Annette Baumgaertner

Phonological decisions require both the left and right supramarginal gyri

Recent functional imaging studies demonstrated that both the left and right supramarginal gyri (SMG) are activated when healthy right-handed subjects make phonological word decisions. However, lesion studies typically report difficulties with phonological processing after left rather than right hemisphere damage. Here, we used a unique dual-site transcranial magnetic stimulation (TMS) approach to test whether the SMG in the right hemisphere contributes to modality-independent (i.e., auditory and visual) phonological decisions. To test task-specificity, we compared the effect of real or sham TMS during phonological, semantic, and perceptual decisions. To test laterality and anatomical specificity, we compared the effect of TMS over the left, right, or bilateral SMG and angular gyri. The accuracy and reaction times of phonological decisions were selectively disrupted relative to semantic and perceptual decisions when real TMS was applied over the left, right, or bilateral SMG. These effects were not observed for TMS over the angular gyri. A follow-up experiment indicated that the threshold-intensity for inducing a disruptive effect on phonological decisions was identical for unilateral TMS over the right or left SMG. Taken together, these findings provide converging evidence that the right SMG contributes to accurate and efficient phonological decisions in the healthy brain, with no evidence that the left and right SMG can compensate for one another during TMS. Our findings motivate detailed studies of phonological processing in patients with acute or long-term damage of the right SMG.

The neural basis for understanding non-intended actions

We can often understand when actions done by others do or do not reflect their intentions. To investigate the neural basis of this capacity we carried out an fMRI study in which volunteers were presented with video-clips showing actions that did reflect the intention of the agent (intended actions) and actions that did not (non-intended actions). Observation of both types of actions activated a common set of areas including the inferior parietal lobule, the lateral premotor cortex and mesial premotor areas. The contrast non-intended vs. intended actions showed activation in the right temporo-parietal junction, left supramarginal gyrus, and mesial prefrontal cortex. The converse contrast did not show any activation. We conclude that our capacity to understand non intended actions is based on the activation of areas signaling unexpected events in spatial and temporal domains, in addition to the activity of the mirror neuron system. The concomitant activation of mesial prefrontal areas, known to be involved in self-referential processing, might reflect how deeply participants are involved in the observed scenes.

Polymodal conceptual processing of human biological actions in the left inferior frontal lobe

Apart from being increasingly implicated in higher motor control, Brocas area is considered to play an important role in action understanding by coding the motor goal of an action. Moreover, recent findings suggest that parts of Brocas area may be able to code action content in a more abstract fashion, independent of modality, specific movement parameters or effector used. We performed functional magnetic resonance imaging to examine whether in humans processing object‐directed hand actions presented either visually as video clips or verbally as spoken sentences relies on the same neural substrates. To control for action specificity, we included videos and sentences depicting inanimate motion events. In order to induce conceptual processing, we asked participants to make judgements about the acceptability of the stimuli. Results show that processing object‐directed hand actions presented both visually and verbally leads to common activation of areas in parietal and frontal regions, most prominently in the pars opercularis of Brocas region. We conclude that the pars opercularis of Brocas area is endowed with polymodal capabilities, allowing the processing of higher‐level conceptual aspects of action understanding.

The right posterior inferior frontal gyrus contributes to phonological word decisions in the healthy brain: Evidence from dual-site TMS

There is consensus that the left hemisphere plays a dominant role in language processing, but functional imaging studies have shown that the right as well as the left posterior inferior frontal gyri (pIFG) are activated when healthy right-handed individuals make phonological word decisions. Here we used online transcranial magnetic stimulation (TMS) to examine the functional relevance of the right pIFG for auditory and visual phonological decisions. Healthy right-handed individuals made phonological or semantic word judgements on the same set of auditorily and visually presented words while they received stereotactically guided TMS over the left, right or bilateral pIFG (n = 14) or the anterior left, right or bilateral IFG (n = 14). TMS started 100 ms after word onset and consisted of four stimuli given at a rate of 10 Hz and intensity of 90% of active motor threshold. Compared to TMS of aIFG, TMS of pIFG impaired reaction times and accuracy of phonological but not semantic decisions for visually and auditorily presented words. TMS over left, right or bilateral pIFG disrupted phonological processing to a similar degree. In a follow-up experiment, the intensity threshold for delaying phonological judgements was identical for unilateral TMS of left and right pIFG. These findings indicate that an intact function of right pIFG is necessary for accurate and efficient phonological decisions in the healthy brain with no evidence that the left and right pIFG can compensate for one another during online TMS. Our findings motivate detailed studies of phonological processing in patients with acute and chronic damage of the right pIFG.

Perturbation of the left inferior frontal gyrus triggers adaptive plasticity in the right homologous area during speech production

Significance The role of the right hemisphere in aphasia recovery is unclear. We demonstrate that a virtual lesion of left inferior frontal gyrus (IFG) decreased activity in the targeted area and increased activity in the contralateral homologous area during pseudoword repetition. This was associated with a stronger facilitatory drive from the right IFG to the left IFG. Importantly, responses became faster with increased influence of the right IFG on the left IFG. Our results shed new light on the dynamic regulation of interhemispheric interactions in the human brain. Particularly, these findings are of potential importance for understanding language recovery after left-hemispheric stroke, indicating that homologous right hemisphere areas actively contribute to language function after a left hemisphere lesion. The role of the right hemisphere in aphasia recovery after left hemisphere damage remains unclear. Increased activation of the right hemisphere has been observed after left hemisphere damage. This may simply reflect a release from transcallosal inhibition that does not contribute to language functions. Alternatively, the right hemisphere may actively contribute to language functions by supporting disrupted processing in the left hemisphere via interhemispheric connections. To test this hypothesis, we applied off-line continuous theta burst stimulation (cTBS) over the left inferior frontal gyrus (IFG) in healthy volunteers, then used functional MRI to investigate acute changes in effective connectivity between the left and right hemispheres during repetition of auditory and visual words and pseudowords. In separate sessions, we applied cTBS over the left anterior IFG (aIFG) or posterior IFG (pIFG) to test the anatomic specificity of the effects of cTBS on speech processing. Compared with cTBS over the aIFG, cTBS over the pIFG suppressed activity in the left pIFG and increased activity in the right pIFG during pseudoword vs. word repetition in both modalities. This effect was associated with a stronger facilitatory drive from the right pIFG to the left pIFG during pseudoword repetition. Critically, response became faster as the influence of the right pIFG on left pIFG increased, indicating that homologous areas in the right hemisphere actively contribute to language function after a focal left hemisphere lesion. Our findings lend further support to the notion that increased activation of homologous right hemisphere areas supports aphasia recovery after left hemisphere damage.

Neural circuitry of the bilingual mental lexicon: Effect of age of second language acquisition

Numerous studies have proposed that changes of the human language faculty caused by neural maturation can explain the substantial differences in ultimate attainment of grammatical competences between first language (L1) acquirers and second language (L2) learners. However, little evidence on the effect of neural maturation on the attainment of lexical knowledge in L2 is available. The present functional magnetic resonance study addresses this question via a cross-linguistic neural adaptation paradigm. Age of acquisition (AoA) of L2 was systematically manipulated. Concrete nouns were repeated across language (e.g., French-German, valise(suitcase)-Koffer(suitcase)). Whereas early bilinguals (AoA of L2<3years) showed larger repetition enhancement (RE) effects in the left superior temporal gyrus, the bilateral superior frontal gyrus and the right posterior insula, late bilinguals (AoA of L2>10years) showed larger RE effects in the middle portion of the left insula and in the right middle frontal gyrus (MFG). We suggest that, as for grammatical knowledge, the attainment of lexical knowledge in L2 is affected by neural maturation. The present findings lend support to neurocognitive models of bilingual word recognition postulating that, for both early and late bilinguals, the two languages are interconnected at the conceptual level.

Unintended imitation in nonword repetition

Verbal repetition is conventionally considered to require motor-reproduction of only the phonologically relevant content of a perceived linguistic stimulus, while imitation of incidental acoustic properties of the stimulus is not an explicit part of this task. Exemplar-based theories of speech processing, however, would predict that imitation beyond linguistic reproduction may occur in word repetition. Five experiments were conducted in which verbal audio-motor translations had to be performed under different conditions. Nonwords varying in phonemic content, in vocal pitch (F(0)), and in speaking style (schwa-syllable expression) were presented. We experimentally varied the factors response delay (repetition vs. shadowing), intention-to-repeat (repetition vs. pseudo-naming), and phonological load (repetition vs. transformation). The responses of ten healthy participants were examined for phonemic accuracy and for traces of para-phonological imitation. Two aphasic patients with phonological impairments were also included, to find out if lesions to left anterior or posterior perisylvian cortex interfere with imitation. In the healthy participants, significant imitation of both F(0) and phonetic style was observed, with markedly stronger effects for the latter. Strong imitation was also found in an aphasic patient with a lesion to left anterior perisylvian cortex, whereas almost no imitation occurred in a patient with a lesion to the posterior language area. The degree of unintended imitation was modulated by each of the three independent factors introduced here. The results are discussed on the background of cognitive and neurolinguistic theories of imitation.

Intensive speech and language therapy in patients with chronic aphasia after stroke: a randomised, open-label, blinded-endpoint, controlled trial in a health-care setting

BACKGROUND Treatment guidelines for aphasia recommend intensive speech and language therapy for chronic (≥6 months) aphasia after stroke, but large-scale, class 1 randomised controlled trials on treatment effectiveness are scarce. We aimed to examine whether 3 weeks of intensive speech and language therapy under routine clinical conditions improved verbal communication in daily-life situations in people with chronic aphasia after stroke. METHODS In this multicentre, parallel group, superiority, open-label, blinded-endpoint, randomised controlled trial, patients aged 70 years or younger with aphasia after stroke lasting for 6 months or more were recruited from 19 inpatient or outpatient rehabilitation centres in Germany. An external biostatistician used a computer-generated permuted block randomisation method, stratified by treatment centre, to randomly assign participants to either 3 weeks or more of intensive speech and language therapy (≥10 h per week) or 3 weeks deferral of intensive speech and language therapy. The primary endpoint was between-group difference in the change in verbal communication effectiveness in everyday life scenarios (Amsterdam-Nijmegen Everyday Language Test A-scale) from baseline to immediately after 3 weeks of treatment or treatment deferral. All analyses were done using the modified intention-to-treat population (those who received 1 day or more of intensive treatment or treatment deferral). This study is registered with ClinicalTrials.gov, number NCT01540383. FINDINGS We randomly assigned 158 patients between April 1, 2012, and May 31, 2014. The modified intention-to-treat population comprised 156 patients (78 per group). Verbal communication was significantly improved from baseline to after intensive speech and language treatment (mean difference 2·61 points [SD 4·94]; 95% CI 1·49 to 3·72), but not from baseline to after treatment deferral (-0·03 points [4·04]; -0·94 to 0·88; between-group difference Cohens d 0·58; p=0·0004). Eight patients had adverse events during therapy or treatment deferral (one car accident [in the control group], two common cold [one patient per group], three gastrointestinal or cardiac symptoms [all intervention group], two recurrent stroke [one in intervention group before initiation of treatment, and one before group assignment had occurred]); all were unrelated to study participation. INTERPRETATION 3 weeks of intensive speech and language therapy significantly enhanced verbal communication in people aged 70 years or younger with chronic aphasia after stroke, providing an effective evidence-based treatment approach in this population. Future studies should examine the minimum treatment intensity required for meaningful treatment effects, and determine whether treatment effects cumulate over repeated intervention periods. FUNDING German Federal Ministry of Education and Research and the German Society for Aphasia Research and Treatment.

Increased facilitatory connectivity from the pre-sma to the left dorsal premotor cortex during pseudoword repetition

Previous studies have demonstrated that the repetition of pseudowords engages a network of premotor areas for articulatory planning and articulation. However, it remains unclear how these premotor areas interact and drive one another during speech production. We used fMRI with dynamic causal modeling to investigate effective connectivity between premotor areas during overt repetition of words and pseudowords presented in both the auditory and visual modalities. Regions involved in phonological aspects of language production were identified as those where regional increases in the BOLD signal were common to repetition in both modalities. We thus obtained three seed regions: the bilateral pre-SMA, left dorsal premotor cortex (PMd), and left ventral premotor cortex that were used to test 63 different models of effective connectivity in the premotor network for pseudoword relative to word repetition. The optimal model was identified with Bayesian model selection and reflected a network with driving input to pre-SMA and an increase in facilitatory drive from pre-SMA to PMd during repetition of pseudowords. The task-specific increase in effective connectivity from pre-SMA to left PMd suggests that the pre-SMA plays a supervisory role in the generation and subsequent sequencing of motor plans. Diffusion tensor imaging-based fiber tracking in another group of healthy volunteers showed that the functional connection between both regions is underpinned by a direct cortico-cortical anatomical connection.

Right-hemispheric processing of non-linguistic word features: implications for mapping language recovery after stroke.

Verbal stimuli often induce right‐hemispheric activation in patients with aphasia after left‐hemispheric stroke. This right‐hemispheric activation is commonly attributed to functional reorganization within the language system. Yet previous evidence suggests that functional activation in right‐hemispheric homologues of classic left‐hemispheric language areas may partly be due to processing nonlinguistic perceptual features of verbal stimuli. We used functional MRI (fMRI) to clarify the role of the right hemisphere in the perception of nonlinguistic word features in healthy individuals. Participants made perceptual, semantic, or phonological decisions on the same set of auditorily and visually presented word stimuli. Perceptual decisions required judgements about stimulus‐inherent changes in font size (visual modality) or fundamental frequency contour (auditory modality). The semantic judgement required subjects to decide whether a stimulus is natural or man‐made; the phonologic decision required a decision on whether a stimulus contains two or three syllables. Compared to phonologic or semantic decision, nonlinguistic perceptual decisions resulted in a stronger right‐hemispheric activation. Specifically, the right inferior frontal gyrus (IFG), an area previously suggested to support language recovery after left‐hemispheric stroke, displayed modality‐independent activation during perceptual processing of word stimuli. Our findings indicate that activation of the right hemisphere during language tasks may, in some instances, be driven by a “nonlinguistic perceptual processing” mode that focuses on nonlinguistic word features. This raises the possibility that stronger activation of right inferior frontal areas during language tasks in aphasic patients with left‐hemispheric stroke may at least partially reflect increased attentional focus on nonlinguistic perceptual aspects of language. Hum Brain Mapp, 2013.