Processing references in context: when the polar bear does not meet a polar bear.

Discourse understanding is hampered when missing or conflicting context information is given. In four experiments, we investigated what happens (a) when the definite determiner "the," which presupposes existence and uniqueness, does not find a unique referent in the context or (b) when the appropriate use of the indefinite determiner is violated by the presence of a unique referent (Experiment 1 and Experiment 2). To focus on the time-course of processing the uniqueness presupposition of the definite determiner, we embedded the determiner in different sentence structures and varied the context (Experiment 3 and Experiment 4). Reading time served as an index of processing difficulty in a word-by-word self-paced reading task and acceptability judgments provided hints for a possible repair of a presupposition violation. Our results showed that conflicting and missing context information lowered acceptability ratings and was associated with prolonged reading times. The pattern of results differed depending on the nature of the presupposition (Experiments 1 and 2) and whether supplementing missing context information was possible (Experiment 3 and Experiment 4). Our findings suggest that different cognitive processes come into play when interpreting presuppositions in order to get a meaningful interpretation of a discourse.

The Role of the Dorsolateral Prefrontal Cortex for Speech and Language Processing.

This review article summarizes various functions of the dorsolateral prefrontal cortex (DLPFC) that are related to language processing. To this end, its connectivity with the left-dominant perisylvian language network was considered, as well as its interaction with other functional networks that, directly or indirectly, contribute to language processing. Language-related functions of the DLPFC comprise various aspects of pragmatic processing such as discourse management, integration of prosody, interpretation of nonliteral meanings, inference making, ambiguity resolution, and error repair. Neurophysiologically, the DLPFC seems to be a key region for implementing functional connectivity between the language network and other functional networks, including cortico-cortical as well as subcortical circuits. Considering clinical aspects, damage to the DLPFC causes psychiatric communication deficits rather than typical aphasic language syndromes. Although the number of well-controlled studies on DLPFC language functions is still limited, the DLPFC might be an important target region for the treatment of pragmatic language disorders.

Discourse management during speech perception: A functional magnetic resonance imaging (fMRI) study.

Discourse structures enable us to generate expectations based upon linguistic material that has already been introduced. We investigated how the required cognitive operations such as reference processing, identification of critical items, and eventual handling of violations correlate with neuronal activity within the language network of the brain. To this end, we conducted a functional magnetic resonance imaging (fMRI) study in which we manipulated spoken discourse coherence by using presuppositions (PSPs) that either correspond or fail to correspond to items in preceding context sentences. Definite and indefinite determiners were used as PSP triggers, referring to (non-) uniqueness or (non-) existence of an item. Discourse adequacy was tested by means of a behavioral rating during fMRI. Discourse violations yielded bilateral hemodynamic activation within the inferior frontal gyrus (IFG), the inferior parietal lobe including the angular gyrus (IPL/AG), the pre-supplementary motor area (pre-SMA), and the basal ganglia (BG). These findings illuminate cognitive aspects of PSP processing: (1) a reference process requiring working memory (IFG), (2) retrieval and integration of semantic/pragmatic information (IPL/AG), (3) cognitive control of inconsistency management (pre-SMA/BG) in terms of "successful" comprehension despite PSP violations at the surface. These results provide the first fMRI evidence needed to develop a functional neuroanatomical model for context-dependent sentence comprehension based on the example of PSP processing.

Cortical phase locking to accelerated speech in blind and sighted listeners prior to and after training.

Cross-correlation of magnetoencephalography (MEG) with time courses derived from the speech signal has shown differences in phase-locking between blind subjects able to comprehend accelerated speech and sighted controls. The present training study contributes to disentangle the effects of blindness and training. Both subject groups (baseline: n = 16 blind, 13 sighted; trained: 10 blind, 3 sighted) were able to enhance speech comprehension up to ca. 18 syllables per second. MEG responses phase-locked to syllable onsets were captured in five pre-defined source locations comprising left and right auditory cortex (A1), right visual cortex (V1), left inferior frontal gyrus (IFG) and left pre-supplementary motor area. Phase locking in A1 was consistently increased while V1 showed opposite training effects in blind and sighted subjects. Also the IFG showed some group differences indicating enhanced top-down strategies in sighted subjects while blind subjects may have a more fine-grained bottom-up resolution for accelerated speech.

Reduced Performance During a Sentence Repetition Task by Continuous Theta-Burst Magnetic Stimulation of the Pre-supplementary Motor Area.

The pre-supplementary motor area (pre-SMA) is engaged in speech comprehension under difficult circumstances such as poor acoustic signal quality or time-critical conditions. Previous studies found that left pre-SMA is activated when subjects listen to accelerated speech. Here, the functional role of pre-SMA was tested for accelerated speech comprehension by inducing a transient "virtual lesion" using continuous theta-burst stimulation (cTBS). Participants were tested (1) prior to (pre-baseline), (2) 10 min after (test condition for the cTBS effect), and (3) 60 min after stimulation (post-baseline) using a sentence repetition task (formant-synthesized at rates of 8, 10, 12, 14, and 16 syllables/s). Speech comprehension was quantified by the percentage of correctly reproduced speech material. For high speech rates, subjects showed decreased performance after cTBS of pre-SMA. Regarding the error pattern, the number of incorrect words without any semantic or phonological similarity to the target context increased, while related words decreased. Thus, the transient impairment of pre-SMA seems to affect its inhibitory function that normally eliminates erroneous speech material prior to speaking or, in case of perception, prior to encoding into a semantically/pragmatically meaningful message.

Neuromagnetic signatures of syllable processing in fetuses and infants provide no evidence for habituation.

BACKGROUND: Habituation, as a basic form of learning, is characterized by decreasing amplitudes of neuronal reaction following repeated stimuli. Recent studies indicate that habituation to pure tones of different frequencies occurs in fetuses and infants. AIMS: Neural processing of different syllables in fetuses and infants was investigated. STUDY DESIGN: An auditory habituation paradigm including two different sequences of syllables was presented to each subject. Each sequence consisted of eight syllables (sequence /ba/: 5× /ba/, 1× /bi/ (dishabituator), 2× /ba/; sequence /bi/: 5× /bi/, 1× /ba/ (dishabituator), 2× /bi/). Each subject was stimulated with 140 sequences. Neuromagnetic signatures of auditory-evoked responses (AER) were recorded by fetal magnetoencephalography (fMEG). SUBJECTS: Magnetic brain signals of N=30 fetuses (age: 28-39weeks of gestation) and N=28 infants (age: 0-3months) were recorded. Forty-two of the 60 fetal recordings and 29 of the 58 infant recordings were included in the final analysis. OUTCOME MEASURES: AERs were recorded and amplitudes were normalized to the amplitude of the first stimulus. RESULTS: In both fetuses and infants, the amplitudes of AERs were found not to decrease with repeated stimulation. In infants, however, amplitude of syllable 6 (dishabituator) was significantly increased compared to syllable 5 (p=0.026). CONCLUSIONS: Fetuses and infants showed AERs to syllables. Unlike fetuses, infants showed a discriminative neural response to syllables. Habituation was not observed in either fetuses or infants. These findings could be important for the investigation of early cognitive competencies and may help to gain a better understanding of language acquisition during child development.

The role of the supplementary motor area for speech and language processing.

Apart from its function in speech motor control, the supplementary motor area (SMA) has largely been neglected in models of speech and language processing in the brain. The aim of this review paper is to summarize more recent work, suggesting that the SMA has various superordinate control functions during speech communication and language reception, which is particularly relevant in case of increased task demands. The SMA is subdivided into a posterior region serving predominantly motor-related functions (SMA proper) whereas the anterior part (pre-SMA) is involved in higher-order cognitive control mechanisms. In analogy to motor triggering functions of the SMA proper, the pre-SMA seems to manage procedural aspects of cognitive processing. These latter functions, among others, comprise attentional switching, ambiguity resolution, context integration, and coordination between procedural and declarative memory structures. Regarding language processing, this refers, for example, to the use of inner speech mechanisms during language encoding, but also to lexical disambiguation, syntax and prosody integration, and context-tracking.

Context-dependent impact of presuppositions on early magnetic brain responses during speech perception.

Discourse structure enables us to generate expectations based upon linguistic material that has already been introduced. The present magnetoencephalography (MEG) study addresses auditory perception of test sentences in which discourse coherence was manipulated by using presuppositions (PSP) that either correspond or fail to correspond to items in preceding context sentences with respect to uniqueness and existence. Context violations yielded delayed auditory M50 and enhanced auditory M200 cross-correlation responses to syllable onsets within an analysis window of 1.5s following the PSP trigger words. Furthermore, discourse incoherence yielded suppression of spectral power within an expanded alpha band ranging from 6 to 16Hz. This effect showed a bimodal temporal distribution, being significant in an early time window of 0.0-0.5s following the PSP trigger and a late interval of 2.0-2.5s. These findings indicate anticipatory top-down mechanisms interacting with various aspects of bottom-up processing during speech perception.

Network Modeling for Functional Magnetic Resonance Imaging (fMRI) Signals during Ultra-Fast Speech Comprehension in Late-Blind Listeners.

In many functional magnetic resonance imaging (fMRI) studies blind humans were found to show cross-modal reorganization engaging the visual system in non-visual tasks. For example, blind people can manage to understand (synthetic) spoken language at very high speaking rates up to ca. 20 syllables/s (syl/s). FMRI data showed that hemodynamic activation within right-hemispheric primary visual cortex (V1), bilateral pulvinar (Pv), and left-hemispheric supplementary motor area (pre-SMA) covaried with their capability of ultra-fast speech (16 syllables/s) comprehension. It has been suggested that right V1 plays an important role with respect to the perception of ultra-fast speech features, particularly the detection of syllable onsets. Furthermore, left pre-SMA seems to be an interface between these syllabic representations and the frontal speech processing and working memory network. So far, little is known about the networks linking V1 to Pv, auditory cortex (A1), and (mesio-) frontal areas. Dynamic causal modeling (DCM) was applied to investigate (i) the input structure from A1 and Pv toward right V1 and (ii) output from right V1 and A1 to left pre-SMA. As concerns the input Pv was significantly connected to V1, in addition to A1, in blind participants, but not in sighted controls. Regarding the output V1 was significantly connected to pre-SMA in blind individuals, and the strength of V1-SMA connectivity correlated with the performance of ultra-fast speech comprehension. By contrast, in sighted controls, not understanding ultra-fast speech, pre-SMA did neither receive input from A1 nor V1. Taken together, right V1 might facilitate the "parsing" of the ultra-fast speech stream in blind subjects by receiving subcortical auditory input via the Pv (= secondary visual pathway) and transmitting this information toward contralateral pre-SMA.

Experience-related structural changes of degenerated occipital white matter in late-blind humans - a diffusion tensor imaging study.

Late-blind humans can learn to understand speech at ultra-fast syllable rates (ca. 20 syllables/s), a capability associated with hemodynamic activation of the central-visual system. Thus, the observed functional cross-modal recruitment of occipital cortex might facilitate ultra-fast speech processing in these individuals. To further elucidate the structural prerequisites of this skill, diffusion tensor imaging (DTI) was conducted in late-blind subjects differing in their capability of understanding ultra-fast speech. Fractional anisotropy (FA) was determined as a quantitative measure of the directionality of water diffusion, indicating fiber tract characteristics that might be influenced by blindness as well as the acquired perceptual skills. Analysis of the diffusion images revealed reduced FA in late-blind individuals relative to sighted controls at the level of the optic radiations at either side and the right-hemisphere dorsal thalamus (pulvinar). Moreover, late-blind subjects showed significant positive correlations between FA and the capacity of ultra-fast speech comprehension within right-hemisphere optic radiation and thalamus. Thus, experience-related structural alterations occurred in late-blind individuals within visual pathways that, presumably, are linked to higher order frontal language areas.

Consensus paper: Language and the cerebellum: an ongoing enigma.

In less than three decades, the concept "cerebellar neurocognition" has evolved from a mere afterthought to an entirely new and multifaceted area of neuroscientific research. A close interplay between three main strands of contemporary neuroscience induced a substantial modification of the traditional view of the cerebellum as a mere coordinator of autonomic and somatic motor functions. Indeed, the wealth of current evidence derived from detailed neuroanatomical investigations, functional neuroimaging studies with healthy subjects and patients and in-depth neuropsychological assessment of patients with cerebellar disorders shows that the cerebellum has a cardinal role to play in affective regulation, cognitive processing, and linguistic function. Although considerable progress has been made in models of cerebellar function, controversy remains regarding the exact role of the "linguistic cerebellum" in a broad variety of nonmotor language processes. This consensus paper brings together a range of different viewpoints and opinions regarding the contribution of the cerebellum to language function. Recent developments and insights in the nonmotor modulatory role of the cerebellum in language and some related disorders will be discussed. The role of the cerebellum in speech and language perception, in motor speech planning including apraxia of speech, in verbal working memory, in phonological and semantic verbal fluency, in syntax processing, in the dynamics of language production, in reading and in writing will be addressed. In addition, the functional topography of the linguistic cerebellum and the contribution of the deep nuclei to linguistic function will be briefly discussed. As such, a framework for debate and discussion will be offered in this consensus paper.

Training of ultra-fast speech comprehension induces functional reorganization of the central-visual system in late-blind humans.

Individuals suffering from vision loss of a peripheral origin may learn to understand spoken language at a rate of up to about 22 syllables (syl) per seconds (s)-exceeding by far the maximum performance level of untrained listeners (ca. 8 syl/s). Previous findings indicate the central-visual system to contribute to the processing of accelerated speech in blind subjects. As an extension, the present training study addresses the issue whether acquisition of ultra-fast (18 syl/s) speech perception skills induces de novo central-visual hemodynamic activation in late-blind participants. Furthermore, we asked to what extent subjects with normal or residual vision can improve understanding of accelerated verbal utterances by means of specific training measures. To these ends, functional magnetic resonance imaging (fMRI) was performed while subjects were listening to forward and reversed sentence utterances of moderately fast and ultra-fast syllable rates (8 or 18 syl/s) prior to and after a training period of ca. 6 months. Four of six participants showed-independently from residual visual functions-considerable enhancement of ultra-fast speech perception (about 70% points correctly repeated words) whereas behavioral performance did not change in the two remaining participants. Only subjects with very low visual acuity displayed training-induced hemodynamic activation of the central-visual system. By contrast, participants with moderately impaired or even normal visual acuity showed, instead, increased right-hemispheric frontal or bilateral anterior temporal lobe responses after training. All subjects with significant training effects displayed a concomitant increase of hemodynamic activation of left-hemispheric SMA. In spite of similar behavioral performance, trained "experts" appear to use distinct strategies of ultra-fast speech processing depending on whether the occipital cortex is still deployed for visual processing.

How can audiovisual pathways enhance the temporal resolution of time-compressed speech in blind subjects?

In blind people, the visual channel cannot assist face-to-face communication via lipreading or visual prosody. Nevertheless, the visual system may enhance the evaluation of auditory information due to its cross-links to (1) the auditory system, (2) supramodal representations, and (3) frontal action-related areas. Apart from feedback or top-down support of, for example, the processing of spatial or phonological representations, experimental data have shown that the visual system can impact auditory perception at more basic computational stages such as temporal signal resolution. For example, blind as compared to sighted subjects are more resistant against backward masking, and this ability appears to be associated with activity in visual cortex. Regarding the comprehension of continuous speech, blind subjects can learn to use accelerated text-to-speech systems for "reading" texts at ultra-fast speaking rates (>16 syllables/s), exceeding by far the normal range of 6 syllables/s. A functional magnetic resonance imaging study has shown that this ability, among other brain regions, significantly covaries with BOLD responses in bilateral pulvinar, right visual cortex, and left supplementary motor area. Furthermore, magnetoencephalographic measurements revealed a particular component in right occipital cortex phase-locked to the syllable onsets of accelerated speech. In sighted people, the "bottleneck" for understanding time-compressed speech seems related to higher demands for buffering phonological material and is, presumably, linked to frontal brain structures. On the other hand, the neurophysiological correlates of functions overcoming this bottleneck, seem to depend upon early visual cortex activity. The present Hypothesis and Theory paper outlines a model that aims at binding these data together, based on early cross-modal pathways that are already known from various audiovisual experiments on cross-modal adjustments during space, time, and object recognition.

Ultra-fast speech comprehension in blind subjects engages primary visual cortex, fusiform gyrus, and pulvinar - a functional magnetic resonance imaging (fMRI) study.

BACKGROUND: Individuals suffering from vision loss of a peripheral origin may learn to understand spoken language at a rate of up to about 22 syllables (syl) per second - exceeding by far the maximum performance level of normal-sighted listeners (ca. 8 syl/s). To further elucidate the brain mechanisms underlying this extraordinary skill, functional magnetic resonance imaging (fMRI) was performed in blind subjects of varying ultra-fast speech comprehension capabilities and sighted individuals while listening to sentence utterances of a moderately fast (8 syl/s) or ultra-fast (16 syl/s) syllabic rate. RESULTS: Besides left inferior frontal gyrus (IFG), bilateral posterior superior temporal sulcus (pSTS) and left supplementary motor area (SMA), blind people highly proficient in ultra-fast speech perception showed significant hemodynamic activation of right-hemispheric primary visual cortex (V1), contralateral fusiform gyrus (FG), and bilateral pulvinar (Pv). CONCLUSIONS: Presumably, FG supports the left-hemispheric perisylvian "language network", i.e., IFG and superior temporal lobe, during the (segmental) sequencing of verbal utterances whereas the collaboration of bilateral pulvinar, right auditory cortex, and ipsilateral V1 implements a signal-driven timing mechanism related to syllabic (suprasegmental) modulation of the speech signal. These data structures, conveyed via left SMA to the perisylvian "language zones", might facilitate - under time-critical conditions - the consolidation of linguistic information at the level of verbal working memory.

Tracking the speech signal--time-locked MEG signals during perception of ultra-fast and moderately fast speech in blind and in sighted listeners.

Blind people can learn to understand speech at ultra-high syllable rates (ca. 20 syllables/s), a capability associated with hemodynamic activation of the central-visual system. To further elucidate the neural mechanisms underlying this skill, magnetoencephalographic (MEG) measurements during listening to sentence utterances were cross-correlated with time courses derived from the speech signal (envelope, syllable onsets and pitch periodicity) to capture phase-locked MEG components (14 blind, 12 sighted subjects; speech rate=8 or 16 syllables/s, pre-defined source regions: auditory and visual cortex, inferior frontal gyrus). Blind individuals showed stronger phase locking in auditory cortex than sighted controls, and right-hemisphere visual cortex activity correlated with syllable onsets in case of ultra-fast speech. Furthermore, inferior-frontal MEG components time-locked to pitch periodicity displayed opposite lateralization effects in sighted (towards right hemisphere) and blind subjects (left). Thus, ultra-fast speech comprehension in blind individuals appears associated with changes in early signal-related processing mechanisms both within and outside the central-auditory terrain.

Neural processing of acoustic duration and phonological German vowel length: time courses of evoked fields in response to speech and nonspeech signals.

Recent experiments showed that the perception of vowel length by German listeners exhibits the characteristics of categorical perception. The present study sought to find the neural activity reflecting categorical vowel length and the short-long boundary by examining the processing of non-contrastive durations and categorical length using MEG. Using disyllabic words with varying /a/-durations and temporally-matched nonspeech stimuli, we found that each syllable elicited an M50/M100-complex. The M50-amplitude to the second syllable varied along the durational continuum, possibly reflecting the mapping of duration onto a rhythm representation. Categorical length was reflected by an additional response elicited when vowel duration exceeded the short-long boundary. This was interpreted to reflect the integration of an additional timing unit for long in contrast to short vowels. Unlike to speech, responses to short nonspeech durations lacked a M100 to the first and M50 to the second syllable, indicating different integration windows for speech and nonspeech signals.

Neurophonetics.

Neurophonetics aims at the elucidation of the brain mechanisms underlying speech communication in our species. Clinical observations in patients with speech impairments following cerebral disorders provided the initial vantage point of this research area and indicated distinct functional-neuroanatomic systems to support human speaking and listening. Subsequent approaches-considering speech production a motor skill-investigated vocal tract movements associated with spoken language by means of kinematic and electromyographic techniques-allowing, among other things, for the evaluation of computational models suggesting elementary phonological gestures or a mental syllabary as basic units of speech motor control. As concerns speech perception, the working characteristics of auditory processing were first investigated based upon psychoacoustic techniques such as dichotic listening and categorical perception designs. More recently, functional hemodynamic neuroimaging and electrophysiological methods opened the door to the delineation of multiple stages of central auditory processing related to signal detection, classification, sensory memory processes, and, finally, lexical access. Beyond the control mechanisms in a stricter sense, both speech articulation and auditory processing represent examples of 'grounded cognition'. For example, both domains cannot be restricted to text-to-speech translation processes, but are intimately interwoven with neuropsychological aspects of speech prosody, including the vocal expression of affects and the actual performance of speech acts, transforming propositional messages to 'real' utterances. Furthermore, during language acquisition, the periphery of language-i.e., hearing and speaking behavior-plays a dominant role for the construction of a language-specific mental lexicon as well as language-specific action plans for the production of a speech message. WIREs Cogn Sci 2013, 4:191-200. doi: 10.1002/wcs.1211 For further resources related to this article, please visit the WIREs website.

Mood modulates auditory laterality of hemodynamic mismatch responses during dichotic listening.

Hemodynamic mismatch responses can be elicited by deviant stimuli in a sequence of standard stimuli even during cognitive demanding tasks. Emotional context is known to modulate lateralized processing. Right-hemispheric negative emotion processing may bias attention to the right and enhance processing of right-ear stimuli. The present study examined the influence of induced mood on lateralized pre-attentive auditory processing of dichotic stimuli using functional magnetic resonance imaging (fMRI). Faces expressing emotions (sad/happy/neutral) were presented in a blocked design while a dichotic oddball sequence with consonant-vowel (CV) syllables in an event-related design was simultaneously administered. Twenty healthy participants were instructed to feel the emotion perceived on the images and to ignore the syllables. Deviant sounds reliably activated bilateral auditory cortices and confirmed attention effects by modulation of visual activity. Sad mood induction activated visual, limbic and right prefrontal areas. A lateralization effect of emotion-attention interaction was reflected in a stronger response to right-ear deviants in the right auditory cortex during sad mood. This imbalance of resources may be a neurophysiological correlate of laterality in sad mood and depression. Conceivably, the compensatory right-hemispheric enhancement of resources elicits increased ipsilateral processing.

Brief report: impaired differentiation of vegetative/affective and intentional nonverbal vocalizations in a subject with Asperger syndrome (AS).

The Asperger syndrome (AS) includes impaired recognition of other people's mental states. Since language-based diagnostic procedures may be confounded by cognitive-linguistic compensation strategies, nonverbal test materials were created, including human affective and vegetative sounds. Depending on video context, each sound could be interpreted either as direct expression of an agent's affective/vegetative state or as result of intentional-executive mental operations. "Situational relevance" and "intentionality" ratings by a group of twelve healthy subjects nicely differentiated between context types. By contrast, an AS subject showed a systematic over-interpretation of vegetative/affective signals in terms of planned activities. Such overestimation of intentional motivation, leading to impaired social cognition, might be due to the inability to utilize "affective resonance" mechanisms for the interpretation of an individual's internal state.

Magnetic brain activity phase-locked to the envelope, the syllable onsets, and the fundamental frequency of a perceived speech signal.

During speech perception, acoustic correlates of syllable structure and pitch periodicity are directly reflected in electrophysiological brain activity. Magnetoencephalography (MEG) recordings were made while 10 participants listened to natural or formant-synthesized speech at moderately fast or ultrafast rate. Cross-correlation analysis was applied to show brain activity time-locked to the speech envelope, to an acoustic marker of syllable onsets, and to pitch periodicity. The envelope yielded a right-lateralized M100-like response, syllable onsets gave rise to M50/M100-like fields with an additional anterior M50 component, and pitch (ca. 100 Hz) elicited a neural resonance bound to a central auditory source at a latency of 30 ms. The strength of these MEG components showed differential effects of syllable rate and natural versus synthetic speech. Presumingly, such phase-locking mechanisms serve as neuronal triggers for the extraction of information-bearing elements.