Contributions of memory circuits to language: The declarative/procedural model

The structure of the brain and the nature of evolution suggest that, despite its uniqueness, language likely depends on brain systems that also subserve other functions. The declarative/procedural (DP) model claims that the mental lexicon of memorized word-specific knowledge depends on the largely temporal-lobe substrates of declarative memory, which underlies the storage and use of knowledge of facts and events. The mental grammar, which subserves the rule-governed combination of lexical items into complex representations, depends on a distinct neural system. This system, which is composed of a network of specific frontal, basal-ganglia, parietal and cerebellar structures, underlies procedural memory, which supports the learning and execution of motor and cognitive skills, especially those involving sequences. The functions of the two brain systems, together with their anatomical, physiological and biochemical substrates, lead to specific claims and predictions regarding their roles in language. These predictions are compared with those of other neurocognitive models of language. Empirical evidence is presented from neuroimaging studies of normal language processing, and from developmental and adult-onset disorders. It is argued that this evidence supports the DP model. It is additionally proposed that "language" disorders, such as specific language impairment and non-fluent and fluent aphasia, may be profitably viewed as impairments primarily affecting one or the other brain system. Overall, the data suggest a new neurocognitive framework for the study of lexicon and grammar

Children's computation of complex linguistic forms: a study of frequency and imageability effects.

This study investigates the storage vs. composition of inflected forms in typically-developing children. Children aged 8-12 were tested on the production of regular and irregular past-tense forms. Storage (vs. composition) was examined by probing for past-tense frequency effects and imageability effects--both of which are diagnostic tests for storage--while controlling for a number of confounding factors. We also examined sex as a factor. Irregular inflected forms, which must depend on stored representations, always showed evidence of storage (frequency and/or imageability effects), not only across all children, but also separately in both sexes. In contrast, for regular forms, which could be either stored or composed, only girls showed evidence of storage. This pattern is similar to that found in previously-acquired adult data from the same task, with the notable exception that development affects which factors influence the storage of regulars in females: imageability plays a larger role in girls, and frequency in women. Overall, the results suggest that irregular inflected forms are always stored (in children and adults, and in both sexes), whereas regulars can be either composed or stored, with their storage a function of various item- and subject-level factors

The neurocognition of developmental disorders of language

Developmental disorders of language include developmental language disorder, motor-speech disorders such as articulation disorder and stuttering, and dyslexia. These disorders have been explained by various accounts, which generally 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 relies 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 multiple 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

Sleep has no critical role in implicit motor sequence learning in young and old adults

The influence of sleep on motor skill consolidation has been a research topic of increasing interest. In this study we distinguished general skill learning from sequence-specific learning in a probabilistic implicit sequence learning task (Alternating Serial Reaction Time) in young and old adults before and after a 12-hour offline interval which did or did not contain sleep (pm-am and am-pm groups respectively). The results showed that general skill learning, as assessed via overall RT, improved offline in both the young and older groups, with the young group improving more than the old. However, the improvement was not sleep-dependent, in that there was no difference between the am-pm and pm-am groups. We did not find sequence-specific offline improvement in either age group for either the am-pm or pm-am groups, suggesting that consolidation of this kind of implicit motor sequence learning may not be influenced by sleep

Procedural learning in Tourette syndrome, ADHD, and comorbid Tourette-ADHD: Evidence from a probabilistic sequence learning task

Procedural memory, which is rooted in the basal ganglia, plays an important role in the implicit learning of motor and cognitive skills. Few studies have examined procedural learning in either Tourette syndrome (TS) or Attention Deficit Hyperactivity Disorder (ADHD), despite basal ganglia abnormalities in both of these neurodevelopmental disorders. We aimed to assess procedural learning in children with TS (n = 13), ADHD (n = 22), and comorbid TS-ADHD (n = 20), as well as in typically developing children (n = 21). Procedural learning was measured with a well-studied implicit probabilistic sequence learning task, the alternating serial reaction time task. All four groups showed evidence of sequence learning, and moreover did not differ from each other in sequence learning. This result, from the first study to examine procedural memory across TS, ADHD and comorbid TS-ADHD, is consistent with previous findings of intact procedural learning of sequences in both TS and ADHD. In contrast, some studies have found impaired procedural learning of non-sequential probabilistic categories in TS. This suggests that sequence learning may be spared in TS and ADHD, while at least some other forms of learning in procedural memory are impaired, at least in TS. Our findings indicate that disorders associated with basal ganglia abnormalities do not necessarily show procedural learning deficits, and provide a possible path for more effective diagnostic tools, and educational and training programs

A Neural Dissociation Within Language: Evidence that the Mental Dictionary is Part of Declarative Memory, and that Grammatical Rules are Processed by the Procedural System

Language comprises a lexicon for storing words and a grammar for generating rule-governed forms. Evidence is presented that the lexicon is part of a temporal-parietalhnedial-temporal “declarative memory” system and that granlmatical rules are processed by a frontamasal-ganglia “procedural” system. Patients produced past tenses of regular and novel verbs (looked and plagged), which require an -ed-suffixation rule, and irregular verbs (dug), which are retrieved from memory. Word-finding difficulties in posterior aphasia, and the general declarative memory impairment in Alzheimer's disease, led to more errors with irregular than regular and novel verbs. Grammatical difficulties in anterior aphasia, and the general impairment of procedures in Parkinson's disease, led to the opposite pattern. In contrast to the Parkinson's patients, who showed suppressed motor activity and rule use, Huntington's disease patients showed excess motor activity and rule use, underscoring a role for the basal ganglia in grammatical processing.Psycholog

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

Working memory in older adults declines with age, but is modulated by sex and education

Working memory (WM), which underlies the temporary storage and manipulation of information, is critical for multiple aspects of cognition and everyday life. Nevertheless, research examining WM specifically in older adults remains limited, despite the global rapid increase in human life expectancy. We examined WM in a large sample (N=754) of healthy older adults (aged 58-89) in a non-Western population (Chinese speakers) in Taiwan, on a digit n-back task. We tested the influence not only of age itself and of load (1-back vs. 2-back), but also effects of both sex and education, which have been shown to modulate WM abilities. Mixed-effects regression revealed that, within older adulthood, age negatively impacted WM abilities (with linear, not nonlinear, effects), as did load (worse performance at 2-back). In contrast, education level was positively associated with WM. Moreover, both age and education interacted with sex. With increasing age, males showed a steeper WM decline than females; with increasing education, females showed greater WM gains than males. Together with other findings, the evidence suggests that age, sex, and education all impact WM in older adults, but interact in particular ways. The results have both basic research and translational implications, and are consistent with particular benefits from increased education for women

Learning and overnight retention in declarative memory in specific language impairment

We examined learning and retention in nonverbal and verbal declarative memory in Hungarian children with (n = 21) and without (n = 21) SLI. Recognition memory was tested both 10 minutes and one day after encoding. On nonverbal items, only the children with SLI improved overnight, with no resulting group differences in performance. In the verbal domain, the children with SLI consistently showed worse performance than the typically-developing children, but the two groups showed similar overnight changes. The findings suggest the possibility of spared or even enhanced declarative memory consolidation in SLI