JOURNAL OF COGNITIVE NEUROSCIENCE

Neural and Computational Mechanisms of Motivation and Decision-making
Yee DM
Motivation is often thought to enhance adaptive decision-making by biasing actions toward rewards and away from punishment. Emerging evidence, however, points to a more nuanced view whereby motivation can both enhance and impair different aspects of decision-making. Model-based approaches have gained prominence over the past decade for developing more precise mechanistic explanations for how incentives impact goal-directed behavior. In this Special Focus, we highlight three studies that demonstrate how computational frameworks help decompose decision processes into constituent cognitive components, as well as formalize when and how motivational factors (e.g., monetary rewards) influence specific cognitive processes, decision-making strategies, and self-report measures. Finally, I conclude with a provocative suggestion based on recent advances in the field: that organisms do not merely seek to maximize the expected value of extrinsic incentives. Instead, they may be optimizing decision-making to achieve a desired internal state (e.g., homeostasis, effort, affect). Future investigation into such internal processes will be a fruitful endeavor for unlocking the cognitive, computational, and neural mechanisms of motivated decision-making.
Neural Basis of Perceptual Surface Qualities and Materials: Evidence from EEG Decoding
Orima T, Wakita S and Motoyoshi I
The human visual system can easily recognize object material categories and estimate surface properties such as glossiness and smoothness. Recent psychophysical and computational studies suggest that the material perception depends on global feature statistics. To elucidate dynamic neural representations underlying surface property and material perception in humans, we measured visual evoked potentials (VEPs) for 191 natural images consisting of 20 categories of materials and then classified material categories and surface properties from the VEPs. As a result, we found that material categories were correctly classified by the VEPs even at latencies of 150 msec or less. The apparent surface properties were also significantly classified within 175 msec (lightness, colorfulness, and smoothness) and after 200 msec (glossiness, hardness, and heaviness). The subsequent reverse-correlation analysis revealed that the VEPs at these latencies are highly correlated with low- and high-level global feature statistics of the surface images, indicating that neural activities about such global features are reflected in the VEPs. Moreover, by using deep generative models (multimodal variational autoencoder models) to reconstruct surface images from the VEPs via style information, we demonstrated that the reconstructed surface images are judged by observers to have very similar material categories and surface properties as the original natural surfaces. These results support the notion that neural representations of statistical features in the early cortical response play a crucial role in the perception and recognition of surface materials in humans.
Transcranial Direct-current Stimulation of the Dorsolateral Prefrontal Cortex Modulates Voluntary Task-order Coordination in Dual-task Situations
Kübler S, Langsdorf L, Meyer M and Schubert T
Dual tasks (DTs) require additional control processes that temporally coordinate the processing of the two component tasks. Previous studies employing imaging as well as noninvasive stimulation techniques have demonstrated that the dorsolateral prefrontal cortex (dlPFC) is causally involved in these task-order coordination processes. However, in these studies, participants were instructed to match their processing order to an externally provided and mandatory order criterion during DT processing. Hence, it is still unknown whether the dlPFC is also recruited for rather voluntary order control processes, which are required in situations that allow for intentional and internally generated order choices. To address this issue, in two experiments, we applied anodal (Experiment 1) and cathodal (Experiment 2) transcranial direct-current stimulation during a random-order DT in which participants could freely decide about their order of task processing. In our results, we found facilitatory and inhibitory effects on voluntary task-order coordination because of anodal and cathodal transcranial direct-current stimulation, respectively. This was indicated by shorter RTs when participants intentionally switched the task order relative to the preceding trial during anodal as well as a reduced tendency to switch the task order relative to the preceding trial during cathodal stimulation compared with the sham stimulation. Overall, these findings indicate that the dlPFC is also causally involved in voluntary task-order coordination processes. In particular, we argue that the dlPFC is recruited for intentionally updating and implementing task-order information that is necessary for scheduling the processing of two temporally overlapping tasks.
Movement Strategy Moderates the Effect of Spatially Congruent Cues on the Stability of Rhythmic Bimanual Finger Movements
Denyer R and Boyd LA
Spatially congruent cues increase the speed of bimanual reach decisions compared with abstract symbolic cues, particularly for asymmetric reaches. Asymmetric rhythmic bimanual movements are less stable than symmetric rhythmic movements, but it is not well understood if spatially congruent cues similarly increase the stability of asymmetric rhythmic bimanual movements. To address this question, in Experiment 1, participants performed symmetric and asymmetric bimanual rhythmic finger tapping movements at different movement frequencies in time with flickering spatially congruent and abstract symbolic stimuli. As expected, symmetric movements were more stable. Spatially congruent cues similarly increased the stability of symmetric and asymmetric movements compared with abstract symbolic cues. The benefits of spatial congruence and movement symmetry were restricted to high movement frequencies (>2 Hz). To better understand if the emergence of these effects at high movement frequencies was driven by a change in movement strategy, in Experiment 2, video of the hands was concurrently recorded during task performance. Markerless motion tracking software revealed that participants switched from discontinuous to continuous movement strategies with increasing movement frequency. Because discontinuous and continuous movements are thought to be controlled by distinct neurocognitive systems, this might explain why the beneficial effects of spatial congruence and response symmetry emerged only at high movement frequencies. Overall, results from the current study indicate that the perceptual quality of the stimulus use to cue movement frequency can have powerful effects on the stability of rhythmic bimanual movements, but that these effects may depend on whether discontinuous or continuous movement strategies are selected.
Optogenetic Manipulation of Covert Attention in the Nonhuman Primate
Katz LN, Bohlen MO, Yu G, Mejias-Aponte C, Sommer MA and Krauzlis RJ
Optogenetics affords new opportunities to interrogate neuronal circuits that control behavior. In primates, the usefulness of optogenetics in studying cognitive functions remains a challenge. The technique has been successfully wielded, but behavioral effects have been demonstrated primarily for sensorimotor processes. Here, we tested whether brief optogenetic suppression of primate superior colliculus can change performance in a covert attention task, in addition to previously reported optogenetic effects on saccadic eye movements. We used an attention task that required the monkey to detect and report a stimulus change at a cued location via joystick release, while ignoring changes at an uncued location. When the cued location was positioned in the response fields of transduced neurons in the superior colliculus, transient light delivery coincident with the stimulus change disrupted the monkey's detection performance, significantly lowering hit rates. When the cued location was elsewhere, hit rates were unaltered, indicating that the effect was spatially specific and not a motor deficit. Hit rates for trials with only one stimulus were also unaltered, indicating that the effect depended on selection among distractors rather than a low-level visual impairment. Psychophysical analysis revealed that optogenetic suppression increased perceptual threshold, but only for locations matching the transduced site. These data show that optogenetic manipulations can cause brief and spatially specific deficits in covert attention, independent of sensorimotor functions. This dissociation of effect, and the temporal precision provided by the technique, demonstrates the utility of optogenetics in interrogating neuronal circuits that mediate cognitive functions in the primate.
A Functional Magnetic Resonance Imaging Study of Phrase Structure and Subject Island Violations
Matchin W, Almeida D, Hickok G and Sprouse J
In principle, functional neuroimaging provides uniquely informative data in addressing linguistic questions, because it can indicate distinct processes that are not apparent from behavioral data alone. This could involve adjudicating the source of unacceptability via the different patterns of elicited brain responses to different ungrammatical sentence types. However, it is difficult to interpret brain activations to syntactic violations. Such responses could reflect processes that have nothing intrinsically related to linguistic representations, such as domain-general executive function abilities. To facilitate the potential use of functional neuroimaging methods to identify the source of different syntactic violations, we conducted a functional magnetic resonance imaging experiment to identify the brain activation maps associated with two distinct syntactic violation types: phrase structure (created by inverting the order of two adjacent words within a sentence) and subject islands (created by extracting a wh-phrase out of an embedded subject). The comparison of these violations to control sentences surprisingly showed no indication of a generalized violation response, with almost completely divergent activation patterns. Phrase structure violations seemingly activated regions previously implicated in verbal working memory and structural complexity in sentence processing, whereas the subject islands appeared to activate regions previously implicated in conceptual-semantic processing, broadly defined. We review our findings in the context of previous research on syntactic and semantic violations using ERPs. Although our results suggest potentially distinct underlying mechanisms underlying phrase structure and subject island violations, our results are tentative and suggest important methodological considerations for future research in this area.
The Impact of Selective Attention and Musical Training on the Cortical Speech Tracking in the Delta and Theta Frequency Bands
Schüller A, Mücke A, Riegel J and Reichenbach T
Oral communication regularly takes place amidst background noise, requiring the ability to selectively attend to a target speech stream. Musical training has been shown to be beneficial for this task. Regarding the underlying neural mechanisms, recent studies showed that the speech envelope is tracked by neural activity in auditory cortex, which plays a role in the neural processing of speech, including speech in noise. The neural tracking occurs predominantly in two frequency bands, the delta and the theta bands. However, much regarding the specifics of these neural responses, as well as their modulation through musical training, still remain unclear. Here, we investigated the delta- and theta-band cortical tracking of the speech envelope of attended and ignored speech using magnetoencephalography (MEG) recordings. We thereby assessed both musicians and nonmusicians to explore potential differences between these groups. The cortical speech tracking was quantified through source-reconstructing the MEG data and subsequently relating the speech envelope in a certain frequency band to the MEG data using linear models. We thereby found the theta-band tracking to be dominated by early responses with comparable magnitudes for attended and ignored speech, whereas the delta band tracking exhibited both earlier and later responses that were modulated by selective attention. Almost no significant differences emerged in the neural responses between musicians and nonmusicians. Our findings show that only the speech tracking in the delta but not in the theta band contributes to selective attention, but that this mechanism is essentially unaffected by musical training.
Experimental Manipulation of the Bilateral Posterior Parietal Cortex Strengthens Associative Memory in Healthy Participants: A Continuous Theta-burst Stimulation
Cheng L, Li X, Hao Z, Li J, Zhao M, Zhan L, Li M, Gu H and Jia X
To test whether targeting left and right posterior parietal cortex (PPC) with continuous theta-burst stimulation (cTBS) in healthy adults would strengthen associative memory (AM) performance. This study consisted of two experiments (a behavioral experiment and a formal experiment during each of the two experimental sessions). In Experiment 1, 18 adults (one male, age = 22.83 ± 3.92 years) were included in the behavioral phase and 18 adults (seven males, age = 40.11 ± 12.27 years) in the stimulation phase. There were 120 neutral facial images paired with 120 two-character nouns and then divided into six test versions (10 male faces and 10 female faces paired with 20 different nouns were considered as one version). In the behavioral experiment, participants were tested by six-version tests to assess memory materials, and in the formal experiment, participants' face-word AM performance was measured by certified tests based on a cued recall paradigm. Furthermore, 30 adults (seven males, age = 20.97 ± 1.85 years) and 15 adults (five males, age = 22.27 ± 1.29 years) participated in Experiment 2, respectively. Stimuli and procedure were the same as in Experiment 1, but the AM test was based on a forced-choice paradigm. Experiment 1 did not yield anticipated outcomes; Experiment 2 showed that cTBS of left and right PPC strengthened the AM performance compared with the control condition. In conclusion, cTBS to left and right PPC improved AM in healthy adults, which provided further experimental evidence for strengthening AM by cTBS.
Neural Correlates of Retrieval Success and Precision: A Functional Magnetic Resonance Imaging Study
Hou M, Hill PF, Aktas ANZ, Ekstrom AD and Rugg MD
Prior studies examining the neural mechanisms underlying retrieval success and precision have yielded inconsistent results. Here, the neural correlates of success and precision were examined with a memory task that assessed precision for spatial location. A sample of healthy young adults underwent functional magnetic resonance imaging scanning during a single study-test cycle. At study, participants viewed a series of object images, each placed at a randomly selected location on an imaginary circle. At test, studied images were intermixed with new images and presented to the participants. The requirement was to move a cursor to the location of the studied image, guessing if necessary. Participants then signaled whether the presented image had been studied. Memory precision was quantified as the angular difference between the studied location and the location selected by the participant. A precision effect was evident in the left angular gyrus, where BOLD activity covaried with location accuracy. In addition, multivoxel pattern analysis revealed a significant item-level reinstatement effect in the angular gyrus for high-precision trials. There was no evidence of a retrieval success effect in this region. BOLD activity in the hippocampus was insensitive to both success and precision. These findings are partially consistent with prior evidence that success and precision are dissociable features of memory retrieval.
How Can Graph Theory Inform the Dual-stream Model of Speech Processing? A Resting-state Functional Magnetic Resonance Imaging Study of Stroke and Aphasia Symptomology
Zhu H, Fitzhugh MC, Keator LM, Johnson L, Rorden C, Bonilha L, Fridriksson J and Rogalsky C
The dual-stream model of speech processing describes a cortical network involved in speech processing. However, it is not yet known if the dual-stream model represents actual intrinsic functional brain networks. Furthermore, it is unclear how disruptions after a stroke to the functional connectivity of the dual-stream model's regions are related to speech production and comprehension impairments seen in aphasia. To address these questions, in the present study, we examined two independent resting-state fMRI data sets: (1) 28 neurotypical matched controls and (2) 28 chronic left-hemisphere stroke survivors collected at another site. We successfully identified an intrinsic functional network among the dual-stream model's regions in the control group using functional connectivity. We then used both standard functional connectivity analyses and graph theory approaches to determine how this connectivity may predict performance on clinical aphasia assessments. Our findings provide evidence that the dual-stream model of speech processing is an intrinsic network as measured via resting-state MRI and that functional connectivity of the hub nodes of the dual-stream network defined by graph theory methods, but not overall average network connectivity, is weaker in the stroke group than in the control participants. In addition, the functional connectivity of the hub nodes predicted linguistic impairments on clinical assessments. In particular, the relative strength of connectivity of the right hemisphere's homologues of the left dorsal stream hubs to the left dorsal hubs, versus to the right ventral stream hubs, is a particularly strong predictor of poststroke aphasia severity and symptomology.
Neurophysiological and Autonomic Dynamics of Threat Processing during Sustained Social Fear Generalization
Pouliot JJ, Ward RT, Traiser CM, Chiasson P, Gilbert FE and Keil A
Survival in dynamic environments requires that organisms learn to predict danger from situational cues. One key facet of threat prediction is generalization from a predictive cue to similar cues, ensuring that a cue-outcome contingency is applied beyond the original learning environment. Generalization has been observed in laboratory studies of aversive conditioning: Behavioral and physiological processes generalize responses from a stimulus paired with threat (the conditioned stimulus [CS+]) to unpaired stimuli, with response magnitudes varying with CS+ similarity. In contrast, work focusing on sensory responses in visual cortex has found a sharpening pattern, in which responses to stimuli closely resembling the CS+ are maximally suppressed, potentially reflecting lateral inhibitory interactions with the CS+ representation. Originally demonstrated with simple visual cues, changes in visuocortical tuning have also been observed in threat generalization learning across facial identities. It is unclear to what extent these visuocortical changes represent transient or sustained effects and if generalization learning requires prior conditioning to the CS+. The present study addressed these questions using EEG and pupillometry in an aversive generalization paradigm involving hundreds of trials using a gradient of facial identities. Visuocortical steady-state visual evoked potential sharpening occurred after dozens of trials of generalization learning without prior differential conditioning, but diminished as learning continued. By contrast, generalization of alpha power suppression, pupil dilation, and self-reported valence and arousal was seen throughout the experiment. Findings are consistent with threat processing models emphasizing the role of changing visucocortical and attentional dynamics when forming, curating, and shaping fear memories as observers continue learning about stimulus-outcome contingencies.
How Linguistic and Nonlinguistic Vocalizations Shape the Perception of Emotional Faces-An EEG Study
Liang J, Zhang M, Yang L, Li Y, Li Y, Wang L, Li H, Chen J and Luo W
Vocal emotions are crucial in guiding visual attention toward emotionally significant environmental events, such as recognizing emotional faces. This study employed continuous electroencephalography (EEG) recordings to examine the impact of linguistic and nonlinguistic vocalizations on facial emotion processing. Participants completed a facial emotion discrimination task while viewing fearful, happy, and neutral faces. The behavioral and ERP results indicated that fearful nonlinguistic vocalizations accelerated the recognition of fearful faces and elicited a larger P1 amplitude, whereas happy linguistic vocalizations accelerated the recognition of happy faces and similarly induced a greater P1 amplitude. In recognition of fearful faces, a greater N170 component was observed in the right hemisphere when the emotional category of the priming vocalization was consistent with the face stimulus. In contrast, this effect occurred in the left hemisphere while recognizing happy faces. Representational similarity analysis revealed that the temporoparietal regions automatically differentiate between linguistic and nonlinguistic vocalizations early in face processing. In conclusion, these findings enhance our understanding of the interplay between vocalization types and facial emotion recognition, highlighting the importance of cross-modal processing in emotional perception.
Neural Correlates of Different Randomization Tasks
Guseva M, Bogler C, Allefeld C, Ziya EB and Haynes JD
In some cases, when we are making decisions, the available choices can appear to be equivalent. When this happens, our choices appear not to be constrained by external factors and, instead, we can believe to be selecting "randomly." Furthermore, randomness is sometimes even explicitly required by task conditions such as in random sequence generation tasks. This is a challenging task that involves the coordination of multiple cognitive processes, which can include the inhibition of habitual choice patterns and monitoring of the running choice sequence. It has been shown that random choices are strongly influenced by the way they are instructed. This raises the question whether the brain mechanisms underlying random selection also differ between different task instructions. To assess this, we measured brain activity while participants were engaging in three different variations of a sequence generation task: On the basis of previous work, participants were instructed to either (1) "generate a random sequence of choices," (2) "simulate a fair coin toss," or (3) "choose freely." Our results reveal a consistent frontoparietal activation pattern that is shared across all tasks. Specifically, increased activity was observed in bilateral inferior and right middle frontal gyrus, left pre-SMA, bilateral inferior parietal lobules, and portions of anterior insular cortex in both hemispheres. Activity in the mental coin toss condition was higher in right dorsolateral prefrontal cortex, left (pre-) SMA, a portion of right inferior frontal gyrus, bilateral superior parietal lobules, and bilateral anterior insula. In addition, our multivariate analysis revealed a distinct region in the right frontal pole to be predictive of the outcome of choices, but only when randomness was explicitly instructed. These results emphasize that different randomization tasks involve both shared and unique neural mechanisms. Thus, even seemingly similar randomization behavior can be produced by different neural pathways.
Frontal Deactivation and the Efficacy of Statistical Learning: Neural Mechanisms Accompanying Exposure to Visual Statistical Sequences
Brylka M, Wojciechowski J, Wolak T and Cygan HB
Statistical learning is the cognitive ability to rapidly identify structure and meaning in unfamiliar streams of sensory experience, even in the absence of feedback. Despite extensive studies, the neurocognitive mechanisms underlying this phenomenon still require further clarification under varying cognitive conditions. Here, we examined neural mechanisms during the first exposure to visually presented sequences in 47 healthy participants. We used two types of visual objects: abstract symbols and pictures of cartoon-like animals. This allowed us to compare informational processing mechanisms with defined distinguishing features. Participants achieved better performance for sequences with easy-to-name than difficult-to-name abstract stimuli. fMRI results revealed greater activation in widespread brain regions in response to random versus statistical sequences for all stimuli types. Behavioral accuracy was associated with increased deactivation of the ventromedial pFC for easy-to-name statistical versus random sequences. For difficult-to-name statistical versus random sequences, performance correlated with dmPFC deactivation. ROI analysis showed a generally positive involvement of the caudate head in sequence processing with significantly stronger activity during the first run of performing the task. Functional connectivity analysis of prefrontal deactivation regions revealed significant connectivity with nodes of the salience network for both object types and inverse connectivity with the caudate head only for easy-to-name objects. The results indicated that distinct subregions of pFC modulate task performance depending on the visual stimulus characteristic. They also showed that among striatal regions, only the head of the caudate was sensitive to initial exposure to visual statistical information.
Tool Representations in Human Visual Cortex
Cortinovis D, Peelen MV and Bracci S
Tools such as pens, forks, and scissors play an important role in many daily-life activities, an importance underscored by the presence in visual cortex of a set of tool-selective brain regions. This review synthesizes decades of neuroimaging research that investigated the representational spaces in the visual ventral stream for objects, such as tools, that are specifically characterized by action-related properties. Overall, results reveal a dissociation between representational spaces in ventral and lateral occipito-temporal cortex (OTC). While lateral OTC encodes both visual (shape) and action-related properties of objects, distinguishing between objects acting as end-effectors (e.g., tools, hands) versus similar noneffector manipulable objects (e.g., a glass), ventral OTC primarily represents objects' visual features such as their surface properties (e.g., material and texture). These areas act in concert with regions outside of OTC to support object interaction and tool use. The parallel investigation of the dimensions underlying object representations in artificial neural networks reveals both the possibilities and the difficulties in capturing the action-related dimensions that distinguish tools from other objects. Although artificial neural networks offer promise as models of visual cortex computations, challenges persist in replicating the action-related dimensions that go beyond mere visual features. Taken together, we propose that regions in OTC support the representation of tools based on a behaviorally relevant action code and suggest future paths to generate a computational model of this object space.
Attentional Refreshing in Working Memory and Its Interplay with Long-term Memory: A Behavioral and EEG Study
Labaronne M, Caclin A and Plancher G
Despite the growing interest in the study of attentional refreshing, the functioning of this working memory maintenance mechanism, including its cerebral underpinnings, is still debated. In particular, it remains unclear whether refreshing promotes long-term memory and whether it, in return, depends on long-term memory content to operate. Here, we used direct maintenance instructions and measured brain activity to investigate working memory maintenance with two objectives: (1) test if different behavioral and oscillatory patterns could be observed when participants were instructed to use attentional refreshing versus verbal rehearsal, and (2) observe whether and how refreshing is modulated when maintaining novel (pseudowords) versus familiar (words) memoranda. We conducted an EEG experiment using a modified Brown-Peterson task, in which we manipulated the type of maintenance engaged through explicit instructions (verbal rehearsal vs. refreshing), the type of memoranda (words vs. pseudowords), and the memory load (2 vs. 6). Using scalp EEG, we measured both neural oscillations during working memory maintenance and ERPs during the concurrent parity judgment task. For words, we showed that verbal rehearsal benefited more short-term recall whereas refreshing benefited more delayed recall. In keeping with these behavioral differences between maintenance instructions, frontal-midline theta power increased with memory load only when using verbal rehearsal, whereas occipito-parietal alpha desynchronization was larger with refreshing than verbal rehearsal. When maintaining pseudowords, verbal rehearsal also benefitted short-term recall more than refreshing. However, no long-term memory benefit of refreshing was observed for pseudowords, and oscillatory activity was not different under the two maintenance instructions. Our results provide new evidence supporting the independence between attentional refreshing and verbal rehearsal, and bring new insight into refreshing functioning. We discuss the possible interpretations of these results and the implications for the attentional refreshing literature.
Theta Transcranial Alternating Current Stimulation Is Not Effective in Improving Working Memory Performance
Kasanov D, Dorogina O, Mushtaq F and Pavlov YG
There is an extensive body of research showing a significant relationship between frontal midline theta activity in the 4- to 8-Hz range and working memory (WM) performance. Transcranial alternating current stimulation (tACS) is recognized for inducing lasting changes in brain oscillatory activity. Across two experiments, we tested whether WM could be improved through tACS of dorsomedial pFC and ACC, by affecting executive control networks associated with frontal midline theta. In Experiment 1, after either a 20-min verum or sham stimulation applied to Fpz-CPz at 1 mA and 6 Hz, 31 participants performed WM tasks, while EEG was recorded. The tasks required participants to either mentally manipulate memory items or retain them in memory as they were originally presented. No significant effects were observed in behavioral performance, and we found no change in theta activity during rest and task after stimulation. However, alpha activity during retention or manipulation of information in WM was less strongly enhanced during the delay period after verum stimulation as compared with sham. In Experiment 2 (n = 25), tACS was administered during the task in two separate sessions. Here, we changed the order of the stimulation blocks: A 25-min task block was either accompanied first by sham stimulation and then by verum stimulation, or vice versa. Again, we found no improvements in WM through either tACS after-effects or online stimulation. Taken together, our results demonstrate that theta frequency tACS applied at the midline is not an effective method for enhancing WM.
Shared Patterns of Cognitive Control Behavior and Electrophysiological Markers in Adolescence
Wiker T, Alnæs D, Pedersen ML, Norbom LB, Boer OD, Kjelkenes R, Voldsbekk I, Karl V, Bukhari SH, Moberget T, Westlye LT, Huster RJ and Tamnes CK
Behavioral parameters obtained from cognitive control tasks have been linked to electrophysiological markers. Yet, most previous research has investigated only a few specific behavioral parameters at a time. An integrated approach with simultaneous consideration of multiple aspects of behavior may better elucidate the development and function of cognitive control. Here, we aimed to identify shared patterns between cognitive control behavior and electrophysiological markers using stop-signal task data and EEG recordings from an adolescent sample (n = 193, aged 11-25 years). We extracted behavioral variables covering various aspects of RT, accuracy, inhibition, and decision-making processes, as well as amplitude and latency of the ERPs N1, N2, and P3. To identify shared patterns between the two sets of variables, we employed a principal component analysis and a canonical correlation analysis. First, we replicated previously reported associations between various cognitive control behavioral parameters. Next, results from the canonical correlation analysis showed that overall good task performance was associated with fast and strong neural processing. Furthermore, the canonical correlation was affected by age, indicating that the association varies depending on age. The present study suggests that although distributional and computational methods can be applied to extract specific behavioral parameters, they might not capture specific patterns of cognitive control or electrophysiological brain activity in adolescents.
Similarity Distractors Increase the Burden of Chinese Character Selection and Updating in Working Memory
Li H and Zhao X
Attentional mechanisms are the primary processes for performing working memory (WM) tasks and can prevent distractors from interfering with the content representations stored in WM. However, our understanding of the mechanisms by which attention affects WM remains limited. As such, we analyzed ERPs of the character n-back task to investigate Chinese character selection, updating, and maintenance in WM. In Experiment 1, we collected electroencephalography data from 27 participants aged 18-25 years to explore the influence of false-character interference and symbol interference on a neural activity in the character n-back task. The results suggest that RT was longer in the false-character interference condition. The N2pc and P300 amplitudes were smaller; however, the slow wave amplitude did not differ significantly. In Experiment 2, we used a single-symbol interference and a multiple-symbol interference to establish whether the number of interferences affected the neural activity in the character n-back task. Thirty participants (aged 19-25 years) took part in the experiment. The findings imply that a longer RT and a larger N2pc amplitude occurred in the multiple-symbol interference condition, but not in the P300 and slow wave conditions. Our findings indicate that distractors that are similar to characters may produce greater interference in character recognition and affect the subsequent updating, whereas the number of distractors may only interfere with early character selection, but not with updating and maintenance phases.
The Role of Sleep in Memory Consolidation and Reading in Dyslexia
Solbi A and Earle FS
Dyslexia is a neurodevelopmental disorder characterized by reading difficulty, which has long been attributed to a phonological processing deficit. However, recent research suggests that general difficulties with learning and memory, but also in memory consolidation, may underlie disordered reading. This review article provides an overview of the relationship between learning and memory, memory consolidation during sleep, and reading and explores the emerging literature on consolidation during sleep in individuals with dyslexia. We consider evidence that sleep appears to be less effective for memory consolidation in children with dyslexia and how this may be related to their deficits in reading. This discussion highlights the need for further research to determine the extent to which atypical sleep patterns may contribute to learning deficits associated with disordered reading.
Semantic Context Effects in Picture and Sound Naming: Evidence from Event-related Potentials and Pupillometric Data
Gruner M, Widmann A, Wöhner S, Schröger E and Jescheniak JD
When a picture is repeatedly named in the context of semantically related pictures, (homogeneous context) responses are slower than when the picture is repeatedly named in the context of unrelated pictures (heterogeneous context). This semantic interference effect in blocked-cyclic naming plays an important role in devising theories of word production. Wöhner, Mädebach, and Jescheniak [2021; Wöhner, S., Mädebach, A., & Jescheniak, J. D. Naming pictures and sounds: Stimulus type affects semantic context effects. Journal of Experimental Psychology: Human Perception and Performance, 47, 716-730, 2021] have shown that the effect is substantially larger when participants name environmental sounds than when they name pictures. We investigated possible reasons for this difference, using EEG and pupillometry. The behavioral data replicated Wöhner and colleagues. ERPs were more positive in the homogeneous compared with the heterogeneous context over central electrode locations between 140-180 msec and 250-350 msec for picture naming and between 250 and 350 msec for sound naming, presumably reflecting semantic interference during semantic and lexical processing. The later component was of similar size for pictures and sounds. ERPs were more negative in the homogeneous compared with the heterogeneous context over frontal electrode locations between 400 and 600 msec only for sounds. The pupillometric data showed a stronger pupil dilation in the homogeneous compared with the heterogeneous context only for sounds. The amplitudes of the late ERP negativity and pupil dilation predicted naming latencies for sounds in the homogeneous context. The latency of the effects indicates that the difference in semantic interference between picture and sound naming arises at later, presumably postlexical processing stages closer to articulation. We suggest that the processing of the auditory stimuli interferes with phonological response preparation and self-monitoring, leading to enhanced semantic interference.
Chunk Duration Limits the Learning of Multiword Chunks: Behavioral and Electroencephalogram Evidence from Statistical Learning
Henke L and Meyer L
Language comprehension involves the grouping of words into larger multiword chunks. This is required to recode information into sparser representations to mitigate memory limitations and counteract forgetting. It has been suggested that electrophysiological processing time windows constrain the formation of these units. Specifically, the period of rhythmic neural activity (i.e., slow-frequency neural oscillations) may set an upper limit of 2-3 sec. Here, we assess whether learning of new multiword chunks is also affected by this neural limit. We applied an auditory statistical learning paradigm of an artificial language while manipulating the duration of to-be-learnt chunks. Participants listened to isochronous sequences of disyllabic pseudowords from which they could learn hidden three-word chunks based on transitional probabilities. We presented chunks of 1.95, 2.55, and 3.15 sec that were created by varying the pause interval between pseudowords. In a first behavioral experiment, we tested learning using an implicit target detection task. We found better learning for chunks of 2.55 sec as compared to longer durations in line with an upper limit of the proposed time constraint. In a second experiment, we recorded participants' electroencephalogram during the exposure phase to use frequency tagging as a neural index of statistical learning. Extending the behavioral findings, results show a significant decline in neural tracking for chunks exceeding 3 sec as compared to both shorter durations. Overall, we suggest that language learning is constrained by endogenous time constraints, possibly reflecting electrophysiological processing windows.
Electrophysiological Correlates of Visual Memory Search
Williams LH, Wiegand I, Lavelle M, Wolfe JM, Fukuda K, Peelen MV and Drew T
In everyday life, we frequently engage in 'hybrid' visual and memory search, where we look for multiple items stored in memory (e.g., a mental shopping list) in our visual environment. Across three experiments, we used event-related potentials to better understand the contributions of visual working memory (VWM) and long-term memory (LTM) during the memory search component of hybrid search. Experiments 1 and 2 demonstrated that the FN400 - an index of LTM recognition - and the CDA -an index of VWM load - increased with memory set size (target load), suggesting that both VWM and LTM are involved in memory search, even when target load exceeds capacity limitations of VWM. In Experiment 3, we used these electrophysiological indices to test how categorical similarity of targets and distractors affects memory search. The CDA and FN400 were modulated by memory set size only if items resembled targets. This suggests that dissimilar distractor items can be rejected before eliciting a memory search. Together, our findings demonstrate the interplay of VWM and LTM processes during memory search for multiple targets.
Rhythm-based Temporal Expectations: Unique Contributions of Predictability and Periodicity
Solli S, Danielsen A, Leske S, Blenkmann AO, Doelling KB, Solbakk AK and Endestad T
Anticipating events and focusing attention accordingly are crucial for navigating our dynamic environment. Rhythmic patterns of sensory input offer valuable cues for temporal expectations and facilitate perceptual processing. Rhythm-based temporal expectations may rely on oscillatory entrainment, where neural activity and perceptual sensitivity synchronize with periodic stimuli. However, whether entrainment models can account for aperiodic predictable rhythms remains unclear. Our study aimed to delineate the distinct roles of predictability and periodicity in rhythm-based expectations. Participants performed a pitch-identification task preceded by periodic predictable, aperiodic predictable, or aperiodic unpredictable temporal sequences. By manipulating the temporal position of the target sound, we observed how auditory perceptual performance was modulated by the target position's relative phase relationship to the preceding sequences. Results revealed a significant performance advantage for predictable sequences, both periodic and aperiodic, compared with unpredictable ones. However, only the periodic sequence induced an entrained modulation pattern, with performance peaking in synchrony with the inherent sequence continuation. Event-related brain potentials corroborated these findings. The target-evoked P3b, possibly a neural marker of attention allocation, mirrored the behavioral performance patterns. This supports our hypothesis that temporal attention guided by rhythm-based expectations modulates perceptual performance. Furthermore, the predictive sequences were associated with enhanced target-preceding negativity (akin to the contingent negative variation), indicating enhanced target preparation. The periodic-specific modulation likely reflects more precise temporal expectations, potentially involving neural entrainment and/or more focused attention. Our findings suggest that predictability and periodicity influence perception through distinct mechanisms.
The Neural Specificity of Interference Resolution in Phonological, Semantic, and Visual Domains at Different Ages
Grégoire C, Attout L, Phillips C, Rifon L, Hody L and Majerus S
The question of whether cognitive control is specific to certain domains or domain-general remains an extensively debated question at both cognitive and neural levels. This study examined the neural substrates associated with resistance to interference (RI) in phonological, semantic, and visual domains by using strictly matched tasks and determining the domain-general or domain-specific manner in which aging affects the neural substrates associated with RI. In an fMRI experiment, young and older participants performed a similarity judgment task with phonological, semantic, or visual interference buildup. For both age groups, domain-specific RI effects were observed at the univariate level, with increased involvement in the phonological domain of the right angular gyrus and the right lingual gyrus, in the semantic domain of the bilateral inferior frontal gyrus, the bilateral superior parietal and angular gyri and the left middle temporal gyrus, and in the visual domain of the middle/superior frontal gyri and occipital gyri. At the multivariate level, although RI effects could be decoded from neural patterns in the bilateral inferior frontal gyrus for all domains and age groups, between-domain prediction of RI conditions was associated with Bayesian evidence for the null hypothesis. This study supports the domain specificity of neural substrates associated with RI while stressing its age independency.
EEG Responses to the Number of Objects in Partially Occluded and Uncovered Scenes
Baykan C and Schütz AC
Perceptual completion is ubiquitous when estimating properties such as the shape, size, or number of objects in partially occluded scenes. Behavioral experiments showed that the number of hidden objects is underestimated in partially occluded scenes compared with an estimation based on the density of visible objects and the amount of occlusion. It is still unknown at which processing level this (under)estimation of the number of hidden objects occurs. We studied this question using a passive viewing task in which observers viewed a game board that was initially partially occluded and later was uncovered to reveal its hidden parts. We simultaneously measured the electroencephalographic responses to the partially occluded board presentation and its uncovering. We hypothesized that if the underestimation is a result of early sensory processing, it would be observed in the activities of P1 and N1, whereas if it is because of higher level processes such as expectancy, it would be reflected in P3 activities. Our data showed that P1 amplitude increased with numerosity in both occluded and uncovered states, indicating a link between P1 and simple stimulus features. The N1 amplitude was highest when both the initially visible and uncovered areas of the board were completely filled with game pieces, suggesting that the N1 component is sensitive to the overall Gestalt. Finally, we observed that P3 activity was reduced when the density of game pieces in the uncovered parts matched the initially visible parts, implying a relationship between the P3 component and expectation mismatch. Overall, our results suggest that inferences about the number of hidden items are reflected in high-level processing.
The Mechanisms and Neural Signature of Time-averaged Numerosity Perception
Togoli I, Collignon O, Bueti D and Fornaciai M
The animal brain is endowed with an innate sense of number allowing to intuitively perceive the approximate quantity of items in a scene, or "numerosity." This ability is not limited to items distributed in space, but also to events unfolding in time and to the average numerosity of dynamic scenes. How the brain computes and represents the average numerosity over time, however, remains unclear. Here, we investigate the mechanisms and EEG signature of the perception of average numerosity over time. To do so, we used stimuli composed of a variable number (3-12) of briefly presented dot arrays (50 msec each) and asked participants to judge the average numerosity of the sequence. We first show that the weight of different portions of the stimuli in determining the judgment depends on how many arrays are included in the sequence itself: the longer the sequence, the lower the weight of the latest arrays. Second, we show systematic adaptation effects across stimuli in consecutive trials. Importantly, the EEG results highlight two processing stages whereby the amplitude of occipital ERPs reflects the adaptation effect (∼300 msec after stimulus onset) and the accuracy and precision of average numerosity judgments (∼450-700 msec). These two stages are consistent with processes involved with the representation of perceived average numerosity and with perceptual decision-making, respectively. Overall, our findings provide new evidence showing how the visual system computes the average numerosity of dynamic visual stimuli, and support the existence of a dedicated, relatively low-level perceptual mechanism mediating this process.
Relevance of Prosodic Focus and Lexical Stress for Discourse Comprehension in Turkish: Evidence from Psychometric and Electrophysiological Data
Zora H, Kabak B and Hagoort P
Prosody underpins various linguistic domains ranging from semantics and syntax to discourse. For instance, prosodic information in the form of lexical stress modifies meanings and, as such, syntactic contexts of words as in Turkish kaz-má "pickaxe" (noun) versus káz-ma "do not dig" (imperative). Likewise, prosody indicates the focused constituent of an utterance as the noun phrase filling the wh-spot in a dialogue like What did you eat? I ate----. In the present study, we investigated the relevance of such prosodic variations for discourse comprehension in Turkish. We aimed at answering how lexical stress and prosodic focus mismatches on critical noun phrases-resulting in grammatical anomalies involving both semantics and syntax and discourse-level anomalies, respectively-affect the perceived correctness of an answer to a question in a given context. To that end, 80 native speakers of Turkish, 40 participating in a psychometric experiment and 40 participating in an EEG experiment, were asked to judge the acceptability of prosodic mismatches that occur either separately or concurrently. Psychometric results indicated that lexical stress mismatch led to a lower correctness score than prosodic focus mismatch, and combined mismatch received the lowest score. Consistent with the psychometric data, EEG results revealed an N400 effect to combined mismatch, and this effect was followed by a P600 response to lexical stress mismatch. Conjointly, these results suggest that every source of prosodic information is immediately available and codetermines the interpretation of an utterance; however, semantically and syntactically relevant lexical stress information is assigned more significance by the language comprehension system compared with prosodic focus information.
The Suppression Mechanisms of Passive Memory in Visual Working Memory: The Evidence from Electrophysiology
Li Z, Guo W, Zhao N and Liu Q
Recent studies of visual working memory (VWM) underscore a structured hierarchy of storage states. Memories that are not immediately relevant to the task at hand but are essential for later use are transferred to a passive state, which operates independently of actively maintaining and manipulating current memories. Note that stimulating passive memory forcefully can reactivate it into an active state, resulting in a competition with active memory. Thus, to remain stable representations for both states within VWM, passive memory might involve sustained suppression during activity-silent maintenance to prevent reactivation from disrupting the current active storage. To investigate this, we analyzed lateralized electrophysiology signals while human participants (both women and men) were engaged in a sequential presentation memory task across two experiments. The results revealed positive contralateral delayed activity components and lateralized alpha enhancement for passive memory, neural indicative of suppression on passive storage. In addition, the suppression effect was independent of the memory load in both the active and the passive states. These findings support the notion of sustained suppression during activity-silent maintenance of passive memory, facilitating the stable maintenance of distinct storage states and advancing our understanding of the dynamic coding framework in VWM.
Locus Coeruleus Is Associated with Higher Openness to Experience and IQ: Implications for the Noradrenergic System for Novelty Seeking in Daily Life
Plini ERG, Robertson IH, Brosnan MB and Dockree PM
Novelty exposure and the upregulation of the noradrenergic (NA) system have been suggested as crucial for developing cognitive reserve and resilience against neurodegeneration. Openness to experience (OE), a personality trait associated with interest in novel experiences, may play a key role in facilitating this process. High-OE individuals tend to be more curious and encounter a wider range of novel stimuli throughout their lifespan. To investigate the relationship between OE and the main core of the NA system, the locus coeruleus (LC), as well as its potential mediation of IQ-a measure of cognitive reserve-MRI structural analyses were conducted on 135 healthy young adults. Compared with other neuromodulators' seeds, such as dorsal and median raphe-5-HT, ventral tegmental area-DA-, and nucleus basalis of Meynert-Ach-, the results indicated that higher LC signal intensity correlated with greater OE and IQ. Furthermore, mediation analyses revealed that only the LC played a mediating role between OE and IQ. These findings shed light on the neurobiology of personality and emphasize the importance of LC-NA system integrity in a novelty-seeking behavior. They provide a psychobiological explanation for how OE expression can contribute to the maintenance of the NA system, enhancing cognitive reserve and resilience against neurodegeneration.