Neural sensitivity to phonological characteristics is stable across the lifespan.

Aging is often associated with declines in language production. For example, compared to younger adults, older adults experience more tip-of-the-tongue (TOT) states, show decreased speed and accuracy in naming objects, and have more pauses and fillers in speech, all of which indicate age-related increases in retrieval difficulty. While prior work has suggested that retrieval difficulty may be phonologically based, it is unclear whether there are age-related differences in the organization of phonological information per se or whether age-related difficulties may arise from accessing that information. Here we used fMRI to investigate the neural and behavioral basis of phonological neighborhood denisty (PND) effects on picture naming across the lifespan (N=91, ages 20-75). Consistent with prior work, behavioral results revealed that higher PND led to faster picture naming times and higher accuracies overall, and that older adults were less accurate in their responses. Consistent with the behavioral analyses, fMRI analyses showed that increasing PND was associated with decreased activation in auditory and motor language regions, including bilateral superior temporal gyri and bilateral precentral gyri. Interestingly, although there were age-related increases in functional activation to picture naming, there were no age-related modulations of neural sensitivity to PND. Overall, these results suggest that having a large cohort of phonological neighbors facilitates language production, and although aging is associated with increases in language production difficulty, sensitivity to phonological features during language production is stable across the lifespan.

Age-related differences in the neural bases of phonological and semantic processes in the context of task-irrelevant information.

As we age we have increasing difficulty with phonological aspects of language production. Yet semantic processes are largely stable across the life span. This suggests a fundamental difference in the cognitive and potentially neural architecture supporting these systems. Moreover, language processes such as these interact with other cognitive processes that also show age-related decline, such as executive function and inhibition. The present study examined phonological and semantic processes in the presence of task-irrelevant information to examine the influence of such material on language production. Older and younger adults made phonological and semantic decisions about pictures in the presence of either phonologically or semantically related words, which were unrelated to the task. FMRI activation during the semantic condition showed that all adults engaged typical left-hemisphere language regions, and that this activation was positively correlated with efficiency across all adults. In contrast, the phonological condition elicited activation in bilateral precuneus and cingulate, with no clear brain-behavior relationship. Similarly, older adults exhibited greater activation than younger adults in several regions that were unrelated to behavioral performance. Our results suggest that as we age, brain-behavior relations decline, and there is an increased reliance on both language-specific and domain-general brain regions that are seen most prominently during phonological processing. In contrast, the core semantic system continues to be engaged throughout the life span, even in the presence of task-irrelevant information.

Task difficulty modulates brain-behavior correlations in language production and cognitive control: Behavioral and fMRI evidence from a phonological go/no-go picture-naming paradigm.

Language production and cognitive control are complex processes that involve distinct yet interacting brain networks. However, the extent to which these processes interact and their neural bases have not been thoroughly examined. Here, we investigated the neural and behavioral bases of language production and cognitive control via a phonological go/no-go picture-naming task. Naming difficulty and cognitive control demands (i.e., conflict monitoring and response inhibition) were manipulated by varying the proportion of naming trials (go trials) and inhibition trials (no-go trials) across task runs. The results demonstrated that as task demands increased, participants' behavioral performance declined (i.e., longer reaction times on naming trials, more commission errors on inhibition trials) whereas brain activation generally increased. Increased activation was found not only within the language network but also in domain-general control regions. Additionally, right superior and inferior frontal and left supramarginal gyri were sensitive to increased task difficulty during both language production and response inhibition. We also found both positive and negative brain-behavior correlations. Most notably, increased activation in sensorimotor regions, such as precentral and postcentral gyri, was associated with better behavioral performance, in both successful picture naming and successful inhibition. Moreover, comparing the strength of correlations across conditions indicated that the brain-behavior correlations in sensorimotor regions that were associated with improved performance became stronger as task demands increased. Overall, our results suggest that cognitive control demands affect language production, and that successfully coping with increases in task difficulty relies on both language-specific and domain-general cognitive control regions.

Frontoparietal activation during visual conjunction search: Effects of bottom-up guidance and adult age.

We conducted functional magnetic resonance imaging (fMRI) with a visual search paradigm to test the hypothesis that aging is associated with increased frontoparietal involvement in both target detection and bottom-up attentional guidance (featural salience). Participants were 68 healthy adults, distributed continuously across 19 to 78 years of age. Frontoparietal regions of interest (ROIs) were defined from resting-state scans obtained prior to task-related fMRI. The search target was defined by a conjunction of color and orientation. Each display contained one item that was larger than the others (i.e., a size singleton) but was not informative regarding target identity. Analyses of search reaction time (RT) indicated that bottom-up attentional guidance from the size singleton (when coincident with the target) was relatively constant as a function of age. Frontoparietal fMRI activation related to target detection was constant as a function of age, as was the reduction in activation associated with salient targets. However, for individuals 35 years of age and older, engagement of the left frontal eye field (FEF) in bottom-up guidance was more prominent than for younger individuals. Further, the age-related differences in left FEF activation were a consequence of decreasing resting-state functional connectivity in visual sensory regions. These findings indicate that age-related compensatory effects may be expressed in the relation between activation and behavior, rather than in the magnitude of activation, and that relevant changes in the activation-RT relation may begin at a relatively early point in adulthood. Hum Brain Mapp 38:2128-2149, 2017. © 2017 Wiley Periodicals, Inc.

Memory-guided force control in healthy younger and older adults.

Successful performance of a memory-guided motor task requires participants to store and then recall an accurate representation of the motor goal. Further, participants must monitor motor output to make adjustments in the absence of visual feedback. The goal of this study was to examine memory-guided grip force in healthy younger and older adults and compare it to performance on behavioral tasks of working memory. Previous work demonstrates that healthy adults decrease force output as a function of time when visual feedback is not available. We hypothesized that older adults would decrease force output at a faster rate than younger adults, due to age-related deficits in working memory. Two groups of participants, younger adults (YA: N = 32, mean age 21.5 years) and older adults (OA: N = 33, mean age 69.3 years), completed four 20-s trials of isometric force with their index finger and thumb, equal to 25% of their maximum voluntary contraction. In the full-vision condition, visual feedback was available for the duration of the trial. In the no vision condition, visual feedback was removed for the last 12 s of each trial. Participants were asked to maintain constant force output in the absence of visual feedback. Participants also completed tasks of word recall and recognition and visuospatial working memory. Counter to our predictions, when visual feedback was removed, younger adults decreased force at a faster rate compared to older adults and the rate of decay was not associated with behavioral performance on tests of working memory.

Age-related differences in the neural bases of phonological and semantic processes.

Changes in language functions during normal aging are greater for phonological compared with semantic processes. To investigate the behavioral and neural basis for these age-related differences, we used fMRI to examine younger and older adults who made semantic and phonological decisions about pictures. The behavioral performance of older adults was less accurate and less efficient than younger adults' in the phonological task but did not differ in the semantic task. In the fMRI analyses, the semantic task activated left-hemisphere language regions, and the phonological task activated bilateral cingulate and ventral precuneus. Age-related effects were widespread throughout the brain and most often expressed as greater activation for older adults. Activation was greater for younger compared with older adults in ventral brain regions involved in visual and object processing. Although there was not a significant Age × Condition interaction in the whole-brain fMRI results, correlations examining the relationship between behavior and fMRI activation were stronger for younger compared with older adults. Our results suggest that the relationship between behavior and neural activation declines with age, and this may underlie some of the observed declines in performance.

Age mediation of frontoparietal activation during visual feature search.

Activation of frontal and parietal brain regions is associated with attentional control during visual search. We used fMRI to characterize age-related differences in frontoparietal activation in a highly efficient feature search task, detection of a shape singleton. On half of the trials, a salient distractor (a color singleton) was present in the display. The hypothesis was that frontoparietal activation mediated the relation between age and attentional capture by the salient distractor. Participants were healthy, community-dwelling individuals, 21 younger adults (19-29 years of age) and 21 older adults (60-87 years of age). Top-down attention, in the form of target predictability, was associated with an improvement in search performance that was comparable for younger and older adults. The increase in search reaction time (RT) associated with the salient distractor (attentional capture), standardized to correct for generalized age-related slowing, was greater for older adults than for younger adults. On trials with a color singleton distractor, search RT increased as a function of increasing activation in frontal regions, for both age groups combined, suggesting increased task difficulty. Mediational analyses disconfirmed the hypothesized model, in which frontal activation mediated the age-related increase in attentional capture, but supported an alternative model in which age was a mediator of the relation between frontal activation and capture.

Multiple sclerosis is associated with differences in semantic memory structure.

OBJECTIVE: Although language is often considered to be largely intact in multiple sclerosis (MS), word-finding difficulties are a common complaint. Recent work suggests that declines in language are not solely the result of motoric and cognitive slowing that is most strongly associated with MS. Network science approaches have been effectively used to examine network structure as it relates to clinical conditions, aging, and language. The present study utilizes a network science approach to investigate whether individuals with MS exhibit less interconnected and resilient semantic networks compared to age-matched neurotypical peers. METHOD: We used semantic fluency data from 89 participants with MS and 88 neurotypical participants to estimate and analyze the semantic network structure for each participant group. Additionally, we conducted a percolation analysis to examine the resilience of each network. RESULTS: Network measures showed that individuals with MS had lower local and global clustering coefficients, longer average shortest path lengths, and higher modularity values compared to neurotypical peers. Small-worldness, network portrait divergence measures, and community detection analyses were consistent with these results and indicated that macroscopic properties of the two networks differed and that the semantic network for individuals with MS was more fractured than the neurotypical peer network. Moreover, a spreading activation simulation and percolation analysis suggested that the semantic networks of individuals with MS are less flexible and activation degrades faster than those of age-matched neurotypical participants. CONCLUSIONS: These differing semantic network structures suggest that language retrieval difficulties in MS partially result from decline in language-specific factors. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

What we mean when we say semantic: Toward a multidisciplinary semantic glossary.

Tulving characterized semantic memory as a vast repository of meaning that underlies language and many other cognitive processes. This perspective on lexical and conceptual knowledge galvanized a new era of research undertaken by numerous fields, each with their own idiosyncratic methods and terminology. For example, "concept" has different meanings in philosophy, linguistics, and psychology. As such, many fundamental constructs used to delineate semantic theories remain underspecified and/or opaque. Weak construct specificity is among the leading causes of the replication crisis now facing psychology and related fields. Term ambiguity hinders cross-disciplinary communication, falsifiability, and incremental theory-building. Numerous cognitive subdisciplines (e.g., vision, affective neuroscience) have recently addressed these limitations via the development of consensus-based guidelines and definitions. The project to follow represents our effort to produce a multidisciplinary semantic glossary consisting of succinct definitions, background, principled dissenting views, ratings of agreement, and subjective confidence for 17 target constructs (e.g., abstractness, abstraction, concreteness, concept, embodied cognition, event semantics, lexical-semantic, modality, representation, semantic control, semantic feature, simulation, semantic distance, semantic dimension). We discuss potential benefits and pitfalls (e.g., implicit bias, prescriptiveness) of these efforts to specify a common nomenclature that other researchers might index in specifying their own theoretical perspectives (e.g., They said X, but I mean Y).

Resting State Network Segregation Modulates Age-Related Differences in Language Production.

Older adults typically exhibit decline in language production. However, how the brain supports or fails to support these processes is unclear. Moreover, there are competing hypotheses about the nature of age-related neural changes and whether age-related increases in neural activity reflect compensation or a decline in neural efficiency. In the current study, we investigated the neural bases of language production focusing on resting state functional connectivity. We hypothesized that language production performance, functional connectivity, and their relationship would differ as a function of age. Consistent with prior work, older age was associated with worse language production performance. Functional connectivity analyses showed that network segregation within the left hemisphere language network was maintained across adulthood. However, increased age was associated with lower whole brain network segregation. Moreover, network segregation was related to language production ability. In both network analyses, there were significant interactions with age-higher network segregation was associated with better language production abilities for younger and middle-aged adults, but not for older adults. Interestingly, there was a stronger relationship between language production and the whole brain network segregation than between production and the language network. These results highlight the utility of network segregation measures as an index of brain function, with higher network segregation associated with better language production ability. Moreover, these results are consistent with stability in the left hemisphere language network across adulthood and suggest that dedifferentiation among brain networks, outside of the language network, is a hallmark of aging and may contribute to age-related language production difficulties.

Task difficulty modulates age-related differences in functional connectivity during word production.

Older adults typically report increased difficulty with language production, while its neural bases are less clear. The current study investigated the neural bases of age-related differences in language production at the word level and the modulating effect of task difficulty, focusing on task-based functional connectivity. Using an English phonological Go/No-Go picture naming task, task difficulty was manipulated by varying the proportion of naming trials (Go trials) and inhibition trials (No-Go trials) across runs. Behaviorally, compared to younger adults, older adults performed worse, and showed larger effects of task difficulty. Neurally, older adults had lower within language network connectivity compared to younger adults. Moreover, older adults' language network became less segregated as task difficulty increased. These results are consistent with the Compensation-Related Utilization of Neural Circuits Hypothesis, suggesting that the brain becomes less specified and efficient with increased task difficulty, and that these effects are stronger among older adults (i.e., more dedifferentiated).

Multilingual Language Diversity Protects Native Language Production under Different Control Demands.

The use of multiple languages has been found to influence individuals' cognitive abilities. Although some studies have also investigated the effect of multilingualism on non-native language proficiency, fewer studies have focused on how multilingual experience affects native language production. This study investigated the effect of multilingualism on native language production, specifically examining control demands through a semantic Go/No-Go picture naming task. The multilingual experience was quantified using language entropy, which measures the uncertainty and diversity of language use. Control demands were achieved by manipulating the proportion of Go (i.e., naming) trials in different conditions. Results showed that as control demands increased, multilingual individuals exhibited poorer behavioral performance and greater brain activation throughout the brain. Moreover, more diverse language use was associated with higher accuracy in naming and more interconnected brain networks with greater involvement of domain-general neural resources and less domain-specific neural resources. Notably, the varied and balanced use of multiple languages enabled multilingual individuals to respond more efficiently to increased task demands during native language production.

Age differences in semantic network structure: Acquiring knowledge shapes semantic memory.

Computational research suggests that semantic memory, operationalized as semantic memory networks, undergoes age-related changes. Previous work suggests that concepts in older adults' semantic memory networks are more separated, more segregated, and less connected to each other. However, cognitive network research often relies on group averages (e.g., young vs. older adults), and it remains unclear if individual differences influence age-related disparities in language production abilities. Here, we analyze the properties of younger and older participants' individual-based semantic memory networks based on their semantic relatedness judgments. We related individual-based network measures-clustering coefficient (CC; connectivity), global efficiency, and modularity (structure)-to language production (verbal fluency) and vocabulary knowledge. Similar to previous findings, we found significant age effects: CC and global efficiency were lower, and modularity was higher, for older adults. Furthermore, vocabulary knowledge was significantly related to the semantic memory network measures: corresponding with the age effects, CC and global efficiency had a negative relationship, while modularity had a positive relationship with vocabulary knowledge. More generally, vocabulary knowledge significantly increased with age, which may reflect the critical role that the accumulation of knowledge within semantic memory has on its structure. These results highlight the impact of diverse life experiences on older adults' semantic memory and demonstrate the importance of accounting for individual differences in the aging mental lexicon. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

Cerebral White Matter Mediation of Age-Related Differences in Picture Naming Across Adulthood.

As people age, one of the most common complaints is difficulty with word retrieval. A wealth of behavioral research confirms such age-related language production deficits, yet the structural neural differences that relate to age-related language production deficits remains an open area of exploration. Therefore, the present study used a large sample of healthy adults across adulthood to investigate how age-related white matter differences in three key left-hemisphere language tracts may contribute to age-related differences in language ability. Specifically, we used diffusion tensor imaging to measure fractional anisotropy (FA) and radial diffusivity (RD) which are indicators of white matter structure. We then used a series of path models to test whether white matter from the superior longitudinal fasciculus (SLF), the inferior longitudinal fasciculus, and the frontal aslant tract (FAT) mediated age-related differences in one form of language production, picture naming. We found that FA, as well as RD from the SLF and FAT mediated the relation between age and picture naming performance, whereas a control tract (corticospinal) was not a mediator. Moreover, differences between mediation of picture naming and a control naming condition suggest that left SLF has a greater role in higher-order aspects of naming, such as semantic and lexical selection whereas left FAT is more sensitive to sensorimotor aspects of fluency or speech motor planning. These results suggest that dorsal white matter contributes to age-related differences in generating speech and may be particularly important in supporting word retrieval across adulthood.

Neural sensitivity to semantic neighbors is stable across the adult lifespan.

As we age, language reflects patterns of both stability and change. On the one hand, vocabulary and semantic abilities are largely stable across the adult lifespan, yet lexical retrieval is often slower and less successful (i.e., slower picture naming times, increased tip of the tongue incidents). Although the behavioral bases of these effects have been well established, less is known about the brain regions that support these age-related differences. We used functional Magnetic Resonance Imaging (fMRI) to examine the neural basis of picture naming. Specifically, we were interested in whether older adults would be equally sensitive to semantic characteristics, specifically the number of semantic near neighbors. Near neighbors, defined here as items with a high degree of semantic feature overlap, were of interest as these are thought to elicit competition among potential candidates and increase naming difficulty. Consistent with prior reports, pictures with more semantic near neighbors were named more slowly and less accurately for all adults. Additionally, this interference for naming times was larger as age increased, starting around 30 years old. In contrast to the age-related behavioral slowing, the neural basis of these effects was stable across adulthood. Across all adults, a number of language-relevant regions including left posterior middle temporal gyrus and left inferior frontal gyrus, pars triangularis were sensitive to the number of near neighbors. Our results suggest that although middle-aged and older adults' picture naming is more slowed by increased semantic competition, the brain regions supporting semantic processes remain stable across the adult lifespan.

The influence of context on hemispheric recruitment during metaphor processing.

Although the left hemisphere's prominence in language is well established, less emphasis has been placed on possible roles for the right hemisphere. Behavioral, patient, and neuroimaging research suggests that the right hemisphere may be involved in processing figurative language. Additionally, research has demonstrated that context can modify language processes and facilitate comprehension. Here we investigated how figurativeness and context influenced brain activation, with a specific interest in right hemisphere function. Previous work in our laboratory indicated that novel stimuli engaged right inferior frontal gyrus (IFG) and that both novel and familiar metaphors engaged right IFG and right temporal pole. The graded salience hypothesis proposes that context may lessen integration demands, increase the salience of metaphors, and thereby reduce right hemisphere recruitment for metaphors. In the present study, fMRI was used to investigate brain function, whereas participants read literal and metaphoric sentences that were preceded by either a congruent or an incongruent literal sentence. Consistent with prior research, all sentences engaged traditional left hemisphere regions. Differences between metaphors and literal sentences were observed, but only in the left hemisphere. In contrast, a main effect of congruence was found in the right IFG, the right temporal pole, and the dorsal medial pFC. Partially consistent with the graded salience hypothesis, our results highlight the strong influence of context on language, demonstrate the importance of the right hemisphere in discourse, and suggest that, in a wider discourse context, congruence has a greater influence on right hemisphere recruitment than figurativeness.

The neural bases of multimodal sensory integration in older adults.

Although hearing often declines with age, prior research has shown that older adults may benefit from multisensory input to a greater extent when compared to younger adults, a concept known as inverse effectiveness. While there is behavioral evidence in support of this phenomenon, less is known about its neural basis. The present fMRI study examined how older and younger adults processed multimodal auditory-visual (AV) phonemic stimuli which were either congruent or incongruent across modalities. Incongruent AV pairs were designed to elicit the McGurk effect. Behaviorally, reaction times were significantly faster during congruent trials compared to incongruent trials for both age groups, and overall older adults responded more slowly. The interaction was not significant suggesting that older adults processed the AV stimuli similarly to younger adults. Although there were minimal behavioral differences, age-related differences in functional activation were identified: Younger adults elicited greater activation than older adults in primary sensory regions including superior temporal gyrus, the calcarine fissure, and left post-central gyrus. In contrast, older adults elicited greater activation than younger adults in dorsal frontal regions including middle and superior frontal gyri, as well as dorsal parietal regions. These data suggest that while there is age-related stability in behavioral sensitivity to multimodal stimuli, the neural bases for this effect differed between older and younger adults. Our results demonstrated that older adults underrecruited primary sensory cortices and had increased recruitment of regions involved in executive function, attention, and monitoring processes, which may reflect an attempt to compensate.

The intensity and connectivity of spontaneous brain activity in a language network relate to aging and language.

Neuroimaging studies often either look at functional activation in response to an explicit task, or functional connectivity (i.e., interregional correlations) during resting-state. Few studies have looked at the intensity of brain activity or its relationship with age, behavior, and language. The current study investigated both intensity (i.e., the Amplitude of Low-Frequency Fluctuations, ALFF) and the functional connectivity of spontaneous brain activity during rest and their relationship with age and language. A life-span sample of individuals (N = 152) completed a battery of neuropsychological tests to assess basic cognitive functions and resting-state functional MRI data to assess spontaneous brain activity. Focusing on an extend language network, the mean ALFF and total degree were calculated for this network. We found that increased age was associated with more intense activity (i.e., higher ALFF) but lower within-network connectivity. Additionally, these increases in activity within the language network during resting-state were related to worse language ability, particularly in younger adults, supporting a dedifferentiation account of cognition. Our results support the utility of using resting-state data as an indicator of cognition and support the role of ALFF as a potential biomarker in characterizing the relationships between resting-state brain activity, age, and cognition.

Language immersion and language training: Two paths to enhanced language regulation and cognitive control.

When bilinguals switch languages they regulate the more dominant language to enable spoken production in the less dominant language. How do they engage cognitive control to accomplish regulation? We examined this issue by comparing the consequences of training on language switching in two different contexts. Chinese-English bilinguals were immersed in English (L2) while studying abroad (this study) or in Chinese (L1) in their native language environment (Zhang et al., 2015). In each study, participants performed the AX-CPT task while EEG was recorded and were then trained on language switching. While Zhang et al. found that training enhanced proactive control in the L1 context, there were no effects of training under L2 immersion conditions. Critically, L2 immersed bilinguals revealed enhanced proactive control at pre-test and greater L1 inhibition on language switching relative to L1 immersed bilinguals. We hypothesize that L2 immersion creates a natural training context that increases reliance on proactive control to enable regulation of the L1.

Age-related differences in resting-state and task-based network characteristics and cognition: a lifespan sample.

Aging is often associated with cognitive and neural decline, but how these factors interact is still not fully understood. Recently, functional connectivity, or the degree to which brain regions are concurrently active, has provided insight into age-related differences. However, functional connectivities during task and rest differ and few studies have examined how these relate to a broad range of cognitive functions. The present study investigated the effect of age on cognition, whole-brain functional connectivity during resting-state and task, and their relationships across the adult lifespan. Cognition was broadly assessed using a battery of cognitive assessments and mean network characteristics were calculated across the whole brain. Behaviorally, increased age was associated with worse recall, executive function, and verbal working memory abilities but better language performance. Neurally, increased age was associated with lower overall within- and between-network functional connectivities during both rest and task, and these age-connectivity relationships were stronger during task performance. Connectivity was also related to cognition, and for all participants, these relationships were strongest during rest. Specifically, higher resting-state between-network functional connectivity was associated with poorer cognition for all adults and poorer language ability among older adults. Collectively, these findings demonstrate that while age effects were strongest during the task, resting-state functional connectivity was most closely tied to cognition. Moreover, these results are theoretically consistent with dedifferentiation accounts of cognition and aging and show that less differentiated functional connectivities are associated with cognitive costs for both older and younger adults.