rotation of objects requires visuospatial functions mediated by the parietal lobes, whereas the mental rotation of hands also engages frontal motor-system processes. Nondemented patients with Parkinson's disease (PD), a frontostriatal disorder, were predicted to be impaired on mentally rotating hands. Side of PD motor symptom onset was investigated because the left motor cortices likely have a causal role in hand mental rotation. The prediction was that patients with right-side onset (RPD, greater left-hemisphere dysfunction) would commit more errors rotating hands than patients with left-side onset (LPD). Fifteen LPD, 12 RPD, and 13 normal control adults (NC) made same/different judgments about pairs of rotated objects or hands. There were no group differences with objects. When rotating hands, RPD, but not LPD, made more errors than the NC group. A control experiment evaluated whether visual field of presentation explained differences between PD subgroups. In the first experiment (1A), the hand to be mentally rotated was presented in the right visual field, but here (1B) it was presented in the left visual field. Only the LPD group made more errors than the NC group. The evidence suggests a double dissociation for the RPD and LPD groups between tasks differing in visual-field presentation. The findings indicate that hemifield location of a to-be-rotated hand stimulus can cause the hemispheric frontoparietal networks to be differentially engaged. Moreover, frontostriatal motor systems and the parietal lobes play a necessary role during the mental rotation of hands, which requires integrating visuospatial cognition with motor imagery.

Evidence for the importance of basal ganglia output nuclei in semantic event sequencing: an fMRI study.

Semantic event sequencing is the ability to plan ahead and order meaningful events chronologically. To investigate the neural systems supporting this ability, an fMRI picture sequencing task was developed. Participants sequenced a series of four pictures presented in random order based on the temporal relationship among them. A control object discrimination task was designed to be comparable to the sequencing task regarding semantic, visuospatial, and motor processing requirements but without sequencing demands. fMRI revealed significant activation in the dorsolateral prefrontal cortex and globus pallidus internal part in the picture sequencing task compared with the control task. The findings suggest that circuits involving the frontal lobe and basal ganglia output nuclei are important for picture sequencing and more generally for the sequential ordering of events. This is consistent with the idea that the basal ganglia output nuclei are critical not only for motor but also for high-level cognitive function, including behaviors involving meaningful information. We suggest that the interaction between the frontal lobes and basal ganglia output nuclei in semantic event sequencing can be generalized to include the sequential ordering of behaviors in which the selective updating of neural representations is the key computation.

Top-down modulation of visual processing and knowledge after 250 ms supports object constancy of category decisions.

People categorize objects more slowly when visual input is highly impoverished instead of optimal. While bottom-up models may explain a decision with optimal input, perceptual hypothesis testing (PHT) theories implicate top-down processes with impoverished input. Brain mechanisms and the time course of PHT are largely unknown. This event-related potential study used a neuroimaging paradigm that implicated prefrontal cortex in top-down modulation of occipitotemporal cortex. Subjects categorized more impoverished and less impoverished real and pseudo objects. PHT theories predict larger impoverishment effects for real than pseudo objects because top-down processes modulate knowledge only for real objects, but different PHT variants predict different timing. Consistent with parietal-prefrontal PHT variants, around 250 ms, the earliest impoverished real object interaction started on an N3 complex, which reflects interactive cortical activity for object cognition. N3 impoverishment effects localized to both prefrontal and occipitotemporal cortex for real objects only. The N3 also showed knowledge effects by 230 ms that localized to occipitotemporal cortex. Later effects reflected (a) word meaning in temporal cortex during the N400, (b) internal evaluation of prior decision and memory processes and secondary higher-order memory involving anterotemporal parts of a default mode network during posterior positivity (P600), and (c) response related activity in posterior cingulate during an anterior slow wave (SW) after 700 ms. Finally, response activity in supplementary motor area during a posterior SW after 900 ms showed impoverishment effects that correlated with RTs. Convergent evidence from studies of vision, memory, and mental imagery which reflects purely top-down inputs, indicates that the N3 reflects the critical top-down processes of PHT. A hybrid multiple-state interactive, PHT and decision theory best explains the visual constancy of object cognition.

Neurophysiological evidence for two processing times for visual object identification.

Event-related brain potentials (ERPs) were recorded to fragmented pictures of objects that were named correctly or were not to investigate the time course of visual object identification. The first ERP difference distinguishing identified from unidentified pictures estimates the upper limit of the time by which human brain regions have begun to activate long-term memory (LTM) representations specifying the identity of a visual object. Data from 15 young adults indicate that this time varies with the extent to which object parts are recoverable from the visual input, being approximately 200 ms earlier with recoverable than unrecoverable parts. Successful identification is evident by approximately 300 ms when object parts and overall structural configuration are readily recoverable but not until approximately 550 ms when object parts are difficult or impossible to recover (i.e. too poorly specified by the available contours to be recovered). In both cases, successful identification is associated with greater relative positivity. However, unidentified recoverable pictures are associated with an enhanced frontal negativity (N350), linked to object matching operations, not seen for non-recoverable pictures. Taken together, these results implicate two distinct processing sequences in the successful identification of visual objects.

Face-specificity is robust across diverse stimuli and individual people, even when interstimulus variance is zero.

The N170 is a brain electrical potential proposed to index the earliest time of categorical perception of faces in occipitotemporal visual areas implicated in face cognition, being more negative for faces than nonface objects between 120 and 200 ms. The interstimulus variance (ISV) account instead explained N170 face-specificity as an artifact due to objects varying more visually than faces. Ganis, Smith, and Schendan (2012) tested this account directly, finding that N170 face-specificity remains even when ISV is eliminated. Here, N170 peak amplitude and face-specificity is quantified for individual stimuli and participants, revealing that the right hemisphere N170 is especially sensitive to stimulus variability. Further, ISV contributes 0 to 37% to N170 face-specificity. These findings provide evidence for optimizing face processing science. The paradigm can apply to any research in which ISV may be uncontrolled (e.g., category comparisons).

Frontostriatal and mediotemporal lobe contributions to implicit higher-order spatial sequence learning declines in aging and Parkinson's disease.

Sequence learning depends on the striatal system, but recent findings also implicate the mediotemporal lobe (MTL) system. Schendan, Searl, Melrose, and Stern (2003) found higher-order associative, learning-related activation in the striatum, dorsolateral prefrontal cortex, and the MTL during the early acquisition phase of both implicit and explicit variants of a serial response time task. This functional MRI (fMRI) study capitalized on this task to determine how changes in MTL function observed in aging and compromised frontostriatal function characteristic of patients with Parkinson's disease (PD) impacts sequence learning and memory under implicit instructions. Brain activity was compared between "sequence" and "random" conditions in 12 nondemented patients with PD and education- and gender-matched healthy control participants of whom 12 were age matched (MC) and 14 were younger (YC). Behaviorally, sequence-specific learning of higher-order associations was reduced with aging and changed further with PD and resulted primarily in implicit knowledge in the older participants. fMRI revealed reduced intensity and extent of sequence learning-related activation in older relative to younger people in frontostriatal circuits and the MTL. This was because signal was greater for the sequence than random condition in younger people, whereas older people, especially those with PD, showed the opposite pattern. Both older groups also showed increased activation to the task itself relative to baseline fixation. In addition, right MTL showed hypoactivation and left MTL hyperactivation in PD relative to the MC group. The results suggest changes in frontostriatal and MTL activity occur during aging that affect task-related activity and the initial acquisition phase of implicit higher-order sequence learning. In addition, the results suggest that Parkinson's disease adversely affects processes in the MTL including sequence learning and memory.

Concealed semantic and episodic autobiographical memory electrified.

Electrophysiology-based concealed information tests (CIT) try to determine whether somebody possesses concealed information about a crime-related item (probe) by comparing event-related potentials (ERPs) between this item and comparison items (irrelevants). Although the broader field is sometimes referred to as "memory detection," little attention has been paid to the precise type of underlying memory involved. This study begins addressing this issue by examining the key distinction between semantic and episodic memory in the autobiographical domain within a CIT paradigm. This study also addresses the issue of whether multiple repetitions of the items over the course of the session habituate the brain responses. Participants were tested in a 3-stimulus CIT with semantic autobiographical probes (their own date of birth) and episodic autobiographical probes (a secret date learned just before the study). Results dissociated these two memory conditions on several ERP components. Semantic probes elicited a smaller frontal N2 than episodic probes, consistent with the idea that the frontal N2 decreases with greater pre-existing knowledge about the item. Likewise, semantic probes elicited a smaller central N400 than episodic probes. Semantic probes also elicited a larger P3b than episodic probes because of their richer meaning. In contrast, episodic probes elicited a larger late positive complex (LPC) than semantic probes, because of the recent episodic memory associated with them. All these ERPs showed a difference between probes and irrelevants in both memory conditions, except for the N400, which showed a difference only in the semantic condition. Finally, although repetition affected the ERPs, it did not reduce the difference between probes and irrelevants. These findings show that the type of memory associated with a probe has both theoretical and practical importance for CIT research.

Electrophysiological potentials reveal cortical mechanisms for mental imagery, mental simulation, and grounded (embodied) cognition.

Grounded cognition theory proposes that cognition, including meaning, is grounded in sensorimotor processing. The mechanism for grounding cognition is mental simulation, which is a type of mental imagery that re-enacts modal processing. To reveal top-down, cortical mechanisms for mental simulation of shape, event-related potentials were recorded to face and object pictures preceded by mental imagery. Mental imagery of the identical face or object picture (congruous condition) facilitated not only categorical perception (VPP/N170) but also later visual knowledge [N3(00) complex] and linguistic knowledge (N400) for faces more than objects, and strategic semantic analysis (late positive complex) between 200 and 700 ms. The later effects resembled semantic congruity effects with pictures. Mental imagery also facilitated category decisions, as a P3 peaked earlier for congruous than incongruous (other category) pictures, resembling the case when identical pictures repeat immediately. Thus mental imagery mimics semantic congruity and immediate repetition priming processes with pictures. Perception control results showed the opposite for faces and were in the same direction for objects: Perceptual repetition adapts (and so impairs) processing of perceived faces from categorical perception onward, but primes processing of objects during categorical perception, visual knowledge processes, and strategic semantic analysis. For both imagery and perception, differences between faces and objects support domain-specificity and indicate that cognition is grounded in modal processing. Altogether, this direct neural evidence reveals that top-down processes of mental imagery sustain an imagistic representation that mimics perception well enough to prime subsequent perception and cognition. Findings also suggest that automatic mental simulation of the visual shape of faces and objects operates between 200 and 400 ms, and strategic mental simulation operates between 400 and 700 ms.

Visual imagery.

Visual mental imagery is our ability to reactivate and manipulate visual representations in the absence of the corresponding visual stimuli, giving rise to the experience of 'seeing with the mind's eye'. Until relatively recently, visual mental imagery had been investigated by philosophy and cognitive psychology. However, these disciplines did not have the tools required to address empirically some of the important questions they had raised, for instance the extent to which visual mental images rely on some of the same representations that support visual perception. During the last two decades, cognitive neuroscience has leveraged the vast amount of knowledge about the neural basis of primate vision to provide new insights into visual mental imagery processes. Such insights enabled the empirical test of key questions about visual mental imagery using the armamentarium of tools provided by cognitive neuroscience, including electrophysiology and neuroimaging. Using a similar logic, we propose that information about the neural basis of memory systems should be used to further enhance understanding of the neural mechanisms of visual mental imagery. WIREs Cogni Sci 2011 2 239-252 DOI: 10.1002/wcs.103 For further resources related to this article, please visit the WIREs website.