Individual neurophysiological signatures of spontaneous rhythm processing.

When sensory input conveys rhythmic regularity, we can form predictions about the timing of upcoming events. Although rhythm processing capacities differ considerably between individuals, these differences are often obscured by participant- and trial-level data averaging procedures in M/EEG research. Here, we systematically assessed neurophysiological variability displayed by individuals listening to isochronous (1.54 Hz) equitone sequences interspersed with unexpected (amplitude-attenuated) deviant tones. Our approach aimed at revealing time-varying adaptive neural mechanisms for sampling the acoustic environment at multiple timescales. Rhythm tracking analyses confirmed that individuals encode temporal regularities and form temporal expectations, as indicated in delta-band (1.54 Hz) power and its anticipatory phase alignment to expected tone onsets. Zooming into tone- and participant-level data, we further characterized intra- and inter-individual variabilities in phase-alignment across auditory sequences. Further, individual modeling of beta-band tone-locked responses showed that a subset of auditory sequences was sampled rhythmically by superimposing binary (strong-weak; S-w), ternary (S-w-w) and mixed accentuation patterns. In these sequences, neural responses to standard and deviant tones were modulated by a binary accentuation pattern, thus pointing towards a mechanism of dynamic attending. Altogether, the current results point toward complementary roles of delta- and beta-band activity in rhythm processing and further highlight diverse and adaptive mechanisms to track and sample the acoustic environment at multiple timescales, even in the absence of task-specific instructions.

ERPs reveal an iconic relation between sublexical phonology and affective meaning.

Classical linguistic theory assumes that formal aspects, like sound, are not internally related to the meaning of words. However, recent research suggests language might code affective meaning such as threat and alert sublexically. Positing affective phonological iconicity as a systematic organization principle of the German lexicon, we calculated sublexical affective values for sub-syllabic phonological word segments from a large-scale affective lexical German database by averaging valence and arousal ratings of all words any phonological segment appears in. We tested word stimuli with either consistent or inconsistent mappings between lexical affective meaning and sublexical affective values (negative-valence/high-arousal vs. neutral-valence/low-arousal) in an EEG visual-lexical-decision task. A mismatch between sublexical and lexical affective values elicited an increased N400 response. These results reveal that systematic affective phonological iconicity - extracted from the lexicon - impacts the extraction of lexical word meaning during reading.

The Evolution of Rhythm Processing.

Behavioral and brain rhythms in the millisecond-to-second range are central in human music, speech, and movement. A comparative approach can further our understanding of the evolution of rhythm processing by identifying behavioral and neural similarities and differences across cognitive domains and across animal species. We provide an overview of research into rhythm cognition in music, speech, and animal communication. Rhythm has received considerable attention within each individual field, but to date, little integration. This review article on rhythm processing incorporates and extends existing ideas on temporal processing in speech and music and offers suggestions about the neural, biological, and evolutionary bases of human abilities in these domains.

The temporal dynamics of processing emotions from vocal, facial, and bodily expressions.

Face-to-face communication works multimodally. Not only do we employ vocal and facial expressions; body language provides valuable information as well. Here we focused on multimodal perception of emotion expressions, monitoring the temporal unfolding of the interaction of different modalities in the electroencephalogram (EEG). In the auditory condition, participants listened to emotional interjections such as "ah", while they saw mute video clips containing emotional body language in the visual condition. In the audiovisual condition participants saw video clips with matching interjections. In all three conditions, the emotions "anger" and "fear", as well as non-emotional stimuli were used. The N100 amplitude was strongly reduced in the audiovisual compared to the auditory condition, suggesting a significant impact of visual information on early auditory processing. Furthermore, anger and fear expressions were distinct in the auditory but not the audiovisual condition. Complementing these event-related potential (ERP) findings, we report strong similarities in the alpha- and beta-band in the visual and the audiovisual conditions, suggesting a strong visual processing component in the perception of audiovisual stimuli. Overall, our results show an early interaction of modalities in emotional face-to-face communication using complex and highly natural stimuli.

A locus for an auditory processing deficit and language impairment in an extended pedigree maps to 12p13.31-q14.3.

Despite the apparent robustness of language learning in humans, a large number of children still fail to develop appropriate language skills despite adequate means and opportunity. Most cases of language impairment have a complex etiology, with genetic and environmental influences. In contrast, we describe a three-generation German family who present with an apparently simple segregation of language impairment. Investigations of the family indicate auditory processing difficulties as a core deficit. Affected members performed poorly on a nonword repetition task and present with communication impairments. The brain activation pattern for syllable duration as measured by event-related brain potentials showed clear differences between affected family members and controls, with only affected members displaying a late discrimination negativity. In conjunction with psychoacoustic data showing deficiencies in auditory duration discrimination, the present results indicate increased processing demands in discriminating syllables of different duration. This, we argue, forms the cognitive basis of the observed language impairment in this family. Genome-wide linkage analysis showed a haplotype in the central region of chromosome 12 which reaches the maximum possible logarithm of odds ratio (LOD) score and fully co-segregates with the language impairment, consistent with an autosomal dominant, fully penetrant mode of inheritance. Whole genome analysis yielded no novel inherited copy number variants strengthening the case for a simple inheritance pattern. Several genes in this region of chromosome 12 which are potentially implicated in language impairment did not contain polymorphisms likely to be the causative mutation, which is as yet unknown.

Lexicality drives audio-motor transformations in Broca's area.

Broca's area is classically associated with speech production. Recently, Broca's area has also been implicated in speech perception and non-linguistic information processing. With respect to the latter function, Broca's area is considered to be a central area in a network constituting the human mirror system, which maps observed or heard actions onto motor programs to execute analogous actions. These mechanisms share some similarities with Liberman's motor theory, where objects of speech perception correspond to listener's intended articulatory gestures. The aim of the current series of behavioral, TMS and fMRI studies was to test if Broca's area is indeed implicated in such audio-motor transformations. More specifically, using a classical phonological rhyme priming paradigm, we investigated whether the role of Broca's area could be purely phonological or rather, is lexical in nature. In the behavioral baseline study, we found a large priming effect in word prime/target pairs (W-W) and no effect for pseudo-words (PW-PW). Online TMS interference of Broca's area canceled the priming difference between W-W and PW-PW by enhancing the effects for PW-PW. Finally, the fMRI study showed activation of Broca's area for W-W pairs, but not for PW-PW pairs. Our data show that Broca's area plays a significant role in speech perception strongly linked to the lexicality of a stimulus.

An electrophysiological response to different pitch contours in words.

A spoken word with more than one syllable contains a specific stress pattern found to be processed during spoken word recognition. The present study investigated the word's pitch contour as a single auditory parameter that marks stress. Event-related brain potentials (ERPs) were recorded while subjects made decisions to artificially pitch manipulated words. ERPs revealed that pitch contours are discriminated already within the first syllable of a word. Furthermore, behavioral responses for words with incorrect pitch contours were longer than for words with correct pitch contours. The results suggest that the pitch contour is an auditory feature of the spoken word that a listener automatically processes during spoken word recognition.

Procedural learning in Broca's aphasia: dissociation between the implicit acquisition of spatio-motor and phoneme sequences.

Procedural learning of spatio-motor and phoneme sequences was investigated in patients with Broca's and Wernicke's aphasia and age-matched controls. In Experiment 1, participants performed a standard serial reaction task (SRT) in which they manually responded to a repeating sequence of stimulus locations. Both Broca's and Wernicke's aphasics showed intact sequence learning, as indicated by a reliable response time (RT) cost when the repeating sequence was switched to a random sequence. In Experiment 2, Broca's aphasics and controls performed a new serial search task (SST), which allowed us to investigate the learning of a spatio-motor sequence and a phoneme sequence independently from each other. On each trial, four letters were presented visually, followed by a single auditorily presented letter. Participants had to press one of four response keys to indicate the location of the auditory letter in the visual display. The arrangement of the visual letters was changed from trial to trial such that either the key-presses or the auditory letters followed a repeating pattern, while the other sequence was random. While controls learned both the key-press and the phoneme sequences, Broca's aphasics were selectively impaired in learning the phoneme sequence. This dissociation between learning of spatio-motor and phoneme sequences supports the assumption that partially separable brain systems are involved in procedural learning of different types of sequential structures.

Lateralization of prosody during language production: a lesion study.

To examine brain lateralization of prosody during speech, the sentence production of six right-hemisphere-lesion patients and five left-hemisphere-lesion patients was compared to that of seven normal controls using a question-answer paradigm. The task required the prosodic realization of two different syntactic structures under conditions of wide and narrow focus. Acoustical analyses were carried out on F0 and time structure. These analyses revealed a preserved ability in patients to express differences in syntactic structure via prosody. However, there were deficits in distinguishing narrow focus from wide focus. Whereas both right- and left-hemisphere lesions caused impairments in the realization of F0, time structure was mainly impaired in left-hemisphere patients. Therefore, the present results from language production support functional as well as cue-dependent hypotheses of the lateralization of prosodic processing in the brain.

On the locus of the semantic satiation effect: evidence from event-related brain potentials.

The present study sought to determine whether semantic satiation is merely a by-product of adaptation or satiation of upstream, nonsemantic perceptual processes or whether the effect can have a locus in semantic memory. This was done by measuring event-related brain potentials (ERPs) in a semantic word-detection task involving multiple presentations of primes and critical related and unrelated words in three experiments involving visual (Experiment 1) and auditory (Experiments 2A and 2B) stimuli. Primes varied in their type case (Experiment 1) or pitch (Experiment 2B) in order to discourage sensory adaptation. Prime satiation and relatedness of the primes to the critical word had interacting effects on ERP amplitude to critical words, particularly within the time-window of the N400 component. Because numerous studies have indicated a role for the N400 in semantic processing, modulation of the N400 relatedness effect by prime satiation (with little or no contribution from perceptual adaptation) suggests that semantic memory can be directly satiated, rather than the cost to semantic processing necessarily resulting from impoverishment of perceptual inputs.

Language related brain potentials in patients with cortical and subcortical left hemisphere lesions.

The role of the basal ganglia in language processing is currently a matter of discussion. Therefore, patients with left frontal cortical and subcortical lesions involving the basal ganglia as well as normal controls were tested in a language comprehension paradigm. Semantically incorrect, syntactically incorrect and correct sentences were presented auditorily. Subjects were required to listen to the sentences and to judge whether the sentence heard was correct or not. Event-related potentials and reaction times were recorded while subjects heard the sentences. Three different components correlated with different language processes were considered: the so-called N400 assumed to reflect processes of semantic integration; the early left anterior negativity hypothesized to reflect processes of initial syntactic structure building; and a late positivity (P600) taken to reflect second-pass processes including re-analysis and repair. Normal participants showed the expected N400 component for semantically incorrect sentences and an early anterior negativity followed by a P600 for syntactically incorrect sentences. Patients with left frontal cortical lesions displayed an attenuated N400 component in the semantic condition. In the syntactic condition only a late positivity was observed. Patients with lesions of the basal ganglia, in contrast, showed an N400 to semantic violations and an early anterior negativity as well as a P600 to syntactic violations, comparable to normal controls. Under the assumption that the early anterior negativity reflects automatic first-pass parsing processes and the P600 component more controlled second-pass parsing processes, the present results suggest that the left frontal cortex might support early parsing processes, and that specific regions of the basal ganglia, in contrast, may not be crucial for early parsing processes during sentence comprehension.