Papers by Emiliano Macaluso
Human Brain Mapping, 2015
The Brief presentation of a complex scene entails that only a few objects can be selected, proces... more The Brief presentation of a complex scene entails that only a few objects can be selected, processed indepth, and stored in memory. Both low‐level sensory salience and high‐level context‐related factors (e.g., the conceptual match/mismatch between objects and scene context) contribute to this selection process, but how the interplay between these factors affects memory encoding is largely unexplored. Here, during fMRI we presented participants with pictures of everyday scenes. After a short retention interval, participants judged the position of a target object extracted from the initial scene. The target object could be either congruent or incongruent with the context of the scene, and could be located in a region of the image with maximal or minimal salience. Behaviourally, we found a reduced impact of saliency on visuospatial working memory performance when the target was out‐of‐context. Encoding‐related fMRI results showed that context–congruent targets activated dorsoparietal r...
RETRIEVAL, 24h later (scanned) • Right ventral fronto-parietal cortices (vFP) are recruited when ... more RETRIEVAL, 24h later (scanned) • Right ventral fronto-parietal cortices (vFP) are recruited when detected stimuli are related to a task goal 1 , even from memory 2. • Set-relevancy of distractors can aid retrieval, and elicits ventral parietal activation 3
Frontiers in Human Neuroscience, 2016
In daily life the brain is exposed to a large amount of external signals that compete for process... more In daily life the brain is exposed to a large amount of external signals that compete for processing resources. The attentional system can select relevant information based on many possible combinations of goal-directed and stimulus-driven control signals. Here, we investigate the behavioral and physiological effects of competition between distinctive visual events during free-viewing of naturalistic videos. Nineteen healthy subjects underwent functional magnetic resonance imaging (fMRI) while viewing short video-clips of everyday life situations, without any explicit goal-directed task. Each video contained either a single semantically-relevant event on the left or right side (Lat-trials), or multiple distinctive events in both hemifields (Multi-trials). For each video, we computed a salience index to quantify the lateralization bias due to stimulus-driven signals, and a gaze index (based on eye-tracking data) to quantify the efficacy of the stimuli in capturing attention to either side. Behaviorally, our results showed that stimulus-driven salience influenced spatial orienting only in presence of multiple competing events (Multitrials). fMRI results showed that the processing of competing events engaged the ventral attention network, including the right temporoparietal junction (R TPJ) and the right inferior frontal cortex. Salience was found to modulate activity in the visual cortex, but only in the presence of competing events; while the orienting efficacy of Multitrials affected activity in both the visual cortex and posterior parietal cortex (PPC). We conclude that in presence of multiple competing events, the ventral attention system detects semantically-relevant events, while regions of the dorsal system make use of saliency signals to select relevant locations and guide spatial orienting.
Human Brain Mapping, 2015
Several methods are available for the identification of functional networks of brain areas using ... more Several methods are available for the identification of functional networks of brain areas using functional magnetic resonance imaging (fMRI) time-series. These typically assume a fixed relationship between the signal of the areas belonging to the same network during the entire time-series (e.g., positive correlation between the areas belonging to the same network), or require a priori information about when this relationship may change (task-dependent changes of connectivity). We present a fully datadriven method that identifies transient network configurations that are triggered by the external input and that, therefore, include only regions involved in stimulus/task processing. Intersubject synchronization with short sliding time-windows was used to identify if/when any area showed stimulus/taskrelated responses. Next, a first clustering step grouped together areas that became engaged concurrently and repetitively during the time-series (stimulus/task-related networks). Finally, for each network, a second clustering step grouped together all the time-windows with the same BOLD signal. The final output consists of a set of network configurations that show stimulus/task-related activity at specific timepoints during the fMRI time-series. We label these configurations: "brain modes" (bModes). The method was validated using simulated datasets and a real fMRI experiment with multiple tasks and conditions. Future applications include the investigation of brain functions using complex and naturalistic stimuli.
Scientific Reports, 2015
The fractionation view holds that distinct cognitive operations are mediated by subregions of the... more The fractionation view holds that distinct cognitive operations are mediated by subregions of the inferior parietal lobule (IPL). Within IPL, we hypothesised that retrieval-related activity in different parts of the right supramarginal gyrus (rSMG) may be modulated differentially by information acquired via different combinations of attention signals at encoding. We had two groups of participants watch a 42-min TV episode and, after a 24-hr delay, perform a temporal-order judgment task during fMRI. Each retrieval trial comprised three images presented sequentially, requiring participants to judge the temporal order between the first and last images while ignoring the second image (“distractor”). We manipulated the bottom-up factor by presenting distractors that were extracted from either an event-boundary or a non-boundary of the movie. The top-down factor was manipulated by instructing one group perform a segmentation task reporting the event-boundaries at encoding, while the other...
Cognition, 2015
Recent demonstrations of scale invariance in cognitive domains prompted us to investigate whether... more Recent demonstrations of scale invariance in cognitive domains prompted us to investigate whether a scale-free pattern might exist in retrieving the temporal order of events from episodic memory. We present four experiments using an encoding-retrieval paradigm with naturalistic stimuli (movies or video clips). Our studies show that temporal order judgement retrieval times were negatively correlated with the temporal separation between two events in the movie. This relation held, irrespective of whether temporal distances were on the order of tens of minutes (Exp 1À2) or just a few seconds (Exp 3À4). Using the SIMPLE model, we factored in the retention delays between encoding and retrieval (delays of 24 h, 15 min, 1.5-2.5 s, and 0.5 s for Exp 1-4, respectively) and computed a temporal similarity score for each trial. We found a positive relation between similarity and retrieval times; that is, the more temporally similar two events, the slower the retrieval of their temporal order. Using Bayesian analysis, we confirmed the equivalence of the RT/similarity relation across all experiments, which included a vast range of temporal distances and retention delays. These results provide evidence for scale invariance during the retrieval of temporal order of episodic memories.
Human brain mapping, Jan 12, 2015
We investigated the neural correlates supporting three kinds of memory judgments after very short... more We investigated the neural correlates supporting three kinds of memory judgments after very short delays using naturalistic material. In two functional magnetic resonance imaging (fMRI) experiments, subjects watched short movie clips, and after a short retention (1.5-2.5 s), made mnemonic judgments about specific aspects of the clips. In Experiment 1, subjects were presented with two scenes and required to either choose the scene that happened earlier in the clip ("scene-chronology"), or with a correct spatial arrangement ("scene-layout"), or that had been shown ("scene-recognition"). To segregate activity specific to seen versus unseen stimuli, in Experiment 2 only one probe image was presented (either target or foil). Across the two experiments, we replicated three patterns underlying the three specific forms of memory judgment. The precuneus was activated during temporal-order retrieval, the superior parietal cortex was activated bilaterally for spat...
Human brain mapping, Jan 17, 2015
The ability to detect changes in the environment is necessary for appropriate interactions with t... more The ability to detect changes in the environment is necessary for appropriate interactions with the external world. Changes in the background go more unnoticed than foreground changes, possibly because attention prioritizes processing of foreground/near stimuli. Here, we investigated the detectability of foreground and background changes within natural scenes and the influence of stereoscopic depth cues on this. Using a flicker paradigm, we alternated a pair of images that were exactly same or differed for one single element (i.e., a color change of one object in the scene). The participants were asked to find the change that occurred either in a foreground or background object, while viewing the stimuli either with binocular and monocular cues (bmC) or monocular cues only (mC). The behavioral results showed faster and more accurate detections for foreground changes and overall better performance in bmC than mC conditions. The imaging results highlighted the involvement of fronto-pa...
PloS one, 2013
The use of naturalistic stimuli to probe sensory functions in the human brain is gaining increasi... more The use of naturalistic stimuli to probe sensory functions in the human brain is gaining increasing interest. Previous imaging studies examined brain activity associated with the processing of cinematographic material using both standard "condition-based" designs, as well as "computational" methods based on the extraction of time-varying features of the stimuli (e.g. motion). Here, we exploited both approaches to investigate the neural correlates of complex visual and auditory spatial signals in cinematography. In the first experiment, the participants watched a piece of a commercial movie presented in four blocked conditions: 3D vision with surround sounds (3D-Surround), 3D with monaural sound (3D-Mono), 2D-Surround, and 2D-Mono. In the second experiment, they watched two different segments of the movie both presented continuously in 3D-Surround. The blocked presentation served for standard condition-based analyses, while all datasets were submitted to computati...
The Journal of Neuroscience, 2004
Two identical stimuli, such as a pair of electrical shocks to the skin, are readily perceived as ... more Two identical stimuli, such as a pair of electrical shocks to the skin, are readily perceived as two separate events in time provided the interval between them is sufficiently long. However, as they are presented progressively closer together, there comes a point when the two separate stimuli are perceived as a single stimulus. Damage to posterior parietal cortex, peri-supplementary motor area (peri-SMA), and basal ganglia can disturb this form of temporal discrimination. Our aim was to establish, in healthy subjects, the brain areas that are involved in this process. During functional magnetic resonance imaging scanning, paired electrical pulses, separated by variable inter-stimulus intervals (5-110 msec), were delivered to different sites on one forearm (8-64 mm from the midline). Subjects were required to simply detect the stimulus (control task) or to identify a stimulus property. For temporal discrimination (TD), subjects reported whether they felt one or two stimuli. For spati...
Neuron, 2011
In everyday life attention operates within complex and dynamic environments, while laboratory par... more In everyday life attention operates within complex and dynamic environments, while laboratory paradigms typically employ simple and stereotyped stimuli. This fMRI study investigated stimulus-driven spatial attention using a virtual-environment video. We explored the influence of bottom-up signals by computing saliency maps of the environment and by introducing attention-grabbing events in the video. We parameterized the efficacy of these signals for the orienting of spatial attention by measuring eye movements and used these parameters to analyze the imaging data. The efficacy of bottom-up signals modulated ongoing activity in dorsal fronto-parietal regions and transient activation of the ventral attention system. Our results demonstrate that the combination of computational, behavioral, and imaging techniques enables studying cognitive functions in ecologically valid contexts. We highlight the central role of the efficacy of stimulus-driven signals in both dorsal and ventral attention systems, with a dissociation of the efficacy of background salience versus distinctive events in the two systems. Neuron Eco-Orienting
NeuroImage, 2006
When two identical stimuli, such as a pair of clicks, are presented with a sufficiently long time... more When two identical stimuli, such as a pair of clicks, are presented with a sufficiently long time-interval between them they are readily perceived as two separate events. However, as they are presented progressively closer together, there comes a point when the two separate stimuli are perceived as one. This phenomenon applies not only to hearing but also to other sensory modalities. Damage to the basal ganglia disturbs this type of temporal discrimination irrespective of sensory modality, suggesting a multimodal process is involved. Our aim was to study the neural substrate of auditory temporal discrimination in healthy subjects and to compare it with structures previously associated with analogous tactile temporal discrimination. During fMRI scanning, paired-clicks separated by variable interstimulus intervals (1-50 ms) were delivered binaurally, with different intensities delivered to each ear, yielding a lateralised auditory percept. Subjects were required (a) to report whether they heard one or two stimuli (TD: temporal discrimination); or (b) to report whether the stimuli were located on the right or left side of the head mid-line (SD: spatial discrimination); or (c) simply to detect the presence of an auditory stimulus (control task). Our results showed that both types of auditory discrimination (TD and SD) compared to simple detection activated a network of brain areas including regions of prefrontal cortex and basal ganglia. Critically, two clusters in pre-SMA and the anterior cingulate cortex were specifically activated by TD. Furthermore, these clusters overlap with regions activated for similar judgments in the tactile modality suggesting that they fulfill a multimodal function in the temporal processing of sensory events.
NeuroImage, 2012
Deduction is the ability to draw necessary conclusions from previous knowledge. Here we propose a... more Deduction is the ability to draw necessary conclusions from previous knowledge. Here we propose a novel approach to understanding the neural basis of deduction, which exploits fine-grained inter-participant variability in such tasks. Participants solved deductive problems and were grouped by the behavioral strategies employed, i.e., whether they were sensitive to the logical form of syllogistic premises, whether the problems were solved correctly, and whether heuristic strategies were employed. Differential profiles of neural activity can predict membership of the first two of these groups. The predictive power of activity profiles is distributed non-uniformly across the brain areas activated by deduction. Activation in left ventro-lateral frontal (BA47) and lateral occipital (BA19) cortices predicts whether logically valid solutions are sought. Activation of left inferior lateral frontal (BA44/45) and superior medial frontal (BA6/8) cortices predicts sensitivity to the logical structure of problems. No specific pattern of activation was associated with the use of a nonlogical heuristic strategy. Not only do these findings corroborate the hypothesis that left BA47, BA44/45 and BA6/8 are critical for making syllogistic deductions, but they also imply that they have different functional roles as components of a dedicated network. We propose that BA44/45 and BA6/8 are involved in the extraction and representation of the formal structure of a problem, while BA47 is involved in the selection and application of relevant inferential rules. Finally, our findings suggest that deductive reasoning can be best described as a cascade of cognitive processes requiring the concerted operation of several, functionally distinct, brain areas.
NeuroImage, 2010
Episodic retrieval is supported by multiple forms of cognitive control that depend on prefrontal ... more Episodic retrieval is supported by multiple forms of cognitive control that depend on prefrontal cortex. However, within prefrontal cortex, the regional specificity for different control processes is still largely underspecified. Here we used fMRI to investigate the processes involved in part-list cueing, a phenomenon entailing the reduction of memory performance when some of the "to-be-remembered items" are presented during retrieval. Retrieval inhibition and strategy disruption have been proposed as possible underlying causes of this surprising effect. Critically, different encoding conditions can dissociate these two hypotheses, as this is thought to have a different impact on strategy-related versus inhibition-related processes. Accordingly, we compared part-list versus no part-list retrieval conditions, following high versus low associative encoding conditions. This revealed activation of the left frontopolar and right dorsolateral prefrontal cortex in conditions of part-list cueing retrieval, but only following low associative encoding. These results are consistent with the inhibitory or interference resolution accounts of part-list cueing, suggesting a critical role of between-items competition and high monitoring demands at retrieval. In addition, we also examined brain activation during the encoding phase, specifying further possible anatomo-functional links between encoding and retrieval processes. The results suggest that different encoding conditions induce distinct patterns of activation at retrieval, corroborating the idea that they promote the adoption of different retrieval strategies. The implications of the results are discussed in relation to the cognitive mechanisms of inhibition, interference resolution, and encoding/retrieval strategies for episodic memory.
Journal of Cognitive Neuroscience, 2010
Selection between competing responses and stimulus-response association strength is thought to af... more Selection between competing responses and stimulus-response association strength is thought to affect performance during verb generation. However, the specific contribution of these two processes remains unclear. Here we used fMRI to investigate the role of selection and association within frontal and BG circuits that are known to be involved in verb production. Subjects were asked to generate verbs from nouns in conditions requiring either high or low selection, but with constant association strength, and in conditions of weak or strong association strength, now with constant selection demands. Furthermore, we examined the role of selection and association during noun generation from noun stimuli. We found that the midpart of the left inferior frontal gyrus was more active in conditions requiring high compared with low selection, with matched association strength. The same left inferior frontal region activated irrespective of verb or noun generation. Results of ROI analyses showed...
Journal of Cognitive Neuroscience, 2014
We investigated the interplay between stimulus-driven attention and memory retrieval with a novel... more We investigated the interplay between stimulus-driven attention and memory retrieval with a novel interference paradigm that engaged both systems concurrently on each trial. Participants encoded a 45-min movie on Day 1 and, on Day 2, performed a temporal order judgment task during fMRI. Each retrieval trial comprised three images presented sequentially, and the task required participants to judge the temporal order of the first and the last images (“memory probes”) while ignoring the second image, which was task irrelevant (“attention distractor”). We manipulated the content relatedness and the temporal proximity between the distractor and the memory probes, as well as the temporal distance between two probes. Behaviorally, short temporal distances between the probes led to reduced retrieval performance. Distractors that at encoding were temporally close to the first probe image reduced these costs, specifically when the distractor was content unrelated to the memory probes. The ima...
Human Brain Mapping, 2013
Previous studies on crossmodal spatial orienting typically used simple and stereotyped stimuli in... more Previous studies on crossmodal spatial orienting typically used simple and stereotyped stimuli in the absence of any meaningful context. This study combined computational models, behavioural measures and functional magnetic resonance imaging to investigate audiovisual spatial interactions in naturalistic settings. We created short videos portraying everyday life situations that included a lateralised visual event and a co-occurring sound, either on the same or on the opposite side of space. Subjects viewed the videos with or without eye-movements allowed (overt or covert orienting). For each video, visual and auditory saliency maps were used to index the strength of stimulus-driven signals, and eye-movements were used as a measure of the efficacy of the audiovisual events for spatial orienting. Results showed that visual salience modulated activity in higher-order visual areas, whereas auditory salience modulated activity in the superior temporal cortex. Auditory salience modulated activity also in the posterior parietal cortex, but only when audiovisual stimuli occurred on the same side of space (multisensory spatial congruence). Orienting efficacy affected activity in the visual cortex, within the same regions modulated by visual salience. These patterns of activation were comparable in overt and covert orienting conditions. Our results demonstrate that, during viewing of complex multisensory stimuli, activity in sensory areas reflects both stimulus-driven signals and their efficacy for spatial orienting; and that the posterior parietal cortex combines spatial information about the visual and the auditory modality.
Human Brain Mapping, 2010
The feeling of guilt is a complex mental state underlying several human behaviors in both private... more The feeling of guilt is a complex mental state underlying several human behaviors in both private and social life. From a psychological and evolutionary viewpoint, guilt is an emotional and cognitive function, characterized by prosocial sentiments, entailing specific moral believes, which can be predominantly driven by inner values (deontological guilt) or by more interpersonal situations (altruistic guilt). The aim of this study was to investigate whether there is a distinct neurobiological substrate for these two expressions of guilt in healthy individuals. We first run two behavioral studies, recruiting a sample of 72 healthy volunteers, to validate a set of stimuli selectively evoking deontological and altruistic guilt, or basic control emotions (i.e., anger and sadness). Similar stimuli were reproduced in a event-related functional magnetic resonance imaging (fMRI) paradigm, to investigate the neural correlates of the same emotions, in a new sample of 22 healthy volunteers. We show that guilty emotions, compared to anger and sadness, activate specific brain areas (i.e., cingulate gyrus and medial frontal cortex) and that different neuronal networks are involved in each specific kind of guilt, with the insula selectively responding to deontological guilt stimuli. This study provides evidence for the existence of distinct neural circuits involved in different guilty feelings. This complex emotion might account for normal individual attitudes and deviant social behaviors. Moreover, an abnormal processing of specific guilt feelings might account for some psychopathological manifestation, such as obsessive-compulsive disorder and depression.
Human Brain Mapping, 2011
Previous studies have indicated that increasing working memory (WM) load can affect the attention... more Previous studies have indicated that increasing working memory (WM) load can affect the attentional selection of signals originating from one object/location. Here we assessed whether WM load affects also the selection of multiple objects/locations (divided attention). Participants monitored either two object-categories (vs. one category; object-based divided attention) or two locations (vs. one location; space-based divided attention) while maintaining in WM either a variable number of objects (object-based WM load) or locations (space-based WM load). Behavioural results showed that WM load affected attentional performance irrespective of divided or focused attention. However, fMRI results showed that the activity associated with object-based divided attention increased linearly with increasing object-based WM load in the left and right intraparietal sulcus (IPS); while, in the same areas, activity associated with space-based divided attention was not affected by any type of WM load. These findings support the hypothesis that WM contributes to the maintenance of resource-demanding attentional sets in a domain-specific manner. Moreover, the dissociable impact of WM load on performance and brain activity suggests that increased IPS activation reflects a recruitment of additional, domain-specific processing resources that enable dual-task performance under conditions of high WM load and high attentional demand.
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Papers by Emiliano Macaluso