Action mechanisms for social cognition: behavioral and neural correlates of developing Theory of Mind

Many psychological theories posit foundational links between two fundamental constructs: (1) our ability to produce, perceive, and represent action; and (2) our ability to understand the meaning and motivation behind the action (i.e. Theory of Mind; ToM). This position is contentious, however, and long‐standing competing theories of social‐cognitive development debate roles for basic action‐processing in ToM. Developmental research is key to investigating these hypotheses, but whether individual differences in neural and behavioral measures of motor action relate to social‐cognitive development is unknown. We examined 3‐ to 5‐year‐old children's (N = 26) EEG mu‐desynchronization during production of object‐directed action, and explored associations between mu‐desynchronization and children's behavioral motor skills, behavioral action‐representation abilities, and behavioral ToM. For children with high (but not low) mu‐desynchronization, motor skill related to action‐representation abilities, and action‐representation mediated relations between motor skill and ToM. Results demonstrate novel foundational links between action‐processing and ToM, suggesting that basic motor action may be a key mechanism for social‐cognitive development, thus shedding light on the origins and emergence of higher social cognition.     

Linking patterns of infant eye movements to a neural network model of the ventral stream using representational similarity analysis

Little is known about the development of higher-level areas of visual cortex during infancy, and even less is known about how the development of visually guided behavior is related to the different levels of the cortical processing hierarchy. As a first step toward filling these gaps, we used representational similarity analysis (RSA) to assess links between gaze patterns and a neural network model that captures key properties of the ventral visual processing stream. We recorded the eye movements of 4- to 12-month-old infants (N = 54) as they viewed photographs of scenes. For each infant, we calculated the similarity of the gaze patterns for each pair of photographs. We also analyzed the images using a convolutional neural network model in which the successive layers correspond approximately to the sequence of areas along the ventral stream. For each layer of the network, we calculated the similarity of the activation patterns for each pair of photographs, which was then compared with the infant gaze data. We found that the network layers corresponding to lower-level areas of visual cortex accounted for gaze patterns better in younger infants than in older infants, whereas the network layers corresponding to higher-level areas of visual cortex accounted for gaze patterns better in older infants than in younger infants. Thus, between 4 and 12 months, gaze becomes increasingly controlled by more abstract, higher-level representations. These results also demonstrate the feasibility of using RSA to link infant gaze behavior to neural network models. A video abstract of this article can be viewed at https://youtu.be/K5mF2Rw98Is     

Cerebral blood flow in 5‐ to 8‐month‐olds: Regional tissue maturity is associated with infant affect

Infancy is marked by rapid neural and emotional development. The relation between brain function and emotion in infancy, however, is not well understood. Methods for measuring brain function predominantly rely on the BOLD signal; however, interpretation of the BOLD signal in infancy is challenging because the neuronal‐hemodynamic relation is immature. Regional cerebral blood flow (rCBF) provides a context for the infant BOLD signal and can yield insight into the developmental maturity of brain regions that may support affective behaviors. This study aims to elucidate the relations among rCBF, age, and emotion in infancy. One hundred and seven mothers reported their infants' (infant age M ± SD = 6.14 ± 0.51 months) temperament. A subsample of infants completed MRI scans, 38 of whom produced usable perfusion MRI during natural sleep to quantify rCBF. Mother‐infant dyads completed the repeated Still‐Face Paradigm, from which infant affect reactivity and recovery to stress were quantified. We tested associations of infant age at scan, temperament factor scores, and observed affect reactivity and recovery with voxel‐wise rCBF. Infant age was positively associated with CBF in nearly all voxels, with peaks located in sensory cortices and the ventral prefrontal cortex, supporting the formulation that rCBF is an indicator of tissue maturity. Temperamental Negative Affect and recovery of positive affect following a stressor were positively associated with rCBF in several cortical and subcortical limbic regions, including the orbitofrontal cortex and inferior frontal gyrus. This finding yields insight into the nature of affective neurodevelopment during infancy. Specifically, infants with relatively increased prefrontal cortex maturity may evidence a disposition toward greater negative affect and negative reactivity in their daily lives yet show better recovery of positive affect following a social stressor.     

Action experience in infancy predicts visual-motor functional connectivity during action anticipation

Despite substantial evidence indicating a close link between action production and perception in early child development, less is known about how action experience shapes the processes of perceiving and anticipating others’ actions. Here, we developed a novel approach to capture functional connectivity specific to certain brain areas to investigate how action experience changes the networks involved in action perception and anticipation. Nine- and-12-month-old infants observed familiar (grasping) and novel (tool-use) actions while their brain activity was measured using EEG. Infants’ motor competence of both actions was assessed. A link between action experience and connectivity patterns was found, particularly during the anticipation period. During action anticipation, greater motor competence in grasping predicted greater functional connectivity between visual (occipital alpha) and motor (central alpha) regions relative to global levels of whole-brain EEG connectivity. Furthermore, visual and motor regions tended to be more coordinated in response to familiar versus novel actions and for older than younger participants. Critically, these effects were not found in the control networks (frontal-central; frontal-occipital; parietal-central; parietal-occipital), suggesting a unique role of visual-motor networks on the link between motor skills and action encoding.

Highlights

• Infants’ motor development predicted functional connectivity patterns during action anticipation.
• Faster graspers, and older infants, showed a stronger ratio of visual-motor neural coherence.
• Overall whole-brain connectivity was modulated by age and familiarity with the actions.
• Measuring inter-site relative to whole-brain connectivity can capture specific brain-behavior links.
• Measures of phase-based connectivity over time are sensitive to anticipatory action.

     

Neural correlates of action observation and execution in 14-month-old infants: an event-related EEG desynchronization study

There is increasing interest in neurobiological methods for investigating the shared representation of action perception and production in early development. We explored the extent and regional specificity of EEG desynchronization in the infant alpha frequency range (6-9 Hz) during action observation and execution in 14-month-old infants. Desynchronization during execution was restricted to central electrode sites, while action observation was associated with a broader desynchronization across frontal, central, and parietal regions. The finding of regional specificity in the overlap between EEG responses to action execution and observation suggests that the rhythm seen in the 6-9 Hz range over central sites in infancy shares certain properties with the adult mu rhythm. The magnitude of EEG desynchronization to action perception and production appears to be smaller for infants than for adults and older children, suggesting developmental change in this measure.     

Microstate analysis in infancy

BackgroundMicrostate analysis is an emerging method for investigating global brain connections using electroencephalography (EEG). Microstates have been colloquially referred to as the “atom of thought,” meaning that from these underlying networks comes coordinated neural processing and cognition. The present study examined microstates at 6-, 8-, and 10-months of age. It was hypothesized that infants would demonstrate distinct microstates comparable to those identified in adults that also parallel resting-state networks using fMRI. An additional exploratory aim was to examine the relationship between microstates and temperament, assessed via parent reports, to further demonstrate microstate analysis as a viable tool for examining the relationship between neural networks, cognitive processes as well as emotional expression embodied in temperament attributes.MethodsThe microstates analysis was performed with infant EEG data when the infant was either 6- (n = 12), 8- (n = 16), or 10-months (n = 6) old. The resting-state task involved watching a 1-minute video segment of Baby Einstein while listening to the accompanying music. Parents completed the IBQ-R to assess infant temperament.ResultsFour microstate topographies were extracted. Microstate 1 had an isolated posterior activation; Microstate 2 had a symmetric occipital to prefrontal orientation; Microstate 3 had a left occipital to right frontal orientation; and Microstate 4 had a right occipital to left frontal orientation. At 10-months old, Microstate 3, thought to reflect auditory/language processing, became activated more often, for longer periods of time, covering significantly more time across the task and was more likely to be transitioned into. This finding is interpreted as consistent with language acquisition and phonological processing that emerges around 10-months. Microstate topographies and parameters were also correlated with differing temperament broadband and narrowband scales on the IBQ-R.ConclusionThree microstates emerged that appear comparable to underlying networks identified in adult and infant microstate literature and fMRI studies. Each of the temperament domains was related to specific microstates and their parameters. These networks also correspond with auditory and visual processing as well as the default mode network found in prior research and can lead to new investigations examining differences across stimulus presentations to further explain how infants begin to recognize, respond to, and engage with the world around them.     

The lasting effects of process‐specific versus stimulus‐specific learning during infancy

The capacity to tell the difference between two faces within an infrequently experienced face group (e.g. other species, other race) declines from 6 to 9 months of age unless infants learn to match these faces with individual‐level names. Similarly, the use of individual‐level labels can also facilitate differentiation of a group of non‐face objects (strollers). This early learning leads to increased neural specialization for previously unfamiliar face or object groups. The current investigation aimed to determine whether early conceptual learning between 6 and 9 months leads to sustained behavioral advantages and neural changes in these same children at 4–6 years of age. Results suggest that relative to a control group of children with no previous training and to children with infant category‐level naming experience, children with early individual‐level training exhibited faster response times to human faces. Further, individual‐level training with a face group – but not an object group – led to more adult‐like neural responses for human faces. These results suggest that early individual‐level learning results in long‐lasting process‐specific effects, which benefit categories that continue to be perceived and recognized at the individual level (e.g. human faces).     

Fronto-parietal mirror neuron system modeling: Visuospatial transformations support imitation learning independently of imitator perspective

Although the human mirror neuron system (MNS) is critical for action observation and imitation, most MNS investigations overlook the visuospatial transformation processes that allow individuals to interpret and imitate actions observed from differing perspectives. This problem is not trivial since accurately reaching for and grasping an object requires a visuospatial transformation mechanism capable of precisely remapping fine motor skills where the observer’s and imitator’s arms and hands may have quite different orientations and sizes. Accordingly, here we describe a novel neural model to investigate the dynamics between the fronto-parietal MNS and visuospatial processes during observation and imitation of a reaching and grasping action. Our model encompasses i) the inferior frontal gyrus (IFG) and inferior parietal lobule (IPL), regions that are postulated to produce neural drive and sensory predictions, respectively; ii) the middle temporal (MT) and middle superior temporal (MST) regions that are postulated to process visual motion of a particular action; and iii) the superior parietal lobule (SPL) and intra-parietal sulcus (IPS) that are hypothesized to encode the visuospatial transformations enabling action observation/imitation based on different visuospatial viewpoints. The results reveal that when a demonstrator executes an action, an imitator can reproduce it with similar kinematics, independently of differences in anthropometry, distance, and viewpoint. As with prior empirical findings, similar model synaptic activity was observed during both action observation and execution along with the existence of both view-independent and view-dependent neural populations in the frontal MNS. Importantly, this work generates testable behavioral and neurophysiological predictions. Namely, the model predicts that i) during observation/imitation the response time increases linearly as the rotation angle of the observed action increases but remain similar when performing both clockwise and counterclockwise rotation and ii) IPL embeds essentially view-independent neurons while SPL/IPS includes both view-independent and view-dependent neurons. Overall, this work suggests that MT/MST visuomotion processes combined with the SPL/IPS allow the MNS to observe and imitate actions independently of demonstrator-imitator spatial relationships.     

Development of infant sustained attention and its relation to EEG oscillations: an EEG and cortical source analysis study

The current study examined the relation between infant sustained attention and infant EEG oscillations. Fifty‐nine infants were tested at 6 (N = 15), 8 (N = 17), 10 (N = 14), and 12 (N = 13) months. Three attention phases, stimulus orienting, sustained attention, and attention termination, were defined based on infants' heart rate changes. Frequency analysis using simultaneously recorded EEG focused on infant theta (2–6 Hz), alpha (6–9 Hz), and beta (9–14 Hz) rhythms. Cortical source analysis of EEG oscillations was conducted with realistic infant MRI models. Theta synchronization was found over fontal pole, temporal, and parietal electrodes during infant sustained attention for 10 and 12 months. Alpha desynchronization was found over frontal, central and parietal electrodes during sustained attention. This alpha effect started to emerge at 10 months and became well established by 12 months. No difference was found for the beta rhythm between different attention phases. The theta synchronization effect was localized to the orbital frontal, temporal pole, and ventral temporal areas. The alpha desynchronization effect was localized to the brain regions composing the default mode network including the posterior cingulate cortex and precuneus, medial prefrontal cortex, and inferior parietal gyrus. The alpha desynchronization effect was also localized to the pre‐ and post‐central gyri. The present study demonstrates a connection between infant sustained attention and EEG oscillatory activities.     

The Relation Between Cognitive Development and Anxiety Phenomena in Children

We examined the relation between cognitive development and fear, anxiety, and behavioral inhibition in a non-clinical sample of 226 Dutch children aged 4–9 years. To assess cognitive development, children were tested with Piagetian conservation tasks and a Theory-of-Mind (TOM) test. Fears were measured by means of a self-report scale completed by the children, while anxiety symptoms and behavioral inhibition were indexed by rating scales that were filled out by parents. Significant age trends were observed for some anxiety phenomena. For example, younger children displayed higher fear scores, whereas older children exhibited higher levels of generalized anxiety. Most importantly, results of regression analyses (in which we controlled for age) indicated that cognitive development, and in particular TOM ability, made a unique and significant contribution to various domains of behavioral inhibition. In all cases, higher levels of TOM were associated with lower levels of behavioral inhibition. In general, percentages of explained variance were rather small (i.e., <6%), indicating that the role of cognitive development in various anxiety phenomena is limited.     

The relation between electroencephalogram asymmetry and attention biases to threat at baseline and under stress

Electroencephalogram (EEG) asymmetry in the alpha frequency band has been implicated in emotion processing and broad approach-withdrawal motivation systems. Questions remain regarding the cognitive mechanisms that may help elucidate the observed links between EEG asymmetry and patterns of socioemotional functioning. The current study observed frontal EEG asymmetry patterns at rest and under social threat among young adults (N = 45, M = 21.1 years). Asymmetries were, in turn, associated with performance on an emotion-face dot-probe attention bias task. Attention biases to threat have been implicated as potential causal mechanisms in anxiety and social withdrawal. Frontal EEG asymmetry at baseline did not predict attention bias patterns to angry or happy faces. However, increases in right frontal alpha asymmetry from baseline to the stressful speech condition were associated with vigilance to angry faces and avoidance of happy faces. The findings may reflect individual differences in the pattern of response (approach or withdrawal) with the introduction of a mild stressor. Comparison analyses with frontal beta asymmetry and parietal alpha asymmetry did not find similar patterns. Thus, the data may reflect the unique role of frontal regions, particularly the dorsolateral prefrontal cortex, in cognitive control and threat detection, coupled with ruminative processes associated with alpha activity.     

Pity or peanuts? Oxytocin induces different neural responses to the same infant crying labeled as sick or bored

The neuropeptide oxytocin plays an important role in mother–infant bonding. However, recent studies indicate that the effects of oxytocin on prosociality are dependent on perceived social context. Using functional magnetic resonance imaging, we examined differential effects of intranasally administered oxytocin on neural responding to 500 and 700 Hz crying that was indicated as emanating from a sick infant and 500 and 700 Hz crying emanating from a bored infant. We found that oxytocin significantly increased insula and inferior frontal gyrus responding to sick infant crying, but decreased activation in these brain regions during exposure to crying of an infant that was labeled as bored. In addition, oxytocin decreased amygdala responding to 500 Hz crying, but increased amygdala responding to 700 Hz crying. These findings indicate that labeling the same infant crying as ‘sick’ or as ‘bored’ drastically changes neural activity in response to intranasal oxytocin administration. Oxytocin increases empathic reactions to sick infants' crying, but lowers the perceived urgency of crying of an infant perceived as bored, thus flexibly adapting adult responses to infant crying labeled in various ways.     

A self-referential default brain state: patterns of coherence,power, and eLORETA sources during eyes-closed rest and Transcendental Meditation practice

Activation of a default mode network (DMN) including frontal and parietal midline structures varies with cognitive load, being more active during low-load tasks and less active during high-load tasks requiring executive control. Meditation practices entail various degrees of cognitive control. Thus, DMN activation patterns could give insight into the nature of meditation practices. This 10-week random assignment study compared theta2, alpha1, alpha2, beta1, beta2 and gamma EEG coherence, power, and eLORETA cortical sources during eyes-closed rest and Transcendental Meditation (TM) practice in 38 male and female college students, average age 23.7 years. Significant brainwave differences were seen between groups. Compared to eyes-closed rest, TM practice led to higher alpha1 frontal log-power, and lower beta1 and gamma frontal and parietal log-power; higher frontal and parietal alpha1 interhemispheric coherence and higher frontal and frontal-central beta2 intrahemispheric coherence. eLORETA analysis identified sources of alpha1 activity in midline cortical regions that overlapped with the DMN. Greater activation in areas that overlap the DMN during TM practice suggests that meditation practice may lead to a foundational or ‘ground’ state of cerebral functioning that may underlie eyes-closed rest and more focused cognitive processes.     

Neurophysiological marker of inhibition distinguishes language groups on a non-linguistic executive function test

Successful interaction with the environment depends on flexible behaviors which require shifting attention, inhibiting primed responses, ignoring distracting information, and withholding motor responses. These abilities, termed executive function (EF), are believed to be mediated by inhibitory processes in the frontal lobes. Superior performance on EF tests (i.e., faster reaction times (RT), and fewer errors) has been shown in bilinguals compared to monolingual speakers. However, findings are inconsistent, and no study has directly linked this bilingual advantage to frontal lobe inhibitory processes. To clarify this uncertainty, we concomitantly tested neural inhibitory processes and behavioral responses on an EF test in bilinguals and monolinguals. Specifically, we compared English monolinguals (N = 15) to Spanish/English bilinguals (N = 13) on event-related brain potentials (ERP) during a non-linguistic, auditory Go/NoGo task, a task linked to non-motor, cognitive inhibition in monolinguals. Participants responded with a button press on trials in which target tone-pairs (Go trials) were presented and withheld their responses on non-target trials (NoGo trials). Results revealed significantly greater inhibition (i.e., greater mean N2 amplitude) in bilinguals compared to monolinguals during NoGo trials even though both groups performed the task equally well (i.e., withheld a motor response). On Go trials where participants pressed a response button, neither ERPs nor RT distinguished the groups. Additionally, scores on a second language proficiency test (i.e., English in our bilingual group) were positively correlated with N2 amplitude. These findings are the first to directly link this bilingual advantage to a neural correlate of inhibition and to reveal that inhibition in bilinguals is moderated by second language proficiency. Results are discussed in the context of plasticity, and we propose that evaluating bilinguals at varying levels of second-language proficiency may serve as a model of human neuroplasticity.     

Brain dynamics of visual anticipation during spatial occlusion tasks in expert tennis players

Stimulus identification and action outcome understanding for a rapid and accurate response selection, play a fundamental role in racquet sports. Here, we investigated the neurodynamics of visual anticipation in tennis manipulating the postural and kinematic information associated with the body of opponents by means of a spatial occlusion protocol. Event Related Potentials (ERPs) were evaluated in two groups of professional tennis players (N = 37) with different levels of expertise, while they observed pictures of opponents and predicted the landing position as fast and accurately as possible. The observed action was manipulated by deleting different body districts of the opponent (legs, ball, racket and arm, trunk). Full body image (no occlusion) was used as control condition. The worst accuracy and the slowest response time were observed in the occlusion of trunk and ball. The former was associated with a reduced amplitude of the ERP components likely linked to body processing (the N1 in the right hemisphere) and visual-motor integration awareness (the pP1), as well as with an increase of the late frontal negativity (the pN2), possibly reflecting an effort by the insula to recover and/or complete the most correct sensory-motor representation. In both occlusions, a decrease in the pP2 may reflect an impairment of decisional processes upon action execution following sensory evidence accumulation. Enhanced amplitude of the P3 and the pN2 components were found in more experienced players, suggesting a greater allocation of resources in the process connecting sensory encoding and response execution, and sensory-motor representation.     

Substance use and mothers’ neural responses to infant cues

Substance use may influence mothers’ responsiveness to their infants and negatively impact the parent–infant relationship. Maternal substance use may co-opt neural circuitry involved in caregiving, thus reducing the salience of infant cues and diminishing the sense of reward experienced by caring for infants. Gaps in understanding exist with regard to the mechanisms by which substance use operates to influence mothers’ processing of infant cues and how this translates to caregiving. Therefore, we examined how substance use might relate to maternal neural responses to infant cues using event-related potentials (ERPs). Substance-using (n = 29) and nonsubstance-using (n = 29) mothers viewed photographs of infant faces and heard recordings of infant vocalizations while electroencephalography was recorded simultaneously. Three specific ERP components were used to examine initial processing of infant faces (N170) and cries (N100), and attentional allocation to infant faces and cries (P300). Substance-using mothers did not discriminate facial affect at early encoding stages (N170), were generally slower to orient to infant cries (N100), showed heightened responses to neutral faces (P300), and failed to adaptively differentiate between high-distress versus low-distress cries (P300). These differences may be important to caregiving behaviors associated with the formation of mother–child attachment. Implications are discussed, as are limitations and future directions.     

Regional and inter-regional theta oscillation during episodic novelty processing

Recent event-related potential (ERP) and functional magnetic resonance imaging (fMRI) studies suggest that novelty processing may be involved in processes that recognize the meaning of a novel sound, during which widespread cortical regions including the right prefrontal cortex are engaged. However, it remains unclear how those cortical regions are functionally integrated during novelty processing. Because theta oscillation has been assumed to have a crucial role in memory operations, we examined local and inter-regional neural synchrony of theta band activity during novelty processing. Fifteen right-handed healthy university students participated in this study. Subjects performed an auditory novelty oddball task that consisted of the random sequence of three types of stimuli such as a target (1000 Hz pure tone), novel (familiar environmental sounds such as dog bark, buzz, car crashing sound and so on), and standard sounds (950 Hz pure tone). Event-related spectra perturbation (ERSP) and the phase-locking value (PLV) were measured from human scalp EEG during task. Non-parametric statistical tests were applied to test for significant differences between stimulus novelty and stimulus targets in ERSP and PLV. The novelty P3 showed significant higher amplitude and shorter latency compared with target P3 in frontocentral regions. Overall, theta activity was significantly higher in the novel stimuli compared with the target stimuli. Specifically, the difference in theta power between novel and target stimuli was most significant in the right frontal region. This right frontal theta activity was accompanied by phase synchronization with the left temporal region. Our results imply that theta phase synchronization between right frontal and left temporal regions underlie the retrieval of memory traces for unexpected but familiar sounds from long term memory in addition to working memory retrieval or novelty encoding.     

Prenatal maternal depression predicts neural maturation and negative emotion in infants

Despite widespread acceptance that prenatal symptoms of depression in mothers are detrimental to infants’ long-term emotional and cognitive development, little is known about the mechanisms that may integrate outcomes across these domains. Rooted in the integrative perspective that emotional development is grounded in developing cognitive processes, we hypothesized that prenatal symptoms of depression in mothers would be associated with delays in neural maturation that support sociocognitive function in infants, leading to more problematic behaviors. We used a prospective longitudinal study of mothers (N = 92) and their infants to test whether self-reported symptoms of depression in mothers during the second and third trimesters were associated with neural development and infant outcomes at 4 months of age. While controlling for postpartum symptoms of depression, more prenatal symptoms of depression in mothers predicted less neural maturation in the parietal region of 4-month-old infants. Less neural maturation, in turn, was associated with greater infant negativity, suggesting neural maturation as a putative mechanism linking maternal symptoms of depression with infant outcomes. Differences in neural regions and developmental timing are also discussed.     

The group as a self-active system: Outline of a theory of group action

Human social systems, and groups in partlcular, are conceived as units which, as a whole, actively strive towards the achievement of external and internal goals. This ‘group action’ consists of simultaneous integrated processes on various individual and social levels. Our theory comprises four groups of constructs, which refer to task structure, group structure, information processing and execution. In an ongoing group action, the task structure is projected on the group structure; according to the resulting pattern, the group processes its action related information and executes the act. The latter two processes proceed on two levels, on an individual and on a group level. There are cognition, emotion and volition on the individual, and communication on the group level of information processing; execution proceeds in individual action and in cooperation. A specific part of the theory concerns analogies between individual cognition and intragroup communication.     

Neurophysiology of sustained attention in early infancy: Investigating longitudinal relations with recognition memory outcomes

The ability to sustain attention is a critical cognitive domain that emerges in infancy and is predictive of a multitude of cognitive processes. Here, we used a heart rate (HR) defined measure of sustained attention to assess corresponding changes in frontal electroencephalography (EEG) power at 3 months of age. Second, we examined how the neural underpinnings of HR-defined sustained attention were associated with sustained attention engagement. Third, we evaluated if neural or behavioral sustained attention measures at 3-months predicted subsequent recognition memory scores at 9 months of age. Seventy-five infants were included at 3 months of age and provided usable attention and EEG data and 25 infants returned to the lab at 9 months and provided usable recognition memory data. The current study focuses on oscillatory power in the theta (4–6 Hz) frequency band during phases of HR-defined sustained attention and inattention phases. Results revealed that theta power was significantly higher during phases of sustained attention. Second, higher theta power during sustained attention was positively associated with proportion of time in sustained attention. Third, longitudinal analyses indicated a significant positive association between theta power during sustained attention on 9-month visual paired comparison scores such that higher theta power predicted higher visual paired comparison scores at 9-months. These results highlight the interrelation of the attention and arousal systems which have longitudinal implications for subsequent recognition memory processes.