What artificial grammar learning reveals about the neurobiology of syntax

In this paper we examine the neurobiological correlates of syntax, the processing of structured sequences, by comparing FMRI results on artificial and natural language syntax. We discuss these and similar findings in the context of formal language and computability theory. We used a simple right-linear unification grammar in an implicit artificial grammar learning paradigm in 32 healthy Dutch university students (natural language FMRI data were already acquired for these participants). We predicted that artificial syntax processing would engage the left inferior frontal region (BA 44/45) and that this activation would overlap with syntax-related variability observed in the natural language experiment. The main findings of this study show that the left inferior frontal region centered on BA 44/45 is active during artificial syntax processing of well-formed (grammatical) sequence independent of local subsequence familiarity. The same region is engaged to a greater extent when a syntactic violation is present and structural unification becomes difficult or impossible. The effects related to artificial syntax in the left inferior frontal region (BA 44/45) were essentially identical when we masked these with activity related to natural syntax in the same subjects. Finally, the medial temporal lobe was deactivated during this operation, consistent with the view that implicit processing does not rely on declarative memory mechanisms that engage the medial temporal lobe. In the context of recent FMRI findings, we raise the question whether Broca’s region (or subregions) is specifically related to syntactic movement operations or the processing of hierarchically nested non-adjacent dependencies in the discussion section. We conclude that this is not the case. Instead, we argue that the left inferior frontal region is a generic on-line sequence processor that unifies information from various sources in an incremental and recursive manner, independent of whether there are any processing requirements related to syntactic movement or hierarchically nested structures. In addition, we argue that the Chomsky hierarchy is not directly relevant for neurobiological systems.     

The Relationship Between Artificial and Second Language Learning

Artificial language learning (ALL) experiments have become an important tool in exploring principles of language and language learning. A persistent question in all of this work, however, is whether ALL engages the linguistic system and whether ALL studies are ecologically valid assessments of natural language ability. In the present study, we considered these questions by examining the relationship between performance in an ALL task and second language learning ability. Participants enrolled in a Spanish language class were evaluated using a number of different measures of Spanish ability and classroom performance, which was compared to IQ and a number of different measures of ALL performance. The results show that success in ALL experiments, particularly more complex artificial languages, correlates positively with indices of L2 learning even after controlling for IQ. These findings provide a key link between studies involving ALL and our understanding of second language learning in the classroom.     

A brief review on the use of functional near-infrared spectroscopy (fNIRS) for language imaging studies in human newborns and adults

Upon stimulation, real time maps of cortical hemodynamic responses can be obtained by non-invasive functional near-infrared spectroscopy (fNIRS) which measures changes in oxygenated and deoxygenated hemoglobin after positioning multiple sources and detectors over the human scalp. The current commercially available transportable fNIRS systems have a time resolution of 1-10 Hz, a depth sensitivity of about 1.5 cm, and a spatial resolution of about 1 cm. The goal of this brief review is to report infants, children and adults fNIRS language studies. Since 1998, 60 studies have been published on cortical activation in the brain’s classic language areas in children/adults as well as newborns using fNIRS instrumentations of different complexity. In addition, the basic principles of fNIRS including features, strengths, advantages, and limitations are summarized in terms that can be understood even by non specialists. Future prospects of fNIRS in the field of language processing imaging are highlighted.     

Functional and connectivity changes during working memory in Huntington’s disease: 18 month longitudinal data from the IMAGE-HD study

Background

This study aimed to characterize, for the first time, 18 month longitudinal changes in both functional activation and functional connectivity during working memory in premanifest Huntington’s disease (pre-HD) and symptomatic HD (symp-HD).

Methods

Functional magnetic resonance imaging (fMRI) was used to investigate longitudinal changes in neuronal activity during working memory performance via an N-BACK task (0-BACK and 1-BACK) in 27 pre-HD, 17 symp-HD, and 23 control participants. Whole-brain analysis of activation and region-of-interest analysis of functional connectivity was applied to longitudinal fMRI data collected at baseline and 18 months follow-up.

Results

Compared with controls, the pre-HD group showed significantly increased activation longitudinally during 1-BACK versus 0-BACK in the lateral and medial prefrontal, anterior cingulate, primary motor, and temporal areas cortically, and caudate and putamen subcortically. Pre-HD far from onset, compared with controls, showed further longitudinal increases in the right and left dorsolateral prefrontal cortex (DLPFC). Longitudinal increased activation in anterior cingulate and medial primary motor areas were associated with disease burden in the pre-HD group. Moreover, in pre-HD increased activation over time in primary motor and putamen regions were associated with average response time during 1-BACK performance. During 1-BACK, functional connectivity between the right DLPFC and posterior parietal, anterior cingulate, and caudate was significantly reduced over 18 months only in the pre-HD group.

Conclusions

Longitudinal reductions in connectivity over 18 months may represent an early signature of cortico-cortical and cortico-striatal functional disconnectivity in pre-HD, whereas the concomitant increased cortical and subcortical activation may reflect a compensatory response to the demands for cognitive resources required during task performance. Our findings demonstrate that functional imaging modalities have the potential to serve as sensitive methods for the assessment of cortical and subcortical responses to future treatment measures.     

Broca’s aphasia, verbs and the mental lexicon

Verb production is notoriously difficult for individuals with Broca’s aphasia, both at the word and at the sentence level. An intriguing question is at which level in the speech production these problems arise. The aim of the present study is to identify the functional locus of the impairment that results in verb production deficits in Broca’s aphasia. Levelt’s (1989) model is used as a theoretical framework for this study. Two experiments have been conducted, one on verb movement and one on verbs with alternating transitivity. The results suggest that the functional impairment in Broca’s aphasia should be located in Levelt’s “grammatical encoder.”     

Linguistic correlates of asymmetric motor symptom severity in Parkinson’s Disease

Asymmetric motor severity is common in Parkinson’s Disease (PD) and provides a method for examining the neurobiologic mechanisms underlying cognitive and linguistic deficits associated with the disorder. In the present research, PD participants (N = 31) were assessed in terms of the asymmetry of their motor symptoms. Interviews with the participants were analyzed with the Linguistic Inquiry and Word Count (LIWC) program. Three measures of linguistic complexity – the proportion of verbs, proportion of function words, and sentence length – were found to be affected by symptom asymmetry. Greater left-side motor severity (and hence greater right-hemisphere dysfunction) was associated with the production of significantly fewer verbs, function words, and shorter sentences. Hence, the production of linguistic complexity in a natural language context was associated with relatively greater right hemisphere involvement. The potential neurobiological mechanisms underlying this effect are discussed.     

Parkinson’s disease disrupts both automatic and controlled processing of action verbs

The problem of how word meaning is processed in the brain has been a topic of intense investigation in cognitive neuroscience. While considerable correlational evidence exists for the involvement of sensory-motor systems in conceptual processing, it is still unclear whether they play a causal role. We investigated this issue by comparing the performance of patients with Parkinson’s disease (PD) with that of age-matched controls when processing action and abstract verbs. To examine the effects of task demands, we used tasks in which semantic demands were either implicit (lexical decision and priming) or explicit (semantic similarity judgment). In both tasks, PD patients’ performance was selectively impaired for action verbs (relative to controls), indicating that the motor system plays a more central role in the processing of action verbs than in the processing of abstract verbs. These results argue for a causal role of sensory-motor systems in semantic processing.     

Affine representations in psychophysics and the near-miss to Weber’s law

A theoretical account for the near-miss to Weber’s law in the form of a power function, with a special emphasis on the interpretation of the exponent, was proposed by Falmagne [Falmagne, J.-C. (1985). Elements of psychophysical theory. New York: Oxford University Press] within the framework of a subtractive representation, P(x,y)=F(u(x)−g(y)). In this paper, we examine a more general affine representation, P(x,y)=F(u(x)h(y)+g(y)). We first obtain a uniqueness theorem for the affine representation. We then study the conditions that force an affine representation to degenerate to a subtractive one. Part of that study involves the case for which two different affine representations co-exist for the same data. We also show that the balance condition P(x,y)+P(y,x)=1 constrains an affine representation to be a special kind of subtractive representation, a Fechnerian one. We further show that Falmagne’s power law takes on a special form for a so-called weakly balanced system of probabilities, in which case the affine representation is Fechnerian. Finally, following Iverson [Iverson, G.J. (2006a). Analytical methods in the theory of psychophysical discrimination I: Inequalities, convexity and integration of just noticeable differences. Journal of Mathematical Psychology, 50, 271-282], we generalize the Fechner method to construct the sensory scales in a weakly balanced affine representation by integrating (derivatives of) just noticeable differences.     

The neural correlates of statistical learning in a word segmentation task: An fMRI study

Functional magnetic resonance imaging (fMRI) was used to assess neural activation as participants learned to segment continuous streams of speech containing syllable sequences varying in their transitional probabilities. Speech streams were presented in four runs, each followed by a behavioral test to measure the extent of learning over time. Behavioral performance indicated that participants could discriminate statistically coherent sequences (words) from less coherent sequences (partwords). Individual rates of learning, defined as the difference in ratings for words and partwords, were used as predictors of neural activation to ask which brain areas showed activity associated with these measures. Results showed significant activity in the pars opercularis and pars triangularis regions of the left inferior frontal gyrus (LIFG). The relationship between these findings and prior work on the neural basis of statistical learning is discussed, and parallels to the frontal/subcortical network involved in other forms of implicit sequence learning are considered.     

Parametric effects of syntactic-semantic conflict in Broca's area during sentence processing

The hypothesized role of Broca’s area in sentence processing ranges from domain-general executive function to domain-specific computation that is specific to certain syntactic structures. We examined this issue by manipulating syntactic structure and conflict between syntactic and semantic cues in a sentence processing task. Functional neuroimaging revealed that activation within several Broca’s area regions of interest reflected the parametric variation in syntactic-semantic conflict. These results suggest that Broca’s area supports sentence processing by mediating between multiple incompatible constraints on sentence interpretation, consistent with this area’s well-known role in conflict resolution in other linguistic and non-linguistic tasks.     

Syntactic reconstruction and reanalysis,semantic dead ends,and prefrontal cortex

The left inferior frontal gyrus (LIFG) is crucially has been found to be involved in syntactic processing of various kinds. This study investigates the cortical effects of two types of syntactic processes: (i) Reconstruction in ellipsis (recovery of left-out material given by context, More people have been to Paris than […] to Oslo), using pseudo-elliptical structures (‘dead ends’) as control (More people have been to Paris than I have). (ii) Reanalysis in the face of structural ambiguity in syntactic ‘garden paths’, where the parser initially assigns an incorrect structure and is forced to reanalyze. Reanalysis and reconstruction require additional syntactic processing and were predicted to increase activation in areas otherwise involved in structural computation: LIFG (BA 44, 45), premotor BA 6, and posterior temporal BA 21, 22. This was borne out. The results showed an interaction effect of the types of construction in all three areas reflecting syntactic processing.     

The "Perceptual Wedge Hypothesis" as the basis for bilingual babies' phonetic processing advantage: new insights from fNIRS brain imaging

In a neuroimaging study focusing on young bilinguals, we explored the brains of bilingual and monolingual babies across two age groups (younger 4-6 months, older 10-12 months), using fNIRS in a new event-related design, as babies processed linguistic phonetic (Native English, Non-Native Hindi) and nonlinguistic Tone stimuli. We found that phonetic processing in bilingual and monolingual babies is accomplished with the same language-specific brain areas classically observed in adults, including the left superior temporal gyrus (associated with phonetic processing) and the left inferior frontal cortex (associated with the search and retrieval of information about meanings, and syntactic and phonological patterning), with intriguing developmental timing differences: left superior temporal gyrus activation was observed early and remained stably active over time, while left inferior frontal cortex showed greater increase in neural activation in older babies notably at the precise age when babies’ enter the universal first-word milestone, thus revealing a first-time focalbrain correlate that may mediate a universal behavioral milestone in early human language acquisition. A difference was observed in the older bilingual babies’ resilient neural and behavioral sensitivity to Non-Native phonetic contrasts at a time when monolingual babies can no longer make such discriminations. We advance the “Perceptual Wedge Hypothesis” as one possible explanation for how exposure to greater than one language may alter neural and language processing in ways that we suggest are advantageous to language users. The brains of bilinguals and multilinguals may provide the most powerful window into the full neural “extent and variability” that our human species’ language processing brain areas could potentially achieve.     

Exploration of Environmental and Genetic Risk Factors for Alzheimer’s Disease: The Value of Cross-Cultural Studies

Advances in molecular genetics have revolutionized epidemiological research. It is now possible to combine the techniques of population genetics with research on risk factors to construct genetic-environmental interactive models that explain geographic-ethnic variations in disease rates. Cross-cultural studies involving populations from developing and developed countries offer a unique opportunity for constructing these models by providing a wide diversity of environmental exposures. Results from a comparative Indianapolis-Ibadan study suggest that Alzheimer’s disease incidence rates are lower in Yoruba than in African Americans and that these lower rates may be due to a combination of genetic and environmental, primarily dietary, influences.     

Time processing in children with Tourette’s syndrome

Background

Tourette syndrome (TS) is characterized by dysfunctional connectivity between prefrontal cortex and sub-cortical structures, and altered meso-cortical and/or meso-striatal dopamine release. Since time processing is also regulated by fronto-striatal circuits and modulated by dopaminergic transmission, we hypothesized that time processing is abnormal in TS.

Methods

We compared time processing abilities between nine children with TS-only (i.e. without major psychiatric comorbidities) and 10 age-matched healthy children, employing a time reproduction task in which subjects actively reproduce different temporal intervals, and a time comparison task in which subjects judge whether a test interval is longer or shorter than a reference interval. IQ, sustained and divided attention, and working memory were assessed in both groups using the Leiter International Performance Scale-Revised, and the Digit Span sub-test of the WISC-R.

Results

Children with TS-only reproduced in an overestimated fashion over-second, but not sub-second, time intervals. The precision of over-second intervals reproduction correlated with tic severity, in that the lower the tic severity, the closer the reproduction of over-second time intervals to their real duration. Time reproduction performance did not significantly correlate with IQ, attention and working memory measures in both groups. No differences between groups were documented in the time comparison task.

Conclusions

The improvement of time processing in children with TS-only seems specific for the over-second range of intervals, consistent with an enhancement in the ‘cognitively controlled’ timing system, which mainly processes longer duration intervals, and depends upon dysfunctional connectivity between the basal ganglia and the dorso-lateral prefrontal cortex. The absence of between-group differences on time comparison, moreover, suggests that TS patients manifest a selective improvement of ‘motor’ timing abilities, rather than of perceptual time abilities. Our data also support an enhancement of cognitive control processes in TS children, probably facilitated by effortful tic suppression.     

The role of mathematical skill in sex differences on Raven’s Matrices

This study replicated a previously reported male advantage on certain items of Raven’s Matrices and found no sex differences in performance on other items. We refer to the latter as analytic (1) items and the former as analytic (2) items. Reasons for the male advantage were investigated by correlating scores obtained by male and female high school students on analytic (1) and analytic (2) items with their scores on tests of spatial, verbal and mathematical ability. There were no sex differences in the magnitude of the correlations between scores on analytic (2) items and the two spatial and verbal tests. In contrast, males but not females showed a significantly higher correlation of maths with analytic (2) than with analytic (1). The results suggest the Raven’s Matrices may engage different, more specific cognitive processes in males and more general cognitive processes in females.     

Speech-associated gestures, Broca’s area, and the human mirror system

Speech-associated gestures are hand and arm movements that not only convey semantic information to listeners but are themselves actions. Broca’s area has been assumed to play an important role both in semantic retrieval or selection (as part of a language comprehension system) and in action recognition (as part of a “mirror” or “observation–execution matching” system). We asked whether the role that Broca’s area plays in processing speech-associated gestures is consistent with the semantic retrieval/selection account (predicting relatively weak interactions between Broca’s area and other cortical areas because the meaningful information that speech-associated gestures convey reduces semantic ambiguity and thus reduces the need for semantic retrieval/selection) or the action recognition account (predicting strong interactions between Broca’s area and other cortical areas because speech-associated gestures are goal-direct actions that are “mirrored”). We compared the functional connectivity of Broca’s area with other cortical areas when participants listened to stories while watching meaningful speech-associated gestures, speech-irrelevant self-grooming hand movements, or no hand movements. A network analysis of neuroimaging data showed that interactions involving Broca’s area and other cortical areas were weakest when spoken language was accompanied by meaningful speech-associated gestures, and strongest when spoken language was accompanied by self-grooming hand movements or by no hand movements at all. Results are discussed with respect to the role that the human mirror system plays in processing speech-associated movements.     

Age of acquisition effects on the functional organization of language in the adult brain

Using functional magnetic resonance imaging (fMRI), we neuroimaged deaf adults as they performed two linguistic tasks with sentences in American Sign Language, grammatical judgment and phonemic-hand judgment. Participants’ age-onset of sign language acquisition ranged from birth to 14 years; length of sign language experience was substantial and did not vary in relation to age of acquisition. For both tasks, a more left lateralized pattern of activation was observed, with activity for grammatical judgment being more anterior than that observed for phonemic-hand judgment, which was more posterior by comparison. Age of acquisition was linearly and negatively related to activation levels in anterior language regions and positively related to activation levels in posterior visual regions for both tasks.     

Understanding cooperation in the Prisoner’s Dilemma game

In the standard one-shot Prisoner’s Dilemma game, participants often choose to cooperate, when the optimal strategy is to defect. This puzzling finding has attracted considerable interest both in terms of possible computational frameworks for modeling human judgment, but also regarding the more general debate of human altruism. In this research, we ask how much of human behavior in this task can be explained by a putative bias for cooperative behavior and whether this, in turn, is influenced by personality. We compared performance on the standard task with performance on a matched neutral one; we manipulated the optimal strategy (defect or cooperate); and we manipulated the amount of payoff for responding optimally. Results revealed little evidence for a bias for cooperative behavior, but significant associations with the personality factors of Behavioural Activation System (BAS) Reward Responsivity and Agreeableness were found. These findings are discussed in terms of the attempt to explain judgment in one-shot, Prisoner’s Dilemma tasks with statistical or probabilistic models.     

Testing the effects of end-goal during reach-to-grasp movements in Parkinson’s disease

Previous evidence suggests that hand shaping during reaching is modulated by the presence and the nature of the end-goal following object’s grasp. Here we test whether such modulation is maintained in Parkinson’s disease (PD). Six participants with PD and six healthy participants took part in the study. Participants were requested to reach towards a bottle filled with water, and then: (1) grasp it without performing any subsequent action; (2) grasp it and place it accurately on a target area; (3) grasp it and pour its contents within a container. The results showed that participants shaped their hand differently depending on the presence or absence of an action following object’s grasp. However, the request to perform an action after grasp determined a modulation of hand kinematics which was delayed for PD than for control participants. Further, whereas for control participants the nature of the end-goal determined a modulation of hand shaping, for PD patients such modulation was not evident. Data are discussed in terms of the role played by basal ganglia in implementing anticipatory mechanisms for the control of manipulative activities. We contend that in PD patients these mechanisms are not totally compromised, but their implementation depends on the action information that has to be anticipated.     

Emotion and object processing in Parkinson’s disease

The neuropsychological literature on the processing of emotions in Parkinson’s disease (PD) reveals conflicting evidence about the role of the basal ganglia in the recognition of facial emotions. Hence, the present study had two objectives. One was to determine the extent to which the visual processing of emotions and objects differs in PD. The other was to assess the impact of cognitive load on the processing of these types of information. Thirty-one patients with idiopathic PD (IPD) under dopamine replacement therapy (DRT) were compared to 30 control subjects on emotion and object recognition tasks. Recognition of objects was more accurate and faster than recognition of facial expressions of emotion, for both groups of subjects. In a second experiment using an N-back procedure with the same stimuli—a more demanding task with a higher cognitive load—patients with IPD were as accurate as control subjects in detecting the correct sequential presentation of stimuli, but were much slower in their decision responses. This indicates that IPD patients under DRT are not impaired in encoding emotion or object information, but that they have difficulty with the processing demands of the N-back task. Thus, patients with IPD appear to be more sensitive to cognitive load than to type of information, whether facial emotions or objects. In this perspective, one must consider that a deafferented dopaminergic system has problems processing more complex information before one can posit the existence of deficits affecting a specific type of information.