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Sequence memory is subject to age-related decline, but the underlying processes are not yet fully understood. We analyzed electroencephalography (EEG) in 21 healthy older (60-80 years) and 26 young participants (20-30 years) and compared time-frequency spectra and theta-gamma phase-amplitude-coupling (PAC) during encoding of the order of visually presented items. In older adults, desynchronization in theta (4-8 Hz) and synchronization in gamma (30-45 Hz) power did not distinguish between subsequently correctly and incorrectly remembered trials, while there was a subsequent memory effect for young adults. Theta-gamma PAC was modulated by item position within a sequence for older but not young adults. Specifically, position within a sequence was coded by higher gamma amplitude for successive theta phases for later correctly remembered trials. Thus, deficient differentiation in theta desynchronization and gamma oscillations during sequence encoding in older adults may reflect neurophysiological correlates of age-related memory decline. Furthermore, our results indicate that sequences are coded by theta-gamma PAC in older adults, but that this mechanism might lose precision in aging., (Copyright © 2024 The Authors. Published by Elsevier Inc. All rights reserved.)

Decoding others' intentions accurately in order to adapt one's own behavior is pivotal throughout life. In this study, we asked how younger and older adults deal with uncertainty in dynamic social environments. We used an advice-taking paradigm together with Bayesian modeling to characterize effects of aging on learning about others' time-varying intentions. We observed age differences when comparing learning on two levels of social uncertainty: the fidelity of the adviser and the volatility of intentions. Older adults expected the adviser to change his/her intentions more frequently (i.e., a higher volatility of the adviser). They also showed higher confidence (i.e., precision) in their volatility beliefs and were less willing to change their beliefs about volatility over the course of the experiment. This led them to update their predictions about the fidelity of the adviser more quickly. Potentially indicative of stereotype effects, we observed that older advisers were perceived as more volatile, but also more faithful than younger advisers. This offers new insights into adult age differences in response to social uncertainty., (Copyright © 2021 Elsevier Inc. All rights reserved.)

Aging attenuates frontostriatal network functioning, which could lead to deficits in value computation when decision-making involves uncertainty. Although it has been shown that visually enhancing information saliency of outcome probability can improve decision-making in old age, mechanisms of this effect are still unclear. In the present study, the saliency of outcome probability was increased using a color-coding scheme as a decision aid in a mixed lottery choice task, and spontaneous eye-blink rate and pupillary responses were assessed in younger and older adults. Older adults showed lower value sensitivity than younger adults; however, increasing information saliency benefitted choice behaviors in both age groups. Furthermore, the decision aid reduced pupil size during decision-making in both age groups, suggesting that enhancing information saliency decreases cognitive demands of value computation. Baseline value sensitivity was negatively correlated with benefit of enhancing information saliency only in older adults. As value representations in older decision makers are less distinctive at baseline, they may have required more environmental compensation than younger adults., (Copyright © 2020 Elsevier Inc. All rights reserved.)

Older adults experience difficulties in daily situations that require flexible information selection in the presence of multiple competing sensory inputs, like for instance multi-talker situations. Modulations of rhythmic neural activity in the alpha-beta (8-30 Hz) frequency range in posterior brain areas have been established as a cross-modal neural correlate of selective attention. However, research linking compromised auditory selective attention to changes in rhythmic neural activity in aging is sparse. We tested younger (n = 25; 22-35 years) and older adults (n = 26; 63-76 years) in an attention modulated dichotic listening task. In this, two streams of highly similar auditory input were simultaneously presented to participants' both ears (i.e., dichotically) while attention had to be focused on the input to only one ear (i.e. target) and the other, distracting information had to be ignored. We here demonstrate a link between severely compromised auditory selective attention in aging and a partial reorganization of attention-related rhythmic neural responses. In particular, in old age we observed a shift from a self-initiated, preparatory modulation of lateralized alpha rhythmic activity to an externally driven response in the alpha-beta range. Critically, moment-to-moment fluctuations in the age-specific patterns of self-initiated and externally driven lateralized rhythmic activity were associated with behavioral performance. We conclude that adult age differences in spatial selective attention likely derive from a functional reorganization of rhythmic neural activity within the aging brain., (Copyright © 2019 Elsevier Inc. All rights reserved.)

Aging and dopamine modulation have both been independently shown to influence the functional connectivity of brain networks during rest. Dopamine modulation is known to decline during the course of aging. Previous evidence also shows that the dopamine transporter gene (DAT1) influences the re-uptake of dopamine and the anyA9 genotype of this gene is associated with higher striatal dopamine signaling. Expanding these two lines of prior research, we investigated potential interactive effects between aging and individual variations in the DAT1 gene on the modular organization of brain acvitiy during rest. The graph-theoretic metrics of modularity, betweenness centrality and participation coefficient were assessed in 41 younger (age 20-30 years) and 37 older (age 60-75 years) adults. Age differences were only observed in the participation coefficient in carriers of the anyA9 genotype of the DAT1 gene and this effect was most prominently observed in the default mode network. Furthermore, we found that individual differences in the values of the participation coefficient correlated with individual differences in fluid intelligence and a measure of executive control in the anyA9 carriers. The correlation between participation coefficient and fluid intelligence was mainly shared with age-related differences, whereas the correlation with executive control was independent of age. These findings suggest that DAT1 genotype moderates age differences in the functional integration of brain networks as well as the relation between network characteristics and cognitive abilities., Competing Interests: The authors have declared that no competing interests exist.

Probabilistic category learning involves complex interactions between the hippocampus and striatum that may depend on whether acquisition occurs via feedback or observation. Little is known about how healthy aging affects these processes. We tested whether age-related behavioral differences in probabilistic category learning from feedback or observation depend on a genetic factor known to influence individual differences in hippocampal function, the KIBRA gene (single nucleotide polymorphism rs17070145). Results showed comparable age-related performance impairments in observational as well as feedback-based learning. Moreover, genetic analyses indicated an age-related interactive effect of KIBRA on learning: among older adults, the beneficial T-allele was positively associated with learning from feedback, but negatively with learning from observation. In younger adults, no effects of KIBRA were found. Our results add behavioral genetic evidence to emerging data showing age-related differences in how neural resources relate to memory functions, namely that hippocampal and striatal contributions to probabilistic category learning may vary with age. Our findings highlight the effects genetic factors can have on differential age-related decline of different memory functions., (Copyright © 2017 Elsevier Inc. All rights reserved.)

Research in younger adults dissociates cognitive from affective facets of social information processing, rather than promoting a monolithic view of social intelligence. An influential theory on adult development suggests differential effects of aging on cognitive and affective functions. However, this dissociation has not been directly tested in the social domain. Employing a newly developed naturalistic paradigm that disentangles facets of the social mind within an individual, we show multi-directionality of age-related differences. Specifically, components of the socio-cognitive route - Theory of Mind and metacognition - are impaired in older relative to younger adults. Nevertheless, these social capacities are still less affected by aging than factual reasoning and metacognition regarding non-social content. Importantly, the socio-affective route is well-functioning, with no decline in empathy and elevated compassion in the elderly. These findings contribute to an integrated theory of age-related change in social functioning and inform interventions tailored to specifically reinstate socio-cognitive skills in old age.

Healthy aging can lead to impairments in learning that affect many laboratory and real-life tasks. These tasks often involve the acquisition of dynamic contingencies, which requires adjusting the rate of learning to environmental statistics. For example, learning rate should increase when expectations are uncertain (uncertainty), outcomes are surprising (surprise) or contingencies are more likely to change (hazard rate). In this study, we combine computational modelling with an age-comparative behavioural study to test whether age-related learning deficits emerge from a failure to optimize learning according to the three factors mentioned above. Our results suggest that learning deficits observed in healthy older adults are driven by a diminished capacity to represent and use uncertainty to guide learning. These findings provide insight into age-related cognitive changes and demonstrate how learning deficits can emerge from a failure to accurately assess how much should be learned.

The hippocampus and striatum are core neural circuits involved in spatial learning and memory. Although both neural systems support spatial navigation, experimental and theoretical evidence indicate that they play different roles. In particular, whereas hippocampal place cells generate allocentric neural representations of space that are sensitive to geometric information, striatum-dependent learning is influenced by local landmarks. How human aging affects these different neural representations, however, is still not well understood. In this paper, we combined virtual reality, computational modeling, and neuroimaging to investigate the effects of age upon the neural computation and representation of space in humans. We manipulated the geometry and local landmarks of a virtual environment and examined the effects on memory performance and brain activity during spatial learning. In younger adults, both behavior and brain activity in the medial-temporal lobe were consistent with predictions of a computational model of hippocampus-dependent boundary processing. In contrast, older adults' behavior and medial-temporal lobe activity were primarily influenced by local cue information, and spatial learning was more associated with activity in the caudate nucleus rather than the hippocampus. Together these results point to altered spatial representations and information processing in the hippocampal-striatal circuitry with advancing adult age, which may contribute to spatial learning and memory deficits associated with normal and pathological aging., (Copyright © 2015 Elsevier Inc. All rights reserved.)

Foresighted decision-making depends on the ability to learn the value of future outcomes and the sequential choices necessary to achieve them. Using a 3-stage Markov decision task and functional magnetic resonance imaging, we investigated age differences in the ability to extract state transition structures while learning to predict future reward. In younger adults learning was associated with enhanced activity in the prefrontal cortex (PFC). In older adults (OA) we found no evidence for PFC recruitment. However, high-performing OA showed enhanced striatal activity, suggesting that they may engage in a model-free (experience-based) learning strategy. Change point analyses revealed that in younger adults learning was characterized by distinct and abrupt shifts in PFC activity, which were predictive of behavioral change points. In OA PFC activity was less pronounced and not predictive of behavior. Our findings suggest that age-related impairments in learning future reward value can be attributed to a deficit in extracting sequential state transition structures. This deficit may lead to myopic decisions in OA if contextual information has to be temporally integrated., (Copyright © 2015 The Authors. Published by Elsevier Inc. All rights reserved.)

Better-performing younger adults typically express greater brain signal variability relative to older, poorer performers. Mechanisms for age and performance-graded differences in brain dynamics have, however, not yet been uncovered. Given the age-related decline of the dopamine (DA) system in normal cognitive aging, DA neuromodulation is one plausible mechanism. Hence, agents that boost systemic DA [such as d-amphetamine (AMPH)] may help to restore deficient signal variability levels. Furthermore, despite the standard practice of counterbalancing drug session order (AMPH first vs. placebo first), it remains understudied how AMPH may interact with practice effects, possibly influencing whether DA up-regulation is functional. We examined the effects of AMPH on functional-MRI-based blood oxygen level-dependent (BOLD) signal variability (SD(BOLD)) in younger and older adults during a working memory task (letter n-back). Older adults expressed lower brain signal variability at placebo, but met or exceeded young adult SD(BOLD) levels in the presence of AMPH. Drug session order greatly moderated change-change relations between AMPH-driven SD(BOLD) and reaction time means (RT(mean)) and SDs (RT(SD)). Older adults who received AMPH in the first session tended to improve in RT(mean) and RT(SD) when SD(BOLD) was boosted on AMPH, whereas younger and older adults who received AMPH in the second session showed either a performance improvement when SD(BOLD) decreased (for RT(mean)) or no effect at all (for RT(SD)). The present findings support the hypothesis that age differences in brain signal variability reflect aging-induced changes in dopaminergic neuromodulation. The observed interactions among AMPH, age, and session order highlight the state- and practice-dependent neurochemical basis of human brain dynamics.

The efficacy of various transmitter systems declines with advancing age. Of particular interest, various pre-synaptic and post-synaptic components of the dopaminergic system change across the human lifespan; impairments in these components play important roles in cognitive deficits commonly observed in the elderly. Here, we review evidence from recent multimodal neuroimaging, pharmacological and genetic studies that have provided new insights for the associations among dopamine functions, aging, functional brain activations and behavioral performance across key cognitive functions, ranging from working memory and episodic memory to goal-directed learning and decision making. Specifically, we discuss these empirical findings in the context of an established neurocomputational theory of aging neuronal gain control. We also highlight gaps in the current understanding of dopamine neuromodulation and aging brain functions and suggest avenues for future research., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

Conditions that render the selection of correct actions difficult require the monitoring of the execution and outcomes of one's own actions. Such performance monitoring abilities undergo maturational and aging-related changes across the lifespan. This review highlights evidence for qualitative differences in behavior and physiological correlates of performance monitoring across the lifespan. Few developmental studies examine both stimulus-locked as well as response-locked components. Here, we examine a lifespan pattern of stimulus- as well as response-locked ERPs during performance monitoring to inform age differences in subprocesses of performance monitoring. Findings from functional magnetic resonance imaging (fMRI) studies that lend further support for the observed age differences in performance monitoring are also reviewed. Together, the evidence suggest that suboptimal performance monitoring during maturation is characterized by a reduced ability to flexibly translate experienced conflicts into top-down control, whereas declined performance monitoring in aging is characterized by difficulties in maintaining task set representations. Such age specific deficits are apparent in performance monitoring related to response conflicts as well as in performance monitoring during reinforcement learning and value-based decision making., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

Children and older adults often show less favorable reward-based learning and decision making, relative to younger adults. It is unknown, however, whether reward-based processes that influence relatively early perceptual and attentional processes show similar lifespan differences. In this study, we investigated whether stimulus-reward associations affect selective visual attention differently across the human lifespan. Children, adolescents, younger adults, and older adults performed a visual search task in which the target colors were associated with either high or low monetary rewards. We discovered that high reward value speeded up response times across all four age groups, indicating that reward modulates attentional selection across the lifespan. This speed-up in response time was largest in younger adults, relative to the other three age groups. Furthermore, only younger adults benefited from high reward value in increasing response consistency (i.e., reduction of trial-by-trial reaction time variability). Our findings suggest that reward-based modulations of relatively early and implicit perceptual and attentional processes are operative across the lifespan, and the effects appear to be greater in adulthood. The age-specific effect of reward on reducing intraindividual response variability in younger adults likely reflects mechanisms underlying the development and aging of reward processing, such as lifespan age differences in the efficacy of dopaminergic modulation. Overall, the present results indicate that reward shapes visual perception across different age groups by biasing attention to motivationally salient events.

Similar to other industrialized countries, Germany's population is ageing. Whereas some people enjoy good physical and cognitive health into old age, others suffer from a multitude of age-related disorders and impairments which reduce life expectancy and affect quality of life. To identify and characterize the factors associated with 'healthy' vs. 'unhealthy' ageing, we have launched the Berlin Aging Study II (BASE-II), a multidisciplinary and multi-institutional project that ascertains a large number of ageing-related variables from a wide range of different functional domains. Phenotypic assessments include factors related to geriatrics and internal medicine, immunology, genetics, psychology, sociology and economics. Baseline recruitment of the BASE-II cohort was recently completed and has led to the sampling of 1600 older adults (age range 60-80 years), as well as 600 younger adults (20-35 years) serving as the basic population for in-depth analyses. BASE-II data are linked to the German Socio-Economic Panel Study (SOEP), a long-running panel survey representative of the German population, to estimate sample selectivity. A major goal of BASE-II is to facilitate collaboration with other research groups by freely sharing relevant phenotypic and genotypic data with qualified outside investigators., (Published by Oxford University Press on behalf of the International Epidemiological Association © The Author 2013; all rights reserved.)

According to a neurocomputational theory of cognitive aging, senescent changes in dopaminergic modulation lead to noisier and less differentiated processing. The authors tested a corollary hypothesis of this theory, according to which genetic predispositions of individual differences in prefrontal dopamine (DA) signaling may affect associations between memory functions, particularly in old age. Latent correlations between factors of verbal episodic memory and spatial working memory were compared between individuals carrying different allelic variants of the Catechol-O-Methyltransferase (COMT) Val158Met polymorphism, which influences DA availability in prefrontal cortex. In younger adults (n = 973), correlations between memory functions did not differ significantly among the 3 COMT genotypes (r = .35); in older adults (n = 1333), however, the correlation was significantly higher in Val homozygotes (r = .70), whose prefrontal DA availability is supposedly the lowest of all groups examined, than in heterozygotes and Met homozygotes (both rs = .29). Latent means of the episodic memory and working memory factors did not differ by COMT status within age groups. However, when restricting the analysis to the low-performing tertile of older adults (n = 443), we found that Val homozygotes showed lower levels of performance in both episodic memory and working memory than heterozygotes and Met homozygotes. In line with the neurocomputational theory, the observed dedifferentiation of memory functions in older Val homozygotes suggests that suboptimal dopaminergic modulation may underlie multiple facets of memory declines during aging. Future longitudinal work needs to test this conjecture more directly., (PsycINFO Database Record (c) 2014 APA, all rights reserved.)

Both the dopaminergic and glutamatergic systems modulate episodic memory consolidation. Evidence from animal studies suggests that these two neurotransmitters may interact in influencing memory performance. Given that individual differences in episodic memory are heritable, we investigated whether variations of the dopamine D2 receptor gene (rs6277, C957T) and the N-methyl-D-aspartate 3A (NR3A) gene, coding for the N-methyl-D-aspartate 3A subunit of the glutamate N-methyl-D-aspartate receptor (rs10989591, Val362Met), interactively modulate episodic memory in large samples of younger (20-31 years; n = 670) and older (59-71 years; n = 832) adults. We found a reliable gene-gene interaction, which was observed in older adults only: older individuals carrying genotypes associated with greater D2 and N-methyl-D-aspartate receptor efficacy showed better episodic performance. These results are in line with findings showing magnification of genetic effects on memory in old age, presumably as a consequence of reduced brain resources. Our findings underscore the need for investigating interactive effects of multiple genes to understand individual difference in episodic memory., (Copyright © 2014 Elsevier Inc. All rights reserved.)

In addition to sensory decline, age-related losses in auditory perception also reflect impairments in attentional modulation of perceptual saliency. Using an attention and intensity-modulated dichotic listening paradigm, we investigated electrophysiological correlates of processing conflicts between attentional focus and perceptual saliency in 25 younger and 26 older adults. Participants were instructed to attend to the right or left ear, and perceptual saliency was manipulated by varying the intensities of both ears. Attentional control demand was higher in conditions when attentional focus and perceptual saliency favored opposing ears than in conditions without such conflicts. Relative to younger adults, older adults modulated their attention less flexibly and were more influenced by perceptual saliency. Our results show, for the first time, that in younger adults a late negativity in the event-related potential (ERP) at fronto-central and parietal electrodes was sensitive to perceptual-attentional conflicts during auditory processing (N450 modulation effect). Crucially, the magnitude of the N450 modulation effect correlated positively with task performance. In line with lower attentional flexibility, the ERP waveforms of older adults showed absence of the late negativity and the modulation effect. This suggests that aging compromises the activation of the fronto-parietal attentional network when processing the competing and conflicting auditory information.

Trial-to-trial reaction time (RT) variability is consistently higher in children and older adults than in younger adults. Converging evidence also indicates that higher RT variability is (a) associated with lower behavioral performance on complex cognitive tasks, (b) distinguishes patients with neurological deficits from healthy individuals, and also (c) predicts longitudinal cognitive decline in older adults. However, so far the processes underlying increased RT variability are poorly understood. Previous evidence suggests that control signals in the medial frontal cortex (MFC) are reflected in theta band activity and may implicate the coordination of distinct brain areas during performance monitoring. We hypothesized that greater trial-to-trial variability in theta power during performance monitoring may be associated with greater behavioral variability in response latencies. We analyzed event-related theta oscillations assessed during a cued-Go/NoGo task in a lifespan sample covering the age range from middle childhood to old age. Our results show that theta inter-trial coherence during NoGo trials increases from childhood to early adulthood, and decreases from early adulthood to old age. Moreover, in all age groups, individuals with higher variability in medial frontal stimulus-locked theta oscillations showed higher trial-to-trial RT variability behaviorally. Importantly, this effect was strongest at high performance monitoring demands and independent of motor response execution as well as theta power. Taken together, our findings reveal that lower theta inter-trial coherence is related to greater behavioral variability within and across age groups. These results hint at the possibility that more variable MFC control may be associated with greater performance fluctuations., (© 2013.)

The striatum and medial temporal lobe play important roles in implicit and explicit memory, respectively. Furthermore, recent studies have linked striatal dopamine modulation to both implicit as well as explicit sequence learning and suggested a potential role of the striatum in the emergence of explicit memory during sequence learning. With respect to aging, previous findings indicated that implicit memory is less impaired than explicit memory in older adults and that genetic effects on cognition are magnified by aging. To understand the links between these findings, we investigated effects of aging and genotypes relevant for striatal dopamine on the implicit and explicit components of sequence learning. Reaction time (RT) and error data from 80 younger (20-30 years) and 70 older adults (60-71 years) during a serial reaction time task showed that age differences in learning-related reduction of RTs emerged gradually over the course of learning. Verbal recall and measures derived from the process-dissociation procedure revealed that younger adults acquired more explicit memory about the sequence than older adults, potentially causing age differences in RT gains in later stages of learning. Of specific interest, polymorphisms of the dopamine- and cAMP-regulated neuronal phosphoprotein (DARPP-32, rs907094) and dopamine transporter (DAT, VNTR) genes showed interactive effects on overall RTs and verbal recall of the sequence in older but not in younger adults. Together our findings show that variations in genotypes relevant for dopamine functions are associated more with aging-related impairments in the explicit than the implicit component of sequence learning, providing support for theories emphasizing the role of dopaminergic modulation in cognitive aging and the magnification of genetic effects in human aging., (© 2013 Elsevier Ltd. All rights reserved.)

Behavioral, cognitive, and motivational development entails co-constructive interactions between the environmental and social influences from the developmental context, on the one hand, and the individual's neurobiological inheritance, on the other hand. Key brain networks underlying cognition, emotion, and motivation are innervated by major transmitter systems (e.g., the catecholamines and acetylcholine). Thus, the maturation and senescence of neurotransmitter systems have direct implications for lifespan development. In addition to reviewing evidence on life age differences in dopaminergic modulation and cognitive development, this brief review selectively highlights recent findings on how important influences from the developmental context, such as reward-mediated motivational processes, transgenerational stress transmission, psychosocial stress, and cognitive interventions, may, in part, exert their effects on brain and behavioral development through their effects on neuromodulatory mechanisms., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

Spatial navigation relies on multiple mnemonic mechanisms and previous work in younger adults has described two separate types of spatial memory. One type uses directional as well as boundary-related information for spatial memory and mainly implicates the hippocampal formation. The other type has been linked to directional and landmark-related information and primarily involves the striatum. Using a virtual reality navigation paradigm, we studied the impacts of aging and a single nucleotide polymorphism (SNP rs17070145) of the KIBRA gene (official name: WWC1) on these memory forms. Our data showed that older adult's spatial learning was preferentially related to processing of landmark information, whereas processing of boundary information played a more prominent role in younger adults. Moreover, among older adults T-allele carriers of the examined KIBRA polymorphism showed better spatial learning compared to C homozygotes. Together these findings provide the first evidence for an effect of the KIBRA rs17070145 polymorphism on spatial memory in humans and age differences in the reliance on landmark and boundary-related spatial information., (Copyright © 2013 Wiley Periodicals, Inc.)

Multitalker situations confront listeners with a plethora of competing auditory inputs, and hence require selective attention to relevant information, especially when the perceptual saliency of distracting inputs is high. This study augmented the classical forced-attention dichotic listening paradigm by adding an interaural intensity manipulation to investigate developmental differences in the interplay between perceptual saliency and attentional control during auditory processing between early and middle childhood. We found that older children were able to flexibly focus on instructed auditory inputs from either the right or the left ear, overcoming the effects of perceptual saliency. In contrast, younger children implemented their attentional focus less efficiently. Direct comparisons of the present data with data from a recently published study of younger and older adults from our group suggest that younger children and older adults show similar levels of performance. Critically, follow-up comparisons revealed that younger children's performance restrictions reflect difficulties in attentional control only, whereas older adults' performance deficits also reflect an exaggerated reliance on perceptual saliency. We conclude that auditory attentional control improves considerably from middle to late childhood and that auditory attention deficits in healthy aging cannot be reduced to a simple reversal of child developmental improvements.

The capacity of visual-spatial working memory (WM) declines from early to late adulthood. Recent attempts at identifying neural correlates of WM capacity decline have focused on the maintenance phase of WM. Here, we investigate neural mechanisms during the encoding phase as another potential mechanism contributing to adult age differences in WM capacity. We used electroencephalography to track neural activity during encoding and maintenance on a millisecond timescale in 35 younger and 35 older adults performing a visual-spatial WM task. As predicted, we observed pronounced age differences in ERP indicators of WM encoding: Younger adults showed attentional selection during item encoding (N2pc component), but this selection mechanism was greatly attenuated in older adults. Conversely, older adults showed more pronounced signs of early perceptual stimulus processing (N1 component) than younger adults. The amplitude modulation of the N1 component predicted WM capacity in older adults, whereas the attentional amplitude modulation of the N2pc component predicted WM capacity in younger adults. Our findings suggest that adult age differences in mechanisms of WM encoding contribute to adult age differences in limits of visual-spatial WM capacity., (Copyright © 2013 Elsevier Inc. All rights reserved.)

Emerging evidence from animal studies suggests that suboptimal dopamine (DA) modulation may be associated with increased forgetting of episodic information. Extending these observations, we investigated the influence of DA-relevant genes on forgetting in samples of younger (n = 433, 20-31 years) and older (n = 690, 59-71 years) adults. The effects of single nucleotide polymorphisms of the DA D2 (DRD2) and D3 (DRD3) receptor genes as well as the DA transporter gene (DAT1; SLC6A3) were examined. Over the course of one week, older adults carrying two or three genotypes associated with higher DA signaling (i.e., higher availability of DA and DA receptors) forgot less pictorial information than older individuals carrying only one or no beneficial genotype. No such genetic effects were found in younger adults. The results are consistent with the view that genetic effects on cognition are magnified in old age. To the best of our knowledge, this is the first report to relate genotypes associated with suboptimal DA modulation to more long-term forgetting in humans. Independent replication studies in other populations are needed to confirm the observed association.

Declines in selective attention are one of the sources contributing to age-related impairments in a broad range of cognitive functions. Most previous research on mechanisms underlying older adults' selection deficits has studied the deployment of visual attention to static objects and features. Here we investigate neural correlates of age-related differences in spatial attention to multiple objects as they move. We used a multiple object tracking task, in which younger and older adults were asked to keep track of moving target objects that moved randomly in the visual field among irrelevant distractor objects. By recording the brain's electrophysiological responses during the tracking period, we were able to delineate neural processing for targets and distractors at early stages of visual processing (~100-300 msec). Older adults showed less selective attentional modulation in the early phase of the visual P1 component (100-125 msec) than younger adults, indicating that early selection is compromised in old age. However, with a 25-msec delay relative to younger adults, older adults showed distinct processing of targets (125-150 msec), that is, a delayed yet intact attentional modulation. The magnitude of this delayed attentional modulation was related to tracking performance in older adults. The amplitude of the N1 component (175-210 msec) was smaller in older adults than in younger adults, and the target amplification effect of this component was also smaller in older relative to younger adults. Overall, these results indicate that normal aging affects the efficiency and timing of early visual processing during multiple object tracking.

Aging compromises dopamine transporter (DAT) and receptor mechanisms in the frontostriatal circuitry. In a sample of 1288 younger and older adults, we investigated (i) whether individual differences in genotypes of the DAT gene (i.e., SLC6A3, the DAT variable number of tandem repeat 9/9, 9/10, and 10/10) and in the D2 receptor (DRD2) gene (i.e., the C957T [rs6277] CC and any T) interactively contribute to phenotype variations in episodic memory performance; and (ii) whether these genetic effects are magnified in older adults, because of considerable declines in the dopamine functions. Our results showed that carrying genotypes associated with higher levels of striatal synaptic dopamine (DAT 9/9) and higher density of extrastriatal D2 receptors (C957T CC) were associated with better backward serial recall, an episodic memory task with high encoding and retrieval demands. Critically, the gene-gene interaction effect was reliably stronger in older than in younger adults. In line with the resource modulation hypothesis, our findings suggest that aging-related decline in brain phenotypes (e.g., dopamine functions) could alter the relations between genotypes and behavioral phenotypes (e.g., episodic memory)., (Copyright © 2013 Elsevier Inc. All rights reserved.)

The reliability, stability, and signal-to-noise ratio (SNR) of event-related potentials (ERPs) were investigated in children, adolescents, younger adults, and older adults in performance monitoring tasks. P2, N2, P3, and P2-N2 peak-to-peak amplitude showed high odd-even split reliabilities in all age groups, ranging from.70 to.90. Multigroup analyses showed that test-retest stabilities (across 2 weeks) of ERP amplitudes did not differ among the four age groups. In contrast, relative to adolescents and younger adults, SNRs were lower in children and older adults, with higher noise levels in children and lower signal power in older adults. We conclude that age differences in the SNR of stimulus-locked ERPs can be successfully compensated by the averaging procedure with about 40 trials in the average. However, age differences in baseline noise and split-half reliability should be considered when comparing age groups in single trial measures or time-varying processes with ERPs., (Copyright © 2012 Society for Psychophysiological Research.)

Executive functions that are dependent upon the frontal-parietal network decline considerably during the course of normal aging. To delineate neuroanatomical correlates of age-related executive impairment, we investigated the relation between cortical thickness and executive functioning in 73 younger (20-32 years) and 56 older (60-71 years) healthy adults. Executive functioning was assessed using the Wisconsin Card Sorting Test (WCST). Cortical thickness was measured at each location of the cortical mantle using surface-based segmentation procedures on high-resolution T1-weighted magnetic resonance images. For regions involved in WCST performance, such as the lateral prefrontal and parietal cortices, we found that thicker cortex was related to higher accuracy. Follow-up ROI-based analyses revealed that these associations were stronger in older than in younger adults. Moreover, among older adults, high and low performers differed in cortical thickness within regions generally linked to WCST performance. Our results indicate that the structural cortical correlates of executive functioning largely overlap with previously identified functional patterns. We conclude that structural preservation of relevant brain regions is associated with higher levels of executive performance in old age, and underscore the need to consider the heterogeneity of brain aging in relation to cognitive functioning., (Copyright © 2011 Wiley-Liss, Inc.)

Attention and working memory are fundamental for selecting and maintaining behaviorally relevant information. Not only do both processes closely intertwine at the cognitive level, but they implicate similar functional brain circuitries, namely the frontoparietal and the frontostriatal networks, which are innervated by cholinergic and dopaminergic pathways. Here we review the literature on cholinergic and dopaminergic modulations of visual-spatial attention and visual working memory processes to gain insights on aging-related changes in these processes. Some extant findings have suggested that the cholinergic system plays a role in the orienting of attention to enable the detection and discrimination of visual information, whereas the dopaminergic system has mainly been associated with working memory processes such as updating and stabilizing representations. However, since visual-spatial attention and working memory processes are not fully dissociable, there is also evidence of interacting cholinergic and dopaminergic modulations of both processes. We further review gene-cognition association studies that have shown that individual differences in visual-spatial attention and visual working memory are associated with acetylcholine- and dopamine-relevant genes. The efficiency of these 2 transmitter systems declines substantially during healthy aging. These declines, in part, contribute to age-related deficits in attention and working memory functions. We report novel data showing an effect of dopamine COMT gene on spatial updating processes in older but not in younger adults, indicating potential magnification of genetic effects in old age.

Among other mechanisms, behavioral and cognitive development entail, on the one hand, contextual scaffolding and, on the other hand, neuromodulation of adaptive neurocognitive representations across the life span. Key brain networks underlying cognition, emotion, and motivation are innervated by major transmitter systems (e.g., the catecholamines and acetylcholine). Thus, the maturation and senescence of neurotransmitter systems have direct implications for life span development. Recent progress in molecular genetics has opened new avenues for investigating neuromodulation of behavioral and cognitive development. This special section features 6 selected reviews of recent cognitive genetic evidence on the roles of dopamine and other transmitters in different domains of behavioral and cognitive development, ranging from temperament, executive control, and working memory to motivation and goal-directed behavior in different life periods.

Older adults often experience hearing difficulties in multitalker situations. Attentional control of auditory perception is crucial in situations where a plethora of auditory inputs compete for further processing. We combined an intensity-modulated dichotic listening paradigm with attentional manipulations to study adult age differences in the interplay between perceptual saliency and attentional control of auditory processing. When confronted with two competing sources of verbal auditory input, older adults modulated their attention less flexibly and were more driven by perceptual saliency than younger adults. These findings suggest that aging severely impairs the attentional regulation of auditory perception.

In this paper, we review the current literature to highlight relations between age-associated declines in dopaminergic and serotonergic neuromodulation and adult age differences in adaptive goal-directed behavior. Specifically, we focus on evidence suggesting that deficits in neuromodulation contribute to older adults' behavioral disadvantages in learning and decision making. These deficits are particularly pronounced when reward information is uncertain or the task context requires flexible adaptations to changing stimulus-reward contingencies. Moreover, emerging evidence points to age-related differences in the sensitivity to rewarding and aversive outcomes during learning and decision making if the acquisition of behavior critically depends on outcome processing. These age-related asymmetries in outcome valuation may be explained by age differences in the interplay of dopaminergic and serotonergic neuromodulation. This hypothesis is based on recent neurocomputational and psychopharmacological approaches, which suggest that dopamine and serotonin serve opponent roles in regulating the balance between approach behavior and inhibitory control. Studying adaptive regulation of behavior across the adult life span may shed new light on how the aging brain changes functionally in response to its diminishing resources., (© 2011 New York Academy of Sciences.)

Individual differences in working memory (WM) performance have rarely been related to individual differences in the functional responsivity of the WM brain network. By neglecting person-to-person variation, comparisons of network activity between younger and older adults using functional imaging techniques often confound differences in activity with age trends in WM performance. Using functional magnetic resonance imaging, we investigated the relations among WM performance, neural activity in the WM network, and adult age using a parametric letter n-back task in 30 younger adults (21-31 years) and 30 older adults (60-71 years). Individual differences in the WM network's responsivity to increasing task difficulty were related to WM performance, with a more responsive BOLD signal predicting greater WM proficiency. Furthermore, individuals with higher WM performance showed greater change in connectivity between left dorsolateral prefrontal cortex and left premotor cortex across load. We conclude that a more responsive WM network contributes to higher WM performance, regardless of adult age. Our results support the notion that individual differences in WM performance are important to consider when studying the WM network, particularly in age-comparative studies.

Intraindividual trial-to-trial reaction time (RT) variability is commonly found to be higher in clinical populations or life periods that are associated with impaired cognition. In the present study, higher within-person trial-to-trial RT variability in a perceptual speed task is related to more forgetting and dedifferentiation of memory functions in older adults (aged 60-71 years). More specifically, our study showed that individuals in a high-variability group (n=175) forgot more memory scenes over a 1-week retention interval than individuals in the low-variability group (n=174). In contrast, slower RT speed was associated with poorer episodic memory in general, but unrelated to the amount of forgetting. Moreover, results from multiple group latent factor analyses showed that episodic memory and working memory functions were more highly correlated in the high-variability (r=.63) than in the low-variability (r=.25) group. Given that deficits in dopamine (DA) modulation may underlie increases in RT variability, the present findings are in line with (i) recent animal studies implicating DA in long-term episodic memory consolidation and (ii) neurocomputational work linking DA modulation of performance variability to dedifferentiation of cognitive functions in old age., (Copyright © 2011 Elsevier Ltd. All rights reserved.)

By recording the feedback-related negativity (FRN) in response to gains and losses, we investigated the contribution of outcome monitoring mechanisms to age-associated differences in probabilistic reinforcement learning. Specifically, we assessed the difference of the monitoring reactions to gains and losses to investigate the monitoring of outcomes according to task-specific goals across the life span. The FRN and the behavioral indicators of learning were measured in a sample of 44 children, 45 adolescents, 46 younger adults, and 44 older adults. The amplitude of the FRN after gains and losses was found to decrease monotonically from childhood to old age. Furthermore, relative to adolescents and younger adults, both children and older adults (a) showed smaller differences between the FRN after losses and the FRN after gains, indicating a less differentiated classification of outcomes on the basis of task-specific goals; (b) needed more trials to learn from choice outcomes, particularly when differences in reward likelihood between the choices were small; and (c) learned less from gains than from losses. We suggest that the relatively greater loss sensitivity among children and older adults may reflect ontogenetic changes in dopaminergic neuromodulation.

The ability to attend to multiple objects that move in the visual field is important for many aspects of daily functioning. The attentional capacity for such dynamic tracking, however, is highly limited and undergoes age-related decline. Several aspects of the tracking process can influence performance. Here, we investigated effects of feature-based interference from distractor objects that appear in unattended regions of the visual field with a hemifield-tracking task. Younger and older participants performed an attentional tracking task in one hemifield while distractor objects were concurrently presented in the unattended hemifield. Feature similarity between objects in the attended and unattended hemifields as well as motion speed and the number of to-be-tracked objects were parametrically manipulated. The results show that increasing feature overlap leads to greater interference from the unattended visual field. This effect of feature-based interference was only present in the slow speed condition, indicating that the interference is mainly modulated by perceptual demands. High-performing older adults showed a similar interference effect as younger adults, whereas low-performing adults showed poor tracking performance overall.

The brain-derived neurotrophic factor (BDNF) plays an important role in activity-dependent synaptic plasticity, which underlies learning and memory. In a sample of 948 younger and older adults, we investigated whether a common Val66Met missense polymorphism (rs6265) in the BDNF gene affects the serial position curve--a fundamental phenomenon of associative memory identified by Hermann Ebbinghaus more than a century ago. We found a BDNF polymorphism effect for backward recall in older adults only, with Met-allele carriers (i.e., individuals with reduced BDNF signaling) recalling fewer items than Val homozygotes. This effect was specific to the primacy and middle portions of the serial position curve, where intralist interference and associative demands are especially high. The poorer performance of older Met-allele carriers reflected transposition errors, whereas no genetic effect was found for omissions. These findings indicate that effects of the BDNF polymorphism on episodic memory are most likely to be observed when the associative and executive demands are high. Furthermore, the findings are in line with the hypothesis that the magnitude of genetic effects on cognition is greater when brain resources are reduced, as is the case in old age.

We recorded Electroencephalograms (EEGs) during a cued Continuous Performance Task (CPT) to investigate lifespan differences in the efficiency of response conflict processing under conditions that put high demands on the ability to suppress a prepotent response. Previous evidence indicates that children and adolescents commit more errors under such conditions than younger adults, whereas older adults are disproportionately slow in responding. We measured event-related potentials (ERPs) in a sample of 45 children, 44 adolescents, 46 younger adults, and 47 older adults to investigate response conflict monitoring (Nogo-N2), cue utilization (Cue-P3), response anticipation (contingent negative variation, CNV), and response suppression (Nogo-P3). In comparison to adolescents and adults, children showed larger ERPs associated with cue utilization. At the same time, children committed more errors and their ERPs reflecting response anticipation and response suppression were smaller and uncorrelated. In contrast, older adults showed ERP indices of attentional distraction (P3a elicited by the infrequent Non-Cue stimuli), reduced conflict monitoring signals (Nogo-N2), and took more time to respond than the other age groups. The present findings reveal marked lifespan differences in processes related to response conflict monitoring. In middle childhood, the readiness to utilize cues for guiding actions is not yet fully matched by the ability to suppress prepotent responses, leading to a relatively large number of commission errors. In older adults, higher indices of attentional distraction as well as lower conflict monitoring signals were observed. This might reflect a dampened build-up of response tendencies, thereby leading to slower responding and relatively low error rates., (Copyright 2010 Elsevier Ltd. All rights reserved.)

The structural and functional brain circuitries supporting episodic memory undergo profound reorganization in childhood and old age. We propose a two-component framework that combines and integrates evidence from child development and aging. It posits that episodic memory builds on two interacting components: (a) the strategic component, which refers to memory control operations, and (b) the associative component, which refers to mechanisms that bind different features of a memory episode into a compound representation. We hypothesize that: (a) children's difficulties in episodic memory primarily originate from low levels of strategic operations, and reflect the protracted development of the prefrontal cortex (PFC); (b) deficits in episodic memory performance among older adults originate from impairments in both strategic and associative components, reflecting senescent changes in the PFC and the medio-temporal lobes (MTL). Initial behavioral and neural evidence is consistent with both hypotheses. The two-component framework highlights the specificities of episodic memory in different age periods, helps to identify and dissociate its components, and contributes to understanding the interplay among maturation, learning, and senescence., (Copyright (c) 2009 Elsevier Ltd. All rights reserved.)

Molecular-imaging studies of dopaminergic neurotransmission measure biomarkers of dopamine (DA), such as the DA transporter and D(1) and D(2) receptor densities in the living brain. These studies indicate that individual differences in DA functions are linked to cognitive performance irrespective of age, and serve as powerful mediators of age-related decline in executive functioning, episodic memory, and perceptual speed. This focused review targets several recent findings pertaining to these relationships. Specifically, we discuss novel evidence concerning (a) the role of DA in within-person cognitive variability; (b) age-related differences in DA release during cognitive processing; (c) DA release following cognitive training in younger and older adults; and (d) the relationship between DA and task-induced functional brain activity. Based on these lines of empirical inquiry, we outline a series of avenues for future research on aging, DA, and cognition., (Copyright 2009 Elsevier Ltd. All rights reserved.)

Economic decision making is a complex process of integrating and comparing various aspects of economically relevant choice options. Neuroeconomics has made important progress in grounding these aspects of decision making in neural systems and the neurotransmitters therein. The dopaminergic and serotoninergic brain systems have been identified as key neurotransmitter systems involved in economic behavior. Both are known to be prone to significant changes during the adult lifespan. Similarly, economic behavior undergoes significant age-related changes over the course of the adult lifespan. Here we propose a triadic relationship between (a) economic decision making, (b) dopaminergic and serotonergic neuromodulation, and (c) aging. In this review, we describe the different relationships around this triad in detail and summarize current evidence that supports them. Based on the reviewed evidence, we propose new research agendas that take the entire triad into account., (Copyright 2009 Elsevier Ltd. All rights reserved.)

Using a population-based sample of 263 individuals ranging from 6 to 89 years of age, we investigated the gains and losses in the abilities to (a) use exogenous cues to shift attention covertly and (b) ignore conflicting information across the life span. The participants' ability to shift visual attention was tested by a typical Posner-type orienting task with valid and invalid peripheral cues. To tap conflict resolution, we asked participants to perform a color version of the Eriksen-type flanker task. The observed cross-sectional age differences in our data indicate that the ability to deal with conflicting information and the ability to covertly orient attention show different cross-sectional age gradients during childhood and that only conflict resolution mechanisms show a marked negative age difference in old age. Moreover, the data suggest that although the overall performance of the participants can, in part, be accounted for by individual differences in information processing speed, performance in the orienting and conflict task depends on factors related to the specific development of the two attentional systems in question.

Working memory (WM) shows pronounced age-related decline. Functional magnetic resonance imaging (fMRI) studies have revealed age differences in task-related brain activation. Evidence based primarily on episodic memory studies suggests that brain activation patterns can be modulated by task difficulty in both younger and older adults. In most fMRI aging studies on WM, however, performance level has not been considered, so that age differences in activation patterns are confounded with age differences in performance level. Here, we address this issue by comparing younger and older low and high performers in an event-related fMRI study. Thirty younger (20-30 years) and 30 older (60-70 years) healthy adults were tested with a spatial WM task with three load levels. A region-of-interest analysis revealed marked differences in the activation patterns between high and low performers in both age groups. Critically, among the older adults, a more "youth-like" load-dependent modulation of the blood oxygen level-dependent signal was associated with higher levels of spatial WM performance. These findings underscore the need of taking performance level into account when studying changes in functional brain activation patterns from early to late adulthood.

Age gradient of the mechanism of stimulus-response conflict cost was investigated in a population-based representative sample of 291 individuals, covering the age range from 6 to 89 years. Stimulus-response conflict cost, indicated by the amount of additional processing time required when there is a conflict between stimulus and response options, follows a U-shaped function across the lifespan. Lifespan age gradient of conflict cost parallels closely those of processing fluctuation and fluid intelligence. Individuals at both ends of the lifespan displayed a greater amount of processing fluctuation and at the same time a larger amount of conflict cost and a lower level of fluid intelligence. After controlling for chronological age and baseline processing speed, conflict cost continues to correlate significantly with fluid intelligence in adulthood and old age and with processing fluctuation in old age. The relation between processing fluctuation and conflict cost in old age lends further support for the neuromodulation of neuronal noise theory of cognitive aging as well as for theories of dopaminergic modulation of conflict monitoring.

Objective: To investigate lifespan age differences in neuronal mechanisms of visual coding in the context of perceptual discrimination., Methods: We recorded EEG from 17 children (10-12 years), 16 younger adults (20-26 years), and 17 older adults (70-76 years) during a simple choice-reaction task requiring discrimination of squares and circles of different sizes. We examined age-group differences in the effect of stimulus size on early ERP components, evoked gamma-band power, and inter-trial phase-stability in the gamma band as assessed by the phase-locking index (PLI)., Results: In the absence of age differences in discrimination accuracy, we observed reliable age differences in patterns of ERP, evoked gamma power, and PLI. P1 and N1 peak amplitudes were larger and the peak latencies longer in children than in adults. Children also showed lower levels of evoked power and PLI than adults. Older adults showed smaller increments in evoked power with increasing stimulus size than younger adults, but similar amounts of phase locking for small- and medium-sized stimuli as younger adults., Conclusions: The relative importance of different coding mechanisms in early visual areas changes from childhood to old age. Due to synaptic overproduction and immature myelination, the visual system of children is less entrained by incoming information, resulting in less synchronized neuronal responses. Adults primarily rely on sparse representations formed through experience-dependent temporally synchronized neuronal interactions. In old age, senescent decline in neuronal density and neurotransmitter availability further increase the reliance on temporally synchronized processing., Significance: Findings from this study defy the notion that sensory aging consists in a reversal of sensory development in childhood, and point to a high degree of age specificity in mechanisms of visual coding.

Working memory (WM) declines prominently during normal aging. The mechanisms underlying this decline are not fully understood. The authors analyzed performance on 2 versions of a 2-back spatial WM task to assess younger and older adults' responses to lures (i.e., nontarget items that match an item earlier in the sequence but not at the current target lag). Results demonstrate lure interference effects that are particularly pronounced among older adults. At the same time, however, older adults showed facilitation for targets. Taken together, these findings suggest that the contribution of familiarity signals to WM performance increases during normal aging.

We investigated lifespan differences of confidence calibration in episodic memory, particularly the susceptibility to high-confidence errors within samples of children, teenagers, younger adults, and older adults. Using an associative recognition memory paradigm, we drew a direct link between older adults' associative deficit and high-confidence errors. We predicted that only older adults would show high-confidence error even though their memory performance was at a similar level to that of children. Participants of all ages showed higher confidence following correct responses compared to incorrect responses, demonstrating the ability to calibrate subjective confidence in relation to memory accuracy. However, older adults were disproportionately more likely to indicate high confidence following erroneously remembered word pairs than participants of the other three age groups. Results are discussed in relation to the misrecollection account of high-confidence errors and ageing-related decline in hippocampus-dependent episodic memory functions.

Previous studies have indicated that older adults have a special deficit in the encoding and retrieval of associations. The current study assessed this deficit using ecologically valid name-face pairs. In two experiments, younger and older participants learned a series of name-face pairs under intentional and incidental learning instructions, respectively, and were then tested for their recognition of the faces, the names, and the associations between the names and faces. Under incidental encoding conditions older adults' performance was uniformly lower than younger adults in all three tests, indicating age-related impairments in episodic memory representations. An age-related deficit specific to associations was found under intentional but not under incidental learning conditions, highlighting the importance of strategic associative processes and their decline in older adults. Separate analyses of hits and false alarms indicate that older adults' associative deficit originated from high false alarm rates in the associative test. Older adults' high false alarm rates potentially reflect their reduced ability to recollect the study-phase name-face pairs in the presence of intact familiarity with individual names and faces.