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

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

Neurology of affective prosody and its functional-anatomic organization in right hemisphere

Unlike the aphasic syndromes, the organization of affective prosody in brain has remained controversial because affective-prosodic deficits may occur after left or right brain damage. However, different patterns of deficits are observed following left and right brain damage that suggest affective prosody is a dominant and lateralized function of the right hemisphere. Using the Aprosodia Battery, which was developed to differentiate left and right hemisphere patterns of affective-prosodic deficits, functional-anatomic evidence is presented in patients with focal ischemic strokes to support the concepts that (1) affective prosody is a dominant and lateralized function of the right hemisphere, (2) the intrahemispheric organization of affective prosody in the right hemisphere, with the partial exception of Repetition, is analogous to the organization of propositional language in the left hemisphere and (3) the aprosodic syndromes are cortically based as part of evolutionary adaptations underlying human language and communication.     

Oscillatory gamma activity mediates the pathway from socioeconomic status to language acquisition in infancy

BackgroundThe predictable path to child language acquisition is largely constrained by both brain maturation and environmental experience. The synchronized activity of large numbers of neurons gives rise to macroscopic brain oscillations on an electroencephalogram (EEG). It has been found that neural oscillations at rest in the gamma frequency band (25–45 Hz) are associated with development of different cognitive systems, including language. Although the etiology of language is explained by genetically driven brain maturation factors, environment plays a significant role. Specifically, candidate pathways from environment to language development include sociodemographic factors, primarily socioeconomic status (SES) which is likely to exert its effects on language development through other factors, such as parenting style.Despite these assumptions, no studies have so far examined the interrelation between brain maturation factors such as gamma frequency oscillatory activity, environmental factors such as SES, and language acquisition.Aim and methodIn a longitudinal study of 84 Italian typically developing infants, we measured the power of oscillatory gamma activity as mediator between SES and language acquisition. Baseline EEG and information about SES were collected when infants were aged 6 months. Children were followed-up longitudinally to measure expressive vocabulary and Mean Length of Utterance (MLU) at 24 months.ResultsThe mediation model showed that SES is associated with gamma power which, in turn, is associated with expressive language at age 24 months. A higher SES predicted an increase in left central gamma power which, in turn, predicted better language scores.ConclusionsThese results confirm the predictive role of gamma activity oscillatory activity on later language acquisition, suggesting a specific role for these oscillatory mechanisms in language development milestones such as vocabulary development and early word combination. Furthermore, they suggest that SES differences in brain activity may be apparent at early stages of life and affect later language skills. If replicated, our findings could contribute to identifying highest-risk children and may prompt cost-effective preventive/treatment strategies.     

Stuttering: a dynamic motor control disorder

The purpose of this review is to determine what neural mechanisms may be dysfunctional in stuttering. Three sources of evidence are reviewed. First, studies of dynamic inter-relationships among brain regions during normal speech and in persons who stutter (PWS) suggest that the timing of neural activity in different regions may be abnormal in PWS. Second, the brain lesions associated with acquired stuttering are reviewed. These indicate that in a high percentage of cases, the primary speech and language regions are not affected but lesions involve other structures, such as the basal ganglia, which may modulate the primary speech and language regions. Third, to characterize the motor control disorder in stuttering, similarities and differences from focal dystonias such as spasmodic dysphonia (SD) and Tourette’s syndrome (TS) are reviewed. This review indicates that the central control abnormalities in stuttering are not due to disturbance in one particular brain region but rather a system dysfunction that interferes with rapid and dynamic speech processing for production.

Educational objectives: The reader will be able to describe: (1) the similarities and differences between stuttering and other speech motor control disorders, (2) which brain lesions are most likely to produce acquired stuttering in adults, and (3) what type of brain abnormality most likely underlies stuttering.     

A fMRI study of word retrieval in aphasia

The neural mechanisms underlying recovery of cognitive functions are incompletely understood. Aim of this study was to assess, using functional magnetic resonance (fMRI), the pattern of brain activity during covert word retrieval to letter and semantic cues in five aphasic patients after stroke, in order to assess the modifications of brain function which may be related to recovery. Four out of five patients had undergone language recovery, according to standard testing, after at least 6 months of rehabilitation. The cerebral activation of each patient was evaluated and compared with the activation pattern of normal controls studied with the same fMRI paradigm. In the patients, the pattern of brain activation was influenced by the site and extent of the lesion, by the degree of recovery of language, as reflected by task performance outside the scanner, and by task requirements. In the case of word retrieval to letter cues, a good performance was directly related to the activation in Broca's area, or in the right-sided homologue. On the other hand, in the case of semantic fluency, the relationship between performance level and activation was less clear-cut, because of extensive recruitment of frontal areas in patients with defective performance. These findings suggest that the performance in letter fluency is dependent on the integrity of the left inferior frontal cortex, with the participation of the homologous right hemispheric region when the left inferior frontal cortex is entirely of partially damaged. Semantic fluency, which engages the distributed network of semantic memory, is also associated with more extensive patterns of cerebral activation, which however appear to reflect retrieval effort rather than retrieval success.     

From the left to the right: How the brain compensates progressive loss of language function

In normal right-handed subjects language production usually is a function oft the left brain hemisphere. Patients with aphasia following brain damage to the left hemisphere have a considerable potential to compensate for the loss of this function. Sometimes, but not always, areas of the right hemisphere which are homologous to language areas of the left hemisphere in normal subjects are successfully employed for compensation but this integration process may need time to develop. We investigated right-handed patients with left hemisphere brain tumors as a model of continuously progressive brain damage to left hemisphere language areas using functional neuroimaging and transcranial magnetic stimulation (TMS) to identify factors which determine successful compensation of lost language function. Only patients with slowly progressing brain lesions recovered right-sided language function as detected by TMS. In patients with rapidly progressive lesions no right-sided language function was found and language performance was linearly correlated with the lateralization of language related brain activation to the left hemisphere. It can thus be concluded that time is the factor which determines successful integration of the right hemisphere into the language network for compensation of lost left hemisphere language function.     

On the neurobiological investigation of language understanding in context

There are two significant problems in using functional neuroimaging methods to study language. Improving the state of functional brain imaging will depend on understanding how the dependent measure of brain imaging differs from behavioral dependent measures (the "dependent measure problem") and how the activation of the motor system may be confounded with non-motor aspects of processing in certain experimental designs (the "motor output problem"). To address these problems, it may be necessary to shift the focus of language research from the study of linguistic competence to the understanding of language use. This will require investigations of language processing in full multi-modal and environmental context, monitoring of natural behaviors, novel experimental design, and network-based analysis. Such a combined naturalistic approach could lead to tremendous new insights into language and the brain.     

Neural Systems Mediating American Sign Language: Effects of Sensory Experience and Age of Acquisition

ERPs were recorded from deaf and hearing native signers and from hearing subjects who acquired ASL late or not at all as they viewed ASL signs that formed sentences. The results were compared across these groups and with those from hearing subjects reading English sentences. The results suggest that there are constraints on the organization of the neural systems that mediate formal languages and that these are independent of the modality through which language is acquired. These include different specializations of anterior and posterior cortical regions in aspects of grammatical and semantic processing and a bias for the left hemisphere to mediate aspects of mnemonic functions in language. Additionally, the results suggest that the nature and timing of sensory and language experience significantly impact the development of the language systems of the brain. Effects of the early acquisition of ASL include an increased role for the right hemisphere and for parietal cortex and this occurs in both hearing and deaf native signers. An increased role of posterior temporal and occipital areas occurs in deaf native signers only and thus may be attributable to auditory deprivation.     

Disentangling narrow and coarse semantic networks in the brain: The role of computational models of word meaning

There has been a recent boom in research relating semantic space computational models to fMRI data, in an effort to better understand how the brain represents semantic information. In the first study reported here, we expanded on a previous study to examine how different semantic space models and modeling parameters affect the abilities of these computational models to predict brain activation in a data-driven set of 500 selected voxels. The findings suggest that these computational models may contain distinct types of semantic information that relate to different brain areas in different ways. On the basis of these findings, in a second study we conducted an additional exploratory analysis of theoretically motivated brain regions in the language network. We demonstrated that data-driven computational models can be successfully integrated into theoretical frameworks to inform and test theories of semantic representation and processing. The findings from our work are discussed in light of future directions for neuroimaging and computational research.     

The bilingual brain: cerebral representation of languages.

The present article deals with theoretical and experimental aspects of language representation in the multilingual brain. Two general approaches were adopted in the study of the bilingual brain. The study of bilingual aphasics allows us to describe dissociations and double dissociations between the different subcomponents of the various languages. Furthermore, symptoms peculiar to bilingual aphasia were reported (pathological mixing and switching and translations disorders) which allowed the correlation of some abilities specific to bilinguals with particular neurofunctional systems. Another approach to the study of the bilingual brain is of the experimental type, such as electrophysiological investigations (electrocorticostimulation during brain surgery and event-related potentials) and functional neuroanatomy studies (positron emission tomography and functional magnetic resonance imaging). Functional neuroanatomy studies investigated the brain representation of languages when processing lexical and syntactic stimuli and short stories. Neurophysiologic and neuroimaging studies evidenced a similar cerebral representation of L1 and L2 lexicons both in early and late bilinguals. The representation of grammatical aspects of languages seems to be different between the two languages if L2 is acquired after the age of 7, with automatic processes and correctness being lower than those of the native language. These results are in line with a greater representation of the two lexicons in the declarative memory systems, whereas morphosyntactic aspects may be organized in different systems according to the acquisition vs learning modality.     

Subjective experience is probably not limited to humans: the evidence from neurobiology and behavior

In humans, conscious perception and cognition depends upon the thalamocortical (T-C) complex, which supports perception, explicit cognition, memory, language, planning, and strategic control. When parts of the T-C system are damaged or stimulated, corresponding effects are found on conscious contents and state, as assessed by reliable reports. In contrast, large regions like cerebellum and basal ganglia can be damaged without affecting conscious cognition directly. Functional brain recordings also show robust activity differences in cortex between experimentally matched conscious and unconscious events. This basic anatomy and physiology is highly conserved in mammals and perhaps ancestral reptiles. While language is absent in other species, homologies in perception, memory, and motor cortex suggest that consciousness of one kind or another may be biologically fundamental and phylogenetically ancient. In humans we infer subjective experiences from behavioral and brain evidence. This evidence is quite similar in other mammals and perhaps some non-mammalian species. On the weight of the biological evidence, therefore, subjectivity may be conserved in species with human-like brains and behavior.     

Cortical memory mechanisms and language origins

We have previously proposed that cortical auditory-vocal networks of the monkey brain can be partly homologized with language networks that participate in the phonological loop. In this paper, we suggest that other linguistic phenomena like semantic and syntactic processing also rely on the activation of transient memory networks, which can be compared to active memory networks in the primate. Consequently, short-term cortical memory ensembles that participate in language processing can be phylogenetically tracked to more simple networks present in the primate brain, which became increasingly complex in hominid evolution. This perspective is discussed in the context of two current interpretations of language origins, the "mirror-system hypothesis" and generativist grammar.     

Semantic feature representations for normal and aphasic language

Aphasic and non-neurological patients grouped nouns on the basis of similarity of meaning. These word groupings served as input matrices for hierarchical clustering and multidimensional scaling analyses. The emergent structures suggest that, while the normal adult has a number of levels upon which to organize his lexicon, the adult aphasic's lexicon can be characterized as a set of partial entries that are tied to affective and situational data. The results also suggest that semantic feature representations derived from similarity-of-meaning judgments are of relevance in the study of factors which influence actual language performance.     

Language representation in the human brain: evidence from cortical mapping

The manner in which the human brain processes grammatical-syntactic and lexical-semantic functions has been extensively debated in neurolinguistics. The discreteness and selectivity of the representation of syntactic-morphological properties in the dominant frontal cortex and the representation of the lexical-semantics in the temporo-parietal cortex have been questioned. Three right-handed adult male neurosurgical patients undergoing left craniotomy for intractable seizures were evaluated using various grammatical and semantic tasks during cortical mapping. The sampling of language tasks consisted of trials with stimulation (experimental) and without stimulation (control) from sites in the dominant fronto-temporo-parietal cortex The sampling of language implicated a larger cortical area devoted to language (syntactic-morphological and lexical-semantic) tasks. Further, a large part of the fronto-parieto-temporal cortex was involved with syntactic-morphological functions. However, only the parieto-temporal sites were implicated with the ordering of lexicon in sentence construction. These observations suggest that the representation of language in the human brain may be columnar or multilayered.     

The neural underpinnings of prosody in autism

This study examines the processing of prosodic cues to linguistic structure and to affect, drawing on fMRI and behavioral data from 16 high-functioning adolescents with autism spectrum disorders (ASD) and 11 typically developing controls. Stimuli were carefully matched on pitch, intensity, and duration, while varying systematically in conditions of affective prosody (angry versus neutral speech) and grammatical prosody (questions versus statement). To avoid conscious attention to prosody, which normalizes responses in young people with ASD, the implicit comprehension task directed attention to semantic aspects of the stimuli. Results showed that when perceiving prosodic cues, both affective and grammatical, activation of neural regions was more generalized in ASD than in typical development, and areas recruited reflect heightened reliance on cognitive control, reading of intentions, attentional management, and visualization. This broader recruitment of executive and “mind-reading” brain areas for a relative simple language-processing task may be interpreted to suggest that speakers with high-functioning autism (HFA) have developed less automaticity in language processing and may also suggest that “mind-reading” or theory of mind deficits are intricately bound up in language processing. Data provide support for both a right-lateralized as well as a bilateral model of prosodic processing in typical individuals, depending upon the function of the prosodic information.     

The neural underpinnings of prosody in autism

This study examines the processing of prosodic cues to linguistic structure and to affect, drawing on fMRI and behavioral data from 16 high-functioning adolescents with autism spectrum disorders (ASD) and 11 typically developing controls. Stimuli were carefully matched on pitch, intensity, and duration, while varying systematically in conditions of affective prosody (angry versus neutral speech) and grammatical prosody (questions versus statement). To avoid conscious attention to prosody, which normalizes responses in young people with ASD, the implicit comprehension task directed attention to semantic aspects of the stimuli. Results showed that when perceiving prosodic cues, both affective and grammatical, activation of neural regions was more generalized in ASD than in typical development, and areas recruited reflect heightened reliance on cognitive control, reading of intentions, attentional management, and visualization. This broader recruitment of executive and "mind-reading" brain areas for a relative simple language-processing task may be interpreted to suggest that speakers with high-functioning autism (HFA) have developed less automaticity in language processing and may also suggest that "mind-reading" or theory of mind deficits are intricately bound up in language processing. Data provide support for both a right-lateralized as well as a bilateral model of prosodic processing in typical individuals, depending upon the function of the prosodic information.     

语言理解中句法加工的脑机制

大量研究表明左侧额叶和颞叶的多个脑区与句法加工有关, 已有的理论模型分别从不同的角度对句法加工的脑机制进行了解释。通过文献比较与分析发现, 对句法加工的操作定义、实验任务、被试个体差异以及语言差异等均可能导致研究结果出现差异。在将来的研究中, 可以通过跨语言、多任务比较, 不同层面的句法加工的操作, 在方法上将个体功能定位分析与传统的组分析相结合, 以及从神经网络的角度来更准确和全面地揭示句法加工的脑机制。     

Cognitive processing deficits in reading disabilities: A prefrontal cortical hypothesis

Much research investigating the neuropsychological underpinnings of reading disabilities has emphasized posterior brain regions. However, recent evidence indicates that prefrontal cortex may also play a role. This study investigated cognitive processes that are associated with prefrontal and posterior brain functions. Subjects were 12-year-old reading disabled and nondisabled boys. Discriminant analysis procedures indicated that measures of prefrontal functions distinguished between the two groups better than measures of posterior functions. The results suggest that reading disabled boys have difficulty with cognitive processes involving selective and sustained attention, inhibition of routinized responses, set maintenance, flexibility in generating and testing alternative hypotheses, and phonemically based language production.     

Linguistic recovery after closed head injury

Recovery from acute aphasia after closed head injury was studied in 21 young adults at least 6 months after injury. Three profiles of scores were found using standardized language tests. Persistent expressive and receptive impairment was present in 6 patients who sustained severe diffuse brain injury resulting in global cognitive deficit. Residual expressive impairment, primarily of naming, was associated with mild diffuse brain injury though focal left-hemisphere injury was present in 3 of 6 cases. Patients who recovered to normal levels on all language tests generally had acute neurologic findings consistent with mild diffuse brain injury. Cranial computed tomography at the time of followup frequently disclosed ventricular enlargement.