The neuroanatomy of developmental language disorder: a systematic review and meta-analysis.

Developmental language disorder (DLD) is a common neurodevelopmental disorder with adverse impacts that continue into adulthood. However, its neural bases remain unclear. Here we address this gap by systematically identifying and quantitatively synthesizing neuroanatomical studies of DLD using co-localization likelihood estimation, a recently developed neuroanatomical meta-analytic technique. Analyses of structural brain data (22 peer-reviewed papers, 577 participants) revealed highly consistent anomalies only in the basal ganglia (100% of participant groups in which this structure was examined, weighted by group sample sizes; 99.8% permutation-based likelihood the anomaly clustering was not due to chance). These anomalies were localized specifically to the anterior neostriatum (again 100% weighted proportion and 99.8% likelihood). As expected given the task dependence of activation, functional neuroimaging data (11 peer-reviewed papers, 414 participants) yielded less consistency, though anomalies again occurred primarily in the basal ganglia (79.0% and 95.1%). Multiple sensitivity analyses indicated that the patterns were robust. The meta-analyses elucidate the neuroanatomical signature of DLD, and implicate the basal ganglia in particular. The findings support the procedural circuit deficit hypothesis of DLD, have basic research and translational implications for the disorder, and advance our understanding of the neuroanatomy of language.

Knowledge of Statistics or Statistical Learning? Readers Prioritize the Statistics of their Native Language Over the Learning of Local Regularities.

A large body of evidence suggests that people spontaneously and implicitly learn about regularities present in the visual input. Although theorized as critical for reading, this ability has been demonstrated mostly with pseudo-fonts or highly atypical artificial words. We tested whether local statistical regularities are extracted from materials that more closely resemble one's native language. In two experiments, Italian speakers saw a set of letter strings modelled on the Italian lexicon and guessed which of these strings were words in a fictitious language and which were foils. Unknown to participants, words could be distinguished from foils based on their average bigram frequency. Surprisingly, in both experiments, we found no evidence that participants relied on this regularity. Instead, lexical decisions were guided by minimal bigram frequency, a cue rooted in participants' native language. We discuss the implications of these findings for accounts of statistical learning and visual word processing.

Subcortical Cognition: The Fruit Below the Rind.

Cognitive neuroscience has highlighted the cerebral cortex while often overlooking subcortical structures. This cortical proclivity is found in basic and translational research on many aspects of cognition, especially higher cognitive domains such as language, reading, music, and math. We suggest that, for both anatomical and evolutionary reasons, multiple subcortical structures play substantial roles across higher and lower cognition. We present a comprehensive review of existing evidence, which indeed reveals extensive subcortical contributions in multiple cognitive domains. We argue that the findings are overall both real and important. Next, we advance a theoretical framework to capture the nature of (sub)cortical contributions to cognition. Finally, we propose how new subcortical cognitive roles can be identified by leveraging anatomical and evolutionary principles, and we describe specific methods that can be used to reveal subcortical cognition. Altogether, this review aims to advance cognitive neuroscience by highlighting subcortical cognition and facilitating its future investigation.

Evidence that ageing yields improvements as well as declines across attention and executive functions.

Many but not all cognitive abilities decline during ageing. Some even improve due to lifelong experience. The critical capacities of attention and executive functions have been widely posited to decline. However, these capacities are composed of multiple components, so multifaceted ageing outcomes might be expected. Indeed, prior findings suggest that whereas certain attention/executive functions clearly decline, others do not, with hints that some might even improve. We tested ageing effects on the alerting, orienting and executive (inhibitory) networks posited by Posner and Petersen's influential theory of attention, in a cross-sectional study of a large sample (N = 702) of participants aged 58-98. Linear and nonlinear analyses revealed that whereas the efficiency of the alerting network decreased with age, orienting and executive inhibitory efficiency increased, at least until the mid-to-late 70s. Sensitivity analyses indicated that the patterns were robust. The results suggest variability in age-related changes across attention/executive functions, with some declining while others improve.

Maternal depression is the predominant persistent risk for child cognitive and social-emotional problems from early childhood to pre-adolescence: A longitudinal cohort study.

RATIONALE: Brain development occurs rapidly during early childhood and continues throughout middle childhood. Early and later windows of opportunity exist to alter developmental trajectories. Few studies in low- and middle-income countries have examined the importance of the timing of exposure to risks for poor pre-adolescent cognitive and social-emotional outcomes. METHODS: We assessed 359 children who participated in two follow-up studies of the Supplementation with Multiple Micronutrients Intervention Trial conducted in Indonesia in 2001-2004: at 3.5 years in 2006 and 9-12 years in 2012-2014. Using structural equation models, we examined indicators of early childhood (3.5 y) and pre-adolescent (9-12 y) exposure to risks (child height-for-age z-score [HAZ], hemoglobin [Hb], maternal depressive symptoms [MDS], home environment [HOME]), with two developmental outcomes: cognitive ability and social-emotional problems. We characterized patterns of change by calculating residuals of indicators measured earlier (3.5 y) predicting the same indicators measured later (9-12 y), for example, the residual of 3.5 y MDS predicting 9-12 y MDS (rMDS). RESULTS: Three early risk indicators (HOME, Hb, and MDS) were indirectly associated with pre-adolescent cognitive scores through early cognitive scores (HOME: 0.15, [95% CI 0.09, 0.21]; Hb: 0.08 [0.04, 0.12], MDS: -0.07 [-0.12, -0.02]). Pre-adolescent cognitive scores were also associated with change in MDS (rMDS: -0.13 [-0.23, -0.02]) and Hb (rHb: 0.10 [0.00, 0.20]) during middle childhood. For pre-adolescent social-emotional problems, both early childhood MDS (0.31 [0.19, 0.44]) and change in MDS during middle childhood (rMDS: 0.48 [0.37, 0.60]) showed strong direct associations with this outcome. CONCLUSIONS: Our findings confirm those of previous studies that prevention of risk exposures during early childhood is likely to support long-term child development. It also adds evidence to a previously scarce literature for the middle childhood period. Prevention of maternal depressive symptoms and child anemia during middle childhood should be assessed for effectiveness to support child development.

Declarative Memory Predicts Phonological Processing Abilities in Adulthood.

Individual differences in phonological processing abilities have often been attributed to perceptual factors, rather than to factors relating to learning and memory. Here, we consider the contribution of individual differences in declarative and procedural memory to phonological processing performance in adulthood. We examined the phonological processing, declarative memory, and procedural memory abilities of 79 native English-speaking young adults with typical language and reading abilities. Declarative memory was assessed with a recognition memory task of real and made-up objects. Procedural memory was assessed with a serial reaction time task. For both tasks, learning was assessed shortly after encoding, and again after a 12-h, overnight delay. We regressed phonological processing ability with memory performance on both days. We found that declarative memory, but not procedural memory, was highly predictive of phonological processing abilities. Specifically, declarative memory scores obtained shortly after learning were associated with non-word repetition performance, whereas declarative memory scores obtained after the overnight delay were associated with phonological awareness. Procedural memory was not associated with either of the phonological processing measures. We discuss these findings in the context of adult participants with mature phonological systems. We examine possible implications for the relationship between declarative memory and phonological processing in adulthood.

Aging affects steaks more than knives: Evidence that the processing of words related to motor skills is relatively spared in aging.

Lexical-processing declines are a hallmark of aging. However, the extent of these declines may vary as a function of different factors. Motivated by findings from neurodegenerative diseases and healthy aging, we tested whether 'motor-relatedness' (the degree to which words are associated with particular human body movements) might moderate such declines. We investigated this question by examining data from three experiments. The experiments were carried out in different languages (Dutch, German, English) using different tasks (lexical decision, picture naming), and probed verbs and nouns, in all cases controlling for potentially confounding variables (e.g., frequency, age-of-acquisition, imageability). Whereas 'non-motor words' (e.g., steak) showed age-related performance decreases in all three experiments, 'motor words' (e.g., knife) yielded either smaller decreases (in one experiment) or no decreases (in two experiments). The findings suggest that motor-relatedness can attenuate or even prevent age-related lexical declines, perhaps due to the relative sparing of neural circuitry underlying such words.

Deficits of Learning in Procedural Memory and Consolidation in Declarative Memory in Adults With Developmental Language Disorder.

Purpose This study examined procedural and declarative learning and consolidation abilities in adults with developmental language disorder (DLD) relative to their typical language (TD) peers. Method A total of 100 young adults (age 18-24 years) with (n = 21) and without (n = 79) DLD participated across two sites. Performance measures on a recognition memory task and a serial reaction time task were used to assess declarative and procedural memory, respectively. Performance was measured shortly after learning (8 a.m.) and again after a 12-hr, overnight delay (8 a.m.). Results Linear mixed-effects modeling was used to examine the effects of time and group membership on task performance. For the serial reaction time task, there were significant effects of group (TD > DLD) and time (Day 1 > Day 2), but no interaction between them. For the recognition memory task, there was a significant interaction between group and time, driven by overnight gains in the TD group, combined with stable performance across days by those with DLD. Conclusions In procedural memory, adults with DLD demonstrate a learning deficit relative to adults without DLD, but appear to have comparable retention of learned information. In declarative memory, adults with DLD demonstrate a deficit in the overnight enhancement of memory retrieval, despite typical-like learning exhibited when tested shortly after encoding. Supplemental Material https://doi.org/10.23641/asha.13626485.

Early-life education may help bolster declarative memory in old age, especially for women.

Although declarative memory declines with age, sex and education might moderate these weaknesses. We investigated effects of sex and education on nonverbal declarative (recognition) memory in 704 older adults (aged 58-98, 0-17 years of education). Items were drawings of real and made-up objects. Age negatively impacted declarative memory, though this age effect was moderated by sex and object-type: it was steeper for males than females, but only for real objects. Education was positively associated with memory, but also interacted with sex and object-type: education benefited women more than men (countering the age effects, especially for women), and remembering real more than made-up objects. The findings suggest that nonverbal memory in older adults is associated negatively with age but positively with education; both effects are modulated by sex, and by whether learning relates to preexisting or new information. The study suggests downstream benefits from education, especially for girls.

Can sex influence the neurocognition of language? Evidence from Parkinson's disease.

Parkinson's disease (PD), which involves basal ganglia degeneration, affects language as well as motor function. However, which aspects of language are impaired in PD and under what circumstances remains unclear. We examined whether lexical and grammatical aspects of language are differentially affected in PD, and whether this dissociation is moderated by sex as well as the degree of basal ganglia degeneration. Our predictions were based on the declarative/procedural model of language. The model posits that grammatical composition, including in regular inflection, depends importantly on left basal ganglia procedural memory circuits, whereas irregular and other lexicalized forms are memorized in declarative memory. Since females tend to show declarative memory advantages as compared to males, the model further posits that females should tend to rely on this system for regulars, which can be stored as lexicalized chunks. We tested non-demented male and female PD patients and healthy control participants on the intensively studied paradigm of English regular and irregular past-tense production. Mixed-effects regression revealed PD deficits only at regular inflection, only in male patients. The degree of left basal ganglia degeneration, as reflected by right-side hypokinesia, predicted only regular inflection, and only in male patients. Left-side hypokinesia did not show this pattern. Past-tense frequency effects suggested that the female patients retrieved regular as well as irregular past-tense forms from declarative memory, whereas the males retrieved only irregulars. Sensitivity analyses showed that the pattern of findings was robust. The results, which are consistent with the declarative/procedural model, suggest a grammatical deficit in PD due to left basal ganglia degeneration, with a relative sparing of lexical retrieval. Female patients appear to compensate for this deficit by relying on chunks stored in declarative memory. More generally, the study elucidates the neurocognition of inflectional morphology and provides evidence that sex can influence how language is computed in the mind and brain.

The effect of bilingualism on brain development from early childhood to young adulthood.

Bilingualism affects the structure of the brain in adults, as evidenced by experience-dependent grey and white matter changes in brain structures implicated in language learning, processing, and control. However, limited evidence exists on how bilingualism may influence brain development. We examined the developmental patterns of both grey and white matter structures in a cross-sectional study of a large sample (n = 711 for grey matter, n = 637 for white matter) of bilingual and monolingual participants, aged 3-21 years. Metrics of grey matter (thickness, volume, and surface area) and white matter (fractional anisotropy and mean diffusivity) were examined across 41 cortical and subcortical brain structures and 20 tracts, respectively. We used generalized additive modelling to analyze whether, how, and where the developmental trajectories of bilinguals and monolinguals might differ. Bilingual and monolingual participants manifested distinct developmental trajectories in both grey and white matter structures. As compared to monolinguals, bilinguals showed: (a) more grey matter (less developmental loss) starting during late childhood and adolescence, mainly in frontal and parietal regions (particularly in the inferior frontal gyrus pars opercularis, superior frontal cortex, inferior and superior parietal cortex, and precuneus); and (b) higher white matter integrity (greater developmental increase) starting during mid-late adolescence, specifically in striatal-inferior frontal fibers. The data suggest that there may be a developmental basis to the well-documented structural differences in the brain between bilingual and monolingual adults.

Learning and Consolidation of Declarative Memory in Good and Poor Readers of English as a Second Language.

Declarative memory abilities may be important for children who are learning to read in a second language. In the present study, we investigated declarative memory in a recognition memory task in 7-to-13-year-old, Kannada native-speaking, good (n = 22) and poor (n = 22) readers of English, in Karnataka, India. Recognition memory was tested shortly (∼10 min) after encoding (day 1) and again on the next (day 2). Analyses revealed that the two groups did not differ in recognition memory performance on day 1. On day 2, the good readers improved from day 1, whereas poor readers did not. A partial correlation analysis suggests that consolidation - the change in performance in recognition memory between the 2 days - is associated with reading skills in good readers, but not in poor readers. Taken together, these results suggest that children who struggle to read in a second language may have deficits in declarative memory consolidation.

Data and analysis script for infant and adult eye movement in an adapted ocular-motor serial reaction time task assessing procedural memory.

This article provides a description of eye movement data collected during an ocular-motor serial reaction time task. Raw gaze data files for 63 infants and 24 adults along with the data processing and analysis script for extracting saccade latencies, summarizing participants' performance, and testing statistical differences, are hosted on Open Science Framework (OSF). Files (in Matlab format) available for download allow for replication of the results reported in "Procedural memory in infancy: Evidence from implicit sequence learning in an eye-tracking paradigm" [1].

The Neurocognition of Developmental Disorders of Language.

Developmental disorders of language include developmental language disorder, dyslexia, and motor-speech disorders such as articulation disorder and stuttering. These disorders have generally been explained by accounts that focus on their behavioral rather than neural characteristics; their processing rather than learning impairments; and each disorder separately rather than together, despite their commonalities and comorbidities. Here we update and review a unifying neurocognitive account-the Procedural circuit Deficit Hypothesis (PDH). The PDH posits that abnormalities of brain structures underlying procedural memory (learning and memory that rely on the basal ganglia and associated circuitry) can explain numerous brain and behavioral characteristics across learning and processing, in multiple disorders, including both commonalities and differences. We describe procedural memory, examine its role in various aspects of language, and then present the PDH and relevant evidence across language-related disorders. The PDH has substantial explanatory power, and both basic research and translational implications.

Procedural memory in infancy: Evidence from implicit sequence learning in an eye-tracking paradigm.

Procedural memory underpins the learning of skills and habits. It is often tested in children and adults with sequence learning on the serial reaction time (SRT) task, which involves manual motor control. However, due to infants' slowly developing control of motor actions, most procedures that require motor control cannot be examined in infancy. Here, we investigated procedural memory using an SRT task adapted for infants. During the task, images appeared at one of three locations on a screen, with the location order following a five-item recurring sequence. Three blocks of recurring sequences were followed by a random-order fourth block and finally another block of recurring sequences. Eye movement data were collected for infants (n = 35) and adults (n = 31). Reaction time was indexed by calculating the saccade latencies for orienting to each image as it appeared. The entire protocol took less than 3 min. Sequence learning in the SRT task can be operationalized as an increase in latencies in the random block as compared with the preceding and following sequence blocks. This pattern was observed in both the infants and adults. This study is the first to report learning in an SRT task in infants as young as 9  months. This SRT protocol is a promising procedure for measuring procedural memory in infants.

Sequence learning in the human brain: A functional neuroanatomical meta-analysis of serial reaction time studies.

Sequence learning underlies numerous motor, cognitive, and social skills. Previous models and empirical investigations of sequence learning in humans and non-human animals have implicated cortico-basal ganglia-cerebellar circuitry as well as other structures. To systematically examine the functional neuroanatomy of sequence learning in humans, we conducted a series of neuroanatomical meta-analyses. We focused on the serial reaction time (SRT) task. This task, which is the most widely used paradigm for probing sequence learning in humans, allows for the rigorous control of visual, motor, and other factors. Controlling for these factors (in sequence-random block contrasts), sequence learning yielded consistent activation only in the basal ganglia, across the striatum (anterior/mid caudate nucleus and putamen) and the globus pallidus. In contrast, when visual, motor, and other factors were not controlled for (in a global analysis with all sequence-baseline contrasts, not just sequence-random contrasts), premotor cortical and cerebellar activation were additionally observed. The study provides solid evidence that, at least as tested with the visuo-motor SRT task, sequence learning in humans relies on the basal ganglia, whereas cerebellar and premotor regions appear to contribute to aspects of the task not related to sequence learning itself. The findings have both basic research and translational implications.

A dissociation between syntactic and lexical processing in Parkinson's disease.

Parkinson's disease (PD), which involves the degeneration of dopaminergic neurons in the basal ganglia, has long been associated with motor deficits. Increasing evidence suggests that language can also be impaired, including aspects of syntactic and lexical processing. However, the exact pattern of these impairments remains somewhat unclear, for several reasons. Few studies have examined and compared syntactic and lexical processing within subjects, so their relative deficits remain to be elucidated. Studies have focused on earlier stages of PD, so syntactic and lexical processing in later stages are less well understood. Research has largely probed English and a handful of other European languages, and it is unclear whether findings generalize more broadly. Finally, few studies have examined links between syntactic/lexical impairments and their neurocognitive substrates, such as measures of basal ganglia degeneration or dopaminergic processes. We addressed these gaps by investigating multiple aspects of Farsi syntactic and lexical processing in 40 Farsi native-speaking moderate-to-severe non-demented PD patients, and 40 healthy controls. Analyses revealed equivalent impairments of syntactic comprehension and syntactic judgment, across different syntactic structures. Lexical processing was impaired only for motor function-related objects (e.g., naming 'hammer', but not 'mountain'), in line with findings of PD deficits at naming action verbs as compared to objects, without the verb/noun confound. In direct comparisons between lexical and syntactic tasks, patients were better at naming words like 'mountain' (but not words like 'hammer') than at syntactic comprehension and syntactic judgment. Performance at syntactic comprehension correlated with the last levodopa equivalent dose. No other correlations were found between syntactic/lexical processing measures and either levodopa equivalent dose or hypokinesia, which reflects degeneration of basal ganglia motor-related circuits. All critical significant main effects, interactions, and correlations yielded large effect sizes. The findings elucidate the nature of syntactic and lexical processing impairments in PD.

Language learning in the adult brain: A neuroanatomical meta-analysis of lexical and grammatical learning.

Language learning as an adult, though often difficult, is quite common. Nevertheless, the neural substrates of this process remain unclear, even though identifying them should clarify how language is learned and could lead to improved success at this endeavor. We addressed this gap by conducting multiple neuroanatomical meta-analyses to synthesize the functional neuroimaging literature of language learning. We focused on learning lexical and grammatical knowledge, two building blocks of language. Lexical and grammatical learning yielded overlapping activation in frontal (e.g., BA 44/45) and posterior parietal regions. Only lexical learning showed ventral occipito-temporal (ventral stream) activation, while only grammatical learning showed basal ganglia (anterior caudate/putamen) activation. To further elucidate the neurocognition of grammar learning, we also tested specific predictions of the declarative/procedural model of language. Consistent with the model, grammar learning predicted to rely especially on declarative memory (e.g., with explicit training) showed hippocampal involvement, while grammar learning predicted to rely particularly on procedural memory (e.g., with implicit training) showed anterior caudate/putamen involvement. Finally, given the prevalence of research on artificial grammars, we performed separate analyses of artificial grammar and non-artificial grammar (e.g., miniature language) paradigms. These yielded overlapping activation, especially in BA 44, underscoring the validity of artificial grammars as models for grammar learning in natural languages. In sum, the study elucidates the empirical and theoretical landscape of language learning and has applied implications.

Dissociable implicit sequence learning mechanisms revealed by continuous theta-burst stimulation.

The primary motor area (M1) has been implicated in visuomotor sequence learning. However, it has been suggested there are multiple neural networks that undertake visuomotor sequence learning. The role of M1 in sequence learning may be specific to learning simple sequences comprising predictable associations between adjacent movements. This study aimed to investigate the role of M1 in learning simple ("first-order conditional") and more complex ("second-order conditional") sequences. It was hypothesized that continuous theta burst stimulation (cTBS) over M1 would result in poorer learning of the simple sequence only. Forty-eight healthy adults received cTBS to either M1 or the parietal lobe or received sham cTBS before immediately completing 2 visuomotor sequence learning tasks. The tasks only differed in relation to the structure (i.e., simple vs. complex) of the sequence. The group who received cTBS over M1 demonstrated significantly poorer learning of the simple sequence in comparison to the more complex sequence. The parietal lobe stimulation and sham stimulation did not affect learning of either sequence. This is the first study to show differential involvement of M1 in visuomotor sequence learning, dependent on sequence structure. The study provides new evidence that sequence learning might be supported by different networks in the brain. Specifically, M1 sequence learning appears to be important for learning simple item-to-item associations but not for more complex sequences. (PsycINFO Database Record (c) 2019 APA, all rights reserved).