Michael T. Ullman

A neural dissociation within language: Evidence that the mental dictionary is part of declarative memory, and that grammatical rules are processed by the procedural system

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

A neurocognitive perspective on language: The declarative/procedural model

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.

The past and future of the past tense

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.

Specific language impairment is not specific to language: The procedural deficit hypothesis

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 Brocas 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.

The Neural Basis of Lexicon and Grammar in First and Second Language: The Declarative/Procedural Model.

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.

The declarative/procedural model of lexicon and grammar.

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.

Working, declarative and procedural memory in specific language impairment

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.

An Event-Related fMRI Study of Syntactic and Semantic Violations

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.

Neural correlates of lexicon and grammar: Evidence from the production, reading, and judgment of inflection in aphasia

Are the linguistic forms that are memorized in the mental lexicon and those that are specified by the rules of grammar subserved by distinct neurocognitive systems or by a single computational system with relatively broad anatomic distribution? On a dual-system view, the productive -ed-suffixation of English regular past tense forms (e.g., look-looked) depends upon the mental grammar, whereas irregular forms (e.g., dig-dug) are retrieved from lexical memory. On a single-mechanism view, the computation of both past tense types depends on associative memory. Neurological double dissociations between regulars and irregulars strengthen the dual-system view. The computation of real and novel, regular and irregular past tense forms was investigated in 20 aphasic subjects. Aphasics with non-fluent agrammatic speech and left frontal lesions were consistently more impaired at the production, reading, and judgment of regular than irregular past tenses. Aphasics with fluent speech and word-finding difficulties, and with left temporal/temporo-parietal lesions, showed the opposite pattern. These patterns held even when measures of frequency, phonological complexity, articulatory difficulty, and other factors were held constant. The data support the view that the memorized words of the mental lexicon are subserved by a brain system involving left temporal/temporo-parietal structures, whereas aspects of the mental grammar, in particular the computation of regular morphological forms, are subserved by a distinct system involving left frontal structures.

Inflectional morphology in a family with inherited specific language impairment

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 )o r 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 singlemechanism 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.