Aging Enhances Neural Activity in Auditory, Visual, and Somatosensory Cortices: The Common Cause Revisited.

In humans, age-related declines in vision, hearing, and touch coincide with changes in amplitude and latency of sensory-evoked potentials. These age-related differences in neural activity may be related to a common deterioration of supra-modal brain areas (e.g., PFC) that mediate activity in sensory cortices or reflect specific sensorineural impairments that may differ between sensory modalities. To distinguish between these two possibilities, we measured neuroelectric brain activity while 37 young adults (18-30 years, 18 males) and 35 older adults (60-88 years, 20 males) were presented with a rapid randomized sequence of lateralized auditory, visual, and somatosensory stimuli. Within each sensory domain, we compared amplitudes and latencies of sensory-evoked responses, source activity, and functional connectivity (via phase-locking value) between groups. We found that older adults' early sensory-evoked responses were greater in amplitude than those of young adults in all three modalities, which coincided with enhanced source activity in auditory, visual, and somatosensory cortices. Older adults also showed stronger neural synchrony than young adults between superior prefrontal and sensory cortices; and in older adults, the degree of phase synchrony was positively correlated with the magnitude of source activity in sensory areas. Critically, older adults who showed enhanced neural activity in one sensory domain also showed enhanced activity in other modalities. Together, these findings support the common cause hypothesis of aging and highlight the role of prefrontal regions in exerting top-down control over sensory cortices.SIGNIFICANCE STATEMENT A prominent theory of aging posits that age-related declines in sensory processing across domains are related to a single common neurobiological mechanism. However, the neural evidence supporting this common cause hypothesis has remained elusive. Our study revealed robust age-related changes in three sensory domains across a range of neural metrics. Importantly, older adults who showed increased neural activity within one sensory domain also showed enhanced neural activity in the other two sensory modalities. No such relation among activity in sensory cortices was observed in young adults. Age-related increases in neural activity in sensory cortices coincided with enhanced neural synchrony between the PFC and sensory cortices, underlining the importance of the PFC in regulating sensory processing.

Holding On to the Past: Older Adults Show Lingering Neural Activation of No-Longer-Relevant Items in Working Memory.

Goal-relevant information can be maintained in working memory over a brief delay interval to guide an upcoming decision. There is also evidence suggesting the existence of a complementary process: namely, the ability to suppress information that is no longer relevant to ongoing task goals. Moreover, this ability to suppress or inhibit irrelevant information appears to decline with age. In this study, we compared younger and older adults undergoing fMRI on a working memory task designed to address whether the modulation of neural representations of relevant and no-longer-relevant items during a delay interval is related to age and overall task performance. Following from the theoretical predictions of the inhibitory deficit hypothesis of aging, we hypothesized that older adults would show higher activation of no-longer-relevant items during a retention delay compared to young adults and that higher activation of these no-longer-relevant items would predict worse recognition memory accuracy for relevant items. Our results support this prediction and more generally demonstrate the importance of goal-driven modulation of neural activity in successful working memory maintenance. Furthermore, we showed that the largest age differences in the regulation of category-specific pattern activity during working memory maintenance were seen throughout the medial temporal lobe and prominently in the hippocampus, further establishing the importance of "long-term memory" retrieval mechanisms in the context of high-load working memory tasks that place large demands on attentional selection mechanisms.

Inhibitory Control Deficits in Individuals with Amnestic Mild Cognitive Impairment: a Meta-Analysis.

Amnestic mild cognitive impairment (aMCI) is a prodromal stage of Alzheimer's disease that is characterized by impairments in episodic memory. Recent evidence has shown that inhibitory control is also impaired in aMCI. The aim of the present meta-analysis was to quantify inhibitory control ability in individuals with aMCI by examining performance across a range of well-defined inhibition paradigms that tapped into one of three inhibitory control subtypes (i) interference control (e.g., Stroop task), (ii) response inhibition (e.g., Go/Nogo task), or (iii) inhibition of cognitive sets (Wisconsin Card Sort Task). Reference databases (PsychINFO, PubMed, and Web of Science) were searched for studies comparing individuals with aMCI to healthy controls on behavioural measures of inhibition. Across 70 effect sizes involving 2184 adults with aMCI and 3049 controls, overall inhibition deficits of moderate magnitude (g = -0.73) were found among individuals with aMCI. Inhibition deficits were moderate in size regardless of inhibitory control subtype: interference control (g = -0.74), response inhibition (g = -0.71), inhibition of cognitive sets (g = -0.76). Subgroup analyses revealed that Stroop outcome measure (reaction time vs. accuracy) and recruitment source (clinical vs. community) moderated interference control deficits. Together these findings support a generalized inhibition deficit in aMCI, and suggest that inhibition tasks should be included routinely in neuropsychological test batteries to provide a more comprehensive overview of executive dysfunction in aMCI.

Spontaneous Distractor Reactivation With Age: Evidence for Bound Target-Distractor Representations in Memory.

Reduced attentional control with age is associated with the processing and maintenance of task-irrelevant information in memory. Yet the nature of these memory representations remains unclear. We present evidence that, relative to younger adults (n = 48), older adults (n = 48) both (a) store simultaneously presented target and irrelevant information as rich, bound memory representations and (b) spontaneously reactivate irrelevant information when presented with previously associated targets. In a three-stage implicit reactivation paradigm, re-presenting a target picture that was previously paired with a distractor word spontaneously reactivated the previously associated word, making it become more accessible than an unreactivated distractor word in a subsequent implicit memory task. The accessibility of reactivated words, indexed by priming, was also greater than the degree of distractor priming shown by older adults in a control condition (n = 48). Thus, reduced attentional control influences the processing and representation of incoming information.

An Incidental Learning Method to Improve Face-Name Memory in Older Adults With Amnestic Mild Cognitive Impairment.

OBJECTIVE: Forgetting names is a common memory concern for people with amnestic mild cognitive impairment (aMCI) and is related to explicit memory deficits and pathological changes in the medial temporal lobes at the early stages of Alzheimer's disease (AD). In the current experiment, we tested a unique method to improve memory for face-name associations in people with aMCI involving incidental rehearsal of face-name pairs. METHOD: Older adults with aMCI and age- and education-matched controls learned 24 face-name pairs and were tested via immediate cued recall with faces as cues for associated names. During a 25- to 30-min retention interval, 10 of the face-name pairs reappeared as a quarter of the items on a seemingly unrelated 1-back task on faces, with the superimposed names irrelevant to the task. After the delay, surprise delayed cued recall and forced-choice associative recognition tests were administered for the face-name pairs. RESULTS: Both groups showed reduced forgetting of the names that repeated as distraction and enhanced recollection of these pairs. CONCLUSIONS: The results demonstrate that passive methods to prompt automatic retrieval of associations may hold promise as interventions for people with early signs of AD.

Neural Correlates of Enhanced Memory for Meaningful Associations with Age.

Evidence suggests that age differences in associative memory are attenuated for associations that are consistent with prior knowledge. Such knowledge structures have traditionally been associated with the default network (DN), which also shows reduced modulation with age. In the present study, we investigated whether DN activity and connectivity patterns could account for this age-related effect. Younger and older adults underwent functional magnetic resonance imaging as they learned realistic and unrealistic prices of common grocery items. Both groups showed greater activity in the DN during the encoding of realistic, relative to unrealistic, prices. Moreover, DN activity at encoding and retrieval and its connectivity with an attention control network at encoding were associated with enhanced memory for realistic prices. Finally, older adults showed overactivation of control regions during retrieval of realistic prices relative to younger adults. Our findings suggest that DN activity and connectivity patterns (traditionally viewed as indicators of cognitive failure with age), and additional recruitment of control regions, might underlie older adults' enhanced memory for meaningful associations.

Do young adults show conceptual knowledge of previous distractors?

Using implicit tests, older adults have been found to retain conceptual knowledge of previously seen task-irrelevant information. While younger adults typically do not show the same effect, evidence from one study [Gopie, N., Craik, F. I. M., & Hasher, L. (2011). A double dissociation of implicit and explicit memory in younger and older adults. Psychological Science, 22, 634-640. doi: 10.1177/0956797611403321 ] suggests otherwise. In that study, young adults showed greater explicit than implicit memory for previous distractors on a word fragment completion task. This was interpreted as evidence for maintaining access to previous conceptual knowledge of the distractors. Here, we report two failures to replicate that original finding, followed by a third study designed to test directly whether young adults use conceptual-level information that was previously irrelevant. Our findings agree with others that young adults show weak to no evidence of conceptual knowledge of previously irrelevant information.

Cultural differences in distraction processing: influence of context at retrieval.

Cultural differences in information processing affect perceptual judgment, attention, and memory. We investigated whether cultural differences in processing patterns, specifically East Asian participants' tendency to encode holistically, compared to Western tendencies to process analytically, affect performance on an implicit memory test. First, participants completed a 1-back task on pictures with superimposed distracting words. After a delay filled with a computerised Corsi block task, they performed a word fragment task in which some fragments could be completed with the distracting words from the 1-back task. Critically, fragments were presented with the same pictures as previously seen (matched condition), with no pictures (control condition), or with pictures from other trials on the 1-back task (mismatched condition). Non-Asian Canadian participants showed virtually no priming for distraction, independent of the reinstatement of encoding context. East Asian Canadian participants showed superior priming for fragments that had been paired with their original pictures. They did not show evidence of a detriment for the mismatched, relative to control, condition.

Task-linked Diurnal Brain Network Reorganization in Older Adults: A Graph Theoretical Approach.

Testing older adults in the morning generally improves behavioral performance relative to afternoon testing. Morning testing is also associated with brain activity similar to that of young adults. Here, we used graph theory to explore how time of day (TOD) affects the organization of brain networks in older adults across rest and task states. We used nodes from the automated anatomical labeling atlas to construct participant-specific correlation matrices of fMRI data obtained during 1-back tasks with interference and rest. We computed pairwise group differences for key graph metrics, including small-worldness and modularity. We found that older adults tested in the morning and young adults did not differ on any graph metric. Both of these groups differed from older adults tested in the afternoon during the tasks-but not rest. Specifically, the latter group had lower modularity and small-worldness (indices of more efficient network organization). Across all groups, higher modularity and small-worldness strongly correlated with reduced distractibility on an implicit priming task. Increasingly, TOD is seen as important for interpreting and reproducing neuroimaging results. Our study emphasizes how TOD affects brain network organization and executive control in older adults.

On the preservation of vigilant attention to semantic information in healthy aging.

Despite decades of research on younger adults, little is known about the way in which vigilant attention is affected by healthy aging, and the small body of work that does exist has yielded mixed findings. Prior examinations of aging and vigilant attention have focused almost exclusively on sensory/perceptual tasks despite the fact that many real-world vigilance tasks are semantic in nature and it has been shown that older adults exhibit memory and attention deficits in semantic tasks in other domains. Here, we present the first empirical investigation of vigilant attention to verbal stimuli in healthy normal aging. In Experiment 1 we find that older adults are just as able as younger adults to identify critical targets defined by category membership (both overall and over time). In Experiment 2, we increase the difficulty of the task by changing the target category from one block to the next, but again find no age-group effects in accuracy. Response time data, however, show that older adults respond more slowly and subjective ratings indicate that older adults experience higher workload and arousal compared to their younger counterparts. The practical as well as theoretical implications of these findings are discussed.

Optimal testing time for suppression of competitors during interference resolution.

Interference between competing memory traces is a common cause of memory failure. Recent research has demonstrated a suppression mechanism that operates at retrieval to resolve interference. Using an adaptation of the suppression paradigm in Healey, Ngo, and Hasher [(2014). Below-baseline suppression of competitors during interference resolution by younger but not older adults. Psychological Science, 25(1), 145-151. doi: 10.1177/0956797613501169 ], we tested whether the ability to suppress competing memory traces varies with the synchrony between optimal arousal period and time of testing. We replicate the below-baseline suppression effect for young adults tested at optimal times of day, and present novel evidence that they do not show competitor suppression during non-optimal times of day. In fact, competitors are actually strengthened at non-optimal times. Our results suggest that the ability to resolve interference by suppression varies with circadian arousal.

Age differences in the neural correlates of distraction regulation: A network interaction approach.

Older adults show decrements in the ability to ignore or suppress distraction relative to younger adults. However, age differences in the neural correlates of distraction control and the role of large-scale network interaction in regulating distractors are scarcely examined. In the current study, we investigated age differences in how the anticorrelation between an externally oriented dorsal attention network (DAN) and an internally focused default mode network (DMN) is related to inhibiting distractors presented during a 1-back working memory task. For both young and older adults, the extent of DAN-DMN anticorrelation predicted reduced distractibility. Activation in a common set of frontal and insular control regions during the task was, however, associated with opposite patterns of network interaction and distractibility in the age groups. For older adults, recruitment of these regions was associated with greater DAN-DMN anticorrelation and less distractibility (better performance). For younger adults, it was associated with decreased DAN-DMN anticorrelation and more distractibility (worse performance). Our findings demonstrate the age-dependent relationship between DAN-DMN interaction patterns and engagement of control regions during an externally oriented distraction control task. This suggests that engagement of those regions may play a compensatory role for older adults but may be indicative of less efficient neural control mechanisms in younger adults.

Age-related deficits in inhibition in figure-ground assignment.

We assessed age differences in the ability to resolve competition for figural status in stationary displays using small, enclosed, symmetrical silhouettes that participants classified as depicting "novel" or "familiar" shapes. The silhouettes were biased such that the inside was perceived as the shaped figure, and the outside was perceived as a shapeless ground. The critical manipulation was whether a portion of a meaningful object was suggested on the outside of the border of some of the novel silhouettes but not others (M+Ground and M-Ground novel silhouettes, respectively). This manipulation was intended to induce greater inhibitory competition for figural status from the groundside in M+Ground silhouettes than M-Ground silhouettes. In previous studies, young adults classified M+Ground silhouettes as "novel" faster than M-Ground silhouettes (Trujillo, Allen, Schnyer, & Peterson, 2010), suggesting that young adults may recruit more inhibition to resolve figure-ground when there is more competition. We replicated this effect with young adults in the present study, but older adults showed the opposite pattern and were less accurate in classifying M+Ground than M-Ground silhouettes. These results extend the evidence for inhibitory deficits in older adults to figure assignment in stationary displays. The (M+Ground - M-Ground) RT differences were evident in observers' longest responses, consistent with the hypothesis that inhibitory deficits are evident when the need for inhibition is substantial.

Conceptual processing of distractors by older but not younger adults.

Evidence from perceptually based implicit memory tasks demonstrates greater priming from distracting information among older compared with younger adults. We examined whether older adults also show greater conceptually based implicit priming from distracting information. We measured priming using a general-knowledge test that was preceded by an incidental-encoding task (a color-naming Stroop task in one experiment and a 1-back task involving pictures with irrelevant words superimposed in a second experiment). Younger adults showed no priming from the distracting information in either experiment, whereas older adults showed reliable priming in both experiments. Thus, unlike young adults, older adults process irrelevant information conceptually and then can use that information to boost their performance on a subsequent task.

Below-baseline suppression of competitors during interference resolution by younger but not older adults.

Resolving interference from competing memories is a critical factor in efficient memory retrieval, and several accounts of cognitive aging suggest that difficulty resolving interference may underlie memory deficits such as those seen in the elderly. Although many researchers have suggested that the ability to suppress competitors is a key factor in resolving interference, the evidence supporting this claim has been the subject of debate. Here, we present a new paradigm and results demonstrating that for younger adults, a single retrieval attempt is sufficient to suppress competitors to below-baseline levels of accessibility even though the competitors are never explicitly presented. The extent to which individual younger adults suppressed competitors predicted their performance on a memory span task. In a second experiment, older adults showed no evidence of suppression, which supports the theory that older adults' memory deficits are related to impaired suppression.

Visual dominance and multisensory integration changes with age.

Objects comprise of visual and auditory signatures that arrive through distinct sensory channels. Exposure to cross-modal events sets up expectations about what a given object most likely "sounds" like, and vice versa, thereby facilitating detection and recognition. Whereas episodic and working memory functions decline with age, the extent to which multisensory integration processes change with age remains an open question. In the present study, we examined whether multisensory integration processes play a compensatory role in normal aging. Magnetoencephalography recordings of semantically-related cross-modal and unimodal auditory and visual stimuli captured the spatiotemporal dynamics of multisensory responses in young and older adults. Whereas sensory-specific regions showed increased activity in response to cross-modal compared to unimodal stimuli 100 ms after stimulus onset in both age groups, posterior parietal and medial prefrontal regions responded preferentially to cross-modal stimuli between 150 and 300 ms in the older group only. Additionally, faster detection of cross-modal stimuli correlated with increased activity in inferior parietal and medial prefrontal regions 100 ms after stimulus onset in older compared to younger adults. Age-related differences in visual dominance were also observed with older adults exhibiting significantly larger multisensory facilitation effects relative to the auditory modality. Using structural equation modeling, we showed that age-related increases in parietal and medial prefrontal source activity predicted faster detection of cross-modal stimuli. Furthermore, the relationship between performance and source activity was mediated by age-related reductions in gray matter volume in those regions. Thus, we propose that multisensory integration processes change with age such that posterior parietal and medial prefrontal activity underlies the integrated response in older adults.

Distraction can reduce age-related forgetting.

In three experiments, we assessed whether older adults' generally greater tendency to process distracting information can be used to minimize widely reported age-related differences in forgetting. Younger and older adults studied and recalled a list of words on an initial test and again on a surprise test after a 15-min delay. In the middle (Experiments 1a and 2) or at the end (Experiment 3) of the delay, participants completed a 1-back task in which half of the studied words appeared as distractors. Across all experiments, older adults reliably forgot unrepeated words; however, older adults rarely or never forgot the words that had appeared as distractors, whereas younger adults forgot words in both categories. Exposure to distraction may serve as a rehearsal episode for older adults, and thus as a method by which general distractibility may be co-opted to boost memory.

For Whom (and When) the Time Bell Tolls: Chronotypes and the Synchrony Effect.

Circadian rhythms are powerful timekeepers that drive physiological and intellectual functioning throughout the day. These rhythms vary across individuals, with morning chronotypes rising and peaking early in the day and evening chronotypes showing a later rise in arousal, with peaks in the afternoon or evening. Chronotype also varies with age from childhood to adolescence to old age. As a result of these differences, the time of day at which people are best at attending, learning, solving analytical problems, making complex decisions, and even behaving ethically varies. Across studies of attention and memory and a range of allied areas, including academic achievement, judgment and decision-making, and neuropsychological assessment, optimal outcomes are found when performance times align with peaks in circadian arousal, a finding known as the synchrony effect. The benefits of performing in synchrony with one's chronotype (and the costs of not doing so) are most robust for individuals with strong morning or evening chronotypes and for tasks that require effortful, analytical processing or the suppression of distracting information. Failure to take the synchrony effect into consideration may be a factor in issues ranging from replication difficulties to school timing to assessing intellectual disabilities and apparent cognitive decline in aging.

Implicit processes enhance cognitive abilities in mild cognitive impairment.

Previous work has shown that older adults with typical age-related memory changes (i.e., without cognitive impairment) pick up irrelevant information implicitly, and unknowingly use that information when it becomes relevant to a later task. Here, we address the possibility that implicit processes play a similarly beneficial role in the cognitive abilities of individuals with amnestic mild cognitive impairment (aMCI). Twenty-two individuals with aMCI and 22 matched controls participated in a picture judgment task while instructed to ignore distractions in the form of word/non-word letter strings. Memory for the distracting words was later tested with a word-fragment completion task. Both groups showed a priming effect, that is, they were significantly more likely to solve fragments of previously presented than non-presented words. However, the aMCI group had significantly higher scores than the older adults without cognitive impairment, t(42) = 2.16, p < .05, Cohen's d = 0.67. Our findings suggest that individuals with aMCI can enhance their performance on an explicit cognitive task, in this case, word-fragment completion, if previously exposed to the relevant information implicitly, opening up possible interventions aimed at this population.

Circadian rhythms in executive function during the transition to adolescence: the effect of synchrony between chronotype and time of day.

To explore the influence of circadian rhythms on executive function during early adolescence, we administered a battery of executive function measures (including a Go-Nogo task, the Iowa Gambling Task, a Self-ordered Pointing task, and an Intra/Extradimensional Shift task) to Morning-preference and Evening-preference participants (N = 80) between the ages of 11 and 14 years who were tested in the morning or afternoon. Significant Chronotype × Time of Day interactions (controlling for amount of sleep the previous night) revealed that adolescents tested at their optimal times of day performed better than those tested at their nonoptimal times. Implications for our understanding of physiological arousal, sleep, and executive function during adolescence are discussed.