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Allen R Braun

Publications and source records attributed to Allen R Braun.

16 recordsLinked to original sources

Species-specific calls activate homologs of Broca's and Wernicke's areas in the macaque.

The origin of brain mechanisms that support human language-whether these originated de novo in humans or evolved from a neural substrate that existed in a common ancestor-remains a controversial issue. Although the answer is not provided by the fossil record, it is possible to make inferences by studying living species of nonhuman primates. Here we identified neural systems associated with perceiving species-specific vocalizations in rhesus macaques using H(2)(15)O positron emission tomography (PET). These vocalizations evoke distinct patterns of brain activity in homologs of the human perisylvian language areas. Rather than resulting from differences in elementary acoustic properties, this activity seems to reflect higher order auditory processing. Although parallel evolution within independent primate species is feasible, this finding suggests the possibility that the last common ancestor of macaques and humans, which lived 25-30 million years ago, possessed key neural mechanisms that were plausible candidates for exaptation during the evolution of language.

Acoustic Stimulation↗

Left hemispheric lateralization of brain activity during passive rhythm perception in musicians.

The nature of hemispheric specialization of brain activity during rhythm processing remains poorly understood. The locus for rhythmic processing has been difficult to identify and there have been several contradictory findings. We therefore used functional magnetic resonance imaging to study passive rhythm perception to investigate the hypotheses that rhythm processing results in left hemispheric lateralization of brain activity and is affected by musical training. Twelve musicians and 12 nonmusicians listened to regular and random rhythmic patterns. Conjunction analysis revealed a shared network of neural structures (bilateral superior temporal areas, left inferior parietal lobule, and right frontal operculum) responsible for rhythm perception independent of musical background. In contrast, random-effects analysis showed greater left lateralization of brain activity in musicians compared to nonmusicians during regular rhythm perception, particularly within the perisylvian cortices (left frontal operculum, superior temporal gyrus, inferior parietal lobule). These results suggest that musical training leads to the employment of left-sided perisylvian brain areas, typically active during language comprehension, during passive rhythm perception.

Acoustic Stimulation↗

Neural bases of categorization of simple speech and nonspeech sounds.

Categorization is fundamental to our perception and understanding of the environment. However, little is known about the neural bases underlying the categorization of sounds. Using human functional magnetic resonance imaging (fMRI) we compared the brain responses to a category discrimination task with an auditory discrimination task using identical sets of sounds. Our stimuli differed along two dimensions: a speech-nonspeech dimension and a fast-slow temporal dynamics dimension. All stimuli activated regions in the primary and nonprimary auditory cortices in the temporal cortex and in the parietal and frontal cortices for the two tasks. When comparing the activation patterns for the category discrimination task to those for the auditory discrimination task, the results show that a core group of regions beyond the auditory cortices, including inferior and middle frontal gyri, dorsomedial frontal gyrus, and intraparietal sulcus, were preferentially activated for familiar speech categories and for novel nonspeech categories. These regions have been shown to play a role in working memory tasks by a number of studies. Additionally, the categorization of nonspeech sounds activated left middle frontal gyrus and right parietal cortex to a greater extent than did the categorization of speech sounds. Processing the temporal aspects of the stimuli had a greater impact on the left lateralization of the categorization network than did other factors, particularly in the inferior frontal gyrus, suggesting that there is no inherent left hemisphere advantage in the categorical processing of speech stimuli, or for the categorization task itself.

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Alterations in CNS activity induced by botulinum toxin treatment in spasmodic dysphonia: an H215O PET study.

Speech-related changes in regional cerebral blood flow (rCBF) were measured using H(2)(15)O positron-emission tomography in 9 adults with adductor spasmodic dysphonia (ADSD) before and after botulinum toxin (BTX) injection and 10 age- and gender-matched volunteers without neurological disorders. Scans were acquired at rest and during production of continuous narrative speech and whispered speech. Speech was recorded during scan acquisition for offline quantification of voice breaks, pitch breaks, and percentage aperiodicity to assess correlations between treatment-related changes in rCBF and clinical improvement. Results demonstrated that speech-related responses in heteromodal sensory areas were significantly reduced in persons with ADSD, compared with volunteers, before the administration of BTX. Three to 4 weeks after BTX injection, speech-related responses were significantly augmented in these regions and in left hemisphere motor areas commonly associated with oral-laryngeal motor control. This pattern of responses was most strongly correlated with the objective measures of clinical improvement (decreases in the frequency of voice breaks, pitch breaks, and percentage aperiodicity). These data suggest a pathophysiological model for ADSD in which BTX treatment results in more efficient cortical processing of sensory information, making this information available to motor areas that use it to more effectively regulate laryngeal movements.

Adult↗

Temporal dissociation of early lexical access and articulation using a delayed naming task--an FMRI study.

Neuroimaging studies of overt speech hold an important practical advantage allowing monitoring of subject performance, particularly valuable in disorders like aphasia. However, speech production is not a monotonic process but a complex sequence of stages. Levelt and colleagues have described these as roughly corresponding to two originally independent systems--conceptual and sensorimotor--that are linked in the formulation and expression of spoken language. In the initial stages a word is chosen to match a concept (lexical selection); in the later stages the sound and motor patterns are encoded and the word is uttered (articulation). It has been difficult to discriminate these stages using conventional neuroimaging techniques. We designed a functional magnetic resonance imaging study in an attempt to do this, by introducing a latency into a conventional naming paradigm, delaying the articulated response. Our results showed that left hemisphere perisylvian areas were active throughout, interacting with visual and heteromodal areas during early lexical access and with motor and auditory areas during overt articulation. These results are consistent with the broadest version of the Levelt model and with that derived from Chomsky's minimalist program in which a core language system interacts with conceptual-intentional systems and articulatory-perceptual systems during the early and late stages of lexical access respectively.

Adult↗

Planum parietale of chimpanzees and orangutans: a comparative resonance of human-like planum temporale asymmetry.

We have previously demonstrated that leftward asymmetry of the planum temporale (PT), a brain language area, was not unique to humans since a similar condition is present in great apes. Here we report on a related area in great apes, the planum parietale (PP). PP in humans has a rightward asymmetry with no correlation to the L>R PT, which indicates functional independence. The roles of the PT in human language are well known while PP is implicated in dyslexia and communication disorders. Since posterior bifurcation of the sylvian fissure (SF) is unique to humans and great apes, we used it to determine characteristics of its posterior ascending ramus, an indicator of the PP, in chimpanzee and orangutan brains. Results showed a human-like pattern of R>L PP (P = 0.04) in chimpanzees with a nonsignificant negative correlation of L>R PT vs. R>L PP (CC = -0.3; P = 0.39). In orangutans, SF anatomy is more variable, although PP was nonsignificantly R>L in three of four brains (P = 0.17). We have now demonstrated human-like hemispheric asymmetry of a second language-related brain area in great apes. Our findings persuasively support an argument for addition of a new component to the comparative neuroanatomic complex that defines brain language or polymodal communication areas. PP strengthens the evolutionary links that living great apes may offer to better understand the origins of these progressive parts of the brain. Evidence mounts for the stable expression of a neural foundation for language in species that we recently shared a common ancestor with.

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Comparison of continuous overt speech fMRI using BOLD and arterial spin labeling.

Overt speech production in functional magnetic resonance imaging (fMRI) studies is often associated with imaging artifacts, attributable to both movement and susceptibility. Various image-processing methods have been proposed to remove these artifacts from the data but none of these methods has been shown to work with continuous overt speech, at least over periods greater than 3 s. In this study natural, continuous, overt sentence production was evaluated in normal volunteers using both arterial spin labeling (ASL) and conventional echoplanar blood oxygenation level-dependent (BOLD) imaging sequences on the same 1.5-T scanner. We found a high congruency between activation results obtained with ASL and the de facto gold standard in overt language production imaging, positron emission tomography (PET). No task-related artifacts were found in the ASL study. However, the BOLD data showed artifacts that appeared as large bilateral false-positive temporopolar activations; percent signal change estimated in these regions showed signal increases and temporal dynamics that were incongruent with typical BOLD activations. These artifacts were not distributed uniformly, but were aligned at the frontotemporal base, close to the oropharynx. The calculated head movement parameters for overt speech blocks were within the range of the rest blocks, indicating that head movement is unlikely the reason for the artifact. We conclude that ASL is not influenced by overt speech artifacts, whereas BOLD showed significant susceptibility artifacts, especially in the opercular and insular regions, where activation would be expected. ASL may prove to be the method of choice for fMRI investigations of continuous overt speech.

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Neural systems for sign language production: mechanisms supporting lexical selection, phonological encoding, and articulation.

Overt production of ASL signs was evaluated using H(2)(15)O PET to differentiate brain systems that support sign language production at the lexical-selection and phonological-articulatory levels. Subjects were 16 right-handed, congenitally deaf native ASL signers (10 women, six men; age 20 to 29 years). Scans were performed while subjects (1) passively viewed ASL nouns, (2) repeated nouns, (3) generated verbs in response to these nouns, (4) passively viewed videotaped segments depicting transitive actions, and (5) generated a verb to describe these actions. Conjunctions between the two verb-generation tasks revealed left-lateralized activation of perisylvian, frontal, and subcortical regions commonly observed in spoken language generation tasks and implicated in processes of semantic feature binding and lexical selection. Analysis of noun repetition minus viewing condition revealed activation of distinct systems supporting phonological encoding and articulation, including bilateral activation of sensorimotor areas and association cortices in the temporal, parietal, and occipital lobes. In addition, lexical-selection and articulatory processes were associated with activation of different corticostriatal-thalamocortical circuits: articulation with activation of the motor, and lexical-selection with activation of the prefrontal circuits, respectively. The results collectively provide insight into dissociable neural systems underlying these psycholinguistic functions. In addition, activation of regions that are typically associated with the auditory system during sign production suggests that these regions may support modality-independent linguistic processes, or may indicate cross-modal plasticity within the deaf brain.

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Auditory lexical decision, categorical perception, and FM direction discrimination differentially engage left and right auditory cortex.

Recent neuroimaging and neuropsychological data suggest that speech perception is supported in bilaterally auditory areas. We evaluate this issue building on well-known behavioral effects. While undergoing positron emission tomography (PET), subjects performed standard auditory tasks: direction discrimination of frequency-modulated (FM) tones, categorical perception (CP) of consonant-vowel (CV) syllables, and word/non-word judgments (lexical decision, LD). Compared to rest, the three conditions led to bilateral activation of the auditory cortices. However, lateralization patterns differed as a function of stimulus type: the LD task generated stronger responses in the left, the FM task a stronger response in the right hemisphere. Contrasts between either words or syllables versus FM were associated with significantly greater activity bilaterally in superior temporal gyrus (STG) ventro-lateral to Heschl's gyrus. These activations extended into the superior temporal sulcus (STS) and the middle temporal gyrus (MTG) and were greater in the left. The same areas were more active in the LD than the CP task. In contrast, the FM task was associated with significantly greater activity in the right lateral-posterior STG and lateral MTG. The findings argue for a view in which speech perception is mediated bilaterally in the auditory cortices and that the well-documented lateralization is likely associated with processes subsequent to the auditory analysis of speech.

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Brain network interactions in auditory, visual and linguistic processing.

In the paper, we discuss the importance of network interactions between brain regions in mediating performance of sensorimotor and cognitive tasks, including those associated with language processing. Functional neuroimaging, especially PET and fMRI, provide data that are obtained essentially simultaneously from much of the brain, and thus are ideal for enabling one to assess interregional functional interactions. Two ways to use these types of data to assess network interactions are presented. First, using PET, we demonstrate that anterior and posterior perisylvian language areas have stronger functional connectivity during spontaneous narrative production than during other less linguistically demanding production tasks. Second, we show how one can use large-scale neural network modeling to relate neural activity to the hemodynamically-based data generated by fMRI and PET. We review two versions of a model of object processing - one for visual and one for auditory objects. The regions comprising the models include primary and secondary sensory cortex, association cortex in the temporal lobe, and prefrontal cortex. Each model incorporates specific assumptions about how neurons in each of these areas function, and how neurons in the different areas are interconnected with each other. Each model is able to perform a delayed match-to-sample task for simple objects (simple shapes for the visual model; tonal contours for the auditory model). We find that the simulated electrical activities in each region are similar to those observed in nonhuman primates performing analogous tasks, and the absolute values of the simulated integrated synaptic activity in each brain region match human fMRI/PET data. Thus, this type of modeling provides a way to understand the neural bases for the sensorimotor and cognitive tasks of interest.

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Language lateralization in a bimanual language.

Unlike spoken languages, sign languages of the deaf make use of two primary articulators, the right and left hands, to produce signs. This situation has no obvious parallel in spoken languages, in which speech articulation is carried out by symmetrical unitary midline vocal structures. This arrangement affords a unique opportunity to examine the robustness of linguistic systems that underlie language production in the face of contrasting articulatory demands and to chart the differential effects of handedness for highly skilled movements. Positron emission tomography (PET) technique was used to examine brain activation in 16 deaf users of American Sign Language (ASL) while subjects generated verb signs independently with their right dominant and left nondominant hands (compared to the repetition of noun signs). Nearly identical patterns of left inferior frontal and right cerebellum activity were observed. This pattern of activation during signing is consistent with patterns that have been reported for spoken languages including evidence for specializations of inferior frontal regions related to lexical-semantic processing, search and retrieval, and phonological encoding. These results indicate that lexical-semantic processing in production relies upon left-hemisphere regions regardless of the modality in which a language is realized, and that this left-hemisphere activation is stable, even in the face of conflicting articulatory demands. In addition, these data provide evidence for the role of the right posterolateral cerebellum in linguistic-cognitive processing and evidence of a left ventral fusiform contribution to sign language processing.

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Common features of fluency-evoking conditions studied in stuttering subjects and controls: an H(2)15O PET study.

UNLABELLED: We used H(2)15O PET to characterize the common features of two successful but markedly different fluency-evoking conditions -- paced speech and singing -- in order to identify brain mechanisms that enable fluent speech in people who stutter. To do so, we compared responses under fluency-evoking conditions with responses elicited by tasks that typically elicit dysfluent speech (quantifying the degree of stuttering and using this measure as a confounding covariate in our analyses). We evaluated task-related activations in both stuttering subjects and age- and gender-matched controls. Areas that were either uniquely activated during fluency-evoking conditions, or in which the magnitude of activation was significantly greater during fluency-evoking than dysfluency-evoking tasks included auditory association areas that process speech and voice and motor regions related to control of the larynx and oral articulators. This suggests that a common fluency-evoking mechanism might relate to more effective coupling of auditory and motor systems -- that is, more efficient self-monitoring, allowing motor areas to more effectively modify speech. These effects were seen in both PWS and controls, suggesting that they are due to the sensorimotor or cognitive demands of the fluency-evoking tasks themselves. While responses seen in both groups were bilateral, however, the fluency-evoking tasks elicited more robust activation of auditory and motor regions within the left hemisphere of stuttering subjects, suggesting a role for the left hemisphere in compensatory processes that enable fluency. EDUCATIONAL OBJECTIVES: The reader will learn about and be able to: (1) compare brain activation patterns under fluency- and dysfluency-evoking conditions in stuttering and control subjects; (2) appraise the common features, both central and peripheral, of fluency-evoking conditions; and (3) discuss ways in which neuroimaging methods can be used to understand the pathophysiology of stuttering.

Adult↗

Activation of Broca's area during the production of spoken and signed language: a combined cytoarchitectonic mapping and PET analysis.

Broca's area in the inferior frontal gyrus consists of two cytoarchitectonically defined regions-Brodmann areas (BA) 44 and 45. Combining probabilistic maps of these two areas with functional neuroimaging data obtained using PET, it is shown that BA45, not BA44, is activated by both speech and signing during the production of language narratives in bilingual subjects fluent from early childhood in both American Sign Language (ASL) and English when the generation of complex movements and sounds is taken into account. It is BA44, not BA45, that is activated by the generation of complex articulatory movements of oral/laryngeal or limb musculature. The same patterns of activation are found for oral language production in a group of English speaking monolingual subjects. These findings implicate BA45 as the part of Broca's area that is fundamental to the modality-independent aspects of language generation.

Adult↗

fMRI study comparing names versus pictures of objects.

We performed an fMRI one-back recognition study aimed at distinguishing the semantic versus perceptual aspects of how objects and their written forms are processed. There were three types of visually presented items: pictures (schematic drawings of objects); words identifying these objects; and a mixed condition in which pictures were interleaved with words. A semantic decision about object identity was required when pictures were interleaved with words. This condition, contrasted with the other two, invoked a larger signal in multiple areas, including frontal cortex, bilateral occipitotemporal cortex, and the right middle temporal gyrus. We propose that the left occipitotemporal and right temporal activations are indicative of the neural substrate mediating picture-word conversions, whereas the frontal activations reflect the coordinating functions of the central executive.

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The process of awakening: a PET study of regional brain activity patterns mediating the re-establishment of alertness and consciousness.

Awakening from sleep entails rapid re-establishment of consciousness followed by the relatively slow (20-30 min later) re-establishment of alertness--a temporal dissociation that facilitates specification of the physiological underpinnings of each of these facets of the awakening process. H(2)(15)O PET was used to assess changes in regional cerebral blood flow (rCBF) upon awakening from stage 2 sleep. Cerebral blood flow (CBF) was most rapidly re-established in centrencephalic regions (e.g. brainstem and thalamus), suggesting that the reactivation of these regions underlies the re-establishment of conscious awareness. Across the ensuing 15 min of wakefulness, further increases in CBF were evident primarily in anterior cortical regions, suggesting that the dissipation of sleep inertia effects (post-awakening performance and alertness deficits) is effected by reactivation of these regions. Concomitant shifts in correlation patterns of regional brain activity across the post-awakening period [in particular, a waning negative correlation between prefrontal cortex and mesencephalic reticular formation (RF) activity, and a waxing positive correlation between prefrontal cortex and ventromedial caudate nucleus (CAUD) activity] suggest that the post-awakening reversal of sleep inertia effects may be mediated by more than mere reactivation--it may also involve the functional reorganization of brain activity. Conversely, stable post-awakening correlations--such as those found between the anterior cingulate cortex (ACC) and most other brain regions--may denote the pattern of functional connectivity that underlies consciousness itself.

Adult↗

Temporal dynamics of cortical and subcortical responses to apomorphine in Parkinson disease: an H2(15)O PET study.

H2(15)O positron emission tomography (PET) was used to study the temporal course of central nervous system (CNS) responses to apomorphine in patients with idiopathic Parkinson disease (PD). Agonist-induced changes in regional cerebral blood flow (rCBF) were evaluated within corticostriatal-thalamocortical circuits as well as in regions that extend beyond the standard pathophysiological model for PD. Compared with controls, rCBF was increased in PD patients in subcortical regions including the basal ganglia and cerebellum and both increased and decreased in prefrontal, parietal, sensorimotor, and paralimbic cortical areas. Apomorphine reversed many of these effects and had widespread effects throughout the brain. We evaluated the effects of apomorphine as they changed over time, comparing rCBF before the motor response and at later times when the motor response was maximal. Apomorphine's effects on functional connectivity also changed over time; activity in the ventrolateral thalamus was coupled with that in the SMA and cerebellum at the time of maximum motor response, but not at 45 seconds. Apomorphine affected rCBF in regions commonly considered part of the pathophysiological model of PD (eg, basal ganglia, thalamus, SMA), and other effects were seen in regions outside of the model (eg, cerebellum and superior parietal lobule). Results are discussed in light of this model.

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