Klaus Hoenig

The Sound of Concepts: Four Markers for a Link between Auditory and Conceptual Brain Systems

Traditionally, concepts are conceived as abstract mental entities distinct from perceptual or motor brain systems. However, recent results let assume modality-specific representations of concepts. The ultimate test for grounding concepts in perception requires the fulfillment of the following four markers: conceptual processing during (1) an implicit task should activate (2) a perceptual region (3) rapidly and (4) selectively. Here, we show using functional magnetic resonance imaging and recordings of event-related potentials, that acoustic conceptual features recruit auditory brain areas even when implicitly presented through visual words. Fulfilling the four markers, the findings of our study unequivocally link the auditory and conceptual brain systems: recognition of words denoting objects, for which acoustic features are highly relevant (e.g.,“telephone”), ignited cell assemblies in posterior superior and middle temporal gyri (pSTG/MTG) within 150 ms that were also activated by sound perception. Importantly, activity within a cluster of pSTG/MTG increased selectively as a function of acoustic, but not of visual and action-related feature relevance. The implicitness of the conceptual task, the selective modulation of left pSTG/MTG activity by acoustic feature relevance, the early onset of this activity at 150 ms and its anatomical overlap with perceptual sound processing are four markers for a modality-specific representation of auditory conceptual features in left pSTG/MTG. Our results therefore provide the first direct evidence for a link between perceptual and conceptual acoustic processing. They demonstrate that access to concepts involves a partial reinstatement of brain activity during the perception of objects.

Conceptual flexibility in the human brain: Dynamic recruitment of semantic maps from visual, motor, and motion-related areas

Traditionally, concepts are assumed to be situational invariant mental knowledge entities (conceptual stability), which are represented in a unitary brain system distinct from sensory and motor areas (amodality). However, accumulating evidence suggests that concepts are embodied in perception and action in that their conceptual features are stored within modality-specific semantic maps in the sensory and motor cortex. Nonetheless, the first traditional assumption of conceptual stability largely remains unquestioned. Here, we tested the notion of flexible concepts using functional magnetic resonance imaging and event-related potentials (ERPs) during the verification of two attribute types (visual, action-related) for words denoting artifactual and natural objects. Functional imaging predominantly revealed crossover interactions between category and attribute type in visual, motor, and motion-related brain areas, indicating that access to conceptual knowledge is strongly modulated by attribute type: Activity in these areas was highest when nondominant conceptual attributes had to be verified. ERPs indicated that these category-attribute interactions emerged as early as 116 msec after stimulus onset, suggesting that they reflect rapid access to conceptual features rather than postconceptual processing. Our results suggest that concepts are situational-dependent mental entities. They are composed of semantic features which are flexibly recruited from distributed, yet localized, semantic maps in modality-specific brain regions depending on contextual constraints.

Neural correlates of semantic ambiguity processing during context verification.

Understanding the relevant meaning of a word with different meanings (homonym) in a given context requires activation of the neural representations of the relevant meaning and inhibition of the irrelevant meaning. The cognitive demand of such disambiguation is highest when the dominant, yet contextually irrelevant meaning of a polar homonym must be suppressed. This central process (semantic inhibition) for lexico-semantic ambiguity resolution was monitored with fMRI during semantic context verifications. Twenty-two healthy volunteers decided whether congruent or incongruent target words fitted into the contexts established by preceding sentences. Half of the sentences ended with a homonym, thereby allowing to cross the factors ambiguity and semantic congruency. BOLD increases related to the inhibitory attentional control over non-selected meanings during ambiguity processing occurred in a brain network including left dorsolateral prefrontal cortex (DLPFC), bilateral angular gyrus (AG), bilateral anterior superior temporal gyrus (aSTG) as well as right ventromedial temporal lobe. In left DLPFC (BA 46/9) and left AG (BA 39) BOLD activity to target words of the incongruent-ambiguous condition correlated with the individual amount of semantic interference experienced by the subjects. BOLD increases of incongruent versus congruent semantic verifications occurred in bilateral inferior frontal gyrus. The results of the present study suggest a specific role of left DLPFC and AG in the resolution of semantic interference from contextually inappropriate homonym meanings. These fronto-parietal areas might exert inhibitory control over temporal regions in service of attentional selection between relevant and irrelevant homonym meanings, by creating a sufficient activation difference between their respective representations.

Neuroplasticity of semantic representations for musical instruments in professional musicians

Professional musicians constitute a model par excellence for understanding experience-dependent plasticity in the human brain, particularly in the auditory domain. Their intensive sensorimotor experience with musical instruments has been shown to entail plastic brain alterations in cortical perceptual and motor maps. It remains an important question whether this neuroplasticity might extend beyond basic perceptual and motor functions and even shape higher-level conceptualizations by which we conceive our physical and social world. Here we show using functional magnetic resonance imaging (fMRI) that conceptual processing of visually presented musical instruments activates auditory association cortex encompassing right posterior superior temporal gyrus, as well as adjacent areas in the superior temporal sulcus and the upper part of middle temporal gyrus (pSTG/MTG) only in musicians, but not in musical laypersons. These areas in and adjacent to auditory association cortex were not only recruited by conceptual processing of musical instruments during visual object recognition, but also by auditory perception of real sounds. Hence, the unique intensive experience of musicians with musical instruments establishes a link between auditory perceptual and conceptual brain systems. Experience-driven neuroplasticity in musicians is thus not confined to alterations of perceptual and motor maps, but even leads to the establishment of higher-level semantic representations for musical instruments in and adjacent to auditory association cortex. These findings highlight the eminent importance of sensory and motor experience for acquiring rich concepts.

Speaking in Multiple Languages: Neural Correlates of Language Proficiency in Multilingual Word Production.

The human brain has the fascinating ability to represent and to process several languages. Although the first and further languages activate partially different brain networks, the linguistic factors underlying these differences in language processing have to be further specified. We investigated the neural correlates of language proficiency in a homogeneous sample of multilingual native Ladin speakers from a mountain valley in South Tyrol, Italy, who speak Italian as second language at a high level, and English at an intermediate level. In a constrained word production task under functional magnetic resonance imaging (fMRI), participants had to name pictures of objects in Ladin, Italian and English in separate blocks. Overall, multilingual word production activated a common set of brain areas dedicated to known subcomponents of picture naming. In comparison to English, the fluently spoken languages Ladin and Italian were associated with enhanced right prefrontal activity. In addition, the MR signal in right prefrontal cortex correlated with naming accuracy as a measure of language proficiency. Our results demonstrate the significance of right prefrontal areas for language proficiency. Based on the role of these areas for cognitive control, our findings suggest that right prefrontal cortex supports language proficiency by effectively supervising word retrieval.

Mechanisms underlying flexible adaptation of cognitive control: Behavioral and neuroimaging evidence in a flanker task

Cognitive control can be adapted flexibly according to the conflict level in a given situation. In the Eriksen flanker task, interference evoked by flankers is larger in conditions with a higher, rather than a lower proportion of compatible trials. Such compatibility ratio effects also occur for stimuli presented at two spatial locations suggesting that different cognitive control settings can be simultaneously maintained. However, the conditions and the neural correlates of this flexible adaptation of cognitive control are only poorly understood. In the present study, we further elucidated the mechanisms underlying the simultaneous maintenance of two cognitive control settings. In behavioral experiments, stimuli were presented centrally above and below fixation and hence processed by both hemispheres or lateralized to stimulate hemispheres differentially. The different compatibility ratio at two stimulus locations had a differential influence on the flanker effect in both experiments. In an fMRI experiment, blocks with an identical compatibility ratio at two central spatial locations elicited stronger activity in a network of prefrontal and parietal brain areas, which are known to be involved in conflict resolution and cognitive control, as compared with blocks with a different compatibility ratio at the same spatial locations. This demonstrates that the simultaneous maintenance of two conflicting control settings vs. one single setting does not recruit additional neural circuits suggesting the involvement of one single cognitive control system. Instead a crosstalk between multiple control settings renders adaptation of cognitive control more efficient when only one uniform rather than two different control settings has to be simultaneously maintained.

Dissociating the representation of action- and sound-related concepts in middle temporal cortex.

Modality-specific models of conceptual memory propose close links between concepts and the sensory-motor systems. Neuroimaging studies found, in different subject groups, that action-related and sound-related concepts activated different parts of posterior middle temporal gyrus (pMTG), suggesting a modality-specific representation of conceptual features. However, as these different parts of pMTG are close to each other, it is possible that the observed anatomical difference is merely related to interindividual variability. In the present functional magnetic resonance imaging study, we now investigated within the same participant group a possible conceptual feature-specific organization in pMTG. Participants performed lexical decisions on sound-related (e.g., telephone) and action-related (hammer) words. Sound words elicited higher activity in anterior pMTG adjacent to auditory association cortex, but action-related words did so in posterior pMTG close to motion sensitive areas. These results confirm distinct conceptual representations of sound and action in pMTG, just adjacent to the respective modality-specific cortices.

Suggestion-Induced Modulation of Semantic Priming during Functional Magnetic Resonance Imaging

Using functional magnetic resonance imaging during a primed visual lexical decision task, we investigated the neural and functional mechanisms underlying modulations of semantic word processing through hypnotic suggestions aimed at altering lexical processing of primes. The priming task was to discriminate between target words and pseudowords presented 200 ms after the prime word which was semantically related or unrelated to the target. In a counterbalanced study design, each participant performed the task once at normal wakefulness and once after the administration of hypnotic suggestions to perceive the prime as a meaningless symbol of a foreign language. Neural correlates of priming were defined as significantly lower activations upon semantically related compared to unrelated trials. We found significant suggestive treatment-induced reductions in neural priming, albeit irrespective of the degree of suggestibility. Neural priming was attenuated upon suggestive treatment compared with normal wakefulness in brain regions supporting automatic (fusiform gyrus) and controlled semantic processing (superior and middle temporal gyri, pre- and postcentral gyri, and supplementary motor area). Hence, suggestions reduced semantic word processing by conjointly dampening both automatic and strategic semantic processes.

Semantic Ambiguity Resolution in Patients With Bipolar Disorder—An Event-Related Potential Study

Deficits in inhibitory function are assumed to underlie psychopathology in bipolar disorder (BD), especially in states of mania. A subdomain of inhibition, semantic inhibition (SI), referring to the suppression of irrelevant word meanings, may underlie formal thought disorder, such as flights of ideas. In the present study, we investigated SI in patients with BD during semantic ambiguity resolution using behavioral and event-related potential (ERP) measures. We presented 14 patients with BD with current manic, hypomanic, or mixed clinical states and 28 healthy controls sequentially with word triplets containing either a homonym (e.g., “organ”) or a comparable unambiguous word (e.g., “piano”). Participants were instructed to make a decision whether or not the target word was related to the meaning field of the first two words. The inappropriate homonym meaning had to be inhibited to correctly perform the target decision. In addition to reaction times (RT) and error rates (ER), the N400 ERP component to the target, an electrophysiological index of semantic processing, was analyzed as measure of the amount of SI that had taken place. Analyses of the behavioral data revealed that BD patients exhibited an overall worse performance in terms of RT and ER. In the ERP data, we found differences in N400 amplitude to ambiguous and unambiguous conditions over the right hemisphere in patients with BD depending on target congruence: In incongruent trials, N400 amplitude was smaller in ambiguous than in unambiguous words. In congruent trials, in contrast, N400 amplitude was larger in ambiguous than in unambiguous words. Such ERP differences between ambiguous and unambiguous words were absent in controls. We conclude that N400 amplitude differences in the ambiguous and unambiguous conditions of the BD group may reflect insufficient suppression of irrelevant homonym meanings in the right hemisphere. Disturbed SI processes might contribute to formal thought disorder in BD.