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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

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.

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.

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.

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]. (C) 2020 The Authors. Published by Elsevier Inc. Funding Agencies|Swedish Research Council (Vetenskapsradet)Swedish Research Council [2016-01033]

Procedural memory, which is rooted in the basal ganglia, underlies the learning and processing of numerous automatized motor and cognitive skills, including in language. Not surprisingly, disorders with basal ganglia abnormalities have been found to show impairments of procedural memory. However, brain abnormalities could also lead to atypically enhanced function. Tourette syndrome (TS) is a candidate for enhanced procedural memory, given previous findings of enhanced TS processing of grammar, which likely depends on procedural memory. We comprehensively examined procedural learning, from memory formation to retention, in children with TS and typically developing (TD) children, who performed an implicit sequence learning task over two days. The children with TS showed sequence learning advantages on both days, despite a regression of sequence knowledge overnight to the level of the TD children. This is the first demonstration of procedural learning advantages in any disorder. The findings may further our understanding of procedural memory and its enhancement. The evidence presented here, together with previous findings suggesting enhanced grammar processing in TS, underscore the dependence of language on a system that also subserves visuomotor sequencing.

Do the mechanisms underlying language in fact serve general-purpose functions that preexist this uniquely human capacity? To address this contentious and empirically challenging issue, we systematically tested the predictions of a well-studied neurocognitive theory of language motivated by evolutionary principles. Multiple metaanalyses were performed to examine predicted links between language and two general-purpose learning systems, declarative and procedural memory. The results tied lexical abilities to learning only in declarative memory, while grammar was linked to learning in both systems in both child first language and adult second language, in specific ways. In second language learners, grammar was associated with only declarative memory at lower language experience, but with only procedural memory at higher experience. The findings yielded large effect sizes and held consistently across languages, language families, linguistic structures, and tasks, underscoring their reliability and validity. The results, which met the predicted pattern, provide comprehensive evidence that language is tied to general-purpose systems both in children acquiring their native language and adults learning an additional language. Crucially, if language learning relies on these systems, then our extensive knowledge of the systems from animal and human studies may also apply to this domain, leading to predictions that might be unwarranted in the more circumscribed study of language. Thus, by demonstrating a role for these systems in language, the findings simultaneously lay a foundation for potentially important advances in the study of this critical domain.

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).

Objective Parkinson's disease (PD), which involves the degeneration of dopaminergic basal ganglia neurons, appears to affect language. We investigated which aspects of language are impaired in PD and what moderates these impairments. Our predictions were based on the declarative/procedural model of language, which links grammar, including in regular inflection, to procedural memory and left-lateralized basal ganglia dopaminergic circuits but links lexical memory, including irregulars, to declarative memory. Because females tend to show declarative memory advantages as compared to males, the model predicts that females rely more on this system for regulars, which can be stored as chunks. Method We probed regular/irregular Farsi past-tense production in 40 Farsi-speaking patients with moderate-to-severe nondemented PD (half female) and 40 normal controls (half female). Results Consistent with our predictions, we found that male, but not female, PD patients showed greater deficits at regular than irregular past-tense production. The females' impairment was mildest for regulars, likely from compensatory storage, as revealed by regular past-tense frequency effects only in females. Right-side hypokinesia (linked to left basal ganglia degeneration) correlated negatively with accuracy of regulars but not irregulars. Similarly, the levodopa equivalent dose of patients' last medication correlated only with regulars. Conclusions The results suggest that language is impaired in PD, but the impairments are moderated by multiple factors, including the type of linguistic knowledge, the degree of left basal ganglia degeneration, dopamine, and sex. The findings underscore the impact of sex on the neurocognition of language and the roles of left basal ganglia dopaminergic circuits in aspects of rule-governed grammar. (PsycINFO Database Record (c) 2019 APA, all rights reserved).

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. (C) 2019 Elsevier Inc. All rights reserved. Funding Agencies|Swedish Research Council (Vetenskapsradet)Swedish Research Council [2016-01033]

The declarative/procedural (DP) model posits that the learning, storage, and use of language critically depend on two learning and memory systems in the brain: declarative memory and procedural memory. Thus, on the basis of independent research on the memory systems, the model can generate specific and often novel predictions for language. Till now most such predictions and ensuing empirical work have been motivated by research on the neurocognition of the two memory systems. However, there is also a large literature on techniques that enhance learning and memory. The DP model provides a theoretical framework for predicting which techniques should extend to language learning, and in what circumstances they should apply. In order to lay the neurocognitive groundwork for these predictions, here we first summarize the neurocognitive fundamentals of the two memory systems and briefly lay out the resulting claims of the DP model for both first and second language. We then provide an overview of learning and memory enhancement techniques before focusing on two techniques – spaced repetition and retrieval practice – that have been linked to the memory systems. Next, we present specific predictions for how these techniques should enhance language learning, and review existing evidence, which suggests that they do indeed improve the learning of both first and second language. Finally, we discuss areas of future research and implications for second language pedagogy.

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.

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.

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.

Autism is characterized by language and communication deficits. We investigated grammatical and lexical processes in high-functioning autism by contrasting the production of regular and irregular past-tense forms. Boys with autism and typically-developing control boys did not differ in accuracy or error rates. However, boys with autism were significantly faster than controls at producing rule-governed past-tenses (slip-slipped, plim-plimmed, bring-bringed), though not lexically-dependent past-tenses (bring-brought, squeeze-squeezed, splim-splam). This pattern mirrors previous findings from Tourette syndrome attributed to abnormalities of frontal/basal-ganglia circuits that underlie grammar. We suggest a similar abnormality underlying language in autism. Importantly, even when children with autism show apparently normal language (e.g., in accuracy or with diagnostic instruments), processes and/or brain structures subserving language may be atypical in the disorder.

Language is often assumed to rely on domain-specific neurocognitive substrates. However, this human capacity in fact seems to crucially depend on general-purpose memory systems in the brain. Evidence suggests that lexical memory relies heavily on declarative memory, which is specialized for arbitrary associations and is rooted in temporal lobe structures. The mental grammar instead relies largely on procedural memory, a system that underlies rules and sequences, and is rooted in frontal/basal-ganglia structures. Developmental and adult-onset disorders such as Specific Language Impairment, autism, Tourette syndrome, Parkinson’s disease, Huntington’s disease, and non-fluent aphasia each seem to involve particular grammatical deficits and analogous non-linguistic procedural memory impairments, as well as abnormalities of procedural memory brain structures. Lexical and declarative memory remain relatively intact in these disorders, and may play compensatory roles. In contrast, Alzheimer’s disease, semantic dementia, fluent aphasia and amnesia each affect lexical and declarative memory, and involve abnormalities of declarative memory brain structures, while leaving grammar and procedural memory largely intact. Overall, the evidence suggests that declarative and procedural memory play critical roles in language disorders, as well as in language more generally.

It has been proposed that an impairment of procedural memory underlies a range of linguistic, cognitive and motor impairments observed in developmental dyslexia (DD). However, studies designed to test this hypothesis using the implicit sequence learning paradigm have yielded inconsistent results. A fundamental aspect of procedural learning is that it takes place over an extended time-period that may be divided into distinct stages based on both behavioural characteristics and neural correlates of performance. Yet, no study of implicit sequence learning in children with DD has included learning stages beyond a single practice session. The present study was designed to fill this important gap by extending the investigation to include the effects of overnight consolidation as well as those of further practice on a subsequent day. The results suggest that the most pronounced procedural learning impairment in DD may emerge only after extended practice, in learning stages beyond a single practice session.

Mathematical disability (MD) is a neurodevelopmental disorder affecting math abilities. Here, we propose a new explanatory account of MD, the procedural deficit hypothesis (PDH), which may further our understanding of the disorder. According to the PDH of MD, abnormalities of brain structures subserving the procedural memory system can lead to difficulties with math skills learned in this system, as well as problems with other functions that depend on these brain structures. This brain-based account is motivated in part by the high comorbidity between MD and language disorders such as dyslexia that may be explained by the PDH, and in part by the likelihood that learning automatized math skills should depend on procedural memory. Here, we first lay out the PDH of MD, and present specific predictions. We then examine the existing literature for each prediction, while pointing out weaknesses and gaps to be addressed by future research. Although we do not claim that the PDH is likely to fully explain MD, we do suggest that the hypothesis could have substantial explanatory power, and that it provides a useful theoretical framework that may advance our understanding of the disorder.

Affixal inflectional morphology has been intensively examined as a model of productive aspects of language. Nevertheless, little is known about the neurocognition of the learning and generalization of affixal inflection, or the influence of certain factors that may affect these processes. In an event-related fMRI study, we examined the neurocognition of the learning and generalization of plural inflections in an artificial language, as well as the influence of both affix type frequency (the proportion of words receiving a given affix) and affix predictability (based on phonological cues in the stem). Adult participants were trained in three sessions, and were scanned after the first and last sessions while inflecting trained and untrained words. Untrained words yielded more activation than trained words in medial frontal (including pre-SMA) and left inferior frontal cortices, which have previously shown activation in compositional grammatical processing. A reliance on phonological cues for untrained word inflection correlated positively with pre-SMA activation, but negatively with activation in the pars triangularis. Thus, pre-SMA may be involved in phonological cue-based composition, while the pars triangularis underlies alternative processes. Inflecting trained items yielded activation in the caudate head bilaterally, only in the first session, consistent with a role for procedural memory in learning grammatical regularities. The medial frontal and left inferior regions activated by untrained items were also activated by trained items, but more weakly than untrained items, with weakest activation for trained-items taking the high-frequency affix. This suggests less involvement of compositional processes for inflecting trained than untrained items, and least of all for trained inflected forms with high-frequency affixes, consistent with the storage of such forms (e.g., in declarative memory). Overall, the findings further elucidate the neural bases of the learning and generalization of affixal morphology, and the roles of affix type frequency and affix phonological predictability in these processes. Moreover, the results support and further specify the declarative/procedural model, in particular in adult language learning.

Most research on the neurobiology of language focuses on language alone. However, new biological functions commonly make use of previously existing mechanisms. Thus, language likely relies on previously existing neurobiological substrates, whether or not these have become further specialized for language. The declarative/procedural model posits that language learning, storage, and use depend heavily on declarative and procedural memory. After all, most, if not all, of language must be learned, and these are the two most important learning and memory systems in the brain. Crucially, both systems are well-studied at many levels in humans and animals, leading to numerous independent predictions about language. Here, I first present background on the neurobiology of the two systems, then lay out ensuing predictions for language, and finally examine evidence testing these predictions. The evidence suggests that language indeed depends on declarative and procedural memory, moreover in interesting and informative ways that advance our understanding of the neurobiology of language.

According to the Procedural Deficit Hypothesis (PDH), abnormalities of brain structures underlying procedural memory largely explain the language deficits in children with specific language impairment (SLI). These abnormalities are posited to result in core deficits of procedural memory, which in turn explain the grammar problems in the disorder. The abnormalities are also likely to lead to problems with other, non-procedural functions, such as working memory, that rely at least partly on the affected brain structures. In contrast, declarative memory is expected to remain largely intact, and should play an important compensatory role for grammar. These claims were tested by examining measures of working, declarative and procedural memory in 51 children with SLI and 51 matched typically-developing (TD) children (mean age 10). Working memory was assessed with the Working Memory Test Battery for Children, declarative memory with the Children's Memory Scale, and procedural memory with a visuo-spatial Serial Reaction Time task. As compared to the TD children, the children with SLI were impaired at procedural memory, even when holding working memory constant. In contrast, they were spared at declarative memory for visual information, and at declarative memory in the verbal domain after controlling for working memory and language. Visuo-spatial short-term memory was intact, whereas verbal working memory was impaired, even when language deficits were held constant. Correlation analyses showed neither visuo-spatial nor verbal working memory was associated with either lexical or grammatical abilities in either the SLI or TD children. Declarative memory correlated with lexical abilities in both groups of children. Finally, grammatical abilities were associated with procedural memory in the TD children, but with declarative memory in the children with SLI. These findings replicate and extend previous studies of working, declarative and procedural memory in SLI. Overall, we suggest that the evidence largely supports the predictions of the PDH. (C) 2011 Elsevier Srl. All rights reserved.

Tourette's syndrome (TS) is a developmental disorder characterized by motor and verbal tics. The tics, which are fast and involuntary, result from frontal/basal-ganglia abnormalities that lead to unsuppressed behaviors. Language has not been carefully examined in TS. We tested the processing of two basic aspects of language: idiosyncratic and rule-governed linguistic knowledge. Evidence suggests that idiosyncratic knowledge (e.g., in irregular past tense formation; bring-brought) is stored in a mental lexicon that depends on the temporal-lobe-based declarative memory system that also underlies conceptual knowledge. In contrast, evidence suggests that rule-governed combination (e.g., in regular past tenses; walk+-ed) takes place in a mental grammar that relies on the frontal/basal-ganglia-based procedural memory system, which also underlies motor skills such as how to use a hammer. We found that TS children were significantly faster than typically developing control children in producing rule-governed past tenses (slip-slipped, plim-plimmed, bring-bringed) but not irregular and other unpredictable past tenses (bring-brought, splim-splam). They were also faster than controls in naming pictures of manipulated (hammer) but not non-manipulated (elephant) items. These data were not explained by a wide range of potentially confounding subject- and item-level factors. The results suggest that the processing of procedurally based knowledge, both of grammar and of manipulated objects, is particularly speeded in TS. The frontal/basal-ganglia abnormalities may thus lead not only to tics, but also to a wider range of rapid behaviors, including the cognitive processing of rule-governed forms in language and other types of procedural knowledge.

Specific Language Impairment (SLI) has been explained by two broad classes of hypotheses, which posit either a deficit specific to grammar, or a non-linguistic processing impairment. Here we advance an alternative perspective. According to the Procedural Deficit Hypothesis (PDH), SLI can be largely explained by the abnormal development of brain structures that constitute the procedural memory system. This system, which is composed of a network of inter-connected structures rooted in frontal/basal-ganglia circuits, subserves the learning and execution of motor and cognitive skills. Crucially, recent evidence also implicates this system in important aspects of grammar. The PDH posits that a significant proportion of individuals with SLI suffer from abnormalities of this brain network, leading to impairments of the linguistic and non-linguistic functions that depend on it. In contrast, functions such as lexical and declarative memory, which depend on other brain structures, are expected to remain largely spared. Evidence from an in-depth retrospective examination of the literature is presented. It is argued that the data support the predictions of the PDH, and particularly implicate Broca's area within frontal cortex, and the caudate nucleus within the basal ganglia. Finally, broader implications are discussed, and predictions for future research are presented. It is argued that the PDH forms the basis of a novel and potentially productive perspective on SLI.

What is the interaction between storage and computation in language processing? What is the psychological status of grammatical rules? What are the relative strengths of connectionist and symbolic models of cognition? How are the components of language implemented in the brain? The English past tense has served as an arena for debates on these issues. We defend the theory that irregular past-tense forms are stored in the lexicon, a division of declarative memory, whereas regular forms can be computed by a concatenation rule, which requires the procedural system. Irregulars have the psychological, linguistic and neuropsychological signatures of lexical memory, whereas regulars often have the signatures of grammatical processing. Furthermore, because regular inflection is rule-driven, speakers can apply it whenever memory fails.

Objective: This article reports findings of a meta-analysis and meta-regression summarizing research on implicit sequence learning in individuals with Parkinson’s disease (PD), as measured by the Serial Reaction Time (SRT) task. Method: Following a systematic search of the literature, we analyzed a total of 27 studies, representing data from 505 participants with PD and 460 neurologically intact control participants. Results: Overall, the meta-analysis indicated significantly (p .001) worse sequence learning by the PD group than the control group. The average weighted effect size was found to be .531 (95% CI [.332, .470]), which is a medium effect size. However, moderate to high levels of heterogeneity (differences) were found between study effect sizes (I 2 58%). Meta-regression analysis suggested that presentation of the SRT task under dual task conditions coupled with PD severity or characteristics of the sequence might affect study effect sizes. Conclusions: The meta-analysis provides clear support that learning in procedural memory (procedural learning), which underlies implicit sequence learning in the SRT task, is impaired in PD.

Most research on neurodevelopmental disorders has focused on their abnormalities. However, what remains intact may also be important. Increasing evidence suggests that declarative memory, a critical learning and memory system in the brain, remains largely functional in a number of neurodevelopmental disorders. Because declarative memory remains functional in these disorders, and because it can learn and retain numerous types of information, functions, and tasks, this system should be able to play compensatory roles for multiple types of impairments across the disorders. Here, we examine this hypothesis for specific language impairment, dyslexia, autism spectrum disorder, Tourette syndrome, and obsessive–compulsive disorder. We lay out specific predictions for the hypothesis and review existing behavioral, electrophysiological, and neuroimaging evidence. Overall, the evidence suggests that declarative memory indeed plays compensatory roles for a range of impairments across all five disorders. Finally, we discuss diagnostic, therapeutic and other implications.

Theoretical and empirical aspects of the neural bases of the mental lexicon and the mental grammar in first and second language (L1 and L2) are discussed. It is argued that in L1, the learning, representation, and processing of lexicon and grammar depend on two well-studied brain memory systems. According to the declarative/procedural model, lexical memory depends upon declarative memory, which is rooted in temporal lobe structures, and has been implicated in the learning and use of fact and event knowledge. Aspects of grammar are subserved by procedural memory, which is rooted in left frontal/basal-ganglia structures, and has been implicated in the acquisition and expression of motor and cognitive skills and habits. This view is supported by psycholinguistic and neurolinguistic evidence. In contrast, linguistic forms whose grammatical computation depends upon procedural memory in L1 are posited to be largely dependent upon declarative/lexical memory in L2. They may be either memorized or constructed by explicit rules learned in declarative memory. Thus in L2, such linguistic forms should be less dependent on procedural memory, and more dependent on declarative memory, than in L1. Moreover, this shift to declarative memory is expected to increase with increasing age of exposure to L2, and with less experience (practice) with the language, which is predicted to improve the learning of grammatical rules by procedural memory. A retrospective examination of lesion, neuroimaging, and electrophysiological studies investigating the neural bases of L2 is presented. It is argued that the data from these studies support the predictions of the declarative/procedural model.

Our use of language depends upon two capacities: a mental lexicon of memorized words and a mental grammar of rules that underlie the sequential and hierarchical composition of lexical forms into predictably structured larger words, phrases, and sentences. The declarative/procedural model posits that the lexicon/grammar distinction in language is tied to the distinction between two well-studied brain memory systems. On this view, the memorization and use of at least simple words (those with noncompositional, that is, arbitrary form-meaning pairings) depends upon an associative memory of distributed representations that is subserved by temporal-lobe circuits previously implicated in the learning and use of fact and event knowledge. This "declarative memory" system appears to be specialized for learning arbitrarily related information (i.e., for associative binding). In contrast, the acquisition and use of grammatical rules that underlie symbol manipulation is subserved by frontal/basal-ganglia circuits previously implicated in the implicit (nonconscious) learning and expression of motor and cognitive "skills" and "habits" (e.g., from simple motor acts to skilled game playing). This "procedural" system may be specialized for computing sequences. This novel view of lexicon and grammar offers an alternative to the two main competing theoretical frameworks. It shares the perspective of traditional dual-mechanism theories in positing that the mental lexicon and a symbol-manipulating mental grammar are subserved by distinct computational components that may be linked to distinct brain structures. However, it diverges from these theories where they assume components dedicated to each of the two language capacities (that is, domain-specific) and in their common assumption that lexical memory is a rote list of items. Conversely, while it shares with single-mechanism theories the perspective that the two capacities are subserved by domain-independent computational mechanisms, it diverges from them where they link both capacities to a single associative memory system with broad anatomic distribution. The declarative/procedural model, but neither traditional dual- nor single-mechanism models, predicts double dissociations between lexicon and grammar, with associations among associative memory properties, memorized words and facts, and temporal-lobe structures, and among symbol-manipulation properties, grammatical rule products, motor skills, and frontal/basal-ganglia structures. In order to contrast lexicon and grammar while holding other factors constant, we have focused our investigations of the declarative/procedural model on morphologically complex word forms. Morphological transformations that are (largely) unproductive (e.g., in go-went, solemn-solemnity) are hypothesized to depend upon declarative memory. These have been contrasted with morphological transformations that are fully productive (e.g., in walk-walked, happy-happiness), whose computation is posited to be solely dependent upon grammatical rules subserved by the procedural system. Here evidence is presented from studies that use a range of psycholinguistic and neurolinguistic approaches with children and adults. It is argued that converging evidence from these studies supports the declarative/procedural model of lexicon and grammar.

The production of regular and irregular past tense forms was investigated among the members of an English-speaking family with a hereditary disorder of language. Unlike the control subjects, the family members affected by the disorder failed to generate overregularizations (e.g., digged) or novel regular forms (plammed, crived), whereas they did produce novel irregularizations (crive–crove). They showed word frequency effects for regular past tense forms (looked) and had trouble producing regulars and irregulars (looked, dug). This pattern cannot be easily explained by deficits of articulation or of perceptual processing, by previous simulations of impairments to a single-mechanism system, or by the extended optional infinitive hypothesis. We argue that the pattern is consistent with a three-level explanation. First, we posit a grammatical deficit of rules or morphological paradigms. This may be caused by a dysfunction of a frontal/basal-ganglia “procedural memory” system previously implicated in the implicit learning and use of motor and cognitive skills. Second, in contexts requiring inflection in the normal adult grammar, the affected subjects appear to retrieve word forms as a function of their accessibility and conceptual appropriateness (“conceptual selection”). Their acquisition and use of these word forms may rely on a “declarative memory” system previously implicated in the explicit learning and use of facts and events. Third, a compensatory strategy may be at work. Some family members may have explicitly learned a strategy of adding suffix-like endings to forms retrieved by conceptual selection. The morphological errors of young normal children appear to be similar to those of the affected family members, who may have been left stranded with conceptual selection by a specific developmental arrest. The same underlying deficit may also explain the impaired subjects' difficulties with derivational morphology.

Meta-analysis and meta-regression were used to evaluate whether evidence to date demonstrates deficits in procedural memory in individuals with specific language impairment (SLI), and to examine reasons for inconsistencies of findings across studies. The Procedural Deficit Hypothesis (PDH) proposes that SLI is largely explained by abnormal functioning of the frontal-basal ganglia circuits that support procedural memory. It has also been suggested that declarative memory can compensate for at least some of the problems observed in individuals with SLI. A number of studies have used Serial Reaction Time (SRT) tasks to investigate procedural learning in SLI. In this report, results from eight studies that collectively examined 186 participants with SLI and 203 typically-developing peers were submitted to a meta-analysis. The average mean effect size was .328 (CI95: .071, .584) and was significant. This suggests SLI is associated with impairments of procedural learning as measured by the SRT task. Differences among individual study effect sizes, examined with meta-regression, indicated that smaller effect sizes were found in studies with older participants, and in studies that had a larger number of trials on the SRT task. The contributions of age and SRT task characteristics to learning are discussed with respect to impaired and compensatory neural mechanisms in SLI.

Although learning a second language (L2) as an adult is notoriously difficult, research has shown that adults can indeed attain native language-like brain processing and high proficiency levels. However, it is important to then retain what has been attained, even in the absence of continued exposure to the L2--particularly since periods of minimal or no L2 exposure are common. This event-related potential (ERP) study of an artificial language tested performance and neural processing following a substantial period of no exposure. Adults learned to speak and comprehend the artificial language to high proficiency with either explicit, classroom-like, or implicit, immersion-like training, and then underwent several months of no exposure to the language. Surprisingly, proficiency did not decrease during this delay. Instead, it remained unchanged, and there was an increase in native-like neural processing of syntax, as evidenced by several ERP changes--including earlier, more reliable, and more left-lateralized anterior negativities, and more robust P600s, in response to word-order violations. Moreover, both the explicitly and implicitly trained groups showed increased native-like ERP patterns over the delay, indicating that such changes can hold independently of L2 training type. The results demonstrate that substantial periods with no L2 exposure are not necessarily detrimental. Rather, benefits may ensue from such periods of time even when there is no L2 exposure. Interestingly, both before and after the delay the implicitly trained group showed more native-like processing than the explicitly trained group, indicating that type of training also affects the attainment of native-like processing in the brain. Overall, the findings may be largely explained by a combination of forgetting and consolidation in declarative and procedural memory, on which L2 grammar learning appears to depend. The study has a range of implications, and suggests a research program with potentially important consequences for second language acquisition and related fields.

The Procedural Deficit Hypothesis (PDH) posits that Specific Language Impairment (SLI) can be largely explained by abnormalities of brain structures that subserve procedural memory. The PDH predicts impairments of procedural memory itself, and that such impairments underlie the grammatical deficits observed in the disorder. Previous studies have indeed reported procedural learning impairments in SLI, and have found that these are associated with grammatical difficulties. The present study extends this research by examining the consolidation and longer-term procedural sequence learning in children with SLI. The Alternating Serial Reaction Time (ASRT) task was given to children with SLI and typically-developing (TD) children in an initial learning session and an average of three days later to test for consolidation and longer-term learning. Although both groups showed evidence of initial sequence learning, only the TD children showed clear signs of consolidation, even though the two groups did not differ in longer-term learning. When the children were re-categorized on the basis of grammar deficits rather than broader language deficits, a clearer pattern emerged. Whereas both the grammar impaired and normal grammar groups showed evidence of initial sequence learning, only those with normal grammar showed consolidation and longer-term learning. Indeed, the grammar-impaired group appeared to lose any sequence knowledge gained during the initial testing session. These findings held even when controlling for vocabulary or a broad non-grammatical language measure, neither of which were associated with procedural memory. When grammar was examined as a continuous variable over all children, the same relationships between procedural memory and grammar, but not vocabulary or the broader language measure, were observed. Overall, the findings support and further specify the PDH. They suggest that consolidation and longer-term procedural learning are impaired in SLI, but that these impairments are specifically tied to the grammatical deficits in the disorder. The possibility that consolidation and longer-term learning are problematic in the disorder suggests a locus of potential study for therapeutic approaches. In sum, this study clarifies our understanding of the underlying deficits in SLI, and suggests avenues for further research.

Dear Editor, Processing rule-governed knowledge recruits the procedural memory system rooted in frontal/basal-ganglia circuits, whereas processing memorized idiosyncratic knowledge depends on temporal lobe regions involved in declarative memory (Ullman, 2001). These or related systems likely underlie rule- and memory-based processes in both language and music (Langheim et al., 2002; Patel, 2003; Miranda and Ullman, 2007). Miranda and Ullman (2007) found that “out-of-key” notes – constituting violations of musical tonality rules in familiar and unfamiliar melodies – elicited similar neural responses to grammar rule violations in language, whereas unexpected “in-key” notes in familiar melodies elicited a pattern of brain activity associated with lexical–semantic processing. Tasks requiring detection of violations can reveal whether putative deficits in patients are associated with processing rule-based (procedural) or memorized (declarative) knowledge. We explored these distinct music memory processes in a 31-year-old individual with noteworthy musical aptitude against a backdrop of widespread cognitive impairment and psychotic symptoms (auditory and visual hallucinations, thought broadcasting, and paranoia) due to schizophrenia. The patient was treated with aripiprazole, clozapine, lithium, and sertraline. Extensive neuropsychological testing – as part of the “CBDB/NIH Sibling Study” – revealed performance under the 30th percentile in most domains, including intelligence, graphomotor speed, episodic verbal memory, long-term lexical and semantic retrieval, and working memory. Despite these deficits, the patient played guitar and sang dozens of popular tunes without apparent difficulty. To characterize the specific nature of the patient's spared musical ability, we tested responses to musical violations (task adapted from Miranda, 2007). Sixty melodies (5–18 s in length) were presented: thirty well-known and thirty novel tunes, matched for musical complexity. Half of the melodies contained a note violation (out-of-key or in-key) that lasted 600 ms. Out-of-key violations in both well-known and novel melodies were inconsistent with musical tonality rules. In-key violations in well-known melodies followed rhythmic and tonal rules but violated the familiar (memorized) version of the melody. The patient listened to each melody and either pressed a button marked “bad” as soon as a violation was detected, or waited until the end of the tune and pressed a button marked “good.” After each tune, the patient indicated orally whether or not it was familiar. Only well-known melodies rated by the patient as familiar, and novel melodies rated as unfamiliar, were analyzed. The patient's performance was compared to previously collected data from 29 healthy volunteers: 16 “musicians” (as defined by training and/or performance experience) and 13 “non-musicians” (Miranda, 2007). Musicians were significantly more accurate than non-musicians at detecting each violation type (p

The cortex constituting Broca's area does not exist in isolation. Rather, like other cortical regions, Broca's area is connected to other brain structures, which likely play closely related functional roles. This paper focuses on the basal ganglia, a set of subcortical structures that project through topographically organized "channels" via the thalamus to different frontal regions. It is hypothesized that the basal ganglia project to Broca's area. This circuitry is further posited to encompass at least two channels. One channel can be characterized as subserving procedural memory, while the other underlies the retrieval of knowledge from declarative memory. These hypotheses are supported by both anatomical and functional evidence. Implications and issues for further investigation are discussed.

Women are better than men at verbal memory tasks, such as remembering word lists. These tasks depend on declarative memory. The declarative/procedural model of language, which posits that the lexicon of stored words is part of declarative memory, while grammatical composition of complex forms depends on procedural memory, predicts a female superiority in aspects of lexical memory. Other neurocognitive models of language have not made this prediction. Here we examine the prediction in past-tense over-regularizations (e.g. holded) produced by children. We expected that girls would remember irregular past-tense forms (held) better than boys, and thus would over-regularize less. To our surprise, girls over-regularized far more than boys. We investigated potential explanations for this sex difference. Analyses showed that in girls but not boys, over-regularization rates correlated with measures of the number of similar-sounding regulars (folded, molded). This sex difference in phonological neighborhood effects is taken to suggest that girls tend to produce over-regularizations in associative lexical memory, generalizing over stored neighboring regulars, while boys are more likely to depend upon rule-governed affixation (hold+-ed). The finding is consistent with the hypothesis that, likely due to their superior lexical abilities, females tend to retrieve from memory complex forms (walked) that men generally compose with the grammatical system (walk+-ed). The results suggest that sex may be an important factor in the acquisition and computation of language.

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.

What are the psychological, computational and neural underpinnings of language? Are these neurocognitive correlates dedicated to language? Do different parts of language depend on distinct neurocognitive systems? Here I address these and other issues that are crucial for our understanding of two fundamental language capacities: the memorization of words in the mental lexicon, and the rule-governed combination of words by the mental grammar. According to the declarative/procedural model, the mental lexicon depends on declarative memory and is rooted in the temporal lobe, whereas the mental grammar involves procedural memory and is rooted in the frontal cortex and basal ganglia. I argue that the declarative/procedural model provides a new framework for the study of lexicon and grammar.