The implications of cortical recruitment and brain morphology for individual differences in inhibitory function in aging humans

The authors assessed individual differences in cortical recruitment, brain morphology, and inhibitory task performance. Similar to previous studies, older adults tended toward bilateral activity during task performance more than younger adults. However, better performing older adults showed less bilateral activity than poorer performers, contrary to the idea that additional activity is universally compensatory. A review of the results and of extant literature suggests that compensatory activity in prefrontal cortex may only be effective if the additional cortical processors brought to bear on the task can play a complementary role in task performance. Morphological analyses revealed that frontal white matter tracts differed as a function of performance in older adults, suggesting that hemispheric connectivity might impact both patterns of recruitment and cognitive performance.     

Event-related fMRI of category learning: differences in classification and feedback networks

Eighteen healthy young adults underwent event-related (ER) functional magnetic resonance imaging (fMRI) of the brain while performing a visual category learning task. The specific category learning task required subjects to extract the rules that guide classification of quasi-random patterns of dots into categories. Following each classification choice, visual feedback was presented. The average hemodynamic response was calculated across the eighteen subjects to identify the separate networks associated with both classification and feedback. Random-effects analyses identified the different networks implicated during the classification and feedback phases of each trial. The regions included prefrontal cortex, frontal eye fields, supplementary motor and eye fields, thalamus, caudate, superior and inferior parietal lobules, and areas within visual cortex. The differences between classification and feedback were identified as (i) overall higher volumes and signal intensities during classification as compared to feedback, (ii) involvement of the thalamus and superior parietal regions during the classification phase of each trial, and (iii) differential involvement of the caudate head during feedback. The effects of learning were then evaluated for both classification and feedback. Early in learning, subjects showed increased activation in the hippocampal regions during classification and activation in the heads of the caudate nuclei during the corresponding feedback phases. The findings suggest that early stages of prototype-distortion learning are characterized by networks previously associated with strategies of explicit memory and hypothesis testing. However as learning progresses the networks change. This finding suggests that the cognitive strategies also change during prototype-distortion learning.     

Sex-related developmental differences in the lateralized activation of the prefrontal cortex and amygdala during perception of facial affect

The lateralization of cognitive abilities is influenced by a number of factors, including handedness, sex, and developmental maturation. To date, a small number of studies have examined sex differences in the lateralization of cognitive and affective functions, and in only few of these have the developmental trajectories of these lateralized differences been mapped from childhood through early adulthood. In the present study, a cross-sectional design was used with healthy children (n=7), adolescents (n= 12), and adults (n= 10) who underwent functional magnetic resonance imaging (fMRI) during a task that required perceiving fearful faces. Males and females differed in the asymmetry of activation of the amygdala and prefrontal cortex across the three age groups. For males, activation within the dorsolateral prefrontal cortex was bilateral in children, right lateralized in adolescents, and bilateral in adults, whereas females showed a monotonic relationship with age, with older females showing more bilateral activation than younger ones. In contrast, amygdala activation was similar for both sexes, with bilateral activation in children, right-lateralized activation in adolescents, and bilateral activation in adults. These results suggest that males and females show different patterns of lateralized cortical and subcortical brain activation across the period of development from childhood through early adulthood.     

Aging and attentional guidance during visual search: functional neuroanatomy by positron emission tomography

Positron emission tomography (PET) was used to examine adult age differences in neural activation during visual search. Target detection was less accurate for older adults than for younger adults, but both age groups were successful in using color to guide attention to a subset of display items. Increasing perceptual difficulty led to greater activation of occipitotemporal cortex for younger adults than for older adults, apparently as the result of older adults maintaining higher levels of activation within the easier task conditions. The results suggest that compensation for age-related decline in the efficiency of occipitotemporal cortical functioning was implemented by changes in the relative level of activation within this visual processing pathway, rather than by the recruitment of other cortical regions.     

Age-related changes in deterministic learning from positive versus negative performance feedback

Feedback-based learning declines with age. Because older adults are generally biased toward positive information (“positivity effect”), learning from positive feedback may be less impaired than learning from negative outcomes. The literature documents mixed results, due possibly to variability between studies in task design. In the current series of studies, we investigated the influence of feedback valence on reinforcement learning in young and older adults. We used nonprobabilistic learning tasks, to more systematically study the effects of feedback magnitude, learning of stimulus–response (S–R) versus stimulus–outcome (S–O) associations, and working-memory capacity. In most experiments, older adults benefitted more from positive than negative feedback, but only with large feedback magnitudes. Positivity effects were pronounced for S–O learning, whereas S–R learning correlated with working-memory capacity in both age groups. These results underline the context dependence of positivity effects in learning and suggest that older adults focus on high gains when these are informative for behavior.     

Adult age differences in visual word identification: functional neuroanatomy by positron emission tomography

Adult age differences in the neural systems mediating semantic (context-independent) memory were investigated using positron emission tomography (PET). Younger (20-29 years) and older (62-70 years) participants performed lexical decision (word/nonword discrimination) and nonsemantic (simple visual search) baseline tasks during PET scanning. Within the lexical decision task, display duration and presentation rate were varied across scans. The behavioral data suggested that although an age-related slowing was evident in visual feature and response processing, the retrieval of semantic/lexical information was similar for younger and older adults. For both age groups, lexical-related activation occurred in inferior prefrontal and occipitotemporal regions of the left hemisphere. Differential activation, as a function of age group, was observed in the left occipitotemporal pathway as a result of older adults' maintaining higher levels of neural activity in striate cortex (during visual search) and in inferior temporal cortex (during lexical decision). The prefrontal activation was similar for the two age groups. Thus, although this form of semantic memory retrieval does not undergo significant age-related decline, an age-related change in the associated pattern of neural activation is evident. These findings differ from previous neuroimaging studies of episodic (context-dependent) memory retrieval, which have suggested that age-related compensatory mechanisms are expressed primarily by greater activation of prefrontal regions for older adults than for younger adults.     

Dissociations in perceptual learning revealed by adult age differences in adaptation to time-compressed speech

When presented with several time-compressed sentences, young adults' performance improves with practice. Such adaptation has not been studied in older adults. To study age-related changes in perceptual learning, the authors tested young and older adults' ability to adapt to degraded speech. First, the authors showed that older adults, when equated for starting accuracy with young adults, adapted at a rate and magnitude comparable to young adults. However, unlike young adults, older adults failed to transfer this learning to a different speech rate and did not show additional benefit when practice exceeded 20 sentences. Listeners did not adapt to speech degraded by noise, indicating that adaptation to time-compressed speech was not attributable to task familiarity. Finally, both young and older adults adapted to spectrally shifted noise-vocoded speech. The authors conclude that initial perceptual learning is comparable in young and older adults but maintenance and transfer of this learning decline with age.     

Positivity effect in healthy aging in observational but not active feedback-learning

ABSTRACT

The present study investigated the impact of healthy aging on the bias to learn from positive or negative performance feedback in observational and active feedback learning. In active learning, a previous study had already shown a negative learning bias in healthy seniors older than 75 years, while no bias was found for younger seniors. However, healthy aging is accompanied by a ‘positivity effect’, a tendency to primarily attend to stimuli with positive valence. Based on recent findings of dissociable neural mechanisms in active and observational feedback learning, the positivity effect was hypothesized to influence older participants' observational feedback learning in particular. In two separate experiments, groups of young (mean age 27) and older participants (mean age 60 years) completed an observational or active learning task designed to differentially assess positive and negative learning. Older but not younger observational learners showed a significant bias to learn better from positive than negative feedback. In accordance with previous findings, no bias was found for active learning. This pattern of results is discussed in terms of differences in the neural underpinnings of active and observational learning from performance feedback.     

Adult age differences in the perception and learning of artistic style categories

Adult age differences in conceptual behavior were studied using informationally complex stimuli from real-world categories: paintings by two impressionist artists. In Experiment 1 we examined perceptions of category structure by having subjects sort paintings according to style similarity. Young adults were observed to depend more on abstract information in making style judgments, whereas older adults relied more on similarity in content. This resulted in different category structures between age groups, but similarity judgments in both groups appeared to correspond to actual style differences between the two artists. In Experiment 2, learning efficiency was shown to increase with a painting's category centrality, but older adults had particular trouble learning noncentral items. At transfer, both age groups were able to use abstracted central tendency information to categorize new paintings, although young adults appeared to have better access to information about specific category exemplars from acquisition. The results are generally consistent with those from studies using simpler artificial stimuli.     

Age Differences in Multiple Outcome Measures of Time-Based Prospective Memory

ABSTRACT

This study examined time-based prospective memory performance in relation to age, monitoring strategy, response accuracy, and dual-task demands. Young, middle-aged and older adults (N?=?115) completed a prospective memory task, in which they indicated the passing of time every 5 min while listening to a short story (low task demands) or completing a series of cognitive tasks (high task demands). Young and older adults showed similar patterns of monitoring behavior, with low rates of clock checking during the early phase of each 5-min interval, followed by linearly accelerating monitoring functions. However, to obtain the same level of prospective memory performance older adults needed more frequent clock checks than young adults. Furthermore, older adults' compensatory monitoring strategy was associated with an additional cost in primary task performance. Finally, increased primary task demands shifted age differences in prospective memory from monitoring frequency to response accuracy. These findings suggest that goal-directed behavior requires efficient task coordination and resource allocation, and that age-related differences in time-based prospective memory should be evaluated by using multiple outcome measures.     

Aging Affects the Ability to Process the Optic Flow Stimulations: A Functional Near-Infrared Spectrometry Study

Abstract

Optic flow (OF) has been utilized to investigate the sensory integration of visual stimuli during postural control. It is little known how the OF speed affects the aging brain during the sensory integration process of postural control. This study was to examine the effect of OF speeds on the brain activation using functional near-infrared spectroscopy (fNIRS) and postural sway between younger and older adults. Eleven healthy younger adults (5M/6F, age 22?±?1-year-old) and ten healthy older adults (4M/6F, age 71?±?5-year-old) participated in this study. A virtual reality headset was used to provide the OF stimulus at different speeds. A forceplate was used to record the center-of-pressure to compute the amplitude of postural sway (peak-to-peak). Compared with younger adults, older adults showed significantly increased activation in the OF speed of 10?m/s and decreased activation in the OF speed of 20?m/s in the left dorsolateral prefrontal cortex. Older adults also showed decreased activation in the left temporoparietal region (VEST) in the OF speed of 20?m/s. A significant difference in peak-to-peak was found between groups. Our results indicated that age might be associated with the ability to process fast OF stimulation.     

Aging affects acquisition and reversal of reward-based associative learning

Reward-based associative learning is mediated by a distributed network of brain regions that are dependent on the dopaminergic system. Age-related changes in key regions of this system, the striatum and the prefrontal cortex, may adversely affect the ability to use reward information for the guidance of behavior. The present study investigated the effects of healthy aging on different components of reward learning, such as acquisition, reversal, effects of reward magnitude, and transfer of learning. A group of 30 young (mean age = 24.2 yr) and a group of 30 older subjects (mean age = 64.1 yr) completed two probabilistic reward-based stimulus association learning tasks. Older subjects showed poorer overall acquisition and impaired reversal learning, as well as deficits in transfer learning. When only those subjects who showed evidence of significant learning were considered, younger subjects showed equivalently fast learning irrespective of reward magnitude, while learning curves in older subjects were steeper for high compared to low reward magnitudes. Acquired equivalence learning, which requires generalization across stimuli and transfer of learned contingencies to new stimuli, was mildly impaired in older subjects.     

Posterior P1 and early frontal negativity reflect developmental changes in attentional distraction during adolescence

Previous studies in adults have revealed that attentional distraction modulates the late positive potential (LPP) during emotion regulation. To determine whether early visual components reflect developmental changes in attentional distraction during adolescence, we collected event-related potentials from 20 young adolescents, 18 older adolescents, and 18 young adults as they performed a distraction task (counting) while viewing affective images. Consistent with previous findings obtained in distraction studies, the distraction task (counting) reduced emotional modulation of the LPP. At an early stage of processing, counting reduced emotional modulation of P1 and increased the negativity bias of early frontal negativity (eFN) for negatively valenced pictures compared to simple viewing with no distraction. sLORETA analyses further revealed eFN indexing of rostral prefrontal cortical activation, a cortical area that has been shown in recent fMRI studies to be activated by distraction. Moreover, P1 amplitudes in young and older adolescents did not differ but were both larger than the P1s in young adults. In addition, eFN amplitudes significantly decreased with age. The dissociable distraction patterns between the posterior P1 and eFN provide evidence not only for the timing hypothesis of emotion regulation but also for different developmental trajectories of visual processing areas and the prefrontal cortex during affective processing in adolescence.     

Episodic priming and memory for temporal source: event-related potentials reveal age-related differences in prefrontal functioning.

Event-related potentials (ERPs) were recorded from young (M = 25) and older (M = 71) adults during a recognition memory paradigm that assessed episodic priming. Participants studied two temporally distinct lists of sentences (each with two unassociated nouns). At test, in response to the nouns, participants made old-new, followed by remember (context)-know (familiarity) and source (i.e., list) judgments. Both young and older adults showed equivalent episodic priming effects. However, compared to the young adults, the older adults showed a greater source performance decrement than item memory performance decrement. Both age groups showed equivalent posterior-maximal old-new ERP effects. However, only the young produced a frontal-maximal, late onset old-new effect that differed as a function of subsequent list attribution. Because source memory is thought to be mediated by prefrontal cortex, we conclude that age-related memory differences may be due to a deficit in a prefrontal cortical system that underlies source memory and are not likely to be due to an age-related decline in episodic priming.     

Judging meaning improves function in the aging brain

Important new evidence suggests that under-recruitment of frontal cortex under intentional learning conditions in older adults can be repaired when older subjects are instructed to focus on meaning. However, the tendency for older subjects to activate both hemispheres when young adults recruit only one was not reversible. These findings provide important insights with respect to plasticity and compensation in the aging brain.     

Compensatory processing during rule-based category learning in older adults

Healthy older adults typically perform worse than younger adults at rule-based category learning, but better than patients with Alzheimer’s or Parkinson’s disease. To further investigate aging’s effect on rule-based category learning, we monitored event-related potentials (ERPs) while younger and neuropsychologically typical older adults performed a visual category-learning task with a rule-based category structure and trial-by-trial feedback. Using these procedures, we previously identified ERPs sensitive to categorization strategy and accuracy in young participants. In addition, previous studies have demonstrated the importance of neural processing in the prefrontal cortex and the medial temporal lobe for this task. In this study, older adults showed lower accuracy and longer response times than younger adults, but there were two distinct subgroups of older adults. One subgroup showed near-chance performance throughout the procedure, never categorizing accurately. The other subgroup reached asymptotic accuracy that was equivalent to that in younger adults, although they categorized more slowly. These two subgroups were further distinguished via ERPs. Consistent with the compensation theory of cognitive aging, older adults who successfully learned showed larger frontal ERPs when compared with younger adults. Recruitment of prefrontal resources may have improved performance while slowing response times. Additionally, correlations of feedback-locked P300 amplitudes with category-learning accuracy differentiated successful younger and older adults. Overall, the results suggest that the ability to adapt one’s behavior in response to feedback during learning varies across older individuals, and that the failure of some to adapt their behavior may reflect inadequate engagement of prefrontal cortex.     

Neurocognitive ageing of storage and executive processes

Converging behavioural and neuropsychological evidence indicates that age-related changes in working memory contribute substantially to cognitive decline in older adults. Important questions remain about the relationship between working memory storage and executive components and how they are affected by the normal ageing process. In several studies using positron emission tomography (PET), we find age differences in the patterns of frontal activation during working memory tasks. We find that separable age differences can be linked to different cognitive operations underlying short-term information storage, and interference resolution. Some operations are associated with age-related increases in activation, with older adults displaying bilateral activations and recruiting prefrontal areas more than younger adults. Other operations are associated with age-related decreases in activation. We consider the implications of these results for understanding the working memory system and potential compensatory processes in the ageing brain.     

The influence of task characteristics on younger and older adult motor overflow

This study investigated the influence of attentional and motor demands on motor overflow in 17 healthy young (18-35 years) and 17 older adults (60-80 years). Participants performed a finger pressing task by exerting either 33% or 66% of their maximal force output using their dominant or nondominant hand. Overflow was concurrently recorded in the passive hand. Attention was manipulated via a tactile stimulus presented to one or both hands for certain trials. Results showed that older adults exhibited greater overflow than young adults and that the effect of target force was exacerbated in older adults. Further, only older adult overflow was increased when tactile stimulation was directed to one or both hands. Increased overflow in older adults may result from bilateral cortical activation that is influenced by increased task demands. To perform comparatively to younger adults, older adults may compensate for age-related brain changes by recruiting an increased cortical network.     

The influence of task characteristics on younger and older adult motor overflow

This study investigated the influence of attentional and motor demands on motor overflow in 17 healthy young (18–35 years) and 17 older adults (60–80 years). Participants performed a finger pressing task by exerting either 33% or 66% of their maximal force output using their dominant or nondominant hand. Overflow was concurrently recorded in the passive hand. Attention was manipulated via a tactile stimulus presented to one or both hands for certain trials. Results showed that older adults exhibited greater overflow than young adults and that the effect of target force was exacerbated in older adults. Further, only older adult overflow was increased when tactile stimulation was directed to one or both hands. Increased overflow in older adults may result from bilateral cortical activation that is influenced by increased task demands. To perform comparatively to younger adults, older adults may compensate for age-related brain changes by recruiting an increased cortical network.