Jörg Bahlmann

Segregating the core computational faculty of human language from working memory

In contrast to simple structures in animal vocal behavior, hierarchical structures such as center-embedded sentences manifest the core computational faculty of human language. Previous artificial grammar learning studies found that the left pars opercularis (LPO) subserves the processing of hierarchical structures. However, it is not clear whether this area is activated by the structural complexity per se or by the increased memory load entailed in processing hierarchical structures. To dissociate the effect of structural complexity from the effect of memory cost, we conducted a functional magnetic resonance imaging study of German sentence processing with a 2-way factorial design tapping structural complexity (with/without hierarchical structure, i.e., center-embedding of clauses) and working memory load (long/short distance between syntactically dependent elements; i.e., subject nouns and their respective verbs). Functional imaging data revealed that the processes for structure and memory operate separately but co-operatively in the left inferior frontal gyrus; activities in the LPO increased as a function of structural complexity, whereas activities in the left inferior frontal sulcus (LIFS) were modulated by the distance over which the syntactic information had to be transferred. Diffusion tensor imaging showed that these 2 regions were interconnected through white matter fibers. Moreover, functional coupling between the 2 regions was found to increase during the processing of complex, hierarchically structured sentences. These results suggest a neuroanatomical segregation of syntax-related aspects represented in the LPO from memory-related aspects reflected in the LIFS, which are, however, highly interconnected functionally and anatomically.

Hierarchical artificial grammar processing engages Broca's area

The present fMRI study investigates the neural basis of hierarchical processing using two types of artificial grammars: one governed by rules of adjacent dependencies and the other by rules of hierarchical dependencies. The adjacent dependency sequences followed the rule (AB)(n), at which simple transitions between two types of syllable categories were generated (e.g. A(1)B(1)A(2)B(2)). The hierarchical syllable sequences followed the rule A(n)B(n), generating a center-embedded structure (e.g. A(2)A(1)B(1)B(2)) the learning of which required the processing of hierarchical dependencies. When comparing the processing of hierarchical dependencies to adjacent dependencies, significantly higher activations were observed in Brocas area and the adjacent rim of the ventral premotor cortex (BA 44/6) in addition to some several other cortical and sub-cortical regions. These results indicate that Brocas area is part of a neural circuit that is responsible for the processing of hierarchical structures in an artificial grammar.

The role of the posterior superior temporal cortex in sentence comprehension

The study investigates to what extent the posterior superior temporal cortex is involved in processing complex sentences. Using functional MRI, we show that hierarchically structured sentences activate the superior temporal cortex bilaterally to greater extent than sentences with a linear structure. The activation in the left hemisphere comprises the superior temporal gyrus and sulcus, whereas the activation in the right hemisphere is confined to the superior temporal sulcus. As earlier studies using similar syntactic structures in semantic-free grammars did not show activation in the superior temporal cortex but instead only in the prefrontal cortex, we conclude that the role of the posterior superior temporal cortex is to integrate lexical–semantic and syntactic information during sentence comprehension.

Hierarchical and Linear Sequence Processing: An Electrophysiological Exploration of Two Different Grammar Types

The present study investigated the processing of two types of artificial grammars by means of event-related brain potentials. Two categories of meaningless CV syllables were applied in each grammar type. The two grammars differed with regard to the type of the underlying rule. The finite-state grammar (FSG) followed the rule (AB)n, thereby generating local transitions between As and Bs (e.g., n = 2, ABAB). The phrase structure grammar (PSG) followed the rule AnBn, thereby generating center-embedded structures in which the first A and the last B embed the middle elements (e.g., n = 2, [A[AB]B]). Two sequence lengths (n = 2, n = 4) were used. Violations of the structures were introduced at different positions of the syllable sequences. Early violations were situated at the beginning of a sequence, and late violations were placed at the end of a sequence. A posteriorly distributed early negativity elicited by violations was present only in FSG. This effect was interpreted as the possible reflection of a violated local expectancy. Moreover, both grammar-type violations elicited a late positivity. This positivity varied as a function of the violation position in PSG, but not in FSG. These findings suggest that the late positivity could reflect difficulty of integration in PSG sequences.

Predicting vocal emotion expressions from the human brain

Speech is an important carrier of emotional information. However, little is known about how different vocal emotion expressions are recognized in a receivers brain. We used multivariate pattern analysis of functional magnetic resonance imaging data to investigate to which degree distinct vocal emotion expressions are represented in the receivers local brain activity patterns. Specific vocal emotion expressions are encoded in a right fronto‐operculo‐temporal network involving temporal regions known to subserve suprasegmental acoustic processes and a fronto‐opercular region known to support emotional evaluation, and, moreover, in left temporo‐cerebellar regions covering sequential processes. The right inferior frontal region, in particular, was found to differentiate distinct emotional expressions. The present analysis reveals vocal emotion to be encoded in a shared cortical network reflected by distinct brain activity patterns. These results shed new light on theoretical and empirical controversies about the perception of distinct vocal emotion expressions at the level of large‐scale human brain signals. Hum Brain Mapp, 2013.

A rostro-caudal gradient of structured sequence processing in the left inferior frontal gyrus

In this paper, we present two novel perspectives on the function of the left inferior frontal gyrus (LIFG). First, a structured sequence processing perspective facilitates the search for functional segregation within the LIFG and provides a way to express common aspects across cognitive domains including language, music and action. Converging evidence from functional magnetic resonance imaging and transcranial magnetic stimulation studies suggests that the LIFG is engaged in sequential processing in artificial grammar learning, independently of particular stimulus features of the elements (whether letters, syllables or shapes are used to build up sequences). The LIFG has been repeatedly linked to processing of artificial grammars across all different grammars tested, whether they include non-adjacent dependencies or mere adjacent dependencies. Second, we apply the sequence processing perspective to understand how the functional segregation of semantics, syntax and phonology in the LIFG can be integrated in the general organization of the lateral prefrontal cortex (PFC). Recently, it was proposed that the functional organization of the lateral PFC follows a rostro-caudal gradient, such that more abstract processing in cognitive control is subserved by more rostral regions of the lateral PFC. We explore the literature from the viewpoint that functional segregation within the LIFG can be embedded in a general rostro-caudal abstraction gradient in the lateral PFC. If the lateral PFC follows a rostro-caudal abstraction gradient, then this predicts that the LIFG follows the same principles, but this prediction has not yet been tested or explored in the LIFG literature. Integration might provide further insights into the functional architecture of the LIFG and the lateral PFC.

Neural Circuits Subserving the Retrieval of Stems and Grammatical Features in Regular and Irregular Verbs

Many languages, including English and Spanish, feature regular (dance → danced) and irregular (catch → caught) inflectional systems. According to psycholinguistic theories, regular and irregular inflections are instantiated either by a single or by two specialized mechanisms. Those theories differ in their assumptions concerning the underlying information necessary for the processing of regular verbs. Whereas single mechanism accounts have stated an increased involvement of phonological processing for regular verbs, dual accounts emphasize the prominence of grammatical information. Using event‐related functional magnetic resonance imaging, we sought to delineate the brain areas involved in the generation of complex verb forms in Spanish. This language has the advantage of isolating specific differences in the regular–irregular contrasts in terms of the number of stems associated with a verb while controlling for compositionality (regular and irregular verbs apply suffixes to be inflected). The present study showed that areas related to grammatical processing are active for both types of verbs (left opercular inferior frontal gyrus). In addition, major differences between regular and irregular verbs were also observed. Several areas of the prefrontal cortex were selectively active for irregular production, presumably reflecting their role in lexical retrieval (bilateral inferior frontal area and dorsolateral prefrontal cortex). Regular verbs, however, showed increased activation in areas related to grammatical processing (anterior superior temporal gyrus/insular cortex) and in the left hippocampus, the latter possibly related to a greater implication of the phonological loop necessary for the reutilization of the same stem shared across all forms in regular verbs. Hum Brain Mapp, 2006.

An fMRI study of canonical and noncanonical word order in German.

Understanding a complex sentence requires the processing of information at different (e.g., phonological, semantic, and syntactic) levels, the intermediate storage of this information and the unification of this information to compute the meaning of the sentence information. The present investigation homed in on two aspects of sentence processing: working memory and reanalysis. Event‐related functional MRI was used in 12 healthy native speakers of German, while they read sentences. Half of the sentences had unambiguous initial noun‐phrases (masculine nominative, masculine accusative) and thus signaled subject‐first (canonical) or object‐first (noncanonical) sentences. Noncanonical unambiguous sentences were supposed to entail greater demand on working memory, because of their more complex syntactic structure. The other half of the sentences had case‐ambiguous initial noun‐phrases (feminine gender). Only the second unambiguous noun‐phrase (eighth position in the sentences) revealed, whether a canonical or noncanonical word order was present. Based on previous data it was hypothesized that ambiguous noncanonical sentences required a recomputation of the sentence, as subjects would initially commit to a subject first reading. In the respective contrasts two main areas of brain activation were observed. Unambiguous noncanonical sentences elicited more activation in left inferior frontal cortex relative unambiguous canonical sentences. This was interpreted in conjunction with the greater demands on working memory in the former condition. For noncanonical ambiguous relative to canonical ambiguous sentences, an activation of the left supramarginal gyrus was revealed, which was interpreted as a reflection of the reanalysis‐requirements induced by this condition. Hum Brain Mapp, 2007.

Learnability of Embedded Syntactic Structures Depends on Prosodic Cues

The ability to process center-embedded structures has been claimed to represent a core function of the language faculty. Recently, several studies have investigated the learning of center-embedded dependencies in artificial grammar settings. Yet some of the results seem to question the learnability of these structures in artificial grammar tasks. Here, we tested under which exposure conditions learning of center-embedded structures in an artificial grammar is possible. We used naturally spoken syllable sequences and varied the presence of prosodic cues. The results suggest that mere distributional information does not suffice for successful learning. Prosodic cues marking the boundaries of the major relevant units, however, can lead to learning success. Thus, our data are consistent with the hypothesis that center-embedded syntactic structures can be learned in artificial grammar tasks if language-like acoustic cues are provided.

Influence of Motivation on Control Hierarchy in the Human Frontal Cortex

The frontal cortex mediates cognitive control and motivation to shape human behavior. It is generally observed that medial frontal areas are involved in motivational aspects of behavior, whereas lateral frontal regions are involved in cognitive control. Recent models of cognitive control suggest a rostro-caudal gradient in lateral frontal regions, such that progressively more rostral (anterior) regions process more complex aspects of cognitive control. How motivation influences such a control hierarchy is still under debate. Although some researchers argue that both systems work in parallel, others argue in favor of an interaction between motivation and cognitive control. In the latter case it is yet unclear how motivation would affect the different levels of the control hierarchy. This was investigated in the present functional MRI study applying different levels of cognitive control under different motivational states (low vs high reward anticipation). Three levels of cognitive control were tested by varying rule complexity: stimulus-response mapping (low-level), flexible task updating (mid-level), and sustained cue-task associations (high-level). We found an interaction between levels of cognitive control and motivation in medial and lateral frontal subregions. Specifically, flexible updating (mid-level of control) showed the strongest beneficial effect of reward and only this level exhibited functional coupling between dopamine-rich midbrain regions and the lateral frontal cortex. These findings suggest that motivation differentially affects the levels of a control hierarchy, influencing recruitment of frontal cortical control regions depending on specific task demands.