Hermann Ackermann

Opposite hemispheric lateralization effects during speaking and singing at motor cortex, insula and cerebellum.

Aside from spoken language, singing represents a second mode of acoustic (auditory-vocal) communication in humans. As a new aspect of brain lateralization, functional magnetic resonance imaging (fMRI) revealed two complementary cerebral networks subserving singing and speaking. Reproduction of a non-lyrical tune elicited activation predominantly in the right motor cortex, the right anterior insula, and the left cerebellum whereas the opposite response pattern emerged during a speech task. In contrast to the hemodynamic responses within motor cortex and cerebellum, activation of the intrasylvian cortex turned out to be bound to overt task performance. These findings corroborate the assumption that the left insula supports the coordination of speech articulation. Similarly, the right insula might mediate temporo-spatial control of vocal tract musculature during overt singing. Both speech and melody production require the integration of sound structure or tonal patterns, respectively, with a speakers emotions and attitudes. Considering the widespread interconnections with premotor cortex and limbic structures, the insula is especially suited for this task.

Identification of emotional intonation evaluated by fMRI.

During acoustic communication among human beings, emotional information can be expressed both by the propositional content of verbal utterances and by the modulation of speech melody (affective prosody). It is well established that linguistic processing is bound predominantly to the left hemisphere of the brain. By contrast, the encoding of emotional intonation has been assumed to depend specifically upon right-sided cerebral structures. However, prior clinical and functional imaging studies yielded discrepant data with respect to interhemispheric lateralization and intrahemispheric localization of brain regions contributing to processing of affective prosody. In order to delineate the cerebral network engaged in the perception of emotional tone, functional magnetic resonance imaging (fMRI) was performed during recognition of prosodic expressions of five different basic emotions (happy, sad, angry, fearful, and disgusted) and during phonetic monitoring of the same stimuli. As compared to baseline at rest, both tasks yielded widespread bilateral hemodynamic responses within frontal, temporal, and parietal areas, the thalamus, and the cerebellum. A comparison of the respective activation maps, however, revealed comprehension of affective prosody to be bound to a distinct right-hemisphere pattern of activation, encompassing posterior superior temporal sulcus (Brodmann Area [BA] 22), dorsolateral (BA 44/45), and orbitobasal (BA 47) frontal areas. Activation within left-sided speech areas, in contrast, was observed during the phonetic task. These findings indicate that partially distinct cerebral networks subserve processing of phonetic and intonational information during speech perception.

Classical conditioning after cerebellar lesions in humans

We explored classical conditioning in human subjects who had lesions in their cerebellar circuitry. Seven patients with damage to cerebellar structures and matched control subjects underwent simple delay tone-airpuff conditioning. Eyelid conditioned response (CR) acquisition was severely disrupted in the patient group, whereas autonomic CRs and slow cortical potentials developing between conditioned stimulus (CS) and the unconditioned stimulus (UCS) were unaffected. Results are consistent with animal studies and earlier case reports indicating that intact cerebellar structures are necessary for the acquisition of classically conditioned motor responses.

fMRI reveals two distinct cerebral networks subserving speech motor control

Background: There are few data on the cerebral organization of motor aspects of speech production and the pathomechanisms of dysarthric deficits subsequent to brain lesions and diseases. The authors used fMRI to further examine the neural basis of speech motor control. Methods and Results: In eight healthy volunteers, fMRI was performed during syllable repetitions synchronized to click trains (2 to 6 Hz; vs a passive listening task). Bilateral hemodynamic responses emerged at the level of the mesiofrontal and sensorimotor cortex, putamen/pallidum, thalamus, and cerebellum (two distinct activation spots at either side). In contrast, dorsolateral premotor cortex and anterior insula showed left-sided activation. Calculation of rate/response functions revealed a negative linear relationship between repetition frequency and blood oxygen level–dependent (BOLD) signal change within the striatum, whereas both cerebellar hemispheres exhibited a step-wise increase of activation at ∼3 Hz. Analysis of the temporal dynamics of the BOLD effect found the various cortical and subcortical brain regions engaged in speech motor control to be organized into two separate networks (medial and dorsolateral premotor cortex, anterior insula, and superior cerebellum vs sensorimotor cortex, basal ganglia, and inferior cerebellum). Conclusion: These data provide evidence for two levels of speech motor control bound, most presumably, to motor preparation and execution processes. They also help to explain clinical observations such as an unimpaired or even accelerated speaking rate in Parkinson disease and slowed speech tempo, which does not fall below a rate of 3 Hz, in cerebellar disorders.

Cerebral processing of linguistic and emotional prosody: fMRI studies.

During acoustic communication in humans, information about a speakers emotional state is predominantly conveyed by modulation of the tone of voice (emotional or affective prosody). Based on lesion data, a right hemisphere superiority for cerebral processing of emotional prosody has been assumed. However, the available clinical studies do not yet provide a coherent picture with respect to interhemispheric lateralization effects of prosody recognition and intrahemispheric localization of the respective brain regions. To further delineate the cerebral network engaged in the perception of emotional tone, a series of experiments was carried out based upon functional magnetic resonance imaging (fMRI). The findings obtained from these investigations allow for the separation of three successive processing stages during recognition of emotional prosody: (1) extraction of suprasegmental acoustic information predominantly subserved by right-sided primary and higher order acoustic regions; (2) representation of meaningful suprasegmental acoustic sequences within posterior aspects of the right superior temporal sulcus; (3) explicit evaluation of emotional prosody at the level of the bilateral inferior frontal cortex. Moreover, implicit processing of affective intonation seems to be bound to subcortical regions mediating automatic induction of specific emotional reactions such as activation of the amygdala in response to fearful stimuli. As concerns lower level processing of the underlying suprasegmental acoustic cues, linguistic and emotional prosody seem to share the same right hemisphere neural resources. Explicit judgment of linguistic aspects of speech prosody, however, appears to be linked to left-sided language areas whereas bilateral orbitofrontal cortex has been found involved in explicit evaluation of emotional prosody. These differences in hemispheric lateralization effects might explain that specific impairments in nonverbal emotional communication subsequent to focal brain lesions are relatively rare clinical observations as compared to the more frequent aphasic disorders.

The contribution of the cerebellum to speech production and speech perception: Clinical and functional imaging data

A classical tenet of clinical neurology proposes that cerebellar disorders may give rise to speech motor disorders (ataxic dysarthria), but spare perceptual and cognitive aspects of verbal communication. During the past two decades, however, a variety of higher-order deficits of speech production, e.g., more or less exclusive agrammatism, amnesic or transcortical motor aphasia, have been noted in patients with vascular cerebellar lesions, and transient mutism following resection of posterior fossa tumors in children may develop into similar constellations. Perfusion studies provided evidence for cerebellocerebral diaschisis as a possible pathomechanism in these instances. Tight functional connectivity between the languagedominant frontal lobe and the contralateral cerebellar hemisphere represents a prerequisite of such long-distance effects. Recent functional imaging data point at a contribution of the right cerebellar hemisphere, concomitant with languagedominant dorsolateral and medial frontal areas, to the temporal organization of a prearticulatory verbal code (‘inner speech’), in terms of the sequencing of syllable strings at a speaker’s habitual speech rate. Besides motor control, this network also appears to be engaged in executive functions, e.g., subvocal rehearsal mechanisms of verbal working memory, and seems to be recruited during distinct speech perception tasks. Taken together, thus, a prearticulatory verbal code bound to reciprocal right cerebellar/left frontal interactions might represent a common platform for a variety of cerebellar engagements in cognitive functions. The distinct computational operation provided by cerebellar structures within this framework appears to be the concatenation of syllable strings into coarticulated sequences.

The cerebellum and cognitive functions in humans

Recent neuropsychological studies have given rise to the hypothesis that the cerebellum is involved in nonmotor cognitive functions. The interpretation of these findings is, however, restricted by methodological problems, such as heterogenous patient samples. The present study compared patients with pathology confined to the cerebellum and patients with combined cerebellar and brainstem lesions to matched normal controls on a range of memory and learning tasks. Two procedural learning tasks were also conducted, involving perceptual (mirror reading) and conceptual skill acquisition (the Tower of Hanoi task). Patients with damage to both cerebellum and brainstem, but not patients with cerebellar pathology alone, showed impairments on memory and visuoconstructive tasks and evidence of frontal lobe dysfunction. Cerebellar damage had no effect on skill acquisition. These results do not support the hypothesis of cerebellar involvement in procedural learning per se.

Parametric analysis of rate-dependent hemodynamic response functions of cortical and subcortical brain structures during auditorily cued finger tapping: a fMRI study.

A multitude of functional imaging studies revealed a mass activation effect at the level of the sensorimotor cortex during repetitive finger-tapping or finger-to-thumb opposition tasks in terms of either a stepwise or a monotonic relationship between movement rate and hemodynamic response. With respect to subcortical structures of the centralmotor system, there is, by contrast, some preliminary evidence for nonlinear rate/response functions within basal ganglia and cerebellum. To further specify these hemodynamic mechanisms, functional magnetic resonance imaging (fMRI) was performed during a finger-tapping task in response to acoustic stimuli (six different frequencies: 2.0, 2.5, 3.0, 4.0, 5.0 and 6.0 Hz; applied via headphones). Passive listening to the same auditory stimuli served as a control condition. Statistical evaluation of the obtained data considered two approaches: categorical and parametric analysis. As expected, the magnitude of the elicited hemodynamic response within left sensorimotor cortex (plateau phase at frequencies above 4 Hz) and mesiofrontal cortex paralleled movement rate. The observed bipartite mesial response pattern, most presumably, reflects functional compartmentalization of supplementary motor area (SMA) in a rostral component (pre-SMA) and in a caudal (SMA proper) component. At the level of the cerebellum, two significant hemodynamic responses within the hemisphere ipsilateral to the hand engaged into finger tapping (anterior/posterior quadrangular lobule and posterior quadrangular lobule) could be observed. Both activation foci exhibited a stepwise rate/response function. In accordance with clinical data, these data indicate different cerebellar contributions to motor control at frequencies below or above about 3 Hz, respectively. Caudate nucleus, putamen, and external pallidum of the left hemisphere displayed, by contrast, a negative linear rate/response relationship. The physiological significance of these latter findings remains to be clarified.

Emotional processing following cortical and subcortical brain damage: contribution of the fronto-striatal circuitry

The present study examined the differential contribution of cortical and subcortical brain structures in emotional processing by comparing patients with focal cortical lesions (n = 32) to those with primarily subcortical dysregulation of the basal ganglia (Parkinsons disease n = 14). A standardized measure of emotional perception (Tübingen Affect Battery) was used. Only patients in the more advanced stages of Parkinsons disease and patients with focal damage to the (right) frontal lobe differed significantly from controls in both facial expression and affective prosody recognition. The findings imply involvement of the fronto-striatal circuitry in emotional processing.

Speech deficits in ischaemic cerebellar lesions.

SummaryTwelve patients with cerebellar infarction, 8 in the region supplied by the posterior inferior cerebellar artery (PICA) and 4 in the territory of the superior cerebellar artery, underwent formal perceptual examination for speech deficits. With respect to topography the results firstly underline the significance of the paravermal region of the superior cerebellar portion for speech functions. In all patients with dysarthric impairment the lesion extended to this area, whereas patients with PICA infarction sparing the superior portion of the cerebellum showed no speech deficits. Secondly the findings do not corroborate the notion of an exclusively left-sided cerebellar speech motor control, since 3 of the 4 dysarthric subjects had unilateral right-sided ischaemia. This study thirdly demonstrates that lesions of the cerebellar cortex without involvement of the dentate nucleus can cause dysarthric impairment. Phonetic analysis revealed irregularly distributed articulatory deficits and slowed speech tempo as the most common dysarthric features.