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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.

Although sex differences have been observed in various cognitive domains, there has been little work examining sex differences in the cognition of music. We tested the prediction that women would be better than men at recognizing familiar melodies, since memories of specific melodies are likely to be learned (at least in part) by declarative memory, which shows female advantages. Participants were 24 men and 24 women, with half musicians and half non-musicians in each group. The two groups were matched on age, education, and various measures of musical training. Participants were presented with well-known and novel melodies, and were asked to indicate their recognition of familiar melodies as rapidly as possible. The women were significantly faster than the men in responding, with a large effect size. The female advantage held across musicians and non-musicians, and across melodies with and without commonly associated lyrics, as evidenced by an absence of interactions between sex and these factors. Additionally, the results did not seem to be explained by sex differences in response biases, or in basic auditory or motor processes as tested in a control task. Though caution is warranted given that this is the first study to examine sex differences in familiar melody recognition, the results are consistent with the hypothesis motivating our prediction, namely that declarative memory underlies knowledge about music (particularly about familiar melodies), and that the female advantage at declarative memory may thus lead to female advantages in music cognition (particularly at familiar melody recognition). Additionally, the findings argue against the view that female advantages at tasks involving verbal (or verbalizable) material are due solely to a sex difference specific to the verbal domain. Further, the results may help explain previously-reported cognitive commonalities between music and language: since declarative memory also underlies language, such commonalities may be partly due to a common dependence on this memory system. More generally, because declarative memory is well studied at many levels, evidence that music cognition depends on this system may lead to a powerful research program generating a wide range of novel predictions for the neurocognition of music, potentially advancing the field.

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.

This study examined the contribution of verbal and visual memory to performance on the Family Pictures subtest of the Children's Memory Scale. This subtest purports to assess declarative memory functioning in the visual/nonverbal domain. A total of 115 nine-year-old children participated in this study. Fifty-eight had specific language impairment (SLI), whilst the remaining 57 were typically developing (TD), with no history of language difficulties. Results showed that the children with SLI, who had intact declarative memory for visual but not verbal information, obtained significantly lower scores on the Family Pictures subtest when compared to the TD group. Regression analyses revealed that across the entire sample, individual differences on the Family Pictures subtest was best predicted by a measure of verbal working memory. These results question whether the Family Pictures subtest can be considered a measure of visual memory in pediatric populations. These results have implications for the interpretation of scores on this subtest regarding the nature of the types of neurocognitive difficulties children may exhibit.

OBJECTIVES: We investigated the relative benefit of maternal multiple micronutrient (MMN) supplementation during pregnancy and until 3 months postpartum compared with iron/folic acid supplementation on child development at preschool age (42 months). METHODS: We assessed 487 children of mothers who participated in the Supplementation with Multiple Micronutrients Intervention Trial, a cluster-randomized trial in Indonesia, on tests adapted and validated in the local context measuring motor, language, visual attention/spatial, executive, and socioemotional abilities. Analysis was according to intention to treat. RESULTS: In children of undernourished mothers (mid-upper arm circumference CONCLUSIONS: When pregnant women are undernourished or anemic, provision of MMN supplements can improve the motor and cognitive abilities of their children up to 3.5 years later, particularly for both motor function and visual attention/spatial ability. Maternal MMN but not iron/folic acid supplementation protected children from the detrimental effects of maternal undernutrition on child motor and cognitive development.

A limited number of studies have investigated language in Huntington's disease (HD). These have generally reported abnormalities in rule-governed (grammatical) aspects of language, in both syntax and morphology. Several studies of verbal inflectional morphology in English and French have reported evidence of over-active rule processing, such as over-suffixation errors (e.g., walkeded) and over-regularizations (e.g., digged). Here we extend the investigation to noun inflection in Hungarian, a Finno-Ugric agglutinative language with complex morphology, and to genetically proven pre-symptomatic Huntington's disease (pre-HD). Although individuals with pre-HD have no clinical, motor or cognitive symptoms, the underlying pathology may already have begun, and thus sensitive behavioral measures might reveal already-present impairments. Indeed, in a Hungarian morphology production task, pre-HD patients made both over-suffixation and over-regularization errors. The findings suggest the generality of over-active rule processing in both HD and pre-HD, across languages from different families with different morphological systems, and for both verbal and noun inflection. Because the neuropathology in pre-HD appears to be largely restricted to the caudate nucleus and related structures, the findings further implicate these structures in language, and in rule-processing in particular. Finally, the need for effective treatments in HD, which will likely depend in part on the ability to sensitively measure early changes in the disease, suggests the possibility that inflectional morphology, and perhaps other language measures, may provide useful diagnostic, tracking, and therapeutic tools for assessing and treating early degeneration in pre-HD and HD.

We investigated the influence of English proficiency on ERPs elicited by lexical semantic violations in English sentences, in both native English speakers and native Spanish speakers who learned English in adulthood. All participants were administered a standardized test of English proficiency, and data were analyzed using linear mixed effects (LME) modeling. Relative to native learners, late learners showed reduced amplitude and delayed onset of the N400 component associated with reading semantic violations. As well, after the N400 late learners showed reduced anterior negative scalp potentials and increased posterior potentials. In both native and late learners, N400 amplitudes to semantically appropriate words were larger for people with lower English proficiency. N400 amplitudes to semantic violations, however, were not influenced by proficiency. Although both N400 onset latency and the late ERP effects differed between L1 and L2 learners, neither correlated with proficiency. Different approaches to dealing with the high degree of correlation between proficiency and native/late learner group status are discussed in the context of LME modeling. The results thus indicate that proficiency can modulate ERP effects in both L1 and L2 learners, and for some measures (in this case, N400 amplitude), L1–L2 differences may be entirely accounted for by proficiency. On the other hand, not all effects of L2 learning can be attributed to proficiency. Rather, the differences in N400 onset and the post-N400 violation effects appear to reflect fundamental differences in L1–L2 processing.

It remains unclear whether adult-learned second language (L2) depends on similar or different neurocognitive mechanisms as those involved in first language (L1). We examined whether English past tense forms are computed similarly or differently by L1 and L2 English speakers, and what factors might affect this: regularity (regular vs. irregular verbs), length of L2 exposure (length of residence), age of L2 acquisition (age of arrival), L2 learners’ native language (Chinese vs. Spanish), and sex (male vs. female). Past tense frequency effects were used to examine the type of computation (composition vs. storage/retrieval). The results suggest that irregular past tenses are always stored. Regular past tenses, however, are either composed or stored, as a function of various factors: both sexes store regulars in L2, but only females in L1; greater lengths of residence lead to less dependence on storage, but only in females; higher adult ages of arrival lead to more reliance on storage. The findings suggest that inflected forms can rely on either the same or different mechanisms in L2 as they do in L1, and that this varies as a function of multiple interacting factors.

It is widely believed that adults cannot learn a foreign language in the same way that children learn a first language. However, recent evidence suggests that adult learners of a foreign language can come to rely on native-like language brain mechanisms. Here, we show that the type of language training crucially impacts this outcome. We used an artificial language paradigm to examine longitudinally whether explicit training (that approximates traditional grammar-focused classroom settings) and implicit training (that approximates immersion settings) differentially affect neural (electrophysiological) and behavioral (performance) measures of syntactic processing. Results showed that performance of explicitly and implicitly trained groups did not differ at either low or high proficiency. In contrast, electrophysiological (ERP) measures revealed striking differences between the groups' neural activity at both proficiency levels in response to syntactic violations. Implicit training yielded an N400 at low proficiency, whereas at high proficiency, it elicited a pattern typical of native speakers: an anterior negativity followed by a P600 accompanied by a late anterior negativity. Explicit training, by contrast, yielded no significant effects at low proficiency and only an anterior positivity followed by a P600 at high proficiency. Although the P600 is reminiscent of native-like processing, this response pattern as a whole is not. Thus, only implicit training led to an electrophysiological signature typical of native speakers. Overall, the results suggest that adult foreign language learners can come to rely on native-like language brain mechanisms, but that the conditions under which the language is learned may be crucial in attaining this goal.

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.

This study employed an artificial language learning paradigm together with a combined behavioral/event-related potential (ERP) approach to examine the neurocognition of the processing of gender agreement, an aspect of inflectional morphology that is problematic in adult second language (L2) learning. Subjects learned to speak and comprehend an artificial language under either explicit (classroomlike) or implicit (immersionlike) training conditions. In each group, both noun-article and noun-adjective gender agreement processing were examined behaviorally and with ERPs at both low and higher levels of proficiency. Results showed that the two groups learned the language to similar levels of proficiency but showed somewhat different ERP patterns. At low proficiency, both types of agreement violations (adjective, article) yielded N400s, but only for the group with implicit training. Additionally, noun-adjective agreement elicited a late N400 in the explicit group at low proficiency. At higher levels of proficiency, noun-adjective agreement violations elicited N400s for both the explicit and implicit groups, whereas noun-article agreement violations elicited P600s for both groups. The results suggest that interactions among linguistic structure, proficiency level, and type of training need to be considered when examining the development of aspects of inflectional morphology in L2 acquisition.

This study examines the storage vs. composition of Spanish inflected verbal forms in L1 and L2 speakers of Spanish. L2 participants were selected to have mid-to-advanced proficiency, high classroom experience, and low immersion experience, typical of medium-to-advanced foreign language learners. Participants were shown the infinitival forms of verbs from either Class I (the default class, which takes new verbs) or Classes II and III (non-default classes), and were asked to produce either first-person singular present-tense or imperfect forms, in separate tasks. In the present tense, the L1 speakers showed inflected-form frequency effects (i.e., higher frequency forms were produced faster, which is taken as a reflection of storage) for stem-changing (irregular) verb-forms from both Class I (e.g., pensar-pienso) and Classes II and III (e.g., perder-pierdo), as well as for non-stem-changing (regular) forms in Classes II/III (e.g., vender-vendo), in which the regular transformation does not appear to constitute a default. In contrast, Class I regulars (e.g., pescar-pesco), whose non-stem-changing transformation constitutes a default (e.g., it is applied to new verbs), showed no frequency effects. L2 speakers showed frequency effects for all four conditions (Classes I and II/III, regulars and irregulars). In the imperfect tense, the L1 speakers showed frequency effects for Class II/III (-ía-suffixed) but not Class I (-aba-suffixed) forms, even though both involve non-stem-change (regular) default transformations. The L2 speakers showed frequency effects for both types of forms. The pattern of results was not explained by a wide range of potentially confounding experimental and statistical factors, and does not appear to be compatible with single-mechanism models, which argue that all linguistic forms are learned and processed in associative memory. The findings are consistent with a dual-system view in which both verb class and regularity influence the storage vs. composition of inflected forms. Specifically, the data suggest that in L1, inflected verbal forms are stored (as evidenced by frequency effects) unless they are both from Class I and undergo non-stem-changing default transformations. In contrast the findings suggest that at least these L2 participants may store all inflected verb-forms. Taken together, the results support dual-system models of L1 and L2 processing in which, at least at mid-to-advanced L2 proficiency and lower levels of immersion experience, the processing of rule-governed forms may depend not on L1 combinatorial processes, but instead on memorized representations.

Patients with psychiatric disorders such as schizophrenia commonly present with impaired language. Here we investigate language in schizophrenia with a focus on inflectional morphology, using an intensively studied and relatively well-understood linguistic paradigm. Patients with schizophrenia (n=43) and age-matched healthy control subjects (n=42) were asked to produce past tenses of regular (slip), irregular (swim), and novel (plag) English verbs. Patients were impaired at regulars and novels (slipped, plagged), with relative sparing of irregulars (swam), controlling for numerous subject- and item-specific factors (e.g., IQ, phonological complexity). Additionally, patients' thought-disorder scores significantly predicted their performance at regular and novel (but not irregular) past-tense production. The results support grammatical deficits in schizophrenia, with a relative sparing of lexical memory, and suggest that thought disorder may be linked with grammatical impairments in the disorder.

Studies of English have shown that temporal-lobe patients, including those with Alzheimer's disease, are spared at processing real and novel regular inflected forms (e.g., blick → blicked; walk → walked), but impaired at real and novel irregular forms (e.g., spling → splang; dig → dug). Here we extend the investigation cross-linguistically to the more complex system of Italian verbal morphology, allowing us to probe the generality of the previous findings in English, as well as to test different explanatory accounts of inflectional morphology. We examined the production of real and novel regular and irregular past-participle and present-tense forms by native Italian-speaking healthy control subjects and patients with probable Alzheimer's disease. Compared to the controls, the patients were impaired at inflecting real irregular verbs but not real regular verbs both for past-participle and present-tense forms, but were not impaired at real regular verbs either for past-participle or present-tense forms. For novel past participles, the patients exhibited this same pattern of impaired production of class II (irregular) forms but spared class I (regular) production. In the present tense, patients were impaired at the production of class II forms (which are regular in the present tense), but spared at production of class I (regular) forms. Contrary to the pattern observed in English, the errors made by the patients on irregulars did not reveal a predominance of regularization errors (e.g., dig → digged). The findings thus partly replicate prior findings from English, but also reveal new patterns from a language with a more complex morphological system that includes verb classes (which are not possible to test in English). The demonstration of an irregular deficit following temporal-lobe damage in a language other than English reveals the cross-linguistic generality of the basic effect, while also elucidating important language-specific differences in the neuro-cognitive basis of regular and irregular morphological forms.

Language requires both storage and composition. However, exactly what is retrieved from memory and what is assembled remains controversial, especially for inflected words. Here, "imageability effects" is introduced as a new diagnostic of storage and a complement to frequency effects. In 2 studies of past-tense morphology, more reliable imageability and frequency effects were found on irregulars than on regulars. An interaction with sex was also observed: Males but not females showed more reliable frequency and imageability effects for irregulars than for regulars; females but not males showed signs of storage for regulars, particularly among higher frequency forms. Overall, the findings validate imageability effects as another diagnostic of storage and suggest that the line between storage and computation is not a simple function; rather, it depends upon the interplay of both item- and subject-specific factors.

Language and communication deficits are key diagnostic criteria for autism. However, not all aspects of language are equally affected. Here we present evidence of enhanced performance of a critical aspect of language—word processing—in children with autism. The results have implications for explanatory theories of autism and language, and for the development of therapeutic approaches.

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.

Many micronutrients, including zinc, iodine, and B-vitamins, are needed for the rapid brain development that occurs during gestation and infancy, laying a foundation for lifelong brain function. Fe...

This chapter examines the neurocognition of second language, that is, the neural and cognitive (psychological and computational) bases of late-learned second or subsequent language (L2). The chapter focuses on lexical and grammatical processes. We first summarize the historical context of research related to the neurocognition of L2, and then examine current neurocognitive theories and their predictions. Next, we describe two common neurocognitive measures (Event-Related Potentials and functional Magnetic Resonance Imaging-that is, ERPs and fMRI) and review their extant neurocognitive empirical evidence for L2. Finally, we discuss the instructional relevance of such theories and evidence and suggest future directions for research.

Clahsen and Felser (CF) have written a beautiful and important paper. I applaud their integrative empirical approach, and believe that their theoretical account is largely correct, if not in some of its specific claims, at least in its broader assumptions. CF directly compare their shallow structure hypothesis (SSH) with a model that my colleagues and I have proposed for aspects of the neurocognition of first and second language: the “declarative/procedural” (DP) model. Although some of CF's discussion accurately depicts the DP model and its relation to the data, they also make a few critical errors.Here, I first summarize the DP model in both first language (L1) and adult-learned second language (L2), in order to be able to contrast it with the SSH, and then address the relevant problems in CF. For further details on the DP model and L1, see Ullman (2001a, 2001c, 2004) and Ullman et al. (1997). For the model as it applies to L2, see Ullman (2001b, 2005).

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.

Cunnings (Cunnings) offers an interpretation of L2-L1 sentence processing differences in terms of memory principles. We applaud such cross-domain approaches, which seem likely to significantly elucidate the neurocognition of language. Cunnings attributes sentence processing differences between (adult) high proficiency L2 and L1 speakers to an increased susceptibility to similarity-based retrieval interference, rather than to qualitative L2-L1 processing differences (cf. Clahsen & Felser, 2006). On his account, both L1 and L2 sentence processing depend upon a ‘bipartite’ working memory, which involves maintaining items active by focusing attention on long-term memory representations (Cowan, 2001).

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.

This study evaluates the input-processing deficit/single system and the grammar-specific deficit/dual system models to account for past tense formation in impaired and normal language development. We investigated regular and irregular past tense formation of 60 real and novel regular and irregular verbs in "Grammatical (G)-SLI" children (aged 9:3 to 12:10) and morphological- or vocabulary-matched younger control children. The G-SLI children and language ability (LA) controls showed quantitatively and qualitatively different patterns of performance. The LA controls, but not the G-SLI children, showed a significant advantage of regular over irregular past tense marking for real and novel verbs. Past tense frequency affected the GSLI children, but not the controls' production of regular verbs, even with stem access controlled for. The G-SLI children's production of regular forms was significantly lower than that of the control groups. Frequency and phonological properties had a similar and significant effect...

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.

We used event-related functional magnetic resonance imaging to identify brain regions involved in syntactic and semantic processing. Healthy adult males read well-formed sentences randomly intermixed with sentences which either contained violations of syntactic structure or were semantically implausible. Reading anomalous sentences, as compared to well-formed sentences, yielded distinct patterns of activation for the two violation types. Syntactic violations elicited significantly greater activation than semantic violations primarily in superior frontal cortex. Semantically incongruent sentences elicited greater activation than syntactic violations in the left hippocampal and parahippocampal gyri, the angular gyri bilaterally, the right middle temporal gyrus, and the left inferior frontal sulcus. These results demonstrate that syntactic and semantic processing result in nonidentical patterns of activation, including greater frontal engagement during syntactic processing and larger increases in temporal and temporo-parietal regions during semantic analyses.

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.

What are the computational and representational bases of the mental lexicon of words, and of the rules of grammar which productively combine lexical forms into larger words, phrases and sentences? “Dual-system” theories posit that lexical forms with non-compositional (arbitrary) sound-meaning pairings are stored in memory, whereas compositional structures are subserved by a distinct rule-processing system. “Single-system” theories claim that lexicon and grammar are both subserved by a single associative memory. Investigations of English past tense may help to resolve this controversy. On the dual-system view, irregular past-tense forms (e.g. blowblew) are retrieved from memory, whereas regular past-tense forms (e.g. walk-walked) are produced by the application of an -ed -suffixation rule. On the single-system view, both types of past-tense forms are learned and computed in associative memory. To test these competing theories, acceptability ratings were elicited from native English-speaking adults for regu...

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.

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.

Developmental dyslexia (DD) has previously been associated with a number of cognitive deficits. Little attention has been directed to cognitive functions that remain intact in the disorder, though the investigation and identification of such strengths might be useful for developing new, and improving current, therapeutical interventions. In this study, an old/new recognition memory paradigm was used to examine previously untested aspects of declarative memory in children with DD and typically developing control children. The DD group was not only not impaired at the task, but actually showed superior recognition memory, as compared to the control children. These findings complement previous reports of enhanced cognition in other domains (e. g., visuo-spatial processing) in DD. Possible underlying mechanisms for the observed DD advantage in declarative memory, and the possibility of compensation by this system for reading deficits in dyslexia, are discussed., Funding Agencies:Sven Jerring Foundation Foundation Sunnerdahls Disability Fund United States Department of Health & Human ServicesNational Institutes of Health (NIH) - USA R01 HD049347Mable H. Flory Charitable Trust United States Department of Health & Human ServicesNational Institutes of Health (NIH) - USANIH Eunice Kennedy Shriver National Institute of Child Health & Human Development (NICHD) R01HD049347

There are several errors in the author information. The Corresponding Author should be Martina Hedenius. The email address, martina.hedenius@neuro.uu.se, is correct. The third author's name is spelled incorrectly. The correct name is Per A. Alm. Three authors, Michael T. Ullman, Margareta Jennische and Jonas Persson, should be noted as joint senior authors of this work.

The mental lexicon is rooted in the biology of the brain. Therefore, understanding the biological bases of the lexicon is critical for a full understanding of the lexicon itself. The vast majority of research on the biology of the mental lexicon and other aspects of language has thus far focused on the level of structural brain anatomy. However, the roles of many other substrates, from cells to molecules to genes, must also be elucidated. One can ask a number of different questions about the biocognition of the mental lexicon. This article examines four broad issues, most—but not all—of which have been and continue to be major areas of research: biological substrates, separability, redundancy, and domain specificity. It also discusses brain regions other than the temporal lobes that underlie aspects of lexically related knowledge and processing. In addition, the article considers how acetylcholine and estrogen modulate aspects of the lexicon and conceptual semantics. Finally, it analyses the link between lexical memory and declarative memory.

Although deep brain stimulation (DBS) of the basal ganglia improves motor outcomes in Parkinson's disease (PD), its effects on cognition, including language, remain unclear. This study examined the impact of subthalamic nucleus (STN) DBS on two fundamental capacities of language, grammatical and lexical functions. These functions were tested with the production of regular and irregular past-tenses, which contrast aspects of grammatical (regulars) and lexical (irregulars) processing while controlling for multiple potentially confounding factors. Aspects of the motor system were tested by contrasting the naming of manipulated (motor) and non-manipulated (non-motor) objects. Performance was compared between healthy controls and early-stage PD patients treated with either DBS/medications or medications alone. Patients were assessed on and off treatment, with controls following a parallel testing schedule. STN-DBS improved naming of manipulated (motor) but not non-manipulated (non-motor) objects, as compared to both controls and patients with just medications, who did not differ from each other across assessment sessions. In contrast, STN-DBS led to worse performance at regulars (grammar) but not irregulars (lexicon), as compared to the other two subject groups, who again did not differ. The results suggest that STN-DBS negatively impacts language in early PD, but may be specific in depressing aspects of grammatical and not lexical processing. The finding that STN-DBS affects both motor and grammar (but not lexical) functions strengthens the view that both depend on basal ganglia circuitry, although the mechanisms for its differential impact on the two (improved motor, impaired grammar) remain to be elucidated.

Using event-related potentials (ERPs), we examined the neurocognition of late-learned second language (L2) Spanish in two groups of typical university foreign-language learners (as compared to native (L1) speakers): one group with only one year of college classroom experience, and low-intermediate proficiency (L2 Low), and another group with over three years of college classroom experience as well as 1-2 semesters of immersion experience abroad, and advanced proficiency (L2 Advanced). Semantic violations elicited N400s in all three groups, whereas syntactic word-order violations elicited LAN/P600 responses in the L1 and L2 Advanced groups, but not the L2 Low group. Indeed, the LAN and P600 responses were statistically indistinguishable between the L1 and L2 Advanced groups. The results support and extend previous findings. Consistent with previous research, the results suggest that L2 semantic processing always depends on L1-like neurocognitive mechanisms, whereas L2 syntactic processing initially differs from L1, but can shift to native-like processes with sufficient proficiency or exposure, and perhaps with immersion experience in particular. The findings further demonstrate that substantial native-like brain processing of syntax can be achieved even by typical university foreign-language learners.

Maternal caregiving capacity, which is affected in part by cognition and mood, is crucial for the health of mothers and infants. Few interventions aim to improve maternal and infant health through improving such capacity. Multiple micronutrient (MMN) supplementation may improve maternal cognition and mood, since micronutrients are essential for brain function. We assessed mothers who participated in the Supplementation with Multiple Micronutrients Intervention Trial (SUMMIT), a double-blind cluster-randomized trial in Indonesia comparing MMN supplementation to iron and folic acid (IFA) during pregnancy and until three months postpartum. We adapted a set of well-studied tests of cognition, motor dexterity, and mood to the local context and administered them to a random sample of 640 SUMMIT participants after an average of 25 weeks (SD = 9) of supplementation. Analysis was by intention to treat. Controlling for maternal age, education, and socio-economic status, MMN resulted in a benefit of 0.12 SD on overall cognition, compared to IFA (95%CI 0.03-0.22, p = .010), and a benefit of 0.18 SD on reading efficiency (95%CI 0.02-0.35, p = .031). Both effects were found particularly in anemic (hemoglobin

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.

Background: Evaluating the impact of nutrition interventions on developmental outcomes in developing countries can be challenging since most assessment tests have been produced in and for developed country settings. Such tests may not be valid measures of children's abilities when used in a new context. Aims: We present several principles for the selection, adaptation, and evaluation of tests assessing the developmental outcomes of nutrition interventions in developing countries where standard assessment tests do not exist. We then report the application of these principles for a nutrition trial on the Indonesian island of Lombok. Sample: Three hundred children age 22-55 months in Lombok participated in a series of pilot tests for the purpose of test adaptation and evaluation. Four hundred and eighty-seven 42-month-old children in Lombok were tested on the finalized test battery. Methods: The developmental assessment tests were adapted to the local context and evaluated for a number of psychometric properties, including convergent and discriminant validity, which were measured based on multiple regression models with maternal education, depression, and age predicting each test score. Results: The adapted tests demonstrated satisfactory psychometric properties and the expected pattern of relationships with the three maternal variables. Maternal education significantly predicted all scores but one, maternal depression predicted socio-emotional competence, socio-emotional problems, and vocabulary, while maternal age predicted socio-emotional competence only. Conclusion: Following the methodological principles we present resulted in tests that were appropriate for children in Lombok and informative for evaluating the developmental outcomes of nutritional supplementation in the research context. Following this approach in future studies will help to determine which interventions most effectively improve child development in developing countries.

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-h offline interval which did or did not contain sleep (p.m.-a.m. and a.m.-p.m. groups, respectively). The results showed that general skill learning, as assessed via overall reaction time, 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 a.m.-p.m. and p.m.-a.m. groups. We did not find sequence-specific offline improvement in either age group for the a.m.-either p.m. or p.m.-a.m. groups, suggesting that consolidation of this kind of implicit motor sequence learning may not be influenced by sleep.

Logic has been intertwined with the study of language and meaning since antiquity, and such connections persist in present day research in linguistic theory (formal semantics) and cognitive psychology (e.g., studies of human reasoning). However, few studies in cognitive neuroscience have addressed logical dimensions of sentence-level language processing, and none have directly compared these aspects of processing with syntax and lexical/conceptual-semantics. We used ERPs to examine a violation paradigm involving “Negative Polarity Items” or NPIs (e.g., ever/any), which are sensitive to logical/truth-conditional properties of the environments in which they occur (e.g., presence/absence of negation in: John hasn’t ever been to Paris, versus: John has *ever been to Paris). Previous studies examining similar types of contrasts found a mix of effects on familiar ERP components (e.g., LAN, N400, P600). We argue that their experimental designs and/or analyses were incapable of separating which effects are connected to NPI-licensing violations proper. Our design enabled statistical analyses teasing apart genuine violation effects from independent effects tied solely to lexical/contextual factors. Here unlicensed NPIs elicited a late P600 followed in onset by a late left anterior negativity (or “L-LAN”), an ERP profile which has also appeared elsewhere in studies targeting logical semantics. Crucially, qualitatively distinct ERP-profiles emerged for syntactic and conceptual semantic violations which we also tested here. We discuss how these findings may be linked to previous findings in the ERP literature. Apart from methodological recommendations, we suggest that the study of logical semantics may aid advancing our understanding of the underlying neurocognitive etiology of ERP components.

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